These species accounts have been prepared by Dr Ralph Forbes, Vice County Recorder for County Fermanagh and co-author of the Flora of Fermanagh.
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Native, occasional. Circumpolar boreo-arctic montane.
1881-2; Barrington, R.M.; eastern range of Cliffs of Magho.
Throughout the year.
Fermanagh occurrence
In Fermanagh, this small, evergreen perennial is widespread but not abundant, mainly on strongly acidic,nutrient-poor, peaty, high ground on the Western Plateau plus in a few outlying sites in the Carnmorearea. Local habitats include mountain summits and slopes, ledges, screes, open moorland and bogs. It hasbeen recorded in a total of 44 tetrads (8.3% of those in the VC), but it has not been refound in sixprevious sites recorded in the 1940s and early 1970s. The lost sites are: Gadalough, N of KeenaghanLough; Lough Scolban; Mullaghmore (the famous Erica vagans (Cornish Heath) site); Lough Doo nearLittle Dog; Brennan's Rocks, N of Lough Mulderg; and Pollnagollum. While these areas need to be searchedto confirm the local extinction of this species, the evidence suggests that Fir Clubmoss has sufferedhabitat loss mirroring that observed in other parts of its British and Irish range.
The only area where H. selago occurs nowadays in any considerable quantity in Fermanagh is on theexposed summit ridge of Cuilcagh, the highest mountain where the rocks peek through the shallow, acid,nutrient-poor blanket peat. Throughout the year, on the blasted heath, H. selago pokes up itsshort, stiff, yellowish-green, bushy tufts between the stones on bare peaty ground or where surroundedby a carpet of grey-green Racomitrium moss and wind-pruned, flat-growing dwarf woody shrubs, forexample, Erica cinerea (Bell Heather), Calluna vulgaris (Common Heather or Ling),Vaccinium myrtillus (Bilberry), V. vitis-idaea (Cowberry) and Empetrum nigrum(Crowberry).
Ecology
As the Fermanagh tetrad map shows, elsewhere in the VC, Fir Clubmoss is fairly frequent in open areas onscarps and peat bogs on the Western Plateau, and less so in open ground on lowland blanket bogs andheaths whenever competition is reduced. In the latter situation, H. selago usually occurs asindividual plants occupying well-drained sites, often on acid, peaty soil close to large rocks, wherecompetition from heather subshrubs and other species is reduced, probably by the shallowness of thesubstrate. Similar competitive conditions occur when Fir Clubmoss grows on upland cliff and rock ledges.It has also been suggested that the lowland sites of H. selago may lie in damp frost pockets, thecold soils of which would again restrict the growth of competitors (Jermy et al. 1978; Jermy& Camus 1991).
On the other hand, Page (1997) emphasises that all H. selago sites, whether on sandy or peatysoils, seem to be particularly free-draining ones, and that the species is tolerant of very exposedconditions, being able to survive both considerable winter cold and summer sun. However, the speciesdoes appear to be susceptible to heavy grazing pressure and populations are even more vulnerable toheathland fires, whether accidental or used to manage heathers, as the plants are readily eliminated byburning (Page 1997).
British occurrence
In Britain, H. selago has become very rare or extinct in many of its former lowland heathlandsites, indeed in most of these it has not been seen for nigh on a hundred years (Page 1988). While thereasons for this major decline of the species in Britain are not fully understood, undoubtedly changesin the pattern of land use involving habitat loss, changed methods of vegetation management – perhapsinvolving heavier agricultural stocking levels, or using herbicides and fire to manage vegetation – plusincreased levels of air and soil pollution have all been suggested as possibly significant contributoryfactors (Page 1988). While air pollution can be absolutely discounted in our area, the most obvious andnotable examples of this sort of change in the Fermanagh context are the development of extensivehectares of forestry plantation on the Western Plateau and, until recently, the drainage, cutting,ploughing and fertilizer spraying of our lowland bogs.
Irish occurrence
Despite a possible or real decline, H. selago is almost certainly still the most widespread of thefive clubmoss species that occur in Ireland, having been found at least once in all 40 vicecounties (Scannell & Synnott 1987). The 1996 edition of Webb's An Irish Floradescribes Fir Clubmoss as, "frequent but local, mostly above 300 m, but sometimes on lowlandbogs" (Webb et al. 1996).
World distribution
The species is widespread in the northern circumpolar region and numerous varieties also extend worldwideand penetrate far into the southern hemisphere, eg to the Falkland Islands, Tierra del Fuego andTasmania (Hultén 1962). Elsewhere in W Europe, H. selago has a widespread occurrence throughoutwestern and northern parts of Britain and Ireland, Iceland, Scandinavia and western central Europeancountries south to the Alps and the Pyrenees (Jalas & Suominen 1972).
Fossil record
Fossil spore evidence proves the presence of H. selago in Britain and Ireland, especially innorthern parts, from the late phase of the last glaciation onwards (ie from the Late Weichselianonwards, throughout the entire Flandrian period) (Godwin 1975).
Current trends
Historical records show that Fir Clubmoss formerly occurred widespread throughout Britain and Irelandexcept in some counties in S, E and C England. Nowadays, while it is still common in Scotland, H.selago has lost much of its former ground in England and Wales. A count of the pre- andpost-1930 symbols for England and Wales on the BSBI Atlas map indicates that of a total of 140hectads plotted, 73 had pre-1930 records only for H. selago (Perring & Walters 1976; Godwin1975). The 1978 Fern Atlas published a revised hectad map which incorporated considerably morerecords for England and Wales and of the 227 symbols, 127 were for post-1930 records of H.selago, although the overall pattern of losses from the S and E of England was obvious (Jermyet al. 1978). Changes recorded during the BSBI Monitoring Scheme in 1987-88 appeared to indicatea decline in England but an increase in Wales. The authors of the relevant report (Rich & Woodruff1990), however, reckoned these changes were not significant.
In comparison, The New Atlas of the British and Irish Flora which reports survey data upuntil the end of 1999, maps H. selago from a total of 1223 hectads. A visual count of Irishhectads in the New Atlas found a total of 213 squares displayed, 61 of which had pre-1970 recordsonly. The data behind the New Atlas map suggests that most Fir Clubmoss losses took place earlyon (pre-1930), due to habitat changes associated with agricultural intensification. While this processhas continued, the overall distribution of H. selago appears stable in these islands (A.D.Headley, in: Preston et al. 2002).
Reproduction
H. selago plants often reproduce prolifically by vegetative means producing sizeable 'bulbils' or'gemmae' which are budded off from near the tops of stems. In form these are small, leafy,'trident-like' flattened buds formed in rings around the erect stems, about one cm below the largeterminal bud (Page 1997). These abundant propagules are efficiently dispersed by wind in the autumn andthey root and rapidly establish new daughter plantlets which grow between one and two cms tall in theirfirst year (Page 1988).
However, Fir Clubmoss plants also produce vast quantities of pale yellow asexual spores, which again areefficiently wind-dispersed. As far as RSF can discover, nothing is known about the efficiency ofHuperzia spore germination in the field, nor about recruitment of plants from this biologicalsource (Page 1988, 1997). I think this matter could be accurately described as a field of near totalscientific ignorance, which applies not only to clubmosses, but for most pteridophytes and bryophytes.
Even on the question of the longevity of individual plants and their population turnover, little ornothing is definite and everything is qualified; for example, "H. selago appears to be arelatively short-lived plant, which slowly builds up its tufts over several seasons and then probablyreaches an abrupt and fairly rapid demise." (Page 1997). Clearly, if in the future Fir Clubmoss isto be actively conserved, whether under a Biodiversity Action Plan designation or not, closer study willbe urgently required to clarify the reproductive capacity and population dynamics as well as identifyingthe significant environmental pressures affecting the species in typical habitats.
Names
The current author has not been able to discover as yet the derivation of the genus name'Huperzia', which first came to my notice with the first volume of Flora Europaea (firstedition), in 1964. The specific epithet 'selago' is a reuse of a previous generic name: it wasfirst used by the Classical Roman Pliny to refer to a plant resembling Sabina herba, a synonym ofJuniperus sabina (Gilbert-Carter 1964).
Threats
Afforestation of upland areas, overgrazing, the use of fire to manage hillside vegetation and thepossibilities of consequent soil erosion are the main threats.
Native, very rare. Circumpolar boreo-temperate.
1905; Colgan, N.; Altscraghy, on Cuilcagh slopes.
Throughout the year.
Growth form and preferred habitats
Unlike H. selago, individuals of Stag's-horn Clubmoss are regarded as being fairly long-livedplants which slowly develop sparse but wide-spreading colonies with long, horizontally running stems,sprawling through or over a mass of accumulated surface leaf and moss litter. The undulating stem, oftenvivid green in colour, was aptly described by Step and Jackson (1945) as having 'the combined stiffnessand pliancy of copper wire'. These spreading stems occasionally branch and root themselves at intervals,anchoring the clone to the substrate. They also send up occasional erect leafy shoots terminating incharacteristic, perfectly vertical, bare stalks, on which two or three long, slender cones are borne(Page 1997).
The typical habitat of L. clavatum is on N-facing, acidic mountain grasslands or heaths, subjectto high rainfall, but where there is sufficient slope to allow drainage water to flush through theground (Jermy et al. 1978; Page 1997).
Reproduction
L. clavatum does not produce any 'bulbils' or 'gemmae', but relies entirely on its spores forreproduction, a fact that has probably contributed to its decline to extreme rarity, at least inIreland. Like H. selago, Stag's-horn Clubmoss is very liable to destruction by fire. Thus thewidespread practice of maintaining heather vigour on mountain slopes by the establishment of a cyclicalburning regime, has undoubted killed off many local populations, particularly in Scotland and in Irelandwhere this is a regular form of heath management.
Irish occurrence
While Page (1997) described Stag's-horn Clubmoss as the best known clubmoss in the British Isles, it ismuch more local in Ireland, indeed rare and declining for at least 60 years. Although it did not fit thecriteria for inclusion in the vascular plant Irish Red Data book (Curtis & McGough 1988),L. clavatum features in the list of scarce and threatened vascular plants in the 'Biodiversityin Northern Ireland' discussion document (Brown et al. 1997). The scale of the decline ofStag's-horn Clubmoss in Ireland is demonstrated by the fact that Praeger (1934) listed it as occurringor having occurred in 27 Irish VCs, while Scannell and Synnott (1987) suggest it is still present injust 11 VCs (7 of which were northern and did not include Fermanagh).
The statistics available in the Fern Atlas are telling: of the 62 hectads mapped with records forL. clavatum in Ireland, only 19 have post-1930 symbols (Jermy et al. 1978). The NewAtlas of Ferns (2005) confirms the decline, although the map indicates a wider distribution:there now are a total of 81 Irish hectads plotted, 58 of them with pre-1970 records only (Wardlaw &Leonard 2005). The new fern map plots 17 post-1986 hectads in Ireland, all but three in the northernhalf of the island.
Fermanagh occurrence
L. clavatum was not seen in Fermanagh during the 83 years between Colgan's original find on theslopes of Cuilcagh mountain in 1905 and its discovery by H.J. Northridge in 1988 on a peaty roadsidebank on Doon Hill at an altitude of just 250 m. Regrettably, the small population at this new and mostunusual lowland site was subsequently destroyed by the unwitting dumping of earth on top of it sometimeprior to January 1995. Happily, in April 1995 a third station was discovered by Matthew Tickner on aknoll by a burn flowing through blanket bog on the Pettigo Plateau. Previous to this in January 1990 thespecies was rediscovered by RHN surviving on Cuilcagh on the slope below the summit cairn, which wasvery possibly Colgan's original site, bringing the number of extant Fermanagh stations back to two!
The typical habitat of L. clavatum is on N-facing, acidic mountain grasslands or heaths, subjectto high rainfall, but where there is sufficient slope to allow drainage water to flush through theground (Jermy et al. 1978; Page 1997). The Cuilcagh site in Fermanagh exactly fits the typicalhabitat of L. clavatum, ie N-facing, acidic mountain grassland and heath, subject to highrainfall, but on a slope where drainage water flushes the ground. Plants in the site sprawl in a looselyundulating manner over thin peaty soil, covering hidden boulders amid low-growing heather moorlandcomposed mainly of Erica cinerea (Bell Heather), Calluna vulgaris (Common Heather orLing), Vaccinium myrtillus (Bilberry) and Empetrum nigrum (Crowberry). BothHuperzia selago (Fir Clubmoss) and Diphasiastrum alpinum (Alpine Clubmoss) also grow nearthis particular site.
At both its Fermanagh sites L. clavatum is struggling to compete with Calluna. A visit tothe Pettigo Plateau station in October 2010 located only two small patches of the clubmoss.
British occurrence
In Britain, L. clavatum is a mainly northern species, but it has more sites in S England than anyof the three other clubmoss species with which it most nearly overlaps, Huperzia selago,Diphasiastrum alpinum (Alpine Clubmoss) and Selaginella selaginoides (LesserClubmoss)(Jermy et al. 1978).
In Britain and Ireland, and especially in more lowland sites, the species has decreased or disappeareddue to the intensification of agriculture and utilisation of previously ignored rough marginal land(Page 1997).
European and world occurrence
Like Huperzia selago (Fir Clubmoss) L. clavatum is a widespread circumpolar species whichalso extends into the southern hemisphere and worldwide has several named varieties (Hultén 1962). InEurope it displays a mainly western distribution with a relatively continuous range from N Fennoscandiato the Alps and Pyrenees (Jalas & Suominen 1972; Page 1997). In Britain and Ireland it is a mainlynorthern species, but it has more sites in S England than any of the three other clubmoss species withwhich it most nearly overlaps, Huperzia selago, Diphasiastrum alpinum (Alpine Clubmoss) andSelaginella selaginoides (Lesser Clubmoss)(Jermy et al. 1978).
Uses
In former years L. clavatum was collected as a source of 'Lycopodium powder', the dry, light,bright yellow spores being used in school physics experiments to display sound waves. The spores werecollected commercially chiefly in Russia, Germany and Switzerland in July and August, the cones beingcut off and sieved to remove the Lycopodium powder or 'vegetable sulphur', or even 'vegetablebrimstone', as it was sometimes referred to (Grieve 1931). The spores, like the rest of the plant, arevery flammable, and in years gone by they were used in the manufacture of fireworks and for pyrotechnicstage lighting effects in theatres (Mabberley 1997).
In herbal medicine the spores were used alone (ie apart from the rest of the plant), from theseventeenth century onwards, being employed as 'a diuretic for dropsy, a drastic in diarrhoea, dysenteryand suppression of urine, a nervine in spasms and hydrophobia, an aperient in gout and scurvy, acorroborant in rheumatism, and also as an application to wounds' (Grieve 1931). The use of Lycopodiumpowder was never admitted to the British Pharmacopoeia, but herbalists in the British Isles did use itas a dusting powder for treating skin diseases. The main pharmaceutical use was as a pill powder, toenvelope pills and prevent them sticking together when boxed (Grieve 1931). In some rural parts of theBritish Isles there once was a folk-tradition of using garlands of Stag's-horn Clubmoss for 'personaladornment' in some form of ceremony (Page 1988 & 1997).
Names
The genus name 'Lycopodium' is derived from a combination of two Greek words 'lycos', meaning 'wolf', and'podion', 'little foot', a translation of the German, 'Wolfsklauen', first used by the German physicianand botanist, James Theodore Tabernaemontanus. He fancied that the clubmoss shoot resembled the paw of awolf in miniature (Gilbert-Carter 1964; Hyam & Pankhurst 1995; Step & Jackson 1945). Thespecific epithet 'clavatum', is Latin meaning 'club-shaped', which like the general name of the group'Club-moss', refers to the club-like shape of the fruiting cones (Stearn 1992; Step & Jackson 1945).The English common name 'Stag-horn Clubmoss' is a book name, and alternative local folk names do notappear to exist.
Threats
Overgrazing by sheep, shading overgrowth by Calluna, and fire.
Native, very rare. Circumpolar arctic-montane.
1 June 1991; Tickner, M.; Altscraghy, Cuilcagh slopes.
Throughout the year.
Growth form and preferred habitats
This moss-like, low-growing, blue-green, arctic-alpine (or alpine-montane) clubmoss grows in short grassover shallow, well-drained, acid peat, often over rocks on or near mountain summits. The slender, wiry,spreading stems produce distinctive, short, erect, evergreen branches clothed with four tightlyoverlapping ranks of leaves, giving the foliage a rather cypress-like appearance. The shoots forkfrequently and evenly to produce clusters of branches all of identical length which often develop in afan-like, decumbent manner. Fertile branches terminate in a solitary stalkless cone 1-2 cm long, whichturns pale yellow as the asexual spores mature and are released in late July and August (Step &Jackson 1945; Page 1997).
Fermanagh occurrence
In Fermanagh, there is just one quite large, sprawling patch growing amongst Sphagnum andRacomitrium moss and spilling down a rock outcrop on the NE face of Cuilcagh mountain, near thesummit at an altitude of about 580 m. It was found during a survey of the Cuilcagh Plateau for the RoyalSociety for the Protection of Birds. A herbarium voucher was deposited in DBN. The site has beenrevisited several times, most recently in September 2010, when the plant was found in good condition.
Irish occurrence
Elsewhere in Ireland, Alpine Clubmoss is known only from a few scattered localities on the acidic highmountains of Ulster, W Mayo, W Galway, Offaly and Wicklow (Jermy et al. 1978). In NorthernIreland, the two main areas for the species remain the granite Mourne Mountains, Co Down (H38), where inthe past it occurred on six different peaks, and the basaltic Garron Plateau, Co Antrim (H39). There arealso isolated stations in the Sperrin mountains and on Slieve Gallion in Co Londonderry (H40), plus thisrecent Fermanagh discovery (Hackney et al. 1992; NI Vascular Plant Database 2002). The loweraltitude limit for the NI sites is around 300 m, considerably less than the 457 m given for Britishmainland sites (Page 1997).
British occurrence
In Great Britain, Alpine Clubmoss has a pronounced northern and western distribution on the higher hillsof Scotland, N England and Wales, with outliers in the Derbyshire Pennines, the Worcestershire Malvernhills and possibly also in Devon – although the latter records need confirmation (Jermy & Camus1991). D. alpinum is characteristic of (and in Wales, at least, can become locally dominant on)well-drained, shallow, peaty slopes, on exposed and/or heavily grazed sites, where plant competition isreduced by these environmental pressures (Page 1997).
Recent changes
Like other clubmoss species, D. alpinum populations and its distribution have undoubtedly suffereda decline in the recent past, particularly at lower altitudes (Jermy et al. 1978; N IrelandVascular Plant Database 2002). The scale of past losses in Ireland is clearly seen from hectadstatistics in the New Fern Atlas (2005), which show the species mapped in a total of 49 squares,only 25 of them with post-1970 records (Wardlaw & Leonard 2005).
Land-use changes, including the extension of coniferous plantation on ever higher ground, and increasedlivestock stocking densities on upland moorland and heaths has resulted in losses of D. alpinumin both N England and N Ireland, and very probably also in other regions of Britain & Ireland. Likeall clubmosses, D. alpinum is very vulnerable to fire. In addition, sulphur dioxide air pollutionis undoubtedly responsible for the disappearance of the plant from the hills overlooking Belfast in thelate 19th century. Despite the advent of clean air legislation and its enforcement in recent years, thisfactor continues to operate and it certainly must be responsible for some of the losses recorded insites downwind of major cities in Britain (Jermy et al. 1978; Page 1997).
European and world occurrence
Beyond Britain & Ireland, D. alpinum has a classic disjunct arctic-alpine distribution inEurope (Jalas & Suominen 1972, Map 12), and it would be circumpolar were it not for a ratherunexpected absence from much of the arctic region of central Canada, creating a large gap in thedistribution (Hultén & Fries 1986, Map 6).
Uses
In past times, D. alpinum was collected and used as a source of dye mordant in place of the usualchemical fixative, alum. Experiment has shown that plant dyes fixed with mordants from clubmoss speciesproduce softer, more permanent colours than those achievable with alum (Page 1988).
Names
I cannot find a derivation for the genus name 'Diphasiastrum', except that it must in some way connectwith a synonym of Lycopodium, 'Diphasium'. The word element 'diphasia' suggests somethingthat exists in, or exhibits 'two stages'.
Threats
Increased sheep density on upland moorland and heaths has resulted in losses of D. alpinum in bothN England and N Ireland. Air pollution is an increasing threat in some areas. The solitary nature of theFermanagh plant renders it vulnerable, but the site is rather difficult of access, reducing thelikelihood of it being grazed.
Native, occasional. Circumpolar boreal-montane, but rather disjunct in both Eurasia and N America;perhaps better considered mainly arctic-montane.
1882; Stewart, S.A.; Drumbad Scarps, Lough Navar Forest Park.
June to December.
Growth form and preferred habitats
A tiny, delicate, moss-like perennial, Lesser Clubmoss is both montane - found above the notionaltree-line in areas dominated by blanket-bog, and in heathland, and also lowland, growing in wet flushed,base-rich ground beside lakes and streams. Being very small and rather inconspicuous it needs to bepositively searched for in seepage areas, stream sides and near lakeshores, or in open, exposedhabitats. It often occurs in shallow soils where the growth of competing species is limited.
The perennial stems of Lesser Clubmoss are generally extremely inconspicuous, weak-looking, prostrate,branching only occasionally, and they trail over and through mossy vegetation and the stem bases ofother, more vigorous vascular plants. Slightly easier to spot are the annual cone shoots, of which oneor two are produced per stem during July and August.
Pinguicula vulgaris (Common Butterwort) is almost invariably a good indicator of the likelypresence of S. selaginoides and it is interesting to compare the distribution maps of the twospecies. In late August and early September the pinkish-yellow colour of the senescing erect annual coneshoot of Lesser Clubmoss makes it much easier to observe (provided, that is, if one bends right over andlooks closely and diligently in the right sort of habitat!). The small size of the plant and the factthat it has a soft, moss-like texture immediately distinguish S. selaginoides from all otherspecies of clubmoss in Britain and Ireland.
Reproduction
The terminal spore-bearing shoots are shaped like an Indian club, or a fox's tail and are held erect,typically between 2 and 10 cm tall. The actual cone is leafy and ill-defined, bearing both sterileleaves and fertile spore-bearing ones (ie sporophylls), carrying separate spore sacs (ie sporangia). Thelatter structures contain asexual spores of one of two kinds on separate parts of the fertile branch:either four white relatively large female megaspores per megasporangium, or vast numbers of microscopicyellow male microspores per microsporangium (Jermy & Camus 1991; Page 1997). Sexual reproductionfollows on a microscopic prothallus produced by each (or some) of the megaspore(s).
Fermanagh occurrence
This tiny, delicate, moss-like perennial has been recorded in 56 Fermanagh tetrads (10.6% of those in theVC), 46 of them with post-1975 records. It is occasionally found on the shores of Lower Lough Erne, butis quite frequent and widespread on the upland limestones of the Western Plateau. The details of themost outlying stations to the S of the county are: Knockninny, 1900, W.N. Tetley; Kilroosky Lough ASSI,1980, R.S. Weyl; and moorland at Skeaghoge Td, 1989, RHN.
The nature of base-rich habitats
The one characteristic that links all S. selaginoides habitats is the requirement for soilenrichment and aeration associated with the movement of base-rich water. The base (ie the positivelycharged cation) most typical of such waters is calcium, but in the Lough Navar area of Fermanagh theparent rock has become partially dolmitized and thus base-rich water here contains both calcium andmagnesium (Whitten & Brooks 1972).
In other sites, base-yielding rocks may include various mica-schists, volcanic tuffs, lavas, or basalts,some of which may release minerals in quantities that are toxic to certain plants. Base-rich water isnot necessarily nutrient-rich as far as plant growth is concerned, and if it is derived from dissolutionof limestone or dolomite it is always nutritionally unbalanced, being oversupplied with Ca++ions. As ground-water percolates through soil, however, it can accumulate and transport dissolvednitrogen and available phosphate, plus traces of other elements essential for plant development, whichmight otherwise be scarce or absent in a particular site and thus become limiting for plant growth.
The best concise account of the concept of base-rich soil and plant nutrient status known to the currentauthor appears in Page's book, Ferns. Their habitats in the British and Irish landscape (Page1988, pp. 70-3 and 311-2), and this is highly recommended reading for anyone who is puzzled by the usageof this technical term. Page clarifies the essential qualifications associated with understanding andapplying this rather difficult and potentially (and actually), very woolly ecological concept.
Fossil record
Micro-fossil megaspores of S. selaginoides have been found at Derryvree, near Maguiresbridge inFermanagh in a full-glacial freshwater deposit of Middle Midlandian age radio-carbon dated to 30,500 BP(Colhoun et al. 1972). The flora and fauna of this fossil deposit indicated open tundravegetation and a periglacial climate prevailed at the time it was laid down.
Elsewhere in Britain & Ireland, the fossil record for S. selaginoides is well studied, bothmicrospores and megaspores being readily recognised. The sediment studies prove the species has beenpresent in these islands during the last four glacial stages. Lesser Clubmoss has been less frequentlyrecorded during some of the intervening warm interglacial periods. This is not terribly surprising,since it cannot cope with tall, shading vegetation typical of the forest maximum. However, the specieshas been found as fossils throughout the entire current heavily studied interglacial, which is calledthe Flandrian in Britain and the Littletonian in Ireland.
British and Irish occurrence
The fossil record also shows that the plant has contracted in range northwards compared to its lateglacial distribution (Godwin 1975). Although we do not understand what factor(s) caused this rangecontraction, it is particularly well demonstrated in the British Isles, where the species until recenthistorical times occurred north of a line in Britain from Barmouth in Wales to Skegness in Lincolnshire,and in Ireland from Foynes near Limerick, eastwards to Arklow. There is just one exceptional site belowthis demarcation, which is on the coast near Wexford town (Jermy et al. 1978).
The distribution of S. selaginoides has been seriously affected by drainage and theintensification of agriculture in the last 50 to 70 years, with significant losses in the SE of Englandbefore 1930. Approximately 70 additional sites were lost between 1950 and 1990. Over the same period,similar environmental pressures in Ireland have resulted in the loss of around 35 sites of the speciesthroughout the island, but again losses have been particularly concentrated in the south of its range(Jermy et al. 1978; Jermy & Camus 1991).
European and world occurrence
Beyond Britain and Ireland, S. selaginoides has an amphi-atlantic or circumpolar, boreal-montanedistribution (Hultén 1958; Hultén & Fries 1986, Map 8; Preston & Hill 1997). In the view of thecurrent author, the absence of records in Siberia and major gaps in Asia in general, does not warrantdescribing the distribution as circumpolar. The disjunctions are simply too wide. Furthermore, thepublished European distribution is very definitely disjunct and appears to fit the arctic-alpine (orarctic-montane) pattern better than the more continuous boreal-montane picture (Jalas & Suominen1972, Map 13). The distribution of S. selaginoides extends quite far south in Finland, the Balticregion and Denmark. Otherwise its distribution closely resembles that of Diphasiastrum alpinum(Alpine Clubmoss) (Jalas & Suominen 1972, compare Maps 12 & 13). Preston & Hill (1997)regard the latter as circumpolar arctic-montane. Most unexpectedly, Page (1971) has discovered a singleextremely disjunct outlying station of S. selaginoides lying 2100 km south of its nearestEuropean mainland station, on the Canary Island of Hierro.
Names
The genus name 'Selaginella' is the diminutive of 'Selago', an ancient name applied by the classicalRoman, Pliny to a plant resembling Sabina herba, an old name of Juniperus sabina(Gilbert-Carter 1964). The plant Pliny was referring to was a clubmoss, the whole group being then named'Selago', including the genus we know as 'Lycopodium' together with subsequent splits from it (Johnson& Smith 1946). The specific epithet 'selaginoides' is Latin meaning, 'Selago-like' or'clubmoss-like', probably meaning, 'like Lycopodium selago', an earlier name for the currentspecies (Gledhill 1985; Stearn 1992). The English common name 'Lesser Clubmoss' is a typical book name,probably of Victorian origin.
Threats
Afforestation of the species' upland habitat, or improvement of rough or damp ground for agriculturalpurposes.
ISOETACEAE – Quillwort family
Native, occasional. Eurosiberian boreal-montane; close relatives in N America exist, rendering thespecies s.l. as amphi-Atlantic.
1946; MCM & D; Lough Jenkin.
February to October.
Growth form and preferred habitats
In Fermanagh, this small submerged perennial can sometimes be clearly seen growing in sheltered shallowwater on the gravelly bottoms of acidic, nutrient-starved, unproductive lakes, as at Lough Nabrickboy inBig Dog Forest. More often it occurs in deeper, medium-sized, brown-water lakes on the Western Plateau,or in more mesotrophic, ie moderately productive, lowland water bodies. In both these situations, it canform a lawn-like turf and become the dominant bottom-growing plant. The presence of the species is oftenonly betrayed when its stiff, evergreen 'quills' are washed up in the plant debris along the shorelineafter stormy weather, sometimes in considerable quantity.
Like the vegetatively similar Lobelia dortmanna (Water Lobelia), I. lacustris is not acompetitive species and in nutrient-rich waters it is easily overgrown by algae or by faster-growingaquatic macrophytes or both. Probably for this reason, Quillwort tends to occupy rather deeper water,typically from 0.5 to 2 m, but it can survive down to 6 m deep in order to avoid competition from morelight-demanding vascular plants (Page 1997; Jonsell et al. 2000). I. lacustris appears toavoid small lakes and silt- or peat-bottomed mountain tarns, habitats more ecologically suited to I.echinospora (Spring Quillwort).
The four air chambers which traverse the length of each dark green tubular leaf are easily seen when itis sectioned at right angles, and they serve to distinguish the plant from either Littorellauniflora (Shoreweed) or Lobelia dortmanna, two other species with similar stiff 'isoetid'style of leaves, with which it frequently occurs in stony or sandy lake shallows. When Lobeliadortmanna flowers, it becomes a much more conspicuous plant than either of the other two speciesmentioned, and since it is so similar to I. lacustris in its ecological requirements andtolerances, it is a very good indicator of the likely presence of Quillwort at a site.
Another way in which I. lacustris is very readily distinguished from Littorellauniflora is by its brown (not white) roots (Jermy & Camus 1991). The four elongate airchambers in Isoetes leaves are supported and divided by numerous crosswalls (ie septae). Thefunction of the air chambers within the photosynthetic green leaves is associated with the unusualmethod of carbon metabolism of these aquatic species, which is called 'Crassulacean Acid Metabolism(CAM)'. This physiology is most frequently found in desert succulent plants such as Yuccabrevifolia (Joshua tree) (Crawford 1989, pp. 140-2). The extensive roots of I. lacustrisabsorb carbon dioxide gas from the mud sediment and store it along with carbon dioxide from night-timerespiration as malic acid. The plant can thus recycle and store inorganic carbon, a scarce essentialelement in this habitat (and see also the species account of Lobelia dortmanna) (Farmer &Spence 1985; Boston 1986; Preston & Croft 1997).
Fermanagh occurrence
Considering the large number of lakes in Fermanagh, many of them upland, it is not surprising that thisVC is the N Ireland headquarters of this aquatic species in terms of frequency, having been recorded 35of tetrads, 6.6% of those in the VC. The two lakes in Fermanagh where it has not been recorded since the1940s are Lough Scolban in the west of the VC and Lough Skale further east.
Irish occurrence
Compared with Co Fermanagh, the low frequency of Quillwort in Cos Down and Antrim (H38 and H39), and itsvery slight representation in both Cos Armagh and Londonderry (H37 and H40) is rather unexpected. InIreland overall, I. lacustris has been recorded at least once in the past in 19 of the 40 VCs(Scannell & Synnott 1987).
The New Atlas hectad map shows that Quillwort is predominantly distributed in the N and Wof both Britain and Ireland, although in the latter it is also present in eastern sites, for instance inboth the Mourne mountains, Co Down (H38) and the Wicklow mountains south of Dublin (H20). There are alsoa few isolated sites close to the SE coast of Ireland in Co Waterford and South Tipperary (H6 and H7).
British occurrence
The distribution in Britain is very decidedly Scottish, Cumbrian and Cambrian (ie Welsh), but it doeshave a very few southern outliers in S Devon (VC 3). The solitary early 19th century E Englandoccurrence at Prestwick Carr in S Northumberland (VC 67) is long extinct (Swan 1993; Preston & Croft1997).
European and world occurrence
Quillwort has essentially a sub-Atlantic distribution in NW Europe, centred on Scandinavia and theBritish Isles. However, there are isolated, widely scattered outliers as far afield as Iceland, thePyrenees and the Urals (Jalas & Suominen 1972, Map 18). Beyond Europe, I. lacustris is alsofound in S Greenland and in NE North America (Hultén 1958, Map 247; Hultén & Fries 1986, Map 9;Jonsell et al. 2000).
Variation
Hultén (1958) and Hultén & Fries (1986) both map the North American form of this plant as 'I.lacustris var. macrospora' Dur., and it has also at times been elevated to species rankand gone through several synonyms which these authors list. However, many botanists reckon this varietyor species is best submerged back into I. lacustris (Preston & Croft 1997). A number of othervarieties with one or more unusual characters have been described, including an Irish form, var.morei Syme, from a lake near Bray, Co Wicklow, which has very long leaves (Brunker 1950). Aninteresting viviparous form, with vegetative buds instead of sporangia, has been recorded from LakeWindermere (Page 1997). These variants, however, are now simply regarded as the evolving products oflong geographic isolation and the consequent inbreeding of an ancient species.
Herbivory and dispersal
Upland game birds such as grouse are reported to feed on Isoetes species in North America (Fassett1957) and their relatives may also do so in Britain and Ireland, although the current author (RSF)cannot locate any mention of this herbivory in the literature. It is considered feasible that thedistribution of the species may reflect the North Atlantic migration pattern of wildfowl, most probablythat of geese, which might carrying the spores, or much less likely, transport vegetative parts of theplant (Page 1997). The possibility of avian transfer is supported by the existence of the endemic formI. azorica, since it is hardly possible to imagine any mechanism of transport apart from waterbirds visiting the remote island group of the Azores on their regular migration route (Ridley 1930).
Names
The genus name 'Isoetes' appears to have been coined by the ancient Roman scientist Pliny, combining twoGreek words 'isos', meaning 'equal' and 'etos', meaning 'a year', ie 'equalling one year'. This refersto the idea that the plant did not change with the seasons, meaning that it was evergreen. However, itis also thought that Pliny originally applied the name not to this species, but rather to a member ofthe Crassulaceae (Gilbert-Carter 1964; Johnson & Smith 1946). The specific epithet 'lacustris' isLatin meaning 'associated with lakes' (Gilbert-Carter 1964). The English common name, 'Quill-wort' or'Quillwort' is an 18th century name, given to this inconspicuous and not well recognised plant from itssupposed resemblance to a bunch of quills (Prior 1879; Grigson 1974).
An alternative name, 'Merlin's Grass', is a translation of a Welsh name 'Gwair Merllyns', where 'gwair'means 'hay' and Merllyn was the name of a Welsh prophet. The Welsh name of the plant appears in amanuscript account of Samuel Brewer's botanical journey through Wales in 1726, which is preserved in theBritish Museum, and it is quoted as follows in Britten & Holland (1886), "At Llyn Ogwen(Carnarvonshire) I saw the horses very greedily eating of that which was cast upon the shore and that onthe water; and the people tel [original spelling] me that they wait there every day for it, and leavegood grass growing near it; and that it improves cattle better than any grass; and that the fish like itas well. The fish are larger there than any of the other lakes, which they attribute to the eating of[this plant], which they call Gwair Merllyns.'
Threats
Eutrophication (ie cultural nutrient enrichment) of lowland sites and silting of upland ones, the latterat least generally attributed to forestry operations.
Native, very rare, possibly a mis-identification. Circumpolar boreal-montane.
1946; MCM & D; Castle Caldwell, Lower Lough Erne.
Fermanagh occurrence
Just two Fermanagh records exist from peaty or muddy lake bottoms, at Castle Caldwell, listed above andBunnahone Lough in 1947, made by Meikle and his co-workers. The Revised Typescript Flora notedthat, "These identifications need checking, as do all Irish Isoetes records." (Meikleet al. 1975). In Meikle's Fermanagh Flora card index, the Bunnahone plant was originally recordedas I. lacustris but was later reassigned. As far as we are aware, no vouchers exist for I.echinospora from Fermanagh (Osborne & Doyle 1992).
Nowadays, both sections of Lough Erne are eutrophic to hypertrophic and thus have become toonutrient-rich to support either species of Isoetes. However, in 2006 and 2007, palaeoecologicalstudies associated with water quality assessment took sediment cores from several N Ireland lakesincluding the Trannish region of Upper Lough Erne and Meenatully Lough on the Pettigo Plateau blanketbog. These very different waterbodies contained fossil megaspores of both I. echinospora andI. lacustris. In the case of Meenatully Lough, macrofossils of both species were present in thelower portion of the upper 0-7 cm zone of the core, dated post-1970 (Davidson et al. 2008). The NIreland Lakes Survey (1988-90) recorded only I. lacustris in this lake.
Irish occurrence and status
Although it did not feature or deserve a mention in the Irish Red Data Book. 1. Vascular plants(Curtis & McGough 1988), I. echinospora is now classified as a scarce species byconservationists in the Republic of Ireland. In our view, it is better described as a rare and possiblydeclining species. The known Irish stations are extremely thinly scattered down the W coast from CoDonegal to Co Kerry (Preston & Croft 1997). There are voucher specimens in DBN for six of the40 Irish VCs, but only those of Co Clare (H9) are modern, all the rest being pre-1911.
Only ten of the total of 41 Irish records held at the Biological Records Centre, Wallingford, Oxfordshireare post-1950. They represent occurrences in five Irish VCs, S Kerry, N Kerry, Co Clare, W Galway and WMayo (H1, H2, H9, H16 and H27). [All but the two records from S Kerry (H1) are from sites below 100 m inaltitude (Osborne & Doyle 1992). I. echinospora is probably slightly more frequent in CoClare (H9) and W Galway (H16), since these two VCs share six of the ten post-1950 records between them(Webb & Scannell 1983; Osborne & Doyle 1992).]
Identification difficulties
It is very difficult to distinguish I. echinospora from I. lacustris (Quillwort) in thefield and they can co-exist and hybridise. The fact that specimens require microscopic confirmationdeters the more casual recorder, so that I. echinospora may be overlooked or mistaken by fieldbotanists for the very much more common species (Jermy & Camus 1991; Page 1997). Very sensibly intheir Flora of Connemara and the Burren, Webb & Scannell (1983) were chary of accepting anyreports of Spring Quillwort that had not been verified by microscopic examination of the microspores, aprocedure which Osborne & Doyle (1992) also regard as absolutely essential.
In addition to the absence of vouchers for the 1940s Fermanagh records, the systematic survey of ourlakes made in recent years has failed to find living specimens of I. echinospora. We thereforebelieve that either Meikle and his co-workers misidentified their specimens, or subsequent field workers(including ourselves) have not looked carefully enough at Isoetes material, especially thatoccurring in more oligotrophic waters.
Names
The genus name 'Isoetes' appears to have been coined by the ancient Roman scientist Pliny, combining twoGreek words 'isos', meaning 'equal' and 'etos', meaning 'a year', ie 'equalling one year'. This refersto the idea that the plant did not change with the seasons, meaning that it was evergreen. However, itis also thought that Pliny originally applied the name not to this species, but rather to a member ofthe Crassulaceae (Gilbert-Carter 1964; Johnson & Smith 1946). The Latin specific epithet'echinospora' means 'spiny spored', which is for once, the major defining character of the species.
Threats
None.
Native, rare, but occasionally locally abundant. Circumpolar boreo-temperate.
1872; Smith, T.O.; Colebrooke River (unspecified region).
March to December.
Growth form and preferred habitats
This rare, slow-growing, rhizomatous, evergreen horsetail species, with its distinctive rough texture, isregularly found growing on shady, sloping river banks and streamsides, which represent its predominanthabitat throughout Britain & Ireland. It typically grows in heavy, permanently moist, sandy orclayey soils that are rich in silica and other minerals. It can also be found in base-rich moorlandflushes, and elsewhere in similar flushes on sand dunes (C. Dixon & T.D. Dines In: Preston etal. 2002).
Identification
The erect, unbranched, dark blue-green stems are pencil-thick and the ash-white, toothless sheaths withblack bands around the top and bottom make E. hyemale reasonably easy to distinguish from itshybrid with E. variegatum, E. × trachyodon. When the leafsheaths are young they do appear to bear short teeth. In reality, these are minute scallops where thetrue sheath teeth would normally be attached (Page 1997, p. 450), but they are not observable on maturesheaths (Rose 1989). In the current author's experience, E. hyemale and E. × trachyodon only rarely associate with one another.
Fermanagh occurrence
This distinctive horsetail has been recorded in Fermanagh from 14 thinly scattered tetrads, 2.7% of thosein the VC. Twelve tetrads have post-1975 records in habitats ranging from moist woods and shaded riverbanks, to peat covered limestone in the uplands. In addition to Smith's first record listed above, thereis another early site at Cloncarn near Magheraveely, where it was recorded by Meikle and co-workers in1948.
Although it typically grows in permanently moist, sandy or clayey soils, surprisingly it has never beenfound on any of the many lakeshores in the VC, although further north in Scandinavia it does occur insuch situations, plus in a wide range of other very different, often much drier habitats which it neveroccupies in Britain & Ireland (Jonsell et al. 2000, p. 24).
In Fermanagh, like several other horsetail species, eg E. palustre (Marsh Horsetail) and E.telmateia (Great Horsetail), E. hyemale appears to require some lateral water movement atit* roots, either a slow seepage or a flushing of moderately base- or mineral-enriched spring water(Rose 1989; Brewis et al. 1996).
An unusual variant
In terms of habitat, E. hyemale is not a very variable species but one exceptional site occurs onthe limestone plateau at Legacurragh above Florencecourt. Here a solitary plant of a completelyprostrate form of E. hyemale grows on thin blanket bog peat developed directly over limestonepavement. According to Clive Jermy (pers. comm. 1995), this unusual prostrate form is known fromScottish dune systems and, as here, grows in flushed, shallow peat over limestone.
Reproduction
Fertile stems emerge along with sterile ones in May-June and are similar in appearance, except that theyproduce a small, black cone which bears a short sharp tip (ie an apiculus). Spores are not produceduntil early spring of the second year and, in common with all other Equisetum species in Britain& Ireland, reproduction and spread of E. hyemale is mainly (but not exclusively), vegetative,involving lateral growth of the rhizome and water dispersal of stem fragments (Praeger 1934).
Growth rate, silica content and rough stem texture
E. hyemale is said to grow and spread very slowly, even when well established (Page 1997, p. 451),yet at the site on Manyburns River in Fermanagh, and in similar places, the plant grows in abundance inthick clumps. In these situations it locally dominates the riverbank vegetation, presumably due to itstenacious rhizome and the longevity of the species.
The observed slow growth of this species is probably due to its very high silica requirement, which inturn is associated with the colourless siliceous tubercles and other physical structural features whichgive the plant its characteristic tough, evergreen stems their very rough, abrasive texture.
Uses and English common names
The English common names, 'Rough Horsetail' and 'Dutch Rush', both allude to the fact that from earlydays, at least from the 17th century onwards, the plant was greatly valued as a scourer, the equivalentof our present day wire-wool (Grieve 1931). E. hyemale stems were collected locally and sold inmarkets for scouring cooking pots and were also used by artists for fine polishing metal, wood and bonearticles (Step & Jackson 1945) and thus the range of English common names 'Pewterwort','Shave-grass', 'Scouring-rush', 'Scrubby-grass' and 'Dishwashings' (Prior 1879; Britten & Holland1886).
Rough Horsetail is mentioned by Gerard (1633) as being used by fletchers (arrow makers) and comb-makersto polish their finished articles. Other more frequent and abundant Equisetum species were alsoused for these purposes, eg E. arvense (Field Horsetail) and E. palustre and, undoubtedly,some of these common names (apart from 'Dutch Rush') would also have been locally applied to them aswell. Grigson (1974) and Mabey (1996) both report that bundles of E. hyemale are still sold asscourers in continental European markets.
Being a slow growing, rather scarce species, commercial collecting in B & I must have very quicklyreduced local populations of this horsetail, so that imports from or through the Netherlands becamenecessary to meet the commercial demand for scourers and hence the name 'Dutch Rush'.
Irish occurrence
Long after the commercial use of the species as a scourer ceased, E. hyemale remains a rare, localand apparently declining species in the whole of the British Isles. In Fermanagh, it occurs in just 12scattered post-1975 tetrads. Elsewhere in N Ireland, E. hyemale is rather rare in Cos Antrim,Tyrone and Londonderry (H39, H36 and H40), and very rare in Cos Armagh and Down (H37 and H38) (NIVascular Plant Database 2001). In the Republic of Ireland, Rough Horsetail (often referred to as 'DutchRush') is very rare and scattered, and is apparently declining here also.
British occurrence
In Britain, the species overall has a decidedly northern distribution and, while scarce and local even inthe northern half of the island, it is very much more rare and obviously declining south of theMersey-Humber line (Jermy et al. 1978; Page 1997).
European and world occurrence
The European distribution of E. hyemale is quite similar to that of E. sylvaticum (WoodHorsetail), being essentially northern and boreal and stretching from SE Greenland (where it was firstfound as recently as 1981), through to Iceland, the Faroes (but not the Arctic Isles), and throughoutall of Scandinavia. It also extends south to Gibraltar (although only very thinly represented across theIberian peninsula) and thence eastwards along the northern shores of the Mediterranean to Greece and NTurkey (Jalas & Suominen 1972, Map 30; Daniels & Van Herk 1984). The distribution then continueseast through the Himalaya and much of N Asia to Japan and Central America (ie Mexico and Guatemala)(Hultén 1962, Map 174; Hultén & Fries 1986, Map 11; Jonsell et al. 2000).
Is it circumpolar?: Although E. hyemale is classified by Preston & Hill (1997) withoutqualification as Circumpolar Boreo-temperate, the species only qualifies as circumpolar if we considerthe taxon in its very broadest sense. In Europe and in W & C Asia, the E. hyemale we know inBritain and Ireland is a moderately variable species, but in E Asia and especially in N America, itbecomes a complex of several forms which, while their taxonomy is incompletely worked out and is asubject of disagreement, have been grouped by American taxonomists into two species, E. hyemaleand E. laevigatum and their hybrid (E. × ferrissii). NAmerican E. hyemale is then further subdivided into three varieties and three forma, none ofwhich is identical with our Eurasian plant (Hultén 1962; Scoggan 1978, p. 130). Thus there exists anenormous void in the circumpolar occurrence of the Eurasian form of E. hyemale (ie ourE. hyemale), throughout N America. In terms of the plant's plant geography, its presence furthersouth in C America does absolutely nothing to fill this northern, Boreo-temperate gap.
Names
The genus name 'Equisetum' was coined by the ancient Roman writer, Pliny and is thought to have beenfirst applied by him to E. arvense. It is a combination of two Latin words, 'equus', a horse and'saetum', a bristle or hair, and it is thought to refer to the bristly appearance of the jointed stemswith their whorled branches (Gilbert-Carter 1964; Grieve 1931). The same notion also gave origin to theEnglish common name 'Horsetail', which is a direct translation of the medieval Latin name, 'caudaequina', under which it was sold in apothecary shops (Prior 1879; Grigson 1974).
The Latin specific epithet 'hyemale' or 'hiemale', means 'of winter', that is, 'reproducing in winter'(Gilbert-Carter 1964).
Threats
Clearance of wooded stream banks, or excessive trampling or grazing of the sites by cattle pose the twomajor threats. Drainage might also be significant in other areas of the British Isles.
Native, rare, but locally abundant.
1904; Praeger, R.Ll.; Bunnahone Lough, Lenaghan Td.
Throughout the year.
Growth form and preferred habitats
Plants of this evergreen, rhizomatous hybrid are more robust and more branched than those of E.variegetum, sometimes stretching up to 75 cm tall. This vigour and the fact that the long blackteeth on the nodal sheaths are usually very persistent together help to distinguish the hybrid from bothits parent species.
The hybrid grows on rocky lakeshores and wooded riverbanks and in Fermanagh is twice as frequent as oneof its parents, Equisetum hyemale (Rough Horsetail). It usually occurs some distance fromthe main concentration of sites of the other parent, E. variegatum (Variegated Horsetail), whichseems to require a more calcareous or more base-rich habitat than does the hybrid. E. × trachyodon does occur near E. hyemale at one spot on theupper reaches of the Colebrooke River, and with E. variegatum at Shean Jetty and Magho Jettyalong the S shore of Lower Lough Erne, but these are the only exceptions. In the remaining twelvestations, E. × trachyodon avoids both its parents – or perhaps onaccount of its vegetative vigour, they avoid it!
The Fermanagh plants of this rhizomatous hybrid can form quite large, dense stands, as happens forinstance on the Bannagh River and at one spot on Upper Lough Macnean, but it can also occur as just afew straggling, branched stems as on the shore of Lough Lattone. Alternatively, it may be scattered inclumps, as it is along a km or so of the bank of the Colebrooke River below Littlemount Bridge.
In general, the known occurrence of all forms of hybrid horsetails are far from evenly distributedthroughout the overlapping portions of their parent species ranges in Europe (Jalas & Suominen1972). Rather they tend to be thinly scattered, but with distinct local concentrations in certain areas.This is particular the case in western and northern regions of Britain & Ireland (Page & Barker1985; Page 1997).
Equisetum hybrids in Britain & Ireland
All eight native Equisetum species in the British Isles are involved in producing hybrids, butthey are formed strictly between pairs of species within the same subgenus. Six of the British& Irish horsetail species belong to subgenus Equisetum, and two (E. hyemale and E.variegatum) to subgenus Hippochaete. Perhaps surprisingly, only two hybrids from subgenusEquisetum have been found so far in Fermanagh.
Fermanagh occurrence of E. × trachyodon
In the case of the solitary subgenus Hippochaete hybrid, E. ×trachyodon, N Ireland undoubtedly has the greatest concentration of known stations for thishybrid possibly anywhere. Within the six-county province, Fermanagh with its 13 main sites and theirsometimes many sub-sites has the greatest representation of this hybrid. The Fermanagh tetrad map plotsrecords from 22 post-1975 tetrads, plus two tetrads with older records. The details of the Fermanaghsites where E. x trachyodon has not been refound are: Bunnahone Lough, 1904, Praeger; andLough Vearty, 1949, MCM & D.
Other Irish occurrences
The Fermanagh coverage is followed by Cos Down (H38), Antrim (H39) and Londonderry (H40), with Tyrone(H36) and Co Armagh (H37) trailing with just one or two sites each (NI Vascular Plant Database 2001). Inthe Republic of Ireland, Co Cavan (H30) also has one old 1950s record from Gowland, yet so far neitherparent species has ever been recorded there (Reilly 2001). Elsewhere there are a further 12 tetradsscattered, mainly in the west, across 8 Irish VCs from Monaghan (H32) and Sligo (H28) to Mid Cork (H4),plus two more inland VCs, N Tipperary (H10) and Kildare (H19) (New Atlas).
Vegetative reproduction and spread
Some of the Fermanagh sites are obviously linked, for instance those along riverbanks, or along lakeshores and they may, or may not, represent fragmented clones generated by secondary vegetative spread.Page (1997) reported that small fragments of hybrid shoots root very readily, even after they havefloated around for up to ten days, so this vigorous hybrid definitely has a mechanism for increase andlocal dispersal.
In Fermanagh, we see evidence of local vegetative spread along the Colebrooke River, along the N shore ofUpper Lough Macnean and the S shore of Lower Lough Erne. However, other stations are sufficiently remoteto certainly represent independent parental hybridisations, and these cases form the majority inFermanagh.
British occurrence
In comparison with NW Ireland, E. × trachyodon is very poorlyrepresented in Britain, there being only three or four sites in England and six or seven in Scotland(New Atlas).
A strong westerly trend in hybrid horsetail occurrence
All but one of the British stations are in the extreme west and when one examines the distribution of allhorsetail hybrids in Britain & Ireland, a westerly trend in their occurrence becomes very obvious.Indeed, if we 'zoom out' to view the whole European picture of hybrid horsetails, the westerly trendappears to be mirrored even at this much larger scale (Page & Barker 1985; Page 1997).
The explanation for this strongly marked trend in distribution presumably lies in the prevailing oceanicclimate of the most westerly parts of Britain & Ireland. This is readily summarised as cloudy skies,high rainfall levels that are evenly dispersed throughout the year (ie over 200 wet days) and generallylow temperatures with no extremes (ie mild winters and cool summers) (Page & Barker 1985; Page 1997,Maps 6-16; Porley 2001). The described climatic conditions appear to allow horsetail sporophytes ofdiffering species to grow in close proximity to one another. The damp, mild environmental conditionsalso favour the survival of normally very short-lived horsetail spores, permit prolonged growth of thegametophyte generation and provide a near-constant film of free water, facilitating male gamete transferconducive to cross-fertilisation (Page & Barker 1985).
The Massenerhebung effect in mountain areas
The ecological overlap of related species in Equisetum and in several other plant groups isprobably facilitated by the wonderfully named 'Massenerhebung effect', a German term literally meaning,'mountain mass elevation effect'. This is a meterological concept that was introduced by A. de Quervainin 1904 to account for the observed tendency for temperature-related parameters such as treeline andsnowline to occur at higher elevations in the Central Alps than on their outer limits (Barry 1981). Theconcept stipulates that climatic and vegetation zones occur at lower altitudes on isolated mountainsthan they do in mountain blocks of increasing scale (Wardle 1974; Johns 1985). The energy physicsrelating to this mass-elevation effect are complex, however, and it can only be applied after verycareful consideration of the specific local topographical and meteorological factors involved (Barry1981, pp. 49-50).
Continentality versus Oceanity
The Massenerhebung effect relates to the much more generally applicable climatic concept ofContinentality versus Oceanity. Applying either of these two concepts, we find that the lowelevation, the small, isolated mass, and the maritime position of British and Irish mountains results ina greatly steepened temperature lapse rate and a marked compression and lowering of our vegetation zonesin comparison with continental European uplands, including the Alps (Barry 1981, p. 265). The overalleffect of this pattern of climatic variation allows southern species to migrate northwards along westernAtlantic coasts, avoiding cold winters. This has produced, for instance, the highly unusual and veryfamous mix of phytogeographic elements found in the flora of the Burren region, Co Clare (H9). At thesame time, montane or alpine plants and animals, with their requirement for cool summer conditions, candescend to lower levels and, in W Ireland, some of these come right down slope, very close to sea level(Praeger 1934, section 67; Webb 1983; Page & Barker 1985; Nelson & Walsh 1991).
Equisetum species mingling
Thus plant species, including sporophytes of our eight native Equisetum species, are often foundgrowing much more closely together in the western parts of Britain & Ireland than they would undermore continental growing conditions, where they would commonly be altitudinally, geographically andecologically separated. Indeed, in the case of some other Fermanagh pteridophytes, colonies of four ormore different but closely related species are often found growing intermingled, and ecological andgenetic isolating mechanisms no longer apply. This enables the observed increase in frequency ofhybridisation.
Reduced competition?
Perhaps we should not ignore the fact that in the case of Ireland, being a more ancient island thanBritain, post-glacial species immigration was cut off earlier by sea level rise and thus thespecies-poor Irish flora presumably presents a less competitive environment than otherwise to newlyarrived genetic combinations in the form of the prothalli of Equisetum hybrids. A similarargument would of course apply when comparison is made between the depauperate flora of Britain and thatof mainland Europe.
E. × trachyodon is usually remote from its parent species
A comparative analysis of the geographical stations of species in the genus Equisetum and theirthen known hybrids was made around 1985 by Page and Barker based on updated hectad maps from The FernAtlas (Jermy et al. 1978). This showed quite clearly that hybrids and their parentspecies behave quite differently in the two subgenera. In subgenus Equisetum, for every hybrid,examined over the British Isles as a whole, there is an almost 100% coincidence or association betweenthe distribution of hybrid stations and the presence of both parents in the same or immediately adjacenthectad grid-square. The picture for subgenus Hippochaete is quite the opposite and, for E.× trachyodon, both parents were present in the same or adjacenthectads in less than 20% of its sites.
Our finer scale Fermanagh station analysis mentioned above shows the separation of E. × trachyodon and its parent species is even more pronounced than theseworkers showed. Page & Barker (1985) considered it likely that whilst a small amount ofhybridisation may continue to take place in subgenus Hippochaete, the geographical evidencesuggested to them that the majority of clones found today are long-established in their particularsites. Indeed, some clones may be very ancient and their parent species appear to have locally died out,or have been ousted by competition with their hybrid progeny.
European and world occurrence
Beyond the shores of Britain & Ireland, E. × trachyodon is locallyfairly frequent in Norway, is widely scattered throughout Iceland and occurs in a few provinces inSweden and Finland. It has also been reported from at least one station in S Greenland, although RSF hasheard that other botanists have cast doubt on the identification: one of the parent species, E.hyemale, is said to be unknown there (Böcher et al. 1968; Jonsell et al. 2000).This might not be as significant as it first appears since exactly the same situation applies in Irelandwith respect to E. × moorei Newman (E. hyemale × E. ramosissimum) (Moore's Horsetail), since E. ramosissimumDesf. (Branched Horsetail) is absent. In addition to the foregoing, E. ×trachyodon is also reported from scattered localities in Czechoslovakia, France, Germany,Switzerland, Hungary and in parts of Russia and temperate N America (Duckett & Page 1985).
Names
The name 'trachyodon' is a combination of two Greek words meaning 'rough teeth' (Gilbert-Carter 1964).]
Threats
Clearing of riverbanks for fishing or agricultural purposes. Locally this is especially problematic alongthe Colebrooke River.
Native, scarce and local. Circumpolar boreo-arctic montane.
1939; Praeger, R.Ll.; Spectacle Lough, Dresternan Td.
March to December.
Growth form and preferred habitats
Fertile and sterile stems of Variegated Horsetail are identical except for the presence or absence of thesmall, pointed terminal cone which sheds spores in July and August. When taken together, the veryslender, generally unbranched, often prostrate, evergreen stems of E. variegatum and the whiteteeth on the nodal sheaths, which Page so aptly describes as looking like "a broad gothicarch" (Page 1997), clearly distinguish the species from the hybrid it forms with E. hyemale(Rough Horsetail), E. × trachyodon (Mackay's Horsetail).
Variegated Horsetail is a scarce and local, rather variable calcicole species, which in Britain &Ireland occurs in a remarkably wide variety of more or less open, damp to wet, base-rich or calcareoussites, often by running water, or where there is movement of groundwater, even if only at subsoil level.Once established, its rhizomatous growth enables it to form large, compact stands as it still does atPraeger's original fen site at Spectacle Lough, the first Fermanagh site on record. However, it can alsobe found as scattered individual shoots at some of its more obviously flushed sites, both lowland and athigher elevations. Variegated Horsetail colonises seasonally flooded depressions and damp hollows inseveral disused quarries in Fermanagh, and it also occurs in similar ground under a hedge in thetownland of Clontelaghan near Kinawley.
In other parts of Britain & Ireland, E. variegatum is reported in periodically flooded coastalsand dune slacks and from mountain ledges, neither of which it occupies in Fermanagh (Stewart etal. 1994). As is the case with E. hyemale (Rough Horsetail), in the more northern part ofits range, E. variegatum occupies a much wider range of habitats, becoming more or lessindifferent to base-status and lime, and being found in much drier situations. In Iceland, for instance,it grows on dry scree slopes and in dry heath (Jonsell et al. 2000).
Fermanagh occurrence
In Fermanagh, E. variegatum has been recorded in a total of 21 tetrads, three of which havepre-1975 records only. It is the eighth most frequent horsetail in the VC. As the tetrad distributionmap indicates, the majority of sites are around Lower Lough Erne or on the higher ground to the SW ofit.
Irish occurrence
Compared with the five other VCs in N Ireland, Fermanagh undoubtedly has the greater representation ofthis species. Tyrone (H36) also has three inland sites for Variegated Horsetail (McNeill 2010), but thespecies is completely absent from Co Armagh (H37) and most of the few remaining N Ireland sites arecoastal (NI Vascular Plant Database 2010).
Elsewhere in Ireland, E. variegatum is thinly scattered, occasional to rare down the E coast andacross the Midlands (Jermy et al. 1978; Webb et al. 1996).
Variation
Reflecting the wide variety of quite different habitats E. variegatum occupies in these islands, anumber of ecotypes have evolved within it. These differ from one another in morphology (ie form,appearance and size), ecology (ie habitat and related features of growth) and in their geographicaldistribution. Having said this, in some cases ecotype ranges overlap to a degree not yet properlyunderstood. Since ecotypes maintain their distinctive forms when grown together under identical gardenconditions, the differences they display must be genetically determined, a feature unique amongsthorsetail species in Britain & Ireland (Page 1997).
The most widespread Irish ecotype is var. majus, which as the name suggests is larger than themost typical form found throughout much of Britain, var. variegatum. Var. majus growserect, rather than decumbent (ie leaning over or lying down, at least in part), and it is generallysomewhere between 20 and 80 cm tall, with stems about 3 or 4 mm in diameter, making it half as thickagain, or up to twice the width of var. variegatum (Webb et al. 1996; Page 1997).
The existence of these ecotypes undoubtedly provides a useful categorisation and description of thevariation within the species (Page 1997). They include a coastal sand-dune form, var. arenarium,common enough elsewhere in Britain & Ireland, but only of incidental interest to us since Fermanaghhas no coastline. Another Irish form, but remote from Fermanagh, is var. wilsoni, which isconfined to Co Kerry (H1 & H2).
At the same time, it is important to recognise that E. variegatum is a phenotypically very plasticspecies, each ecotype being capable of displaying considerable modification of form and scale withrespect to levels of a wide range of common local environmental variables (Stark 1991). Environmentalvariables include, for instance, shade, exposure, moisture, competition from other plants, trampling andgrazing pressure.
Similar observations of a more local nature can be drawn from Paul Hackney's comparative study of E.variegatum at four sites in N Ireland. Three of the selected sites were coastal sand dunes, butthe fourth was the fen shore of Carrick Lake in Fermanagh. Here, Hackney found that the VariegatedHorsetail occupied ground with a 90% cover of mosses. It was typically robust, growing up to 50 cm tall,but where the vegetation had been subjected to grazing pressure, the plants only reached 20 cm in height(Hackney 1981). Hackney did not attempt to recognise E. varietatum ecotypes and, since in most ofour survey we used the 1977 edition of Webb's An Irish Flora as our field guide, we have not doneso either.
British occurrence
E. variegatum is a rather scarce plant in Great Britain, mainly concentrated in the N and W,becoming very rare in the C and S of England (Stewart et al. 1994; Stace 1997). The number ofpre-1930 or pre-1970 hectads from which the plant has no recent records suggests the species is indecline and maps indicate that this is happening throughout both Britain and Ireland (Preston etal. 2002). Clearly this is a matter for concern.
European and world occurrence
In terms of geographical distribution, Variegated Horsetail is described as a northern-montane species(Stewart et al. 1994) or a circumpolar boreo-arctic montane species (Preston & Hill 1997).The latter may appear somewhat long-winded but it does summarise a range which stretches in Europe fromIceland and the Arctic Isles, to the northern tip of Scandinavia, thinning markedly towards the S ofboth Norway and Sweden, and present only at the northern tip of Denmark. In southern areas of thesethree Scandinavian countries it is also present as an introduction (Jonsell et al. 2000, Map onp. 25). E. variegatum then extends southwards in a somewhat scattered manner until it reaches afurther centre of distribution in the Pyrenees, the Alps and other C European mountains (Jalas &Suominen 1972, Map 32).
Further east, E. variegatum reaches the Altai Mountains of C Asia, Manchuria and Japan. In NAmerica, it stretches from the S Rocky Mountains to Labrador and around much of Greenland's coast(Hultén 1962, Map 45; Jonsell et al. 2000). Along the NW Pacific States of N America, the speciesis represented by a different form, E. variegatum subsp.alaskanum (A.A. Eaton) Hultén.
Names
The Latin specific epithet 'variegatum', means 'irregularily coloured' (Gledhill 1985), and the referenceto the black and white banded sheath on the slender green stem is obvious. The English common name is asimple book name translation requiring no comment. As the plant is rare or scarce and it is easilyoverlooked, it has not accumulated any English folk names.
Threats
A number of sites are vulnerable to changes in agricultural practices. Locally, one quarry site wasdestroyed when the ground was covered in concrete.
Native, very common, widespread and locally abundant. Circumpolar
boreo-temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This distinctive, erect, emergent aquatic or semi-aquatic rhizomatous, deciduous horsetail is veryvariable in size and in degree of branching. It is most commonly and abundantly found in still orslow-moving, shallow water by lakes, ponds and ditches, a habitat where it quite often represents thedominant colony-forming species. Even when dense pure stand communities of Water Horsetail are found, beadvised by Wolfe-Murphy et al. (1992), who wrote with bitter experience, that the rhizomes areless robust than those of rather larger emergent species such as Schoenoplectus lacustris (CommonClub-rush), and they do NOT form platforms that can bear the weight of the average botanist!
In lakes and ponds, E. fluviatile frequently forms large, dense stands, either pure or accompaniedby a very long list of other common emergent wetland species such as Schoenoplectus lacustris,Cladium mariscus (Sword Sedge), Carex rostrata (Bottle Sedge), C. elata(Tufted-sedge), Sparganium erectum (Branched Bur-reed), Glyceria fluitans (FloatingSweet-grass) and Phragmites australis (Common Reed).
E. fluviatile can also persist, but to a lesser extent, in shallows in bays of larger water bodieswhich by their nature are subject to more water turbulence and wave-induced physical scour. In thesecirc*mstances, where Phragmites australis (Common Reed), Schoenoplectus lacustris orTypha latifolia (Bulrush) often represent the deeper water dominants, E. fluviatileregularly replaces Eleocharis palustris (Common Spike-rush) when water is deeper than about 50 cm(Spence 1964). Water Horsetail can also be the dominant species and regularly forms dense, almost purestands in water 1.5 m deep or deeper (Spence 1964; Wolfe-Murphy et al. 1992; Page 1997).
Like its common relatives, E. arvense (Field Horsetail) and E. palustre (Marsh Horsetail),the rhizome of E. fluviatile runs much deeper in the soil than the underground organs ofassociated species, thus avoiding most root competition with them, if not altogether.
Identification
Individual stems are readily identified by the large central hollow that occupies between 80 and 90% ofthe stem diameter. As with several other Equisetum species, when E. fluviatile is in moreopen, unshaded situations, its stems are typically unbranched, or they bear only short, sparse lateralbranches. The latter are usually irregularly whorled near the middle of the aerial length of the stem,but in more shaded situations, for instance in marsh or reed-swamp among fairly dense tall grasses,sedges and rushes, or under trees or scrub in fen carr, Water Horsetail regularly produces regularlywhorled lateral branches on the emergent portion of its stems (Preston & Croft 1997; Page 1997, p.446; Rose 1989, plate 52).
Fermanagh occurrence
Fermanagh has a huge number and variety of lakes, both very large and small, and they exhibit an enormousdiversity of water chemistry (Gibson 1988). It is therefore not surprising that the shores of thesesupply E. fluviatile with plenty of scope for colonisation, and it is by far the most frequentlyrecorded horsetail in the VC. While it has been recorded in 329 Fermanagh tetrads, 62.3% of those in theVC, E. fluviatile is not the most widespread horsetail, a distinction held by E.arvense (Field Horsetail).
Wide ecological tolerances
Water Horsetail is very remarkable for the extreme width of its ecological tolerances with respect to aspectrum of inter-related environmental factors, restricted levels of which typically curtail the growthand govern the occurrence of most other wetland plant species. The tolerances of E. fluviatileinclude levels of nutrients (from oligotrophic to eutrophic), lime content (from starved to rich),acidity-alkalinity (from pH 4-7.5), light (from full sun to half-shade) and exposure (from stillbackwater to open, moderately wave-beaten shore). The substrate textures it tolerates vary from clean,firm, mineral sand to silty, smelly, organic mud, deep enough to suck the boots off you!
Water Horsetail can also survive in oxygen-depleted, highly anaerobic, hydrogen sulphide-releasingconditions that exclude many other aquatic species (Grime et al. 1988; Wolfe-Murphy et al.1992; Preston & Croft 1997). The very large central cavity in the hollow stem of E.fluviatile is considered to be an adaptation allowing air to diffuse downwards to the rhizome,in which latter organ, unusually among Equisetum species, the wide lumen persists. Very possiblyit is this unusual morphological property which enables the rhizome of E. fluviatile topenetrate, grow and persist in anaerobic layers of mud (Page 1997).
In N Ireland, Wolfe-Murphy et al. (1992) carried out a detailed lake survey for government from1988-1991 covering all six counties in the Province. This generated a macrophyte vegetationclassification of lake vegetation using the computer program 'Twinspan' (unpublished report to DOE, NI,p. 294). The study found that E. fluviatile was abundant or dominant in eight of the 30 shorelineplant communities these workers defined in N Ireland, indicating the exceptionally wide ecological rangeof the species. Water Horsetail is common on exposed mud or shallow water in sheltered backwaters oflarger lakes, and occasionally so in sluggish rivers and streams, particularly if there is only a minorfluctuation in water levels. It also frequents, in a more scattered manner, the closed turf vegetationof marshes, swamps and Salix-Alder fen-carr habitats, where its status in these plant communitiesis that of a minor companion species, except when shade seriously restricts the more competitive speciesaround it: reduced light allows rhizomatous E. fluviatile to reassert its vigour and develop amore significant presence (Grime et al. 1988).
Reproduction
Again as is the case in both E. palustre and E. arvense, although E. fluviatileproduces vast numbers of cones and spores, conditions for the completion of the full sexual life-cycleare stringent, and prothalii are very seldom observed (Page 1967; Duckett & Duckett 1980). It isthus very probable that increase and dispersal of Water Horsetail is heavily dependent on vegetativereproduction, achieved mainly by fragmentation. Propagation involves either free-floating segments ofbroken stem, or rhizome fragments. E. fluviatile does not possess tubers (Page 1997).
Herbivory and incidental vegetative propagation
Ducks, geese and other waterfowl feed on the stems of the plant (Fassett 1957), and Coot have also beenobserved biting off stems and using them for nest building (Praeger 1934). In both these circ*mstances,the birds were observed deliberately breaking excessive numbers of Equisetum shoots, more of themthan they actually used (and in the case of feeding birds, they broke them into many small segments), sothat some of the stem pieces could disperse without being used. These stem fragments can develop rootsand may thus propagate the plant on suitable wet terrain (Praeger 1934; Page 1997).
British and Irish occurrence
E. fluviatile is a very common, widespread and locally abundant species throughout most ofBritain & Ireland occupying a wide variety of habitats. It has declined quite substantially in thelast 50 years, due mainly to drainage of smaller wetlands and poor, unsympathetic management of othersites (C. Dixon & T.D. Dines, In Preston et al. 2002).
Environmental pressures and species change
The distribution of Water Horsetail in both Britain and Ireland has undoubtedly been somewhat curtailedin the past century by agricultural drainage and other forms of development. These habitat pressures aremost obvious in the most heavily populated and intensively farmed areas in the south of England (Grimeet al. 1988; Preston & Croft 1997). At the same time, in other areas of these islands it ispossible that E. fluviatile may have benefited from the suppression or demise of other aquaticand marsh species more sensitive to increased eutrophication than it. This is not to suggest that E.fluviatile tolerates extreme levels of organic pollution or sewage-induced accelerated culturaleutrophication. Study of a polluted lake in S Wales showed the species declined noticeably under suchcirc*mstances (Wade 1999).
In comparison to crop plants and terrestrial herbs, aquatic macrophytes have not been much studied byscientists working on climatic change. However, in a three year study, Ojala et al. (2002) foundgrowth and reproduction of E. fluviatile was sensitive to a rise of around 2.5 to 3°C.
European and world occurrence
Water Horsetail has a widespread and more or less continuous distribution in boreal and temperate partsof Europe that is very similar to the occurrence of E. palustre. However, the distribution ofE. fluviatile thins much more noticeably than E. palustre in the Mediterranean basin(Jalas & Suominen 1972, Map 34). E. fluviatile spreads eastwards from Turkey and theCaucasus, through temperate Asia to N Japan and N America. Only its absence from Greenland prevents itfrom being circumpolar (Hultén 1962, Map 96; Hultén & Fries 1986, Map 15; Jonsell et al.2000).
Names
Since it so frequently occurs in muddy ground, E. fluviatile used to bear the name 'E.limosum', 'Mud Horsetail', the Latin specific epithet being derived from 'limosus', meaning, 'ofmarshy or muddy places'. An earlier name of the plant was 'E. heleocharis', derived from twoGreek words, 'helos', meaning 'marsh', and 'karis', meaning, 'charm, grace, or beauty' (Gilbert-Carter1964; Johnson & Smith 1931). By comparison, the modern specific epithet, 'fluviatile' is derivedfrom the Latin, 'fluviatilis', meaning, 'growing in a river or in running water' (Stearn 1992). In thecurrent writer's opinion, this is an inaccurate and completely misleading indication of the normalhabitat of this horsetail, which more often is characterised by still or slow-moving water.
Additional English common names for E. fluviatile include 'Smooth Horsetail', a feature of thestems, said, in the past, to make them acceptable to cattle as food. However, reference to Linnaeus inthis connection suggests he was really referring to E. telmateia (Great Horsetail), since thelatter was his 'fluviatile' (Grieve 1931; Step & Jackson 1945).
Other interesting local names include 'Paddock Pipes' and variants thereof (eg paddow, paddie andpuddock). 'Paddock' is a Scottish name for frogs, making it rather appropriate for this horsetail, butit is devalued by being used for other horsetails (Britten & Holland 1886). Page (1988) mentions theinteresting name, 'Trowie Spindles', but offers no details of its origin or derivation.]
Threats
Drainage probably poses the only threat likely to affect this species locally in Fermanagh.
Native, occasional.
1939; Praeger, R.Ll.; north of Enniskillen.
May to November.
Growth form and preferred habitats
Typical plants of the deciduous perennial hybrid E. × litorale arefairly well branched in the middle portion of the stem, with the upper half to one third of the stembeing long, curved and unbranched. The internodes, when squeezed gently give slightly, and then 'bounceback' to their original diameter. In comparison, the internodes of E. arvense when squeezed inthis manner do not 'give' at all.
Although like many other horsetails, E. × litorale most frequentlycolonises open, disturbed ground, Shore Horsetail possesses hybrid vigour and is strongly competitive,often forming large colonies. In fact, like both its parents, E. ×litorale can occur in a very wide variety of damp to apparently quite dry habitats (Page 1997).
In sharp contrast to our other most frequent Fermanagh hybrid horsetail, E. × trachyodon (Mackay’s Horsetail), more often than not Shore Horsetailoccurs in close proximity to one or both of its parent species (Page & Barker 1985). However, we dohave at least one site on the Tempo River where E. × litorale occurswith both its parents, together with E. telmateia (Great Horsetail) and E. × trachyodon!
The phenomenon of many horsetail species growing together in close proximity and the relatively highfrequency of hybrids in western parts of British & Ireland has already been discussed at length inthe current authors E. × trachyodon account, so interested readersare requested to look there for more information and opinion.
Variation
Plants of Shore Horsetail are usually intermediate between the parent species in most characters, butthey are extremely variable, the variation undoubtedly being induced by very local environmental growingconditions. As a result, when occupying relatively dry conditions E. × litorale closelyresembles E. arvense and, when the habitat is wetter, its features are most like E.fluviatile.
Fermanagh occurrence
This vigorous, competitive hybrid has now been recorded in a total of 60 Fermanagh tetrads, 11.4% ofthose in the VC, 52 of them containing post-1975 records. Despite figures like the above, Preston &Croft (1997) still regard Shore Horsetail as frequently overlooked and under-recorded in much of Britain& Ireland.
In Fermanagh, this hybrid between a terrestrial and an aquatic horsetail species most often grows ondamp, bare gravelly ground near our larger lakes, in ground that is either permanently wet, or issubject to at least occasional flooding. It is especially frequent around Lower Lough Erne. Very rarelyit is found in open peaty mud on bogs, for example at Rossgweer Bog, or on damp roadside verges. OurFermanagh stations closely fit the types of habitat it occupies elsewhere in Britain & Ireland,described by Preston & Croft (1997) as marshy ground, near bare or disturbed soil.
Of the three hybrid horsetails in Fermanagh, E. × litorale is the morefrequent, being known from 52 post-1975 tetrads compared with 22 for E. ×trachyodon, and just one for the extremely rare, weak and very probably ephemeral E. × dycei.
Reproduction
E. × litorale produces cones in June and July, but the spores andreported to all abort (Duckett & Page 1985).
British and Irish occurrence
It is interesting to note that Praeger, who first found this hybrid horsetail in Fermanagh in 1939, waswriting in 1917 of it having just, "one station in each of the three kingdoms, England, Scotlandand Ireland". The New Atlas records the plant in Ireland from a total of 175 hectads withpost-1970 dates scattered across the island, and 315 post-1970 hectads throughout Britain, butconcentrated mainly in the N & W. The patchy occurrence in isolated VCs however indicates a degreeof recorder bias, eg Surrey and Worcestershire (VCs H17 and H37) (New Atlas).
European and world occurrence
Apart from Britain & Ireland, E. × litorale is widespread inFennoscandia and very likely also occurs throughout the whole northern boreal range of the parentspecies. It probably occurs everywhere in the world the distributions of the parent horsetail species,E. fluviatile and E. arvense, overlap (Jalas & Suominen 1972, Maps 34 & 38; Hultén1962, Maps 96 & 98; Duckett & Page 1985; Hultén & Fries 1986, Maps 15 & 19; Jonsellet al. 2000).
Names
The Latin epithet, 'litorale', is derived from 'litus', meaning 'shore' and obviously refers to the mostfrequent habitat of the hybrid – lake shores (Gilbert-Carter 1964).
Threats
None.
Native, very rare.
8 August 1991; Wolfe-Murphy, S.A.; marshy ground, south shore of Edergole Island, Upper Lough Erne.
Growth form and preferred habitats
There are no less than five hybrid horsetails in Ireland, the most common of which is E. × litorale (Shore Horsetail). However, despite the high frequency ofits parent species, E. × dycei is extremely rare in both Britain& Ireland. This horsetail hybrid is small, slender, with unbranched or sparsely branched shoots anda very long branchless terminal portion. It is very spindly or sickly-looking, lacks hybrid vigour(heterosis) and resembles a smaller, weaker form of E. palustre (Marsh Horsetail) or adebilitated form of the usually very much more vigorous hybrid E. ×litorale (E. arvense × E. fluviatile). E. × dycei is mainly confined to man-made, open, muddy roadside ditchhabitats (Page 1988 & 1997).
Irish occurrence
On a joint Botanical Society and British Pteridological Society outing to NW Ireland in July 1984, MauraScannell of Glasnevin Botanic Garden in Dublin and Page together discovered the first Irish station forE. × dycei along the southern shore of Doon Lough, in Co Leitrim(H29), adjacent to Fermanagh. In this case, the hybrid was growing on the marshy shore of the lake, bythe roadside, along with both parents and it was recognised by Page on site. A voucher of this find isnow in DBN. In July 1986, Scannell found a second station in N Kerry (H2) on the shore ofInisfallen Island, Lough Leane, Killarney. A voucher for this latter discovery was sent to the NationalMuseum of Wales herbarium in Cardiff (NMW), where Page again confirmed the identification(Scannell 1995).
The 1997 second edition of Page's book, Ferns of Britain and Ireland deals with E. × dycei in great detail. The account includes a very small andquite inadequate map, which nevertheless indicates a total of five stations in the Republic of Ireland,the two mentioned, plus one in Kerry, and two additional stations in Co Clare (H9)(Page 1997, p. 492).While one may deplore the poor quality of Page's map, there is no map at all in the BSBI's NewAtlas, nor on the accompanying compact disk (Preston et al. 2002).]
Fermanagh occurrence
Sean Wolfe-Murphy made the solitary Fermanagh record listed above when taking part in the NI Lake Surveyin 1991. The plant was determined by Paul Hackney from a voucher specimen in BEL. It was growingwith E. fluviatile (Water Horsetail) but the other parent was not recorded. This is a newFermanagh County Record and was the first record of the hybrid in NI. The BEL voucher has theaccession number H37907. A second NI station was subsequently discovered in June 1999 by P. Hackney atGrange More Td, just SE of Castlerock on the N coast of Co Londonderry (H40).
Ephemeral nature of the plant
In field notes relating to this hybrid, Page (1997) wrote that from his experience, E. × dycei seems an elusive and possibly short-lived plant and, indeed,by 2008 the Grange More plant had completely disappeared (P. Hackney, pers. comm. 2010). The EdergoleIsland station has never been refound either. A third find of this hybrid in NI was made in July 2008 byDave Riley, at Colm Harkin, near Swatragh, Co Antrim (H39). Vouchers for all three NI records are inBEL.
Earlier, Page (1973) suggested that the weak hybrids in this genus are those formed between ecologicallysimilar, but not very closely related pairs of species, while the vigorous ones are those betweenspecies pairs which are ecologically more different, yet closely interrelated.
In view of the abundance of both parents and their frequent overlap in many Fermanagh sites, we fullyexpect that there must be further stations for this perhaps casual hybrid to be found, but only a veryexperienced eye could detect them.
Names
Chris Page gave E. × dycei its names, both Botanical and English,calling it 'Hebridean Horsetail', after its first discovery in 1962 on these Scottish islands.
Threats
None.
Native, very common, widespread and locally abundant. Circumpolar wide-
boreal and introduced widely in the southern hemisphere.
1881; Stewart, S.A.; Co Fermanagh.
March to December.
Growth form, identification and preferred habitats
As any gardener or farmer will testify, E. arvense is a too frequently found rhizomatous and tuberproducing perennial weed, which grows so deeply and spreads so relentlessly and rapidly that it iswidely regarded as nearly impossible to eradicate (Holm et al. 1977; Grime et al. 1988).The acute, green, spreading teeth on the side branches of E. arvense help distinguish its sterilestems from the rather similar ones of E. palustre (Marsh Horsetail), in which the correspondingbranch teeth are black-tipped and tightly clasp the stem for their whole length. In terms ofidentification it is fortunate that at least some of the characteristics of the side branches areunvaryingly reliable: irrespective of the size of the sterile stem of the plant, the lowest internode oneach side branch of the stem of E. arvense is always equal to or longer than the adjacent stemsheath on the node from which it arises (Jermy & Camus 1991).
Typical habitats include rough grassland and disturbed areas in damp open woods, hedgerows, field-,river- and roadside banks, lakeshores, cliffs, screes, quarries, gardens, gravel paths and waste ground.Although Field Horsetail is able to grow in a very wide variety of soil types, it has a definitepreference for neutral or slightly base-rich conditions. It is really only common and vigorous wherethere is a high water table and the soil is poorly drained (Williams 1979; Cody & Wagner 1981).Having said this, E. arvense is recognised to be the most plastic and adaptable horsetail inBritain & Ireland and it can survive and function over a very wide spectrum of soil moisture andnutrient regimes (Page 1997).
Field Horsetail occurs in a huge range of more or less open habitats and like Ulex europaeus(Gorse), Pteridium aquilinum (Bracken), Tussilago farfara (Coltsfoot) and other vigorousperennial weeds, it particularly avails itself of soil disturbance. Frequently such disturbance is dueto mans' activities, but it can also be naturally occurring, for example in waterside habitats. Indeed,it has been suggested that river banks may have been the original natural habitat of E. arvense(Hauke 1966), and possibly of all Equisetum species.
Whatever the cause of soil disturbance, it creates an opening for the horsetail, providing bare ground ora gap in the vegetation into which the species then readily invades (Salisbury 1964; Grime et al.1988). The last mentioned authors found that in their main study area around Sheffield, E.arvense was absent in 'skeletal habitats' (ie bare rocks, cliffs, scree and walls). However, inFermanagh, we do have records of it colonising cliffs and screes, which might be anotherreflection of our very wet, more or less hyper-oceanic climate.
Fertile sporing and green vegetative shoots
Stems of E. arvense are of two types; the short, stout, pale- or buff-brown, cone-like, fertilesporing branches appearing for a fortnight or so in early April. As the non-photosynthetic sporingshoots appear before the green vegetative ones, they are sometimes referred to as 'precocious'. Thefertile shoots are soon followed by the much more slender, green sterile or barren stems, branched tovarying degrees. The green shoots are deciduous, growing and persisting until the frosts in late autumn,and then dying and disappearing in November, or by early December at the latest.
Rhizome extent and ecology
The extremely persistent, perennial, underground rhizome is long and branched and can reach a depth of1.5 to 2 m., or more (Page 1997). In warmer climates it can exceptionally extend down to around 6 m deep(Cody & Williams 1981). Having said this, Williams (1979) found that in a sandy loam in England, 50%of the rhizome dry weight occurred in the uppermost 25 cm of soil, a further 25% in the next 25 cm andonly 10% of the total rhizome material was found between 75 and 100 cm deep. He also found that ifarable ground containing E. arvense was left fallow for one or two years, rhizomes andstarch-filled tubers occurred much more shallowly, more than 80% of them being resident in the uppermost25 cm of soil. He concluded that this was most likely due to the horsetail being released fromcompetition by the absence of any crop and other work he quotes from different countries appears tosupport this suggestion (Williams 1979). Climatic factors such as the depth of soil warming are known toaffect the depth to which rhizomes and tubers penetrate, as will soil aeration and moisture status(Williams 1979).
The rapidity of rhizome spread reported is both amazing and alarming; for example, a 10 cm length ofrhizome planted in a growth room (presumably, although not specified, in the absence of competition)produced in one year a total of 64 m of growth in a vertical direction through branching (Cody &Wagner 1981). As far as the current author is aware, horizontal rhizome growth rates have not beenaccurately measured under field conditions, but an individual rhizome has been reported achieving aspread of 100 m (Weber 1903, quoted in Cody & Wagner 1981).
Nutrient requirements
The nutrient demands of E. arvense on soil fertility were concisely expressed in classic ‘cropversus weed’ terms by Salisbury (1964) when he wrote, "its filching of soil nutrients is veryappreciable". Hill et al. (1999) placed Field Horsetail among other rathernutrient-demanding weeds such as Cirsium arvense (Creeping Thistle) and Rumex crispus(Curled Dock), which typically require moderately fertile soil. Sinker et al. (1985)characterised E. arvense as requiring a medium to rich supply of phosphate, nitrogen and othermineral nutrients. Field Horsetail, however, appears as variable in this respect as in otherenvironmentally controlled characters, since in his Rothamstead pot experiments, Williams (1979) foundthat the horsetail seemed well adapted to growth in soil with low nitrogen and it produced little growthresponse to additions of it.
Silica content
The horsetails, particularly E. hyemale (Rough Horsetail), but to a lesser extent E.arvense, are well-known for their ability to take up and deposit silica in the walls of theepidermal cells of their shoots. The silica content of dry E. arvense can vary between 1.2 to6.9% and the ash can contain anywhere between 6.2 and 76% silica. This is why these horsetails have inthe past been widely used for scouring pots and as polishing and buffing tools (Cody & Wagner 1981).The low levels of silica in both limestone and deep peat soils probably limits the growth ofEquisetum species where these soils occur (Page 1997).
In mining areas E. arvense and E. palustre (Marsh Horsetail) are relatively frequent onmetalliferous spoil heaps, the soils of which most other plants find too toxic to colonise (Grime etal. 1988). E. arvense has an unusual ability to take up and accumulate heavy metals suchas copper, zinc, lead and cadmium, so that it has been proposed as a biological tool to monitor levelsof pollution (Ray & White 1979). Reports quoted in Holm et al. (1977), that horsetailaccumulates gold in quantities of up to 4.5 ounces per ton of fresh plant material, are exaggerationsbased on mistaken chemical analytic methods (Grime et al. 1988).
Ecological flexibility
It is the degree of ecological flexibility E. arvense displays in terms of both requirements andtolerances, which enables it to become the only horsetail that is a common weed of cultivated ground.Following a detailed systematic study, Hauke (1966) concluded that despite appearances, E.arvense is a hydrophyte! This decision calls into question how we define an aquatic plant, butwe can hardly believe that many biologists or ecologists would entirely agree with Hauke's use of theterm for this species. He supports his contention by pointing out that although Field Horsetail issometimes found growing in very dry habitats (for instance along roadsides and railways), on the basisof his observations and measurements, he believes that it is quite sensitive to moisture stress and thatit is able to survive in places where other plants cannot, simply because its extensive,deeply-penetrating rhizome system always manages to tap groundwater supplies.
Variation
Being versatile in terms of its ecology, E. arvense also demonstrates a very wide range ofenvironmentally induced modification (phenotypic variation) of both plant size and form, some of whichare easily demonstrated even within a single clone (Hauke 1966). Examples of this variation are veryclearly illustrated in Page (1997, pp. 439 & 442), but Hauke (1966) concluded that all thisvariation is only superficial and none of it merits taxonomic recognition.
Asexual reproduction and dispersal
E. arvense dispersal can readily be achieved by its lightweight spores, but without question italso spreads very effectively vegetatively, both by means of rhizome fragments and through its small,oval, root tubers being carried unwittingly in mud attached to machines, animals and boots.Establishment from spores involves the successful negotiation of several high-risk stages ofdevelopment, involving the ready release and transport of asexual spores and the production ofgamete-bearing prothalli, structures which are known to have an extremely narrow habitat tolerance. Theprothalli require bare muddy ground of high nutrient status, neither too wet nor too dry, and the entireabsence of shading and competition, even that from mosses and liverworts. In view of this, the rarity ofprothallus observations in nature is not all that surprising. Nevertheless, the existence ofEquisetum hybrids and the few studies of wild gametophytes that have been made, do prove thatsexual reproduction does happen. Even if it is a rare event, sexual reproduction will be significantsince it maintains genetic variability and heterogeneity within horsetail populations that could nototherwise occur (Duckett & Duckett 1980).
Sexual and vegetative reproduction
As with other hom*osporous (one type of asexual spores only), sporophyte (spore-bearing plants) (ie theferns and horsetails), vast numbers of spores are produced and released in the early spring (generallyfrom early April into May). A small proportion of these asexual spores germinate to produce aprothallus, on which separate male and female sex organs develop. Huge numbers of male gametes (sexcells) swim in a film of water to fertilise the much fewer female ova, and young sporophyte plantssuccessfully produced by this sexual cycle then grow and develop. Some of these juvenile sporophyteplants have been known to produce up to two dozen shoots and several tuber-bearing rhizomes by the endof their first season of growth (Page 1967; Duckett & Duckett 1980).
By comparison, vegetative dispersal and successful subsequent establishment appears very much moreprobable and this is generally regarded as the more usual method of reproduction of the species(Marshall 1986; Grime et al. 1988).
Sporophyte light requirements and colony development
Once the plant has colonised (by whatever means) and a rhizome has been established, growth of thehorsetail can be incredibly rapid, allowing the plant to quickly form a clonal patch. E. arvensehas such a terrible reputation among gardeners, fruit-growers and farmers (both arable andgrass-managing), that along with its relative E. palustre, it appears in Holm et al.(1977) book entitled, The World's Worst Weeds. This might be pitching its status a little higherthan it deserves. The reason for saying this is that, having no leaves, E. arvense entirelyrelies for its photosynthesis on its annual, green, wiry, branching stems, which grow up to 80 cm inheight. Thus most Equisetum species cannot tolerate much deep shade, and the horsetail plantcasts very little shade itself (Salisbury 1964).
E. arvense has been given an Ellenberg's indicator value by Hill et al. (1999) for itslight preference in the British Isles of '7' on a scale from '1' to '9', meaning that it is regarded bythese authors as a plant generally found in well lit places, although it may also sometimes occur inpartial shade. This light requirement limits the species competitive ability, and being relatively lowgrowing and possessing a comparatively insignificant canopy of its own, it never (or rarely ever),becomes a dominant plant after the manner of taller, aggressive and persistent invading weeds, such asgorse and bracken. Grime et al. (1988) draw a very interesting parallel between many of thefeatures of E. arvense and Tussilago farfara (Coltsfoot), both in terms of theirpioneering, colonising ability and weedy persistence.
Toxicity
Pretty well all species of Horsetails are dreaded worldwide, partly for their deep seated growth andtheir enduring survival ability, but also because the plants are so poisonous to cattle, sheep andhorses. Here again, comparison may usefully be made with Pteridium aquilinum (Bracken), sinceboth fern and horsetail contain a toxic enzyme called thiaminase, which destroys thiamine and creates avitamin B1 deficiency in any monogastric animals which graze them (Cody & Wagner 1981).
Equisetum species also contain high levels of a number of toxic alkaloids, of which the best knownis palustrine (Cooper & Johnston 1998). In this case, the poisonous principle is not destroyed bydrying and storage and horses can show clinical signs of poisoning when their hay contains as little as5% horsetail (Cooper & Johnson 1998). Fortunately the plants contain varying quantities ofsilicates, making them harsh to touch and unpalatable, at least in the fresh state, so that animals tendto avoid grazing horsetails and therefore poisoning is rare in Britain & Ireland.
The levels of toxins in Equisetum species suggest the possibility of allelopathy, ie toxicsuppression, directed towards surrounding competing plants. Work on this topic in Russia is regularlyquoted, which showed that when tested along with twelve other species, water extracts of E.arvense displayed the strongest inhibitory effect on seed germination and seedling vigour whenit was applied to 30 species of meadow grasses (Zelenchuk & Gelemei 1967 (in Russian), quoted byCody & Wagner 1981).
Weed control
Control of E. arvense is extremely difficult. Forking out, cutting and burning all prove a uselesswaste of time and effort. Eradication may be achieved in the long term by shading the weed out withtaller plants, which is what normally happens to the species in undisturbed natural vegetation (Page1997). In the garden, even as simple a matter as sowing a patch of Nasturtium has beenrecommended for this purpose (Allan 1978). Mulching with leaf compost is reputed to stop lateralmovement of Field Horsetail, and it has been reported that black plastic sheeting placed over infestedsoil killed rhizomes in the upper 60 cm of soil within three to four years (Cody & Wagner 1981).
Herbicide experiments
The choice of herbicides used against Equisetum infestation is governed by the crop or vegetationtype affected and the scale of the weed problem. For instance, it was found that MCPA applied after thehorsetail had completed emergence gave 100% control of aerial growth for the rest of the season and alsoreduced the number of horsetail stems emerging in the second year (Hoyt & Carder 1962, quoted byCody & Wagner 1981).
Glyphosate and other translocated herbicides, such as Asulam (which in particular is also effective insimilar manner against Bracken), can be used in uncultivated areas. Glyphosate herbicides are mosteffective if the horsetail is allowed to fully emerge and its stems are then crushed before applying thechemical in late summer (late July or August is probably the best time) (Marshall 1986). Crushing isrecommended because the slender stems and branches are covered with microscopic silica spicules andtherefore they are not easily wetted.
Fermanagh occurrence
E. arvense is both the most common and widespread horsetail on a world basis and the mostwidespread horsetail in Fermanagh, present in 433 post-1975 tetrads, 82% of those in the VC. However,since Fermanagh is a county extraordinarily well supplied with lakes and wet marshy or boggy ground, interms of record frequency, Field Horsetail remains second to E. fluviatile (Water Horsetail) by amargin of over a thousand records. E. arvense is found throughout the county, although it isuncommon on high ground and absent from aquatic situations, except the gametophyte, which is a pioneercoloniser of muddy water margins of lakes and reservoirs, very occasionally producing the sporophytegeneration (Duckett & Duckett 1980).
British and Irish occurrence
Field Horsetail is the most common and widespread horsetail in Britain & Ireland, being present inevery VC. The distribution thins slightly in areas of predominant deep peat soils in N Scotland and inthe SW of both islands (Jermy et al. 1978; Page 1997; Preston et al. 2002).
European and world occurrence
E. arvense is very widespread throughout most of Europe. It extends into the far south of Italy,but is more thinly present in the Iberian Peninsula. It occurs in most of the Mediterranean islands, butnot in Cyprus (Jalas & Suominen 1972, Map 38). It stretches eastwards through the Caucasus, theHimalaya, C China and Japan to the S USA. It was introduced into Mexico and New Zealand and may or maynot have persisted (Hultén 1962).
E. arvense is the most common and widespread species of the genus, being circumpolar chiefly inboreal latitudes and present though less common in the Arctic, including Greenland, Iceland (Hultén& Fries 1986, Map 19). It also occurs in a prostrate form, subsp. boreale (Bong.) Á. Löve, inthe Arctic circumpolar region including the mountains of Norway, Sweden, Iceland and the arctic islands(Jonsell et al. 2000).
Uses
The starch-filled rhizome tubers of E. arvense are eaten by ducks in Alaska (Hauke 1966) and inthe past N American Indians both peeled and ate raw stems of the fertile cone-bearing stems and used thedried ashes of the sterile stems to treat sore mouths (Cody & Wagner 1981). Grieve (1931) list manymedicinal uses for the fresh or dried sterile stems and their ashes, the main uses being as a diureticand astringent, to staunch bleeding (including nose bleeds and ulcers) and to counter acidity of thestomach.
Names
The genus name 'Equisetum' was coined by the ancient Roman writer, Pliny and is thought to have beenfirst applied by him to E. arvense. It is a combination of two Latin words, 'equus', a horse and'saetum', a bristle or hair, and it is thought to refer to the bristly appearance of the jointed stemswith their whorled branches (Gilbert-Carter 1964; Grieve 1931). The same notion also gave origin to theEnglish common name 'Horsetail', which is a direct translation of the medieval Latin name, 'caudaequina', under which it was sold in apothecary shops (Prior 1879; Grigson 1974). The Latin specificepithet 'arvense' is a common one, being derived from 'arvum solum', meaning 'arable land', on which theplant is often found growing (Gilbert-Carter 1964).
There are a total of 21 additional English Common names for Equisetum species in general inBritten & Holland’s reference work (1886), many of which carry watery connotations involving pipes,frogs (and tadpoles), toads and rushes, eg Tadpipes, Tad-broom, Toadpipes, Snake Pipes, Water Grass,Cat-rushes. Several other names refer to the bushy appearance of the branched stem, for example,Cat's-tail, Colt's-tail and Bottle-brush (Britten & Holland 1886). The weedy, unwanted nature of theplant, or its malign, pernicious presence is featured in a Welsh name which is translated as 'Evil man'sgarters', the evil man being a standard euphemism for the devil (Awbery 1984).
Threats
None.
Native, very rare. Circumpolar boreal-montane.
1904; Praeger, R.Ll.; scarp behind Poulaphouca cliffs, Western Plateau.
June to November.
Growth form and preferred habitats
E. pratense is a rhizomatous perennial with two forms of aerial shoot (vegetative and sporing),neither of which overwinter. The species is a northern one with an overall distribution markedlycircumpolar boreal-montane or arctic-alpine (Stewart et al. 1994; Page 1982 & 1997). InIreland it is a rare Irish Red Data Book species, confined to the northern province of Ulster,where it is most frequent in the glens of Antrim (H39), although even here it is really rare and local(Praeger & Megaw 1938; Curtis & McGough 1988, pp. 93-4; Hackney et al. 1992). E.pratense is rarely recorded, perhaps in part because it is not very distinctive in appearanceand it often occurs as scattered, diminutive individuals composing small, diffuse colonies. In order tolocate it, field recorders first need to encounter a good specimen colony and get to know its particularhabitat requirements. Thus it could easily enough be overlooked and under-recorded.
In texture and colour, the shoots of E. pratense most closely resemble the delicate ones of E.sylvaticum (Wood Horsetail), although unlike the latter, the slender primary lateral branchesare themselves unbranched. The flat tops of mature shoots are quite distinctive in appearance.
Like the majority of other horsetail species, it is a plant of well-drained slopes (including uplandglens and talus slopes below cliffs), where silty or sandy soil is flushed with, or subject to seepageof, base- and mineral-rich groundwater, and where shade, shelter and proximity to running water issufficient to keep humidity high and prevent any possibility of desiccation (Stewart et al.1994). A certain amount of winter flooding or mild scouring of base-rich E. pratense sites nearstreams and rivers may also be significant, producing surface instability and shallow erosion of the wetsoil, thus minimising the burden of competing vascular plant species while simultaneously providingplentiful silica (Page 1988). In Fermanagh, this could particularly be the case at the Cladagh RiverGlen site. To some extent, therefore, Shade Horsetail may be considered a pioneer species, growing bestin situations where relatively open conditions are maintained by these types of naturally operatingfactors. Both of the Fermanagh sites are relatively lowland, but the species is reputed to reach 850 min Glen Coe in Scotland.
Fermanagh occurrence
In Fermanagh, E. pratense is found in only five tetrads and is the rarest horsetail species in theVC. It is twice as rare as E. hyemale (Rough Horsetail), which is known in twelve post-1975tetrads), and much less frequent than two of our three hybrid horsetails, E. × litorale (Shore Horsetail) and E. ×trachyodon (Mackay's Horsetail). In fact, there are just two main sites (with subsites), in thecounty for Shady Horsetail in shaded, moist woods and scarps, but they are rather different from oneanother.
In the Cladagh River Glen (also referred to as the 'Marble Arch Glen'), E. pratense grows in onelarge patch on a moist, sloping bank in the light shade of an ash dominated woodland. It also has anumber of smaller colonies nearby, along the bank of the river either side of the adjacent path. At thesecond more extensive site on the scarp top of the Magho cliffs and the talus slope below that overlookLower Lough Erne, Shady Horsetail occurs thinly scattered within the canopy of a six km long stretch ofwet, mixed deciduous woodland, growing here amongst Luzula sylvatica (Great Wood-rush) andStellaria holostea (Greater Stitchwort).
Even when failing to cone and restricted to an entirely vegetative condition, the persistence of E.pratense is quite remarkable. Records of Shady Horsetail at Co Antrim sites (H39) extend back upto a century or more. The plant has not, however, been refound in Fermanagh at Praeger's original 1904site, which he (in the days before grid references) rather loosely described as being amongst rankheather on an open moorland scarp, behind (ie south of) Poulaphouca cliffs – a name rather widelyprinted across several cliff ranges on the one inch OS map (Praeger 1904). Nevertheless, the speciespersists on adjacent scarp tops and scarp woodland along the Cliffs of Magho and, indeed, if one cancall it such, the species has its Fermanagh headquarters here.
Sterility and changing winter temperatures
In Fermanagh, in common with elsewhere in Britain & Ireland in recent years, observation suggeststhat E. pratense is sterile and cones only very rarely and spasmodically, if at all. E.pratense appears to rely almost exclusively on vegetative reproduction for its increase andsurvival, and on rhizomatous creep for any local spread that it manages to achieve. The species is,however, long persistent in the vegetative state in Britain and Ireland, most of the sites known acentury or more remaining extant (Stewart et al. 1994).
As Page (1982 & 1997) points out, herbarium specimens from the 19th century indicate that E.pratense used to cone abundantly, and possibly did so regularly. However, it is possible toimagine that field botanists may have favoured or restricted their collecting to 'complete', ie sporingspecimens, thus unintentionally distorting the frequency of cones in the herbarium record.
Since the great majority of Shady Horsetail sites in Britain & Ireland are fairly remote, they areleft relatively undisturbed by man's activities. In the absence of human disturbance, the near sterilityof modern clones suggests some other environmental factor(s) controlling cone induction has changedsufficiently during the last 100 - 200 years to prevent spore production. The most likely rapidlychanging environmental factor that might affect horsetail fertility is winter temperature. Since about1850 AD there has been a gradual climatic warming in Britain & Ireland producing milder winters.This trend began just prior to current CO2 and other greenhouse gas-induced global warmingassociated with the industrial revolution. Even slight climate warming may have provided the significantbiological threshold beyond which E. pratense fails to produce sporing cones (Page 1997).
Other evidence of species decline
In Fermanagh, E. pratense is at the very southern edge of its W European geographical range and,because of its poor to negligible reproductive capacity in modern times, Page (1982 & 1997) fearsthat the species is in the process of dying out in Britain & Ireland. Species living at the extremelimits of their distribution are often represented by small, scattered, isolated populations, whichfrequently are either completely sterile or are reproductively weak. Compared to populations in thecentral parts of the species distribution, they tend to occur, as E. pratense now does here, insmaller numbers and in a much more restricted range of habitat conditions than occurs elsewhere withintheir overall distribution. These features indicate such species are suffering from both poorcompetitive ability and a lack of recruitment into fresh suitable sites.
In genetic terms, small, isolated populations inevitably lose vigour over extended periods of time,gradually losing genetically variability (sometimes referred to as 'genetic erosion'), as a consequenceof restricted gene flow and 'genetic drift' (ie the tendency for gene alleles to fix within smallpopulations at random, or even somewhat against selective forces) (Richards 1997, p. 46-9).
These processes associated with inbreeding lead to the accumulation of hom*ozygous gene alleles anddeleterious recessive mutants but, in polyploid species, such as most pteridophytes, these processes maydevelop very slowly indeed. Thus we cannot involve them in the much more rapidly occurring loss offertility that appears to be the case in E. pratense populations.
As a northern-montane or circumpolar boreal-montane species, E. pratense is also likely to findthe current rather dramatic rapid climatic warming in NW Europe unfavourable to its continued survivalin these islands, a situation leading to slow decline and eventual extinction as Chris Page (1982 &1997) has predicted. Thus E. pratense in Britain & Ireland already demonstrates several ofthe features associated with genetically weakened species populations (reproductively inadequate, small,scattered, more or less isolated populations lacking recruitment, displaying poor competitive ability).When this is compounded with the unsettling effects of an unfavourable warming climate, plus thethinning effects of random mortality inevitable in all small populations, with regret the current writeris forced to agree with Page's depressing verdict.
Along with other arctic-alpine, circumboreal and northern-montane plants, E. pratense must berecognised as a relict species in our latitudes of past cooler environments, and as such it is doomed tolocal extinction, probably in the not very distant future (Briggs & Walters 1997, pp. 411-419).
British occurrence
In England, E. pratense is already completely confined to a few scattered stations in the northernPennines, but it is much more widespread in Scotland, typically occurring as small patches in the lowervalleys in the Highlands and islands, but occasionally found on more open upland moorland where drainageand spring water flushes provide nutrient enrichment of the turf (Jermy & Camus 1991; Stewart etal. 1994; Page 1997).
European and world occurrence
In Europe, E. pratense has a very pronounced circumpolar boreal-montane range that is distinctivefrom any other European horsetail. It ranges from Iceland, north to within the Arctic Circle inScandinavia, and south to the C European mountains and the Caucasus. However, it is completely absentfrom both the French Alps and the Pyrenees (Jalas & Suominen 1972, Map 37; Jonsell et al.2000, p. 21). The distribution then stretches eastwards across most of N Asia to Manchuria, N Japan andacross N America, south to about 40o N. It is absent, however, from Greenland and from muchof W Europe (Hultén 1962, Map 83; Hultén & Fries 1986, Map 18; Jonsell et al. 2000).
Names
E. pratense is neither distinctive enough nor sufficiently common to have been given localEnglish common names, 'Shady Horsetail' or 'Shade Horsetail' being mere invented book names. The Latinspecific epithet, 'pratense', means 'growing in meadows' (Gilbert-Carter 1964), which fits the behaviourof the species in Scandinavia and undoubtedly in other northern parts of its range.
Threats
Both Fermanagh sites are protected; the Cladagh River Glen is a National Nature Reserve and part of theMagho cliffs is a Forest Nature Reserve. They could still be vulnerable to grazing or trampling.
Native, frequent and widely scattered. Circumpolar boreo-temperate.
1881; Stewart, S.A.; Co Fermanagh.
March to November.
Growth form and preferred habitats
The only horsetail with branched lateral branches, this feature, together with the graceful, finetextured, drooping, pale yellow-green stems and branches makes Wood Horsetail quite unmistakable and themost beautiful of our horsetails. Sterile and fertile stems are similarly green and branched, the latterdiffering only in that in May each year they bear a terminal, narrowly oval cone, about 10-20 mm long.
E. sylvaticum is a small to medium-sized, deciduous, colony-forming, rhizomatous horsetail ofmoderately wet, usually deep, acid to neutral, clay or peaty soils, which, since the species iscalcicole, must be more or less constantly flushed with moderately calcium- or base-rich groundwater. Inits preference for half-shade and the root-flushing requirement, Wood Horsetail, at its lowland sites atleast, is rather similar in its ecological demands to E. telmateia (Great Horsetail) and, to alesser extent, E. palustre (Marsh Horsetail), with both of which it frequently associates. Theroots of both E. sylvaticum and E. telmateia have an absolute requirement for some degreeof water movement, while E. palustre can tolerate almost no flow (Sinker et al. 1985; Page1997).
While Wood Horsetail tends to be most prevalent in damp, sheltered, at least partially shaded sites withconstant high humidity (for instance, in fen-carr, woodland margins, glades, hedgerow-, river-, stream-or ditch-banks), it can frequently be found in very much more open and exposed upland positions. InFermanagh, examples of the latter habitat occur on the Drumbad Scarps, Poulaphouca cliffs and ToppedMountain. In these more exposed sites, as might be expected, it is both rather dwarfed and much moresparsely branched, which makes it quite difficult to recognise (eg see illustration in Page 1997, p.470).
In Fermanagh, near the NW coast of Ireland, E. sylvaticum can also be found in open, more sunnyconditions, as well as in more typical semi-shade. Open habitats here include stabilised scree,quarries, flushed heath and moorland, cut-over bog and even in churchyards. The important essentialproviso is that sufficient moisture and the nutrient requirements of the species must be met. The uplandtendency of E. sylvaticum on damp, peaty heath and moorland, as well as on cliffs and moresheltered gullies and stream-sides, together with its known woodland shade preferences, suggests thepossibility that the upland plants of this horsetail might be relicts of former forest in these sites(Jermy et al. 1978). As with several pteridophytes of moist western heathland, Wood Horsetailappears to have swapped the shade, shelter and constant high humidity under woodland canopy, for anoverall heathland region 'Atlantic' climate of overcast, grey, cloudy skies and regular, almost daily,precipitation, evenly spread throughout the year (Gimingham 1972, pp. 11-13; Page 1988, p. 288).
Fermanagh occurrence
E. sylvaticum is the fifth most frequent horsetail in the Fermanagh Flora Database, beingrepresented in 147 tetrads, 27.8% of those in the VC. Eight tetrads contain pre-1976 records only,suggesting some loss of habitat. As the tetrad distribution map indicates, Wood Horsetail is verywidespread, but especially frequent on the western limestones. Typical local habitats are wet woods,shaded meadows, flushed heath and moorland slopes, cliffs, streamside banks, damp, shaded ditches androadsides.
British and Irish occurrence
In Britain, E. sylvaticum is common and widespread in the N & W, but decidedly rare in mostareas of C, E and SW England (New Atlas). There has also been a marked decline in the species inlowland England and Wales, which has been going on since before the first BSBI Plant Atlas (Perring& Walters 1962) and continues to the present (C.Dixon & T.D. Dimes In: Preston et al.2002).
In Ireland, Wood Horsetail is frequent and widespread in the north, but it is much more widely scatteredand only occasional in the southern half of the island (Jermy et al. 1978; An Irish Flora1996; New Atlas).
European and world occurrence
The north-western trend in the British & Irish distribution is mirrored in the horsetail’s Europeanoccurrence, covering almost the whole of N and C Europe south to N Spain, the Alps, N Greece, theBalkans and Turkey, but almost entirely absent from the Mediterranean coasts (Jalas & Suominen 1972,Map 36). The species also stretches, sometimes commonly and with very little taxonomic variation, acrossN Asia to Japan, and it completes its circumpolar range in the higher latitudes of N America, fromAlaska to Labrador and on to W Greenland, Iceland and the Faeroes (Hultén 1962, Map 86; Hultén &Fries 1986, Map 17).
Names
Despite its very distinctive and beautiful appearance, E. sylvaticum has not in the past beenrecognised by most people as anything other than simply another horsetail and for English Common namesit has really only got what we refer to as 'book names', such as 'Wood Horsetail' and 'Bottle-brush'.The latter name it shares with E. arvense and also with Hippuris vulgaris (Mare’s-tail)(Britten & Holland 1886). The Latin specific epithet 'sylvaticum' simply gives us the sameinformation as the English common name, meaning 'of woodland' (Gilbert-Carter 1964).
Threats
None.
Native, very common and widespread. Circumpolar boreo-temperate.
1881; Stewart, S.A.; Co Fermanagh.
March to December.
Growth form and preferred habitats
Marsh Horsetail is a characteristic rhizomatous perennial of wet to moist, moderately to slightlybase-rich, often calcareous or dolomitic (ie magnesium rich limestone) habitats. It tends to occur asscattered individuals or clonal patches on the margins of small streams and ditches in marshy ground,including lakeshore fen-scrub woods, damp to wet meadows that seasonally flood and hedgerow ditches. Inupland areas, Marsh Horsetail frequents calcareous flushes in acid, peaty, heather or grass dominatedmoorland. It grows beside lakeshores and pools at every altitude, including those on cut-over bogs, indisused quarries and gravel pits – always provided there is some lateral water movement and a degree ofbase-enrichment.
As its English common name suggests, Marsh Horsetail is more or less confined to these wetland habitatsand in this respect it differs somewhat from its close relative, the much more weedy E. arvense(Field Horsetail), which is frequently found growing in relatively dry ground, including arable fields –if there were any of those around Fermanagh!
Plants of E. palustre vary greatly in the amount of branching they produce (see below), but theycan readily be distinguished from the more frequently found E. arvense by the first internode ofthe side branches always being shorter than the adjacent stem sheath. Also, when the plant bears itsblack spore-producing cones, they are found on the tips of the slender green branches.
Like E. arvense, E. palustre readily colonises disturbed ground, and on marshy ground andin peatland situations it often becomes abundant after the original surface vegetation has been brokenor trampled, for instance along tracks and beside streams, or along freshly dug boggy ditches. Borg(1971) has reported similar behaviour of E. palustre on Finnish bogs and fens.
In a detailed Dutch study of 1,000 agricultural fields, Marsh Horsetail was found to be more common on29% of the fields surveyed that were regularly mown, than on those which had been under continuousgrazing (8% of the fields surveyed). E. palustre was also much more frequent on soils low inpotassium or phosphorus than when these nutrients were in good supply. In general, the study found MarshHorsetail was a rare weed of well-drained, grazed pastures that were supplied with a manure rich inphosphorus and potassium (Sonneveld 1953 (in Dutch), quoted by Holm et al. 1977). Typically,Marsh Horsetail is a minor component of moderately fertile, grazed, herb-rich wet grassland conditions.
Variation
E. palustre is extremely sensitive to environmental change, varying markedly in size and form withdegree of exposure. In the past, this variability led to numerous growth forms being named which arenowadays regarded as taxonomically insignificant. Plants in sheltered, somewhat shaded positions aremuch taller and are well furnished with whorls of upswept branches compared with the stunted and almostor entirely unbranched forms that occur in more open or windswept upland sites (Page 1988, 1997). Theseoften very small, unbranched horsetails require careful identification, as sparsely scattered shoots ofseveral horsetail species occasionally co-exist, eg E. arvense, E. palustre and E.variegatum (Variegated Horsetail).
Shade, trampling, spring frosts, or even air turbulence associated with road traffic can cause damage tothe delicate growing point of E. palustre, causing the normally erect plant to become prostrateand bushy (Holm et al. 1977).
Spore production, dispersal and sexual reproduction
Dispersal in all horsetail species, including E. palustre, is surrounded by a degree of mystery,but we have to presume that in the case of transport to occupy a freshly created vegetation gap, iteither involves wind dispersal of spores, their germination in a suitable soil and then subsequentdelicate prothallial sexual process or, alternatively, vegetative spread. The sexual production of a newgeneration of the sporophyte plant is probably the least likely, or least frequent mechanism of increaseand dispersal, since spore production and release is seasonal, the spores are short-lived and the manystages of the sex process require very specific environmental conditions. All stages of the sexualreproductive process involve high levels of risk and a failure of any of them negates the whole venture(Duckett & Duckett 1980).
Hybrids
E. palustre forms hybrids with E. arvense ( E. x rothmaleri C.N. Page) andE. fluviatile (E. x dycei C.N. Page). The former has not yet been recorded anywherein Ireland although there are 30 records widely scattered in Britain (Stace et al. 2015). E. xdycei has been found in five sites across Ireland and in 33 well scattered sites, mainly in the N& W of Britain (Stace et al. 2015).
Vegetative reproduction
Vegetative reproduction or spread, on the other hand, may prove less risky and could be achieved inseveral ways, either by the fortuitous arrival of a tuber or rhizome fragment, perhaps transported bywater, or in mud attached to animal or human traffic. Alternatively, if an already established cloneexists nearby, E. palustre may spread laterally by means of normal rhizome growth, bothhorizontal and vertical, eventually to occupy the vegetation gap at the soil surface, a form ofdispersal known as 'diffusion'. In addition, soil disturbance could fragment the hardy and resilientrhizome and attached tubers, allowing the possibility of transport to fresh suitable sites.
Underground stem or rhizome
The rhizome of Marsh Horsetail is remarkably robust and wide in diameter when compared with its slenderaerial stems and while the individual rhizome lives only a few years, it produces buds at its nodeswhich allow it to branch and spread underground and then develop fresh aerial shoots. Occasionally therhizome bears tuberous outgrowths. Individual plants in a favourable habitat are known to have spreadover 100 m in a few years (Holm et al. 1997). The plant does not however tolerate firm, packedsoils, and rhizomes have been known to travel considerable distances in soft subsoil to avoid compactedupper horizons before the photosynthetic shoots re-emerge in more porous substrates (Borg 1971).
A pioneer colonist, competitive ability poor
It appears that Marsh Horsetail only competes effectively if its optimal growing conditions are met. As aresult, it frequently behaves as a pioneer colonist, being abundant in suitable open, disturbed, muddyhabitats in the early more open stages, but it declines thereafter as other species arrive, compete andbecome dominant. However, E. palustre is usually able to survive as a scarce, often diminutivecompanion species, provided the potential dominants are restricted in their vigour by grazing, cuttingor some other destabilising environmental factor(s). Marsh Horsetail has a rather sparse canopy ofslender shoots and this, together with its comparatively poor competitive ability, means it is notnormally a dominant species in any plant community in which it occurs (Borg 1971; Grime et al.1988).
While E. palustre occurs in a very wide range of wetlands, yet in all the vegetation communitiesit frequents, it remains a minor component. It is difficult to typify its role and status beyond that ofa pioneer colonist of gaps and bare ground associated with disturbance. It may be reasoned that sincethe main part of the horsetail species grows well below ground level, it does not really belong to thesame vegetation stratum as most of the more shallow-rooted species with which it associates. Thisrevelation helps explain the wide range of vegetation types in which Marsh Horsetail occurs, albeit inmany instances having a very low presence in them, and its relationship with associated species andvegetation communities are not firmly set. This matter is further complicated, since the rhizome ofE. palustre confers exceptional persistence on the species, even in a changing environment.Possessing a vigorous rhizome allows the horsetail to survive and effectively ignore dynamic vegetationmodifications happening around and above it, so that the species may become a relict of past plantcommunities in a particular spot of ground (Borg 1971).
Plant associates and communities
Having said this, we can identify some of its more regular associates in particular habitats. Forinstance, in moderately fertile marsh grasslands, the common associates of E. palustre includeFilipendula ulmaria (Meadowsweet), Caltha palustris (Marsh Marigold), Geum rivale(Water Avens), Lycopus europaeus (Gypsywort), Angelica sylvestris (Wild Angelica) andCirsium palustre (Marsh Thistle), together with numerous species of rush and sedge. In some ofthe marshy grasslands, swampy fens, woods and heathland, E. palustre overlaps and associates withE. arvense and E. fluviatile (Water Horsetail), and especially in hedgerow ditches, it canassociate with E. sylvaticum (Wood Horsetail) and E. telmateia (Great Horsetail).
In more upland acid grassy blanket peat moorland, E. palustre frequently occurs in sloping flushesand beside small streams where the flow of base-rich spring water enhances both the nutrient supply andthe aeration of thinner layers of peaty or silty soils. Here, and around moorland lakes where there issimilar spring water enrichment, for instance in Fermanagh, around Spectacle Lough and Drumcose Lough,Marsh Horsetail occurs with species such as Potentilla palustris (Marsh Cinquefoil), Galiumuliginosum (Fen Bedstraw), Eriophorum latifolium (Broad-leaved Cottongrass), Parnassiapalustris (Grass-of-Parnassus), Schoenus nigricans (Black Bog-rush or Black Beak-sedge),Pinguicula vulgaris (Common Butterwort), Carex dioica (Dioecious Sedge), Selaginellaselaginoides (Lesser Clubmoss), Cirsium dissectum (Meadow Thistle) and, occasionally,E. variegatum (Variegated Horsetail). In a similar flush, but at lower altitude below the cliffsof Poulaphouca, close to the shore of Lower Lough Erne, E. palustre has again been found growingalong with E. variegatum.
Fermanagh occurrence
In Fermanagh, it is the third most frequent horsetail being found in 248 post-1975 tetrads, 47.0% ofthose in the VC. It is especially frequent around Upper and Lower Lough Erne and on the uplandlimestone, bog and heath in the west of the county.
British & Irish occurrence
A common and widespread deciduous species throughout Britain & Ireland, E. palustre is perhapssomewhat less common in the S and W of Ireland due to the prevalence in this part of the country ofheavily leached, base-poor, excessively acid soils (Jermy et al. 1978).
European and world occurrence
In Europe and beyond, its widespread distribution closely matches that of both E. arvense andE. fluviatile, extending from Iceland to almost all of Europe southwards to Spain, Italy, Greeceand Turkey (Jalas & Suominen 1972, Map 35). The world distribution stretches eastwards through muchof N Asia to almost encircle the arctic, although there is a distinct gap in N America in the centralprovinces of Canada (Hultén 1962, Map 89; Hultén & Fries 1986, Map 16).
Toxicity
In common with other species of Equisetum, E. palustre contains high levels of silica, plusthe thiamine-destroying enzyme thiaminase, and the toxic alkaloid(s), palustrine and/or equisetine. Whenanimals are allowed to graze selectively they avoid horsetails. However, in cut and stored hay, there islittle way they can do so, and the toxins survive drying and storage. Tests in Finland showed that aslittle as 2 gm of dried horsetail in the daily fodder of cattle caused their milk yield to decline.Larger amounts of horsetail caused lack of appetite, diarrhoea and general illness of the animals (Borg1971). Horses are not as sensitive as cattle, but if fed small amounts over long periods, they can alsosuffer serious poisoning (Holm et al. 1977).
Weed control
In spite of numerous and prolonged trials, a really effective method of controlling the growth of E.palustre has not yet been developed. The aerial shoots are easily destroyed by any of severalherbicides, but the deep, wide-ranging rhizome survives and persists, and its starch reserves simplycannot be exhausted by attacking the aerial parts of the colony. The systemic herbicide MCPA gives thebest penetration into the rhizome, and if low rates of application are used, the aerial shoots are notdestroyed as quickly as at higher rates. This allows more time for the herbicide to be translocateddownwards into the rhizome. Using this MCPA herbicide procedure in conjunction with both sub-surfacedrainage and intensive arable cultivation, rather than allowing prolonged grass leys to develop, appearsto be the best available agricultural treatment of the Marsh Horsetail problem in pastures (Borg 1971).
Names
The Latin specific epithet 'palustre' is derived from 'palus', meaning a swamp or bog, and translates as'growing in swampy places' (Gilbert-Carter 1964). E. palustre shares some of the English commonnames applied to horsetail species in general, and they are? similar to those applied to the more commonspecies E. arvense. Example include Cat-whistles, Marshweed, Paddock Pipes and Snake Pipes(Britten & Holland 1886).
Recommended further reading
The detailed review of the biology and ecology of E. palustre published in English by the Finn,Pekka J.V. Borg (1971) is highly recommended reading for anyone requiring further information andspeculation on the properties and behaviour of this species.
Threats:
Drainage and other agricultural improvements and acidification associated with forestry plantation.
Native, frequent and widespread. European southern-temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This imposing rhizomatous species always grows in colonies and the large size of the tall, sterile,annual, deciduous shoots, their abundant lateral branches and ivory-white stems make Great Horsetailboth conspicuous and completely unmistakable. E. telmateia is closely tied to permanently wetsoil, typically either poorly drained clay or shallow peat, which is more or less constantly flushedwith base-rich, generally calcareous ground water springs or seepage emerging from overlying, morepermeable sedimentary rocks (Brewis et al. 1996; Page 1997). It often grows along the edge ofroadside hedges and ditches, on eroding river banks, or at the base of wooded cliffs where it is shadedor semi-shaded by overhanging tree and shrub branches. It is associated with man and some habitats hecreates, and therefore is apophytic on clayey slopes of roadside banks, railway cuttings andembankments, provided there is the necessary degree of base-rich water seepage through the soil (Jonsellet al. 2000).
The ecological requirements and tolerances of Great Horsetail overlap quite closely with those of severalother horsetails, since it commonly associates with E. palustre (Marsh Horsetail), lessfrequently with E. fluviatile (Water Horsetail) and rarely with E. hyemale (RoughHorsetail).
E. telmateia is essentially a lowland plant, unable to tolerate upland exposure and rarely foundover 250 m, but the requirement for moving groundwater rich in calcium or other bases, together with thelaws of gravity, confine it both geographically and geologically in any case.
Plant structure and spore production
The thick, fleshy-looking, white sterile stems of Great Horsetail can reach 2 m or more in height and areheavily furnished with whorls of long lateral branches. Despite their robust appearance, the tall, stoutsterile stems are in fact very brittle, making the species intolerant of any trampling or grazingwhatsoever (Sinker et al. 1985).
As in the case of E. arvense (Field Horsetail), separate, whitish-brown, unbranched fertile stemsup to about 20 cm in height appear before the photosynthetic sterile stems in April and May. Thecone-like fertile spikes are produced most abundantly on the drier, warmer, marginal areas of thetypical habitat (Jonsell et al. 2000). The cone-bearing shoots are very ephemeral, typicallysporing, withering and dying off within two weeks of their initial production (Jermy & Camus 1991;Page 1997).
The deeply running branched rhizomes of Great Horsetail are about 1.0 cm in diameter, have large aircanals embedded in their conducting tissues, similar to those in E. palustre, and they bearnumerous starch storage tubers at their nodes (Page 1997).
Vegetative and sexual reproduction
As with all other horsetails, the main form of reproduction is probably vegetative by means of therhizome tubers and fragmentation of other parts of the plant. Since E. telmateia forms a numberof rare or very rare hybrids with other Equisetum species, some successful spore germination andprothalial sexual reproduction must also occasionally take place (Page 1997).
Hybrids
E. telmateia is capable of forming hybrids, and does so very rarely with E. fluviatile (onerecord from Ireland), E. palustre (2 hexads in Co Sligo), E. sylvaticum (Wood Horsetail)(no Irish records) and E. arvense (again, no Irish records) (Stace et al. 2015).
Fermanagh occurrence
E. telmateia is the fourth most frequent and widespread horsetail in Fermanagh, occurring in 171tetrads, 32.4% of those in the VC. Six tetrads contain pre-1976 records only. As the tetrad distributionmap shows, it is widely distributed, especially in the S & W of the county. The most extensivestands in Fermanagh are along the shore road of Lower Lough Erne, below the Cliffs of Magho (also knownas Poulaphouca). However, it occurs in quite a wide variety of habitats chiefly in limestone areas,including lakeshores, river and stream banks (especially in gullies), cliff bases, below waterfalls, inquarries and even along railway lines disused for over 50 years.
Toxins and uses
Like other horsetails, stands of E. telmateia contain large quantities of silicates and a varietyof toxic principles, including alkaloids and an enzyme called 'thiaminase' that destroys thiamine(vitamin B) (Cooper & Johnson 1998). The plants are usually avoided or ignored by grazing horses,but owing to the smoothness and fleshy softness of its stems, it is said to be acceptable to cattle asfood. The 18th century Swedish botanist Linnaeus commented that in N Sweden E. telmateia was cutand given to cows for fodder and that in Lapland reindeer would eat it, although horses alwaysrefused to do so (Grieve 1931).
There are reports of young stems of Great Horsetail being eaten like asparagus from Roman times onward,particularly by the starving poor. Unfortunately, as a vegetable they are neither palatable nor verynutritious and like other horsetails they undoubtedly contain toxins (Grieve 1931).
British and Irish occurrence
E. telmateia is frequent to locally abundant throughout most of the British Isles, although it ismore occasional in the southern half of Ireland and is rare or absent in many parts of C, E and NScotland (Jermy et al. 1978; Stace 1997; New Atlas).
European and world occurrence
Great Horsetail reaches the most northernly point of its European range at Caithness in Scotland (VC109), and the European distribution extends southwards through W, C and S Europe to N Africa, Turkey,the Caucasus and eastwards to Iran. In the Pacific States of N America, it is represented by a separatesubspecies, subsp. braunii (Milde) Hauke (Jalas & Suominen 1972, Map 39; Hultén 1958, Map258; Hultén & Fries 1986, Map 20; Jonsell et al. 2000).
Names
The Latin specific epithet, 'telmateia' means 'of marshes', or 'of muddy waters' (Gledhill 1985). The fewEnglish common names that are specific to this species include 'Fox-tailed Asparagus' and 'Horse Pipes'.The former obviously refers to the similar appearance of the spikes of the young plants to a foxtail andthe previously mentioned fact that both have been eaten by humans. This name first appeared in notesmade by Lyte when he was preparing his Newe Herball (Lyte 1578), based on his translation of theFrench edition of Dodoens' Dutch language Cruydeboeck, that was originally published in Antwerpin 1554 (Britten & Holland 1886; Anderson 1977).
Threats
None.
Native, occasional. Circumpolar temperate, but widely disjunct in Eurasia, N America and E Asia; also inisolated warmer stations further south.
1860; Smith, Rev Prof R.W.; Ardunshin.
April to September.
Growth form and preferred habitats
As this fern is small and generally occurs as thinly scattered individuals, it is very easily overlooked.It is therefore hard to be certain despite our efforts that O. vulgatum is fully recorded in thecounty. It needs to be deliberately searched for from late April to September when the annually producedaerial parts of this perennial species are visible. The texture of the sterile frond and the fact thatit has no mid-rib, together make it fairly easy to spot once you know the sort of moist, usuallycalcareous or lime-enriched loamy, clay soil it favours, and 'you've got your eye in for it'!
It does, in fact, occur in a wide variety of calcareous or mildly acidic, moist, unimproved grassland androcky habitats, including calcareous lakeshores, limestone pavement, scree and sometimes in scrubwoodland when it is invading such sites. In our experience, however, it is never really abundant, evenon or around grassy turloughs' floors (ie vanishing limestone lakes which drain through their base),which ought to suit its preferences perfectly (Webb & Scannell 1983; Page 1997).
Fermanagh occurrence
In Fermanagh, this little, deciduous, rhizomatous, taxonomically polymorphic fern has been recorded in atotal of 64 tetrads, 12.1% of those in the VC. As the distribution map indicates, Adder's-tongue iswidely distributed in Fermanagh, mainly to the SW of Lough Erne, but with a few scattered stationsfurther east. The most isolated easterly recent station is on a roadside at Knocknalosset Td, where RHNfound it in June 1990.
Vegetative Reproduction
Adder's-tongue is capable of vegetative reproduction, producing shoot buds on its spreading roots, sothat patches of the plant may develop over time. Having said this, like Botrychium lunaria(Moonwort), another fern species closely dependant on a mycorrhizal fungal partner in the soil, O.vulgatum population densities fluctuate from year to year, apparently being very sensitive toprevailing local climate (Page 1997).
Irish occurrence
The second edition of the Census Catalogue of the Flora of Ireland indicatesthat O. vulgatum has been recorded at least once in all 40 Irish VCs (Scannell& Synnott 1987). Careful manual inspection of the Botanical Society Atlas (Perring &Walters 1962, 1987) shows that in many Irish VCs this is literally the case, there being a solitaryplotted symbol. On the hectad map of Ireland, 71 of the symbols represent pre-1930 records. While themost recent Irish Flora continues to describe this easily overlooked fern as 'occasional' (Parnell &Curtis 2012), in reality Adder's-tongue is much more scarce and scattered, and in some areas it is inlong term decline. On the other hand, Irish field recording has greatly improved over the last 50 years,and the New Atlas hectad map plots 110 Irish hectads with post-1987 records, a high proportion ofthese being in the northern province of Ulster (Preston et al. 2002).
Population decline
There is evidence of a long term decline of O. vulgatum populations in lowland areas of Britain& Ireland and the trend appears particularly obvious in Ireland. The species losses are widelyattributed to drainage, agricultural intensification, a changing grazing pattern and the move from hayto silage production (C. Jermy, in: Preston et al. 2002). At the same time, O. vulgatumseems to be holding its ground in the areas of Fermanagh where agriculture is less intensive, whileundoubtedly it is now less frequent in the N & E of the VC, where drainage and disturbanceassociated with pasture improvement will probably eventually eliminate it entirely. Even in areas'improved' in this manner, however, the fern may manage to hold on in adjacent, more rocky marginalground. The decrease in the number of unimproved limestone or lime-enriched pasture sites in Fermanaghis very real, however, a rough measure being given by the fact that there are 15 tetrads where thespecies was recorded prior to 1976, but has not been re-recorded. This compares with 49 tetrads whichhave post-1975 records of the fern.
The decline of this species in Ireland appears obvious from the map in The Fern Atlas (Jermy etal. 1978, p. 28), and this has also been commented upon in other modern Irish VCFloras, notably that of Co Dublin (H21), and of the three VCs in NE Ireland (Cos Down, Antrim andLondonderry (H38, H39 & H40) (Doogue et al. 1998; Hackney et al. 1992). However, tworecently published Floras from the far south of the island indicate that in the areas covered,Adder's-tongue has either been under-recorded in the past, or has recently increased its presence: in CoWaterford (H6), Green (2008) lists the species from eight sites post-1996 and in Co Limerick (H8),Reynolds (2013) lists six sites with modern records.
British occurrence
O. vulgatum appears much more frequent and widespread in lowland England and S Wales than is thecase in Ireland. The New Atlas map, displaying plant data recorded up to the end of 1999,indicates that there has been a considerable improvement in the recording of this fern in Britain overthe last 40 years in comparison with the original Botanical Society Atlas (Perring & Walters1962). However, despite the enhanced recording effort, the presence of O. vulgatum thins andbecomes sparse and declining to the N of a line between Morecambe and Middlesbrough (Preston etal 2002; Wardlaw & Leonard 2005). Drainage and the widespread agricultural use of fertiliserand slurry-spreading in particular, have undoubtedly accelerated the previously gradual process ofAdder's-tongue population decline, a process which in Britain has been operating for more than threecenturies (Page 1997).
European and world occurrence
O. vulgatum is widespread in W & C Europe, becoming more coastal in Scandinavia and veryscattered in the Iberian Peninsula and along the N Mediterranean coast (Jalas & Suominen 1972, Map42). Elsewhere, the species (considered broadly) extends in an extremely widely-spaced, disjointedmanner across boreal Asia to Japan, Burma and N America as a very disjunct circumpolar temperatespecies. It also occurs further south in equatorial W Africa and in Mexico (Hultén 1962, Map 91; Hultén& Fries 1986, Map 21).
Folklore and uses
In earlier centuries Adder's-tongue, like Moonwort, was very much more plentiful in the British Isles,and it was better known than it is today so that it developed a folklore reputation. It was regardedboth as a herbal antidote for snakebite from Adders (another case of 'the Doctrine of Signatures' basedon a fanciful resemblance), and as having a malevolent influence on the health of pasture grasses (Step& Jackson 1945; Page 1997). An ointment called 'Green Oil of Charity' was prepared using the fernand was still in demand as a salve for wounds in the 1940s in parts of Britain (Grieve 1931; Step &Jackson 1945). The expressed juice of the leaves, drunk either alone or with distilled water ofHorsetail, was once widely used by country people for internal wounds and bruises, vomiting or bleedingat the mouth or nose (Grieve 1931). On the other hand, doubts as to the authenticity of the reports offolk use of this small, rather scarce native are expressed by Allen & Hatfield (2004), who suspectthe possibility of imports of plant material and possible 'borrowings' of reported use from olderherbals.
Names
The genus name 'Ophioglossum' is a combination of two Greek words, 'ophis' = a serpent and 'glossa' = atongue, said to be because of the appearance of the fertile branch, although in reality no snake'stongue has any likeness to it. The Latin specific epithet 'vulgatum' means common or ordinary (Step& Jackson 1945).
Additional English common names locally applied to O. vulgatum (and sometimes also to otherunrelated species), include 'Edder's-tongue', 'Serpent's-tongue', 'Dragon's-tongue', 'Adder's Spear','Adder's-grass' and 'co*ck's-comb' (Britten & Holland 1886).
Threats
Disturbance of older pastures, including drainage, agrichemical application and grazing pressure fromsheep and rabbits would all tend to depress the frequency of this small fern.
Native, very rare, probably only casual. Circumpolar boreo-temperate; also scattered in the S Hemisphere.
1901; West, W.; Knockmore Hill.
June and July.
Growth form and preferred habitats
Moonwort is a small, rather fleshy, hairless, perennial but deciduous fern that typically grows onnear-neutral to strongly basic soils. On ericaceous heaths, B. lunaria is limited to ground whichcontains at least a little lime and where the dominant subshrubs, Calluna and Erica areabsent. B. lunaria usually occurs as scattered individual shoots in short, grassy turf. It is asmall plant, the sterile frond rarely being more than 15 cm in height and often only half or one thirdof this figure, so it is very easily overlooked. The sterile fronds appear in early May, each followedby the adjacent but separate fertile frond. When the latter is eventually fully developed in mid-June,it is somewhat taller than the sterile branch and bears a pinnately divided, triangular spike ofglobular orange-brown spore-sacs, aptly likened to a miniature bunch of grapes (Page 1997). This isalluded to in the genus name, see below.
Fermanagh occurrence
The local Victorian field botanists West, Abraham and McCullagh all saw this little fern in Fermanagharound the 1900s, as did Meikle and his co-workers in the 1940s. Yet despite all the surveys of the last30 years, Moonwort has only been seen on four occasions recently, a fact that is difficult to understandas the vegetation at the plant's original sites at Belmore Mountain (recorded in 1946) and Drumkeenagh(recorded in 1902), have probably not changed much during the whole of the 20th century (RevisedTypescript Flora, Meikle et al. 1975). The map shows records occurring in seven scatteredtetrads, four of them with post-1975 dates. None of the records give information on the species degreeof presence, but we recollect that the Legacurragh discovery was a single very small plant, and thelatest late-June 2013 record at Doagh Lough was a solitary plant just 3 cm in height.
Details of the other seven Fermanagh records are: Drumkeenagh, near Black River, 1902, J.T. Abraham &F.R. McCullagh; Belmore Mountain, July 1946, MCM & D; scree slope above Doagh Lough, 1947, MCM &D; Callow, Monawilkin, 11 July 1985, ASSI Team, DOENI; Legacurragh, above Florencecourt, 1991, M.Tickner; Isle Namanfin, Lower Lough Erne, 1990-5, D. Hughes; limestone hill south of Dough Lough, 27June 2013, H. Northridge.
Irish occurrence
In Ireland, while B. lunaria has been found at least once in all but two of the 40 VCs (Scannell& Synnott 1987), it appears to occur only rarely and fleetingly in small numbers as a casual specieson dry heathy grassland, mountain ledges and in old lawns or sand-dunes (Webb & Scannell 1983; Webbet al. 1996). The New Atlas hectad map indicates how very scarce B. lunaria hasbecome in Ireland: the map displays scattered modern records in just 14 VCs (28 hectads), mainly in thenorth of the island.
It could be that unlike earlier generations, modern field botanists do not have an eye for the plant, butmore likely it really has declined or become even more transient than previously was the case. RSF isvery familiar with Moonwort from continental field trips, but he has never found an original site forthe fern in Fermanagh, which does suggest genuine rarity.
In Co Dublin (H21), B. lunaria was already a rare species at the turn of the 19th century, whereasformerly it was more common in upland, usually base-poor pastures. There are only three modern recordsin Co Dublin: one site with a solitary plant and another with only four individuals. Both the mentionedsites are in unimproved pasture, but interestingly, the third Co Dublin site is from an unusual habitat,in woodland beside a reservoir (Doogue et al. 1998).
British occurrence
In Britain, Moonwort is widely scattered, but while it is predominantly a northern and upland species, italso occurs in a wide range of open, exposed, short-grassland habitats, including at low altitudessand-dunes, golf-links, old unimproved meadows (now an exceedingly rare habitat), grassy banks anddowns. Upland habitats in Britain include grassy moors and heaths, old stabilised grassy screes, alpinemeadows and pastures, as well as on cliff ledges (Page 1997).
B. lunaria has suffered a gradual decline in Britain as well as in Ireland as evidenced, forinstance, by the map in the Atlas of Ferns (Jermy et al. 1978), where the number ofpre-1930 stations recorded almost matches those more recently recorded. The New Atlas hectad mapconfirms the continuing decline of Moonwort sites in Britain, although often being very small and eithersolitary or in small populations, it could well be overlooked and under-recorded (A.C. Jermy, in:Preston et al. 2002).
One reason why B. lunaria might fare less well in Irish conditions may be the tendency for it tooccur in conditions of high rainfall on very shallow peaty soils formed over limestone. Englishexperience, on the other hand, suggests growth of the fern really benefits from deep, well-aerated soilswhich allow adequate scope for its deeply running root system and which presumably facilitate thesuccess of the associated mycorrhizal fungi (Page 1997).
European and world occurrence
Beyond our shores B. lunaria is widespread in N, C & S Europe, extending well into the arcticregion and as far south as Sicily and the Greek Peloponnese. The distribution peters out in much of WFrance and in the southern parts of the Iberian Peninsula (Jalas & Suominen 1972, Map 44).
B. lunaria, taken in a broad taxonomic sense, extends in a circumpolar temperate manner aroundnorthern latitudes through Asia to Japan, and across northern N America from Alaska to Labrador and onto S Greenland and Iceland. A typical form of B. lunaria occurs also in southern S America alongwith closely related varieties, and it is also found in SE Australia, Tasmania and New Zealand (Hultén1962, Map 103; Hultén & Fries 1986, Map 23).
Folklore and medicinal use
Moonwort has long had a reputation for being sporadic in making its appearances and disappearances and,perhaps partly for this reason, it has gathered a considerable body of folklore around it. Magicalproperties include the ability to unlock locks and unshoe horses' feet. The former notion perhapsderived from the key-like outline of the sterile frond, the latter maybe somehow too associated with itspower over iron locks (Step & Jackson 1945; Page 1997). Other well known myths describe Moonwort'spowers of alchemy and witches collecting it by moonlight for their magic spells.
In herbal medicine, it was said to heal wounds like Ophioglossum vulgatum (Adder's-tongue) (Grieve1931). The significant weight of folklore that exists suggests that this small fern was once much moreprevalent than now. It was widely known by country people, compared to the few who might recognise ittoday (Page 1988, 1997).
Names
'Botrychium', which is a diminutive based on the Greek 'botrys', meaning 'a bunch of grapes'(Gilbert-Carter 1964). The specific epithet 'lunaria' is derived from the Latin 'luna' meaning the moon,the curved individual lobes of the sterile frond being likened to the crescent moon (Gilbert-Carter1964; see also figure in Arber 1970, p. 257).
'Moonwort' is the general widely accepted English common name of the fern, a translation of its specificepithet, but also linked to the magical folklore attached to the plant. Other common names include'Lunary' or 'Lunarie' (Turner 1568), 'Unshoe the Horse' (Culpeper 1653) and 'Shoeless Horse', anotherreminder of the folklore (Britten & Holland 1886).
Threats
Improvement of upland pastures for agriculture.
Native, occasional, declining or perhaps a casual at some sites. Sub-oceanic southern-temperate; a verywide, disjunct distribution.
1806; Scott, Prof R.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This large, robust, rhizomatous, clump-forming fern with its tall, broad, erect, bipinnately divided,distinctive light-green sterile fronds up to 3 or even 4 m tall also produces separate, smaller, fertilefronds with rusty spike-like clusters of sporangia. O. regalis is a very conspicuous andunmistakable calcifuge species of fens, bogs, lakeshores, streamsides and ditches. While Royal Fern istypically a species of wet, acid, peat conditions, when necessary it can tolerate the considerablebase-enrichment associated with salt-laden coastal winds on sea cliffs and the like (Jermy et al.1978; Page 1997).
Ecology and longevity
In spite of the large robust nature of the sterile fronds and their leathery appearance, the aerial partsshrivel and die-back at the first serious touch of frost (Step & Jackson 1945). While in nature itis relatively slow growing, Royal Fern is probably one of the longest lived native ferns in Britain& Ireland. Large individuals in garden cultivation are known to be over 100 years old, and theirthick, erect, rhizome bases display no signs of decline. Comparisons of massive wild ferns, based on thesize of such garden specimens, suggest some individuals must be many centuries old (Page 1997).
Fermanagh occurrence
The tetrad map of its Fermanagh occurrence gives an over-optimistic picture of Osmunda's frequencyin the VC. While it appears to be widely distributed in the S & W of the county and scatteredelsewhere, at many of the 70 post-1975 tetrads there are only one or two plants recorded. Furthermore,the high frequency of pre-1975 sites where the plant has not recently been seen (20 tetrads on the map),suggests that either there has been a decline of the species in the area, or it was merely casual in itsoccurrence at these sites.
British and Irish occurrence
It has a mainly, but not exclusively, western and southern distribution in Britain & Ireland,although it is more coastal in Britain and more definitely western in Ireland (Jermy et al. 1978;New Atlas).
From at least the time of the Victorian fern craze in the 1850's, plants of this fern have been collectedfor garden use, both as a decorative subject in itself, and because the fibres of its large leaf basesprovide an ideal horticultural medium for growing orchids (Allen 1969). We have no evidence of thiswholesale collecting ever happening in Fermanagh, or if it has, it is at an insignificant frequencycompared with some places in England, for instance in Westmorland where the species has been depleted bycollecting, almost to extinction (Halliday 1997).
As a result of the widespread garden cultivation of the fern, particularly in Britain, it is not a simplematter to separate native occurrences from naturalised garden escapes or discards, a point which must beborne in mind when studying the detail in distribution maps of the species (Jermy et al. 1978;New Atlas).
O. regalis has undoubtedly declined considerably in N Ireland due to habitat loss associated withdrainage and, in the E of the province, from development, for instance in bogs around Donaghadee in CoDown (Hackney et al. 1992). Apart from the Lough Neagh basin, the Fermanagh records probablyconstitute the most easterly large concentration of sites of the fern in Ireland. However, the speciesis very much associated with areas of high rainfall, and thus it is decidedly western in itsdistribution anyway.
European and world occurrence
Royal Fern being very variable and polymorphic, on the world scale it is best examined in the broad senseas an aggregate species. In Europe, O. regalis has its main area of distribution in the W and Sof the continent, from the S Swedish coast to the Azores and it ranges discontinuously across theMediterranean to Crete, Turkey and the Caucasus (Jales & Suominen 1972, Map 50; Page 1997).
On a world basis, Hultén (1958), and again in Hultén & Fries (1986 Map 29), map this polymorphicspecies along with three varieties they recognise, so that in total it stretches in a decidedly disjunctmanner from Europe and C & S Africa to the Cape, a small pocket in N India and the Himalaya, toChina, Burma and Japan, to eastern N America and parts of S America.
Names
The genus name 'Osmunda' is derived either from 'Osmund the Waterman' (an English Common name given byLyte (1578), since it is a fern of bogs and streamsides), or from the Anglo-Saxon equivalent of the name'Thor', the Scandinavian god of thunder (Grigson 1974; Gledhill 1985). The specific epithet 'regalis',Latin meaning 'kingly' or 'royal', apparently refers to the dignified and impressive appearance of theplant, and its great longevity (Gilbert-Carter 1964).
The plant has a long list of local English Common names, some of which date back to Anglo-Saxon, othersof the 16th century. One of the most interesting is 'Herb Christopher' or 'St Christopher's Herb', whichappears in Lyte (1578), Gerard (1597) and later authors. These names allude to the waterside habitat ofthe fern which the saint frequented and where, before he was converted, he exercised his self-imposedtask of carrying people across fords. A very odd name is 'Bog Onion', from Cumberland, the explanationof which entirely defeats the current writer (Britten & Holland 1886; Vickery 1985).
Uses
The root or rhizome of O. regalis has been used not only as a potting compost as mentioned above,but also medicinally. The supposed curative powers are attributed to the salts of lime and potash,amongst others, which it obtains from the bog soil and water in which the fern grows. It was prescribedby herbalists for treating jaundice in its early stages, or for removing alimentary obstructions. Anointment made from the root was also recommended for healing wounds, including bruises, dislocations andlumbago (Grieve 1931).
Threats
Mechanical peat cutting on lowland raised bogs is the main threat, in many of the sites.
Native, extinct. European boreal-montane.
1866; McDonald, J.; Altscraghy, Cuilcagh slopes.
Growth form and preferred habitats
This small, clump-forming, deciduous, polymorphic, montane fern with its distinctive, finely cut sterilefronds and separate, narrower fertile branches is very much rarer in Ireland than in Britain. In bothislands, it is strictly confined to calcium-free, silica-rich, acid soils. The species is so sensitiveto bases that it is absent from otherwise suitable geological areas, if such habitats are subject to anyenrichment with base-rich cations, for instance from salt-laden onshore winds (Jermy et al.1978).
C. crispa prefers well-drained sites on steep, but relatively stable, screes, but it also occupiescrevices on cliff ledges. In some areas it can be found on artificial habitats, namely mortar-free drystone walls (Page 1997; T.D. Dines, in: Preston et al. 2002).
Fermanagh occurrence
The solitary mid-19th century Fermanagh record of John McDonald (of whom we know absolutely nothing atpresent), from "the East side of Caulteach (Cuilcagh) Mountain, near Florencecourt;sparingly", is from a very suitable site well furnished with acidic rock cliffs and steep screes(Cybele Hibernica 1866). Regrettably, this appears to have been one of the fleeting, casualoccurrences that appear to typify the behaviour of this very strict calcifuge species in Ireland. No onehas seen the fern in Fermanagh since, despite diligent searching for it on the appropriate parts of thisvery remote mountain area.
Interestingly, another fleeting occurrence of the species was recorded in 1937 from adjacent Co Cavan(H30): "a solitary tuft of the plant found growing out of a crack in the overhanging face of asmall boulder on the W side of Bruse Hill at about 183 m altitude" (Cole 1938). It was reported asbeing killed the following year, probably by drought (Praeger 1946; Reilly 2001). On account of widergeographical distribution and biodiversity aims, this fern is included on the NIEA list of PrioritySpecies of special concern requiring conservation action.
Irish occurrence
The presence of C. crispa in Ireland has declined during the 20th century from ten VCs to justthree at present. Recent sightings are mentioned in the Irish Red Data Book, from one site in WGalway (H16), three in Down (H38) and two in Co Antrim (H39) (Curtis & McGough 1988). Plants in NIrish sites tend to be small and inconspicuous, growing in rock crevices on cliffs (Hackney etal. 1992). Flora of Connemara and the Burren noted that the fern is, "very rare inIreland, and in some of its stations little more than a casual" (Webb & Scannell 1983).
British occurrence
In Britain, C. crispa has a strongly marked northern and western distribution, and itsheadquarters very obviously occurs in the Scottish Highlands. Losses in England occurred in the SouthernPennines prior to 1930, perhaps as a result of fern collecting, but the distribution now appears stable(New Atlas).
European and world occurrence
This variable, polymorphic fern species has several named lower taxa with separate distribution areas. InEurope, it has an Arctic-alpine distribution, the main areas of occurrence being the mountains of WScandinavia, the British Isles, the Alps and the Pyrenees, with scattered stations south within theIberian Peninsula, Corsica, N Italy and the Macedonian mountains (Jalas & Suominen 1972, Map 61;Page 1997).
In more continental areas of Europe, C. crispa tends to be found in the upper subalpine tomid-alpine belt in sites with reliable blanketing winter snow-cover (Jonsell et al. 2000).Indeed, while Parsley Fern evades severe frost in this manner, the species also chooses sites that allowit to avoid high summer maximum temperatures. The present-day distribution of C. crispa inIreland and the Scottish Highlands correlates closely with the 24°C maximum summer temperature summitisotherm (ie for the highest places in the landscape). In the Scottish Lowlands, England and Wales theequivalent temperature limit is the 26°C isotherm (Conolly & Dahl 1970).
Beyond Europe, C. crispa, in the broad sense, extends to Asia Minor and there are at least fourgeographically differentiated races which carry the plant to the Himalaya, China, Alaska and N America(Hultén 1958; Hultén & Fries 1986, Map 32).
Names
The genus name, 'Cryptogramma', is derived from two Greek words, 'kruptos' = 'hidden', and 'gramme' = 'aline', an allusion to the fact that the lines of the sori are not as evident as on most other fernspecies, being covered by the rolled frond margin (Step & Jackson 1945; Jonsell et al. 2000).The specific epithet 'crispa', meaning 'curly', or 'with wavy margins', refers to the Parsley-likeappearance of the deeply cut sterile fronds (Gilbert-Carter 1964).
The English Common name most commonly used is 'Parsley Fern', which Grigson (1974) reckons dates from the18th century, the plant resembling curled forms of parsley. Other Common names less often heard include'Rock Brakes', and 'Curled Brakes'; 'Brakes', 'Brake' or 'Brake-fern' are general names for the largerferns dating back to Turner, Lyte and Gerard in the mid-16th century (Britten & Holland 1886).'Stone Fir' and 'Mountain Parsley' are two 19th century 'book names' (ie, a derogatory term for plantnames invented by other authors), also listed by Britten & Holland (1886).
Threats
None.
Native and naturalised garden introduction, very rare. Mediterranean-Atlantic.
1939; Praeger, R.Ll.; on the walls of Crom Castle.
May to August.
Growth form and preferred habitats
This is a small to moderate sized, delicate-looking, rhizomatous fern with fronds divided into numeroussmall, fan-shaped segments borne on thin, blackish, wire-like branches. It is frost-tender and a rare orvery rare species mainly of damp, mild, limestone coastal areas of W Ireland and SW Britain. In verysheltered sites fronds are sometimes semi-evergreen, surviving overwinter. Native populations arethreatened by unthinking collectors.
Apart from its scattered natural sites, Maidenhair Fern is frequently cultivated (mainly indoors), andcan sometimes be found 'escaped' on sheltered, damp, lime-rich mortar on garden walls near oldgreenhouses.
Variation
Plants are rather variable in size, degree of frond dissection and margin serration, so that extremeforms have been brought into cultivation and have become named as horticultural cultivars (Page 1997).
Fermanagh occurrence
Praeger's original Fermanagh station was given as, "On outer side of wall opposite main door of CromCastle" (Praeger 1939). It continues to thrive on the boat house at Crom, approximately 100 m fromthe old castle, and it is also abundant on the garden wall of Florencecourt against which a greenhouseonce stood. These naturalised plants are the only recognised records of this fern in N Ireland, several19th century finds in coastal stations in Down (H38) and Co Antrim (H39) having been discarded as eithererrors or unconfirmed and, in any event, now extinct (Hackney et al. 1992).
European and world occurrence
Maidenhair Fern is essentially a Mediterranean Basin and S Atlantic species, although Hultén (1962, Map139) maps it as circumpolar in warm-temperate latitudes of the northern hemisphere. It is alsowidespread around the southern hemisphere, including appearances in some very remote island groups.
Irish occurrence
A. capillus-veneris occurs as a native species in Ireland only in scattered stations along the Wcoast, most abundantly in the Burren, Co Clare (where it is especially luxuriant and impressive onInishmore, Aran Islands) (H9) and in SW Donegal (H35) (Jermy et al. 1978; Scannell & Synnott1987). The latter, since the Antrim stations are discarded, must now be regarded as the most northerlysite of the species anywhere in the world (Hultén 1962; Jalas & Suominen 1972). The New Atlasmap displays nine Irish hectads scattered across the island where this fern has been recorded as anintroduction.
British occurrence
In Britain, A. capillus-veneris is native in scattered sites from the Channel Isles and the SWcoast, up the W coast from Cornwall, through S Wales to Cumbria and the Isle of Man (Jermy et al.1978; Page 1997; New Atlas). This rather delicate-looking fern also crops up in Britain from timeto time as established escapes from cultivation at inland sites in lime-rich mortar or on limestonewalls, exactly as it does in Fermanagh (Jermy & Camus 1991).
Names
The genus name 'Adiantum' is a Greek plant name used by ancient botanical writers, derived from'adiantos' meaning 'dry' or 'unwetted', an allusion to the non-wettable character of the foliage, afeature known to Pliny (Step & Jackson 1945; Gilbert-Carter 1964). The Latin specific epithet'capillus-veneris', means 'Hair of Venus', and hence the English Common name 'Maidenhair Fern'.'Capillus veneris' was the medieval Latin name used by apothecaries for this medicinal fern, a namefirst found in the Herbarius of the fourth century AD ascribed to Apuleius, and it refers to wirypubic hair which are likened to the blackish stalks of the fern fronds (Grigson 1974).
Additional local common names include 'Capillaire', 'Lady's Hair' and 'Dudder-grass', the latter astrange Norfolk usage, apparently making comparison with Briza media (Quaking-grass orDodder-grass), because of the trembling motion of the frond segments resembling the movement of thegrass spikelets (Britten & Holland 1886).
Uses
Medicinal use dates back to Dioscorides, as a remedy in pectoral complaints and pulmonary catarrhs(Grieve 1931).
Threats
The species has died out at a number of previous stations along the coasts of Ireland, England and Wales.Either of the two Fermanagh sites could easily be destroyed by re-plastering of stonework or otherexcessive or unknowing 'tidying' operations.
Native, scarce or occasional, but locally abundant. Oceanic temperate; widespread and extremely disjunct– a preglacial relict species.
1860; Smith, Rev R.W.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Tunbridge Filmy-fern tends to be the only small, moss-like, mat-forming fern species growing on verticalor near-vertical, permanently and deeply shaded rock faces on mountain slopes and in woodlands, on andunder rocks in deep, damp shade.
Identification
The long, flat, overlapping bluish-green fronds easily distinguish H. tunbrigense from thesomewhat more common, blackish fronds of H. wilsonii (Wilson's Filmy-fern). Confirmation is oftenprovided by the irregularly toothed margin of the pocket-like indusium covering the spore-sacs, afeature visible with a good hand-lens. The indusia are not always present however when these species arefound growing in conditions of very moist heavy shade.
Fermanagh occurrence
In Fermanagh, H. tunbrigense has been recorded in a total of 24 tetrads, 23 of which havepost-1975 records. The main areas where it occurs are on Cuilcagh mountain and the sandstone scarps andwooded glens of SW Fermanagh. The isolated 1950 station where it has not been refound was at AnnaghmoreGlebe Lough.
At some Fermanagh sites, H. tunbrigense colonies can cover several square metres of rock, andsometimes the fern is sufficiently profuse to completely fill rock crevices. Locally, TunbridgeFilmy-fern is almost always found growing on acidic rocks, most frequently on sandstone, butoccasionally it may also occur on fairly hard, basic igneous and metamorphic rock types, particularlythose providing crevices and with a texture, location and position that enables them to retain moisturefor prolonged (or relatively long) periods (Richards & Evans 1972).
H. tunbrigense occurs in close physical proximity with its near relative H. wilsonii rightup on the summit ridge of the highest mountain, Cuilcagh (c 600 m). At this site and others on theN-facing slopes of Cuilcagh, on sandstone scarps on the Western Plateau (ie in and around the LoughNavar Forest Park in particular), and in oak and mixed deciduous woodlands nearby (eg the Correl GlenNR), H. tunbrigense is always confined to very sheltered conditions. Typically it grows in deep,shaded hollows under very large overhanging boulders, cliffs or trees, growing on bare rock surfaces orin crevices. It is also found much less frequently on the peaty or uncompacted humus soils of woodlandfloors. Only rarely does it occur as an epiphyte on the bases of oak or old ash trees in very dampwoodland, eg in the Correl Glen, and at the base of the heavily wooded Cliffs of Magho.
Comparison with H. wilsonii
By comparison, H. wilsonii occupies less shaded, somewhat more open and exposed conditions and itis often intermingled and embedded in cushions of moss and leafy liverwort. H. wilsonii does notform large single-species mats to quite the same extent as H. tunbrigense does.
On Cuilcagh, both these species also occur under and around huge, house-sized, rocks on block screes onthe northern slopes just below the long, whale-back summit ridge, and again together on further massiverock falls at Cuilcagh Gap, and in similar situations around Lough Atona lower down these same slopes(at c 500 m). There are no trees on any of these heathy moorland slopes, which at this altitude andexposure are dominated by a canopy of ericaceous subshrubs and upland grassland.
Comparative tolerance of desiccation
The normal belief is that of these two species of perfectly frost-hardy filmy-fern, H. wilsonii isbetter able to tolerate high altitude exposure and the associated risk of desiccation than can H.tunbrigense (Jermy & Camus 1991; Page 1997). In mountain environments in the W of Ireland,rainfall is so very plentiful, frequent and regularly distributed, that droughting of the delicatefronds is not anything like as great a risk as might at first appear. At various sites in Fermanaghdrought affected Hymenophyllum plants are occasionally found, particularly affecting the usuallymore exposed individuals of H. wilsonii, but occasionally also those of H. tunbrigense.The plants may be discovered looking very shrivelled, brown and desiccated, sometimes apparently dead.However, the rhizome and even the fronds have greater powers of recovery than their appearance andstructure might suggest, and they can recover surprisingly well from temporary desiccation, or even fromthe effects of a light heathland fire (Richards & Evans 1972).
Comparative experimental studies by the latter authors have shown that H. tunbrigense suffers theeffects of desiccation more immediately than H. wilsonii, but it also recovers more quickly thanthe latter. However, of the two, H. wilsonii has greater drought resistance, its protoplasmcoping better with desiccation and its cells avoiding severe diurnal mechanical stress during droughtperiods.
A most interesting and rather unexpected finding is that the fronds of both Hymenophyllum speciespossess the capacity for indeterminate apical growth. This allows individual fronds to survive andcontinue growing for several years (perhaps four, five, or even more seasons), and thus both speciesmanage to produce sporangia and new indusia in waves, maybe twice a year under favourable growingconditions (Richards & Evans 1972).
Growing as it does in more sheltered and more deeply shaded conditions, it is not really surprising thatH. tunbrigense consistently has a lower photosynthetic compensation point than H.wilsonii. On Cuilcagh, the two Hymenophyllum species have found adjacent but distincthabitats, and although their microclimates definitely overlap, obviously they are both able tolerate theprevailing environmental conditions and have found ways of avoiding direct competition.
In the past, doubt has been cast on the finding of H. tunbrigense at altitudes above 460 m inBritain and Ireland (Richards & Evans 1972; Page 1997). However, the local botanists who identifiedthis fern on the ridge of our highest mountain, Cuilcagh, are very familiar with both species offilmy-fern, and I am confident that the identifications are correct. Having emphasised this, H.tunbrigense is recorded only half as frequently in Fermanagh as H. wilsonii, so that ofthe two, Tunbridge Filmy-fern still clearly has the narrower ecological range (Richards & Evans1972). While both filmy-fern species grow extremely slowly, c 2.5 cm/yr, H. tunbrigense appearsto possess less biological vigour and suffers more from desiccation than H. wilsonii does.
Intolerance of liquid water wetting
One of the interesting facts to emerge from the study of Evans (1964) is that althoughHymenophyllum species demand humid conditions, they absolutely do not tolerate being directlywetted with liquid water, for instance, by splashes from streams or waterfalls, which often are thesource of the atmospheric humidity they do require. In other words, a rapid change of water content ismuch more harmful to a filmy-fern than a gradual one (Richards & Evans 1972). Evan's finding appearsto directly contradict a statement regarding the habitat of H. tunbrigense made by Page (1997, p.248), where the latter suggests the fern thrives in the splash zone of cool, permanently tumblingstreams. While this is often quoted as one of the preferred habitats of another very much rarer speciesof filmy-fern, Trichomanes speciosum (Killarney Fern), RSF does not believe he has ever observedHymenophyllum growing right in the spray zone anywhere in Ireland.
British and Irish occurrence
H. tunbrigense is restricted to a very discontinuous occurrence in the N, W and S of both Irelandand Britain, extending from Cornwall to Skye. In Britain, apart from an outlying group of sites in EastSussex (where it has markedly declined in recent years) and a couple of very scattered sites in NEYorkshire, S Northumberland and Cheviot (VCs 62, 67 & 68), it is otherwise completely absent fromthe east of the island (Preston et al. 2002).
European and world occurrence
The distribution of H. tunbrigense in continental European is very sparse and disjunct, even incomparison with its representation in the British Isles. It is known only from a few stations each inFrance, Luxembourg, N Spain, Italy and the eastern coast of the Black Sea. Beyond this, it does alsooccur on the Azores, Madeira and the Canaries (Jalas & Suominen 1972, Map 69; Richards & Evans1972).
Fossil history
Fossil spores of H. tunbrigense have been recorded from the Hoxnian interglacial in Ireland byWatts (1959). This fact, taken together with the species' present-day widely disjunct and sporadicEuropean distribution, undoubtedly confirms it is a relict species. Previously the species had a largerand much more continuous range (probably most recently around the current post-glacial climaticoptimum), but it has declined and continues to do so (Richards & Evans 1972). Gradual climaticdeterioration, compounded in recent centuries with habitat destruction by man, has resulted in thefragmented distribution of H. tunbrigense we observe in Europe today.
Name
The specific epithet 'tunbrigense' is a Latinised reference to the first known British site, found byDaire a few years previous to Ray's (1686) published report, at High Rocks, Tunbridge Wells, Sussex (fora full history see Evans & Jermy (1962)).
Threats
Some Fermanagh sites were for a time threatened by being overgrown by coniferous forest plantations, butthe threat has eased since the trees have been felled and they are not being re-planted.
Native, occasional. Oceanic boreo-temperate.
1844; Cole, Hon J.L.; Trien Mountain, above Florencecourt.
Throughout the year.
Growth form and preferred habitat
This moss-like, rhizomatous, cushion-forming perennial fern grows in similar habitats to H.tunbrigense (Tunbridge Filmy-fern) on ± vertically erect, damp, shaded, mountain rock crevicesand ledges, and on mossy rocks and tree boles in damp woodlands, but while usually in shade it alwaysoccupies slightly more open situations than the latter.
Fermanagh occurrence
In Fermanagh, H. wilsonii has been recorded in 34 tetrads (6.4%), 31 of which have post-1975records. As the tetrad map indicates, it is quite widely distributed on the Western Plateau, CuilcaghMountain and Florencecourt. The most isolated recent station is on a scarp at Drumskinny Td in the farnorth of the county, where RHN found it in May 1990.
Comparison with H. tunbrigense
The fronds of H. wilsonii are generally longer, narrower and more upright than those of H.tunbrigense and in overall appearance they are less flattened and of a deeper shade ofolive-green, sometimes rather blackish, or indeed brown if it has recently suffered drought. This littlefern is often found growing amongst mosses where its distinctive colour makes it quite easy to spot.
Wilson's Filmy-fern generally forms rather smaller patches than Tunbridge Filmy-fern and sometimes whenit is growing through cushions of moss, the fronds are few and quite distant from one another, so itcould easily be overlooked or mistaken for one of the larger species of leafy liverwort. The frond isstiffer and of thicker texture than that of H. tunbrigense and, because it is more droughtresistant and can thus thrive better in illuminated sites than the latter, it sometimes competes withlichens as well as with mosses and liverworts.
Although the two filmy-ferns occur in very similar habitats and their micro-environments overlap, in anytype of habitat where they occur together they typically occupy different vegetation zones. Forinstance, H. wilsonii occurs higher on the bark of trees and on block screes on mountainsides itoccurs in more exposed, better lit sites. Like H. tunbrigense, H. wilsonii is usuallyabsent from the spray zone of waterfalls, but while it normally avoids running or dripping water, it isslightly more tolerant of being wetted in this way and occasionally it is found on rocks which areflooded from time to time (Richards & Evans 1972).
Of the two species, H. wilsonii is better able to withstand relatively prolonged drought.Sometimes, however, drought can result in colonies peeling away from rock or soil surfaces, which maylead either to their destruction (Richards & Evans 1972) or, very occasionally in the W of Ireland,this may serve as a method of vegetative dispersal.
Very little is known about the life-history of Hymenophyllum species, save that they are veryslow-growing and that the individual fronds may survive up to about five years. The frequent occurrenceof large colonies of Hymenophyllum suggests that they may be long-lived, perhaps surviving instable environments for centuries. This is clearly an area which would repay further field study.
Irish occurrence
The distribution of Wilson's Filmy-fern overlaps that of Tunbridge Filmy-fern in the Atlantic-influencedN and W of both Britain and Ireland, but it is quite a lot more widespread – recorded in more than twiceas many hectads as H. tunbrigense. In Ireland, H. wilsonii is locally abundant in all themain mountain ranges (which with the exception of the Galtees in Tipperary, all happen to be quitecoastal), and in addition it is thinly scattered in damp woods throughout the island (Richards &Evans 1972; Jermy et al. 1978).
Survey work for Atlas 2000 in the north of Ireland has proven that H. wilsonii is much morewidespread than had been previously considered (NI Flora Web site 2001; New Atlas).
British occurrence
In Britain, the species is again locally abundant in the N & W of the island, but unlike H.tunbrigense it has no south-eastern outlier. Instead, it is confined west of a line from StartPoint in Devon to the mouth of the River Tees. Within this area of Great Britain, it does extend furthereast and much further north than H. tunbrigense does, extending to Shetland (Richards & Evans1972).
World occurrrence
H. wilsonii has a much more confined world distribution than H. tunbrigense, being largelyrestricted to W Europe and N Atlantic islands (Hultén & Fries 1986). It is known only from theFaeroes, one station in Iceland plus the more southern Atlantic islands (Azores, Madeira and theCanaries), which are not really part of Europe but where species populations are generally conspecificwith European flora. On the European mainland, the distribution of H. wilsonii runs from a broadcoastal band of SW Norway plus just four coastal sites in N France (Jalas & Suominen 1972, Map 70;Page 1997; Jonsell et al. 2000).
Names
The genus name 'Hymenopyllum' is a compound of two Greek words, 'umen' meaning 'a membrane or thin skin'and 'phyllon' meaning 'a leaf', a reference of course to the membranous fronds which are a single cellthick (Step & Jackson 1945). The Latinised specific epithet 'wilsonii' commemorates the bryologist,W. Wilson, who in 1830 drew the differences between the two filmy-ferns to the attention of the elderHooker (Professor, and later Sir William). Hooker named the species but subsequently reunited the twospecies under H. tunbrigense once more (Richards & Evans 1972). For a full history, see Evans& Jermy (1962).
Threats
Some sites could be threatened by forestry operations.
Native, sporophyte extremely rare; gametophyte occasional. Oceanic temperate.
1900; Tetley, W.N.; near Carrick.
Throughout the year.
Fermanagh occurrence
The discovery of this very distinctive fern in Fermanagh was announced in the Irish Naturalist ofDecember 1900 when R.Ll. Praeger wrote, "The investigations of Messrs. West and Tetley have addedthe Killarney Fern to the Flora of Co Fermanagh. They sent me a specimen last August, describing theexact locality – a deep crevice in limestone rocks; but I think it better not to publish the station, soterribly has this lovely plant suffered from the depredations of unscrupulous collectors." Thefollowing year Praeger included mention of the station in his monumental Irish TopographicalBotany simply as, "Near Carrick". The site was more clearly identified as the CorrelGlen in the unpublished Typescript Flora of Co Fermanagh of 1951. The rocks in the areaaround the Correl Glen are limestone and calcified or dolomitised sandstone. The glen supports old,moist, mixed deciduous woodland, growing on peaty soils at an altitude of around 100 m.
Growth form and preferred habitat
Killarney Fern is closely related to the genus Hymenophyllum and is another type of filmy-fernhaving rather larger, dark-green, membranous, translucent fronds, 7-60 cm in length and much divided(Ratcliffe et al. 1993). Unlike both of the Hymenophyllum species, however, T.speciosum typically resides near waterfalls or mountain streams in sheltered, deep, shady rockygorges, growing in rock crevices between boulders, in dripping caves or under overhangs where spray orseepage water keeps the plant permanently moist and dripping (Page 1997). Almost all of its British& Irish stations past and present have been in mild oceanic districts in the far west and often atlow altitudes. Evidently it is a warmth-demanding species requiring a great deal of shelter, andprobably it is rather frost sensitive, a feature which contrasts strongly with the much more hardyHymenophyllum species (Wigginton 1999).
Gametophyte generation
The prostrate, sexual, gamete-bearing gametophyte generation of Trichomanes was first recognisedby Mettenius in cultivation as long ago as 1864. Uniquely amongst European ferns, the gametophytereproduces vegetatively in the absence of the asexual spore-bearing generation, the ± upright, or elsedangling, frond-producing sporophyte. The independent gametophyte populations were completely overlookedin Britain until 1989 (Rumsey et al. 1992).
In February 1993, Nick Stewart visited Fermanagh with an international group of bryologists. Whenexamining mosses and liverworts in the Correl Glen and on the overhanging scarps of Lough Navar ForestPark, he twice found the gametophyte of the Killarney Fern. RHN was on the outing and learnt how toidentify this green, thread-like sexual stage of the fern life-cycle. Since 1993 he and his wife Hannahhave found it in twelve tetrads spread across three hexads. Many of these gametophyte finds have beenverified by staff at the Natural History Museum, London. The southerly, most isolated site shown on themap is in a deep, dark hole in a scarp at Aghnahoo, on the slopes of Cuilcagh mountain.
The gametophyte grows in Fermanagh under deep, dark rock overhangs, usually about an arm's length removedfrom the light. It is the last form of plant life to grow before light levels become totally inadequate.The gametophyte forms patches which vary in size from several square centimetres to almost the size ofone's hand in exceptional cases. It is light green, looks like a filamentous alga, and has a spongytexture when pressed down. When viewed under a hand lens it looks like wire wool with small spikessticking out from it like the thorns on certain rose bushes. The inability of all these gametophyteplants around the British Isles to carry out successful sexual reproduction and develop new maturesporophyte frond-bearing plants, means that the high conservation status of T. speciosum, rankingit as vulnerable, must remain in force. Along with other oceanic species, Killarney Fern is consideredby some conservation biologists to be at further risk from the predicted effects of global warming(Plantlife 1991).
Further detailed study of the exact growth requirements of T. speciosum is urgently needed, and aninvestigation as to why the gametophyte fails to produce new sporophyte plants would also assistconservation and rehabilitation efforts associated with increasing biodiversity awareness.
Robert Northridge rediscovers the sporophyte
On 1 May 2005, while searching for the gametophyte under a large rock at a location within the H05hectad, RHN discovered a solitary plant of the sporophyte. It consisted of two fronds each 8 cm long,one frond 6 cm long, another frond 6 cm long but with the top half dead, plus the broken stipe only ofanother frond. There was also a small patch of the gametophyte about 15 cm × 10 cm close by. When thesporophyte site was visited a year later on 26 May 2006, as well as the previously seen fronds, therewas a new frond unfurling which was just over 5 cm long.
This site was in a very sheltered, cave-like area under large fallen rocks through which there is noperceptible flow of air that might dry and damage the wet filmy fronds. The site is under trees and theentrance to the 'cave' is overhung by dense straggling leaves of Luzula sylvatica (GreatWood-rush). Exact details of the location of this site have been lodged with the Northern IrelandEnvironment Agency (NIEA) in Belfast.
The rediscovery of the Fermanagh Trichomanes sporophyte has been the recording highlight of thiswhole VC Flora project.
Gametophyte Irish occurrence
The Trichomanes gametophyte has been recognised in numerous Irish sites and by 1998 was known from13 VCs. In Britain, the same published report listed the gametophyte from a total of 38 VCs and provideda map for both islands (Rumsey et al. 1998, Figure 4). Since then isolated finds of thegametophyte have been made in Cos Tyrone and Antrim (H36 and H39) and we are confident that thisgeneration of the species remains under-recorded.
Previous Irish occurrence of the sporophyte
In the early 19th century, the sporophyte of T. speciosum was found to be frequent and widespreadin and around Killarney and SW Ireland in general (Jermy et al. 1978; Wigginton 1999). On accountof the beauty and comparative rarity of the fern, T. speciosum became the supreme target ofcollectors and gardeners during the long-running Victorian Fern Craze (1830-1920) (Allen 1969; Page1988).
The Victorian Fern Craze in SW Ireland
Kerry and other parts of the west of Ireland attracted many visitors on account of its wealth of fernspecies, and since T. speciosum appeared in such abundance in the region, it probably seemedharmless to remove a little of it in order to grow the lovely fern in one's front room in a glazedWardian Case or in a Conservatory, or to press and dry it for the family herbarium collection. Visitorsfound the fern easily, and there was no difficulty in pulling it down from its wet rock under-hang orcave roof habitat. In fact, the long strands of bristly, black, rhizome came away far too readily, andit is very likely that visitors accidentally removed far more of the plant than they ever wanted totransport. At one stage, tinkers collected Killarney Fern and hawked boxes of it around the Killarneyhotels, selling it to visitors, until eventually the extremely slow-growing plant was reduced to extremerarity and local extinction (Marren 1999).
A lingering collective cultural guilt is the reason why naturalists are now so extremely circ*mspectabout the remaining stations of the Killarney Fern in the British Isles, and it is now recognised as aRed Data Book species, protected by conservation laws in both Britain and Ireland.
Irish occurrence
In Ireland, over the years, the sporophyte T. speciosum has been recorded from a total of 43scattered sites. The Irish Red Data Book (1988) reported that it had been recently found in justten of these sites. A subsequent study of T. speciosum habitats in the British Isles made byRatcliffe and his associates found the sporophyte was still quite widespread in the hill country of CosKerry and Cork, where they located a total of 26 'colonies' (ie individual patches, each representing asingle rhizome system). These workers reckoned they had searched only a small part of the total possibleground in SW Ireland that might support the sporophyte generation. Nevertheless, they concluded thatT. speciosum is a much reduced species, and that some of the more outlying Irish colonies werealso collected out of existence besides the ones previously known about in Cos Kerry and Cork (Ratcliffeet al. 1993).
Similar depredations befell the much more thinly scattered populations in England, Scotland and Wales,where sporophyte T. speciosum is now confined to very few localities. Ratcliffe and hisassociates located a total of just 13 of their individual 'colonies' in Great Britain, details of whichare kept so secret that their map references do not even appear on the confidential computer records atthe Biological Records Centre at Monkswood (Ratcliffe et al. 1993; Marren 1999).
Field observations suggest that under existing environmental conditions Killarney Fern has been incapableof producing any new sporophyte populations for over a hundred years. Again, like other filmy-ferns,growth of the fern is very slow. Each frond can continue growing for five or more seasons, andindividual colonies that have been observed and measured over many years, do not appear to change inappearance or show any significant growth (Ratcliffe et al. 1993).
World occurrence
Reflecting the climatic limitations discussed above, T. speciosum is confined to a very limitedmild, damp oceanic area of Europe and the Atlantic isles (Azores, Madeira and the Canaries). Besides theBritish Isles, it occurs only in Brittany, the Pyrenees, near Gibraltar and near the west coast of NItaly in the Alpi Apuane at between 180-250 m (Jalas & Suominen 1972, Map 71; Ratcliffe etal. 1993; Pignatti 1997, vol. 1, p. 53; Rumsey et al. 1998).
Uses
There does not appear to be any folklore about the fern, nor is it credited with any medicinalproperties. It has, of course, been valued for its showy beauty and rarity and has been cultivated bykeen gardeners since its discovery. Trichomanes is a cosmopolitan genus and contains a total of25 species, most of them tropical, and the majority in horticulture originate from the southernhemisphere (Griffiths 1994).
Names
The genus name 'Trichomanes' was given to an unknown fern species by Theophrastus (possibly Aspleniumtrichomanes, Maidenhair Spleenwort). Gilbert-Carter (1964) regards its origin as obscure, but itis definitely a Greek compound involving 'thrix, thichos', meaning 'a hair', 'a bristle', or 'hairy'(Stearn 1992). One idea for the derivation of the other half of the genus name is the Greek 'manos',meaning 'flexible', but there is some doubt as to the relevance of this suggestion, and the correctderivation therefore remains a mystery (Step & Jackson 1945).
The specific epithet 'speciosus' is Latin meaning 'showy' or 'handsome'.
Threats
All the currently known sites are within ASSIs. Further study of the ecology and biology of the speciesis urgently required in order to properly manage its conservation.
Native, common, widespread and locally abundant. European boreo-temperate.
1860; Smith, Rev Prof R.W.; Brookeborough.
Throughout the year.
Taxonomy
What was for many years treated as one Polypodium species in B & I has been recognised, since1960, as a polyploid complex of two rhizomatous perennial species (tetraploid Polypodium vulgareand diploid P. cambricum (Southern Polypody)) and the fertile allopolyploid hybrid between them(hexaploid P. interjectum (Intermediate Polypody)) (Shivas 1962; Jermy & Camus 1991). Someforms of Polypody require microscopic investigation to identify with certainty and, therefore, fieldrecords are still very frequently made at species aggregate level (ie as P. vulgare agg. ors.l.), and the constituent species and hybrids seem destined to remain under-recorded.
Growth form and preferred habitats
All forms of Polypodium, like Bracken, do not develop a crown, but instead possess a scaly,creeping rhizome, which in this case runs along the surface of the ground rather than being buried inthe soil. From the rhizome, leathery, evergreen, hairless, aerial fronds resistant to both frost anddrought arise at intervals, generally only a few centimetres apart. Fronds vary enormously in lengthfrom just a few centimetres in stunted forms growing in dry or exposed situations as in rock crevices oron walls, but they can develop up to 45 cm in length in the case of hybrids which display 'hybridvigour' or heterosis.
Representatives of the Polypodium vulgare agg. typically grow amongst mosses and leaf-mould onsemi-shaded rock outcrops, under hedges, on walls, on roots and stumps of trees, or on the trunks andthicker branches of mature trees in damp woods.
Reproduction
The sporing sori are formed in pairs on the underside of the herringbone-like frond either side of thestalk (technically called a 'rhachis') in the upper one to two thirds of its length. The sori are large,rounded, naked of any covering indusium tissue and are a conspicuous, golden-orange colour when fullydeveloped, anytime between June to September. The sporangia are so securely attached that indentationsappear on the upper side of the frond opposite the sori positions beneath.
Fermanagh occurrence
In Fermanagh, P. vulgare s.l. is very common and widespread, having been recorded in 401 tetrads,76% of those in the VC. The species aggregate typically grows amongst mosses and leaf-mould onsemi-shaded rock outcrops, under hedges, on walls, on roots and stumps of trees, or on the trunks andthicker branches of mature trees in damp woods.
British and Irish occurrence
The New Atlas hectad map shows P. vulgare s.l. occurring throughout the vast majority of B& I, but absent from the Channel Isles and less prevalent in the English Midlands and up the E coastfrom the Wash to Newcastle-upon-Tyne. The New Atlas map also indicates areas of lesscommon occurrence or absence over parts of the Irish Midland plain, and also in the more exposed, wet,nutrient-starved, boggy ground of Cos Clare (H9), Roscommon (H25), W Mayo (H27) and W Donegal (H35).
World occurrence
On a European basis, the aggregate species occurs widely and commonly throughout continental Europeextending from Gibraltar and Crete in the south, to the northern tip of Norway, Iceland and the southerntip of Greenland, but absent from Jan Mayen and the Arctic Islands (Svalbard, etc.) (Jalas &Suominen 1972). Eastwards, P. vulgare agg. occurs in the Caucasus, Urals, the Himalaya, and, inan even broader definition (P. vulgare sens. lat.), it becomes circumpolar and stretches aroundmiddle latitudes of E Asia and across northern and central regions of N America (Hultén 1962; Hultén& Fries (1986, Map 74).
Medicinal uses
The rhizome of Polypodium had several uses in herbal medicine and was often referred to asPolypody of the Oak, or Oak fern (not our modern, much more delicate Gymnocarpium dryopteris).There was a belief that ferns and flowering plants that grew on oak roots or branches were especiallyefficacious as remedies. Its principal use was as a mild laxative, but it was also considered useful forcoughs and catarrh. An infusion made from the crushed rhizome was drunk like tea as a treatment for theearly stages of consumption or for rheumatism. Polypody was used to treat jaundice, dropsy, scurvy andother skin complaints, and dried powdered rhizome used alone or mixed with honey was also said to removenasal polyps (Grieve 1931; Vickery 1985).
Names
The genus name 'Polypodium' is Latin but is derived from two Greek words 'polus' meaning 'many' and'pous' meaning 'a foot', the notion being either that the plant having a branched rhizome has many feet(Grigson 1974), or more likely, that the comb-like, pectinate frond resembles a centipede (Prior 1879).
In addition to the English common names mentioned above, this easily recognised and well-known fernaggregate has been given at least nine other names including: 'Brake of the Wall', 'Adder's-fern','Everferne', 'Wall Fern', 'Wood Fern', 'Golden Locks', 'Golden Maiden-hair', 'Golden Polypody' and 'MossFern'. The latter, a name proposed by Gerard (1597), is rather apt (Britten & Holland 1886).
Threats
None.
Native, frequent but under-recorded. European boreo-temperate.
1858; Brenan, Rev S.A.; Ardunshin.
Throughout the year.
Growth form and identification
Some of the earlier Fermanagh records of this rhizomatous species were determined from herbarium vouchersin BEL by Paul Hackney. Typically P. vulgare s.s. has narrow, rather leathery, evergreen,parallel-sided fronds with the lowest pair of pinnae not bent forward (ie not inflexed) likethose of P. interjectum (Intermediate Polypody) (the allohexaploid derivative of its hybrid withP. cambricum (Southern Polypody)).
New fronds of P. vulgare s.s. are produced in early summer and the species, which is a tetraploid,has numerous pairs of naked sori in the upper portion of the frond which ripen their spores inmid-summer (Jermy & Camus 1991).
Preferred habitats
In contrast to the two other species of the genus in our survey area, P. vulgare s.s. is adefinite calcifuge. It typically grows on cliff ledges and in rock crevices on steep, peaty banks, inbetween the rocks in old, dry-stone walls or along the tops of such walls, and it also occurs as anepiphyte on the bark of mature deciduous trees in damp woods.
A study of the preferences of a number of ferns with respect to soil reaction carried out in W Europe byKoedam et al. (1992) found that soil taken from the root mass of P. vulgare s.s. had amedian pH of 4.13, and consequently the plant was regarded by these workers as 'acidiphilous' (ie acidloving, or acid tolerant). Such species are adapted to soils with high levels of exchangeable aluminiumand hydrogen, and relatively low levels of exchangeable calcium.
British and Irish occurrence
P. vulgare s.s. is the most common form of Polypody recorded in both Britain & Ireland, andin N Ireland it is recorded in almost every hectad.
Fermanagh occurrence
In Fermanagh, this is the most common and widespread species of polypody, having been recorded sofar in 94 tetrads, 17.8% of those in the VC. Nevertheless, since many field recorders work at thespecies aggregate level and do not distinguish the separate Polypodium species, we regard it asdefinitely under-recorded. As the tetrad distribution map indicates, this form is widely scatteredthroughout Fermanagh, but more prevalent in the wetter, more acid, rocky upland environments of theWestern Plateau.
Threats
None.
Native, very rare, but probably under-recorded.
1969; Jackson, Dr J.S.; Boho Caves.
Throughout the year.
Growth form and preferred habitats
All ten records for this sterile pentaploid hybrid in the Fermanagh Flora Database have been determinedby Paul Hackney working on specimens deposited in the herbarium at BEL. The most recent eightwere collected by RHN either growing in moss on rocks or as epiphytes on tree trunks in damp old woods.In view of the frequency of the parent species and their degree of ecological overlap, this intermediatehybrid is probably quite widespread and is likely to be the most common of the three sterilePolypodium hybrids.
In a W European study assessing the pH preferences of ferns and their root cation-exchange properties andpreferences, Koedam et al. (1992) found that P. × mantoniaefollows its tetraploid parent P. vulgare s.s. in being 'acidiphilous' (ie strongly calcifuge),since it occurred on soils with a median pH value as low as 3.8. Furthermore, no calcium carbonate wasdetected in any of the soil samples taken from the roots of this fern hybrid. On the other hand, Page(1997) reported it occurring in humid conditions on mossy boulders over a wide range of rock typesincluding limestones.
British and Irish occurrence
Page (1997) reported that this hybrid had been found in at least 29 VCs in Britain & Ireland and thatthe distribution was slanted towards western areas. The New Atlas hectad map shows that P.× mantoniae is now known quite widely across Britain and N Ireland,and it occurs on soils derived from both siliceous acid and limestone rocks.
Reproduction
Although this is usually regarded as a highly sterile hybrid, some of the plants appear to have a highproportion of apparently normal spores. The backcross with P. vulgare s.s. has been suspected inat least one of the Fermanagh records. This hybrid is vegetatively vigorous and can form large clones.
Fermanagh occurrence
The details of the other nine Fermanagh records follow with their collector; all were determined by P.Hackney and vouchers exist: Finlane Td, Florencecourt Forest, 1976, P. Hackney; scarp SW of LoughAchork, December 1987, RHN; Castle Archdale, Lower Lough Erne, December 1987, RHN; Ely Lodge Forest,Lower Lough Erne, February 1988, RHN; Sillees Wood, March 1989, RHN; Clonelly, NW of Kesh, April 1989,RHN; Arney River, April 1989, RHN; Crossmurrin NR, December 1989, RHN; Ballindarragh Bridge, ColebrookeRiver, 1988-90, RHN. The last listed record is considered a possible backcross with P. vulgares.s.
Native, frequent but still under-recorded. Suboceanic temperate.
15 February 1969; Jackson, Dr J.S.; Boho Caves.
Throughout the year.
Growth form and preferred habitats
Typical specimens of this species (a fertile hexaploid hybrid formed by allopolyploidy between the othertwo British Isles Polypodium species) have ovate to narrowly oval fronds, generally longer thanthose of P. vulgare s.s. and with at least the lowest pair of pinnae bent forwards (ie inflexed),to form a 'V' shape. The fronds are leathery, evergreen and frost and drought resistant, fresh onesbeing produced in late summer and autumn, ie later in the year than P. vulgare s.s. andbefore those of P. cambricum (Southern Polypody) (Jermy & Camus 1991).
The plant is weakly calcicole or may prefer near-neutral conditions. A study of fern species with respectto root cation-exchange properties carried out in W Europe discovered the pH of soil samples at theroots of P. interjectum had a median value of 6.67, so that Koedam et al. (1992)classified it as 'neutrocline'.
P. interjectum grows in very much the same types of habitat as P. vulgare s.s., ie onrocks, cliffs, stony banks, mortared walls and also as an epiphyte on trees in damp woods.
Fermanagh occurrence
As with P. vulgare s.s. (Polypody), this perennial polypody is definitely still under-recorded inFermanagh. Many of the existing records have been determined or verified by Paul Hackney at BEL.Intermediate Polypody is the second most common species of polypody in Fermanagh having been recorded sofar in 90 tetrads, 17.1% of those in the VC. As the distribution map indicates, it is widelyscattered throughout, but more frequent in the west of the county.
British and Irish occurrence
The Fern Atlas hectad map gave an early picture of the known distribution of P. interjectumin 1978, where the sub-Atlantic influence appeared quite strong in Britain, while the Irish distributionwas then very much more evenly, or randomly scattered, although inland sites in NI appeared almostentirely absent (Jermy et al. 1978). The New Atlas hectad map now shows IntermediatePolypody is common and widespread throughout most of NI, but with apparent gaps in Cos Tyrone and Armagh(H36 and H37). The very obvious patchiness of the plotted distribution on both islands strongly suggeststhat recording of P. interjectum, although greatly improved in comparison with the FernAtlas, remains incomplete.
Name
The Latin specific epithet 'interjectum' means 'intermediate in form', and of course refers to the factthat this species arose as a fertile hybrid (Gledhill 1985).
Threats
None.
Native, very rare, but probably still under-recorded.
1974; Hackney, P.; Boho Caves, voucher in BEL.
September and October.
Growth form and preferred habitats
P. × shivasiae is the rarest of the three Polypodium hybrids inBritain and Ireland. However, since the parent species are becoming quite frequently recorded inFermanagh (particularly P. interjectum (Intermediate Polypody)), and as they are both weaklycalcicole and undoubtedly overlap geographically and ecologically, this hybrid might be rather less rarein the VC than the current few records indicate. Having said that, as Roberts (1970) pointed out,survival of the relatively small, often sporadic populations of P. cambricum might be adverselyaffected by the formation of its hybrids with other species, since the offspring will compete with thediploid parent in a habitat in which it is already rather restricted. Under these circ*mstances anygenetic barrier to hybridisation would tend to be strengthened.
P. × shivasiae is the most spectacular in appearance of the threesterile hybrid polypodies found in the British Isles, combining as it does large size with vigorousgrowth. As to be expected, it is intermediate in its morphological and physiological characteristicswith respect to its parents. It grows on sheltered sections of limestone walls and on equally sheltered,shaded, parts of steep, wet, basic rock cliff faces, usually with both parent species nearby (Page1997).
Fermanagh occurrence
All four Fermanagh records of this rare hybrid have been determined by Paul Hackney, the first recordverified by R.H. Roberts. The details of the other three Fermanagh records, all collected by RHN, are asfollows: Keenaghan Lough, Tievealough Td, September 1988; Levally House, 1 km SE of Roosky, October1990; and W end of Lough Acrussel, 1988-90. This last record is marked "requires checking".
Name
The specific name 'shivasiae' commemorates the work of the British fern taxonomist M.G. Shivas, whocontributed greatly to understanding of the genus in the 1960s and 1970s.
Native, occasional, but probably still under-recorded. Mediterranean-Atlantic.
1975; Hackney, P.; limestone rocks on a steep slope at Hanging Rock NR.
Throughout the year.
Growth form and preferred habitats
The fronds of P. cambricum are broadly triangular and can be up to 50 cm in length, with the edgesof the pinnae often quite serrate. New fronds are produced later than in the other two Polypodiumspecies, in the autumn and winter, and they are reputedly the least frost hardy of the three species. Ithas been suggested that in Britain and Ireland Southern Polypody is mainly confined to lowland siteswithin a few hundred metres of sea-level, often coastal and lying within the 4°C winter minimum isotherm(Page 1997). Spore-producing sori are confined to the upper third of the frond; young sori are more ovalthan in other forms of polypody, and when the plant is vigorously vegetative, no sporangia are formed,rendering the plants sterile.
Species identification
Variation in both vegetative and reproductive characters is so great in the genus Polypodium thatP. cambricum (the diploid) and P. interjectum (the hexaploid allopolyploid fertile hybridderived from a cross between P. cambricum and P. vulgare s.s., followed by spontaneouschromosome doubling), tend to overlap in many respects. The presence of relatively large, branchedcolourless threads (ie paraphyses), within the sorus among the sporangia, are unique to P.cambricum, and taken in conjunction with 'good spores' (ie fully formed, fertile,regular-shaped, non-aborted ones), they provide the best distinguishing character for the species(Roberts 1970; Page 1997).
Having said this, these are very much laboratory microscopic characters, and furthermore care must betaken not to confuse the much larger, more branched paraphyses with the minute glandular hairs whichoccur scattered on the lower surface of the frond in all European species of Polypodium. Thesehairs are much smaller than true paraphyses, being usually only 3 or 4 cells long – with theirglandular, inflated terminal cells mostly coloured a dark reddish-brown (Roberts 1970 – see p. 128, Fig.6 to clarify these differences).
Fermanagh occurrence
While this perennial is the most distinctive and readily identified of the Polypodium species orhybrids in Britain & Ireland, it is also very definitely the least frequent of the three species inFermanagh and elsewhere in these islands (Page 1997). So far, P. cambricum has been recorded injust 37 of Fermanagh tetrads, 7.0% of those in the VC.
Southern Polypody is a strongly calcicole species and, as the Fermanagh tetrad map indicates, ittypically grows in the W & S of the county on limestone natural rock outcrops. Often it occupies themore sheltered parts of cliffs, crevices in old lime-mortared walls, or ground on the steep-slopingrocky floors of hazel woods, where it is capable of forming large stands. Locally, it has never beenfound in artificial habitats such as quarries, nor growing as an epiphytic on the bark of trees,behaviour that may be confined to more maritime districts than occurs in Fermanagh (Jermy & Camus1991).
In drier, more exposed upland sites, which are usually limestone gorges and cave mouths, P.cambricum associates with Asplenium ruta-muraria (Wall-rue) and Ceterachofficinarum (Rustyback). On the other hand, when occurring in shade in Ash and Hazel woodland inconstantly damp and humid conditions, it consorts instead with other shade-tolerant species such asHedera helix (Ivy), Polystichum setiferum (Soft Shield-fern) and Phyllitisscolopendrium (Hart's-tongue).
British and Irish occurrence
The current knowledge of the occurrence of P. cambricum in Britain & Ireland, as displayed atthe hectad level in the New Atlas and the 2005 New Atlas of Ferns, shows it is bothsouthern and western in Britain, although stretching northwards as far as mainland Argyll in W Scotland(VC 98) (Preston et al. 2002; Wardlaw & Leonard 2005). The same two maps show it very muchmore widely scattered across the whole of Ireland. The distribution looks patchy, however, suggestingthat the recording effort is distinctly uneven. More work is required to arrive at an accurate pictureof the real distribution of this fern.
World occurrence
P. cambricum is entirely confined to W and S Europe, being sparse and rather disjunctly spreadfrom the British Isles to Portugal, N Africa and eastwards through the Mediterranean islands to Greeceand Turkey (Jalas & Suominen 1972, Map 140; Page 1997).
Name
The species Latin epithet 'cambricum' means 'of Wales' (Cambria) (Gilbert-Carter 1964).
Hybrids
While diploid P. cambricum is by far the least common of the three Polypodium species inBritain & Ireland, its hybrids with the other two species are even rarer. The hybrid between P.cambricum and P. interjectum (P. × shivasiae) shouldbe actively looked for as both species are calcicole and P. interjectum (being very much the morecommon parent) probably occurs at or near every P. cambricum site. So far only four records ofthis hybrid have been discovered in Fermanagh.
Threats
Clearance of woodland.
Native, common, widespread and locally dominant. Circumpolar temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Introduction
Bracken is a serious opportunistic, invasive, colony-forming weed almost everywhere in Britain andIreland, and indeed world-wide on under-used, under-managed or abandoned farmland (Cody & Crompton1975; Taylor 1990). Current land-use changes in Britain and Northern Ireland induced by socio-economicand political forces, including the policies of 'set-aside' and 'Environmentally Sensitive Area'farmland designation, may very well end up promoting Bracken encroachment. Bracken is such a vigorousinvader, persistent and so extremely difficult to eradicate, that its many researchers have formed anInternational Bracken Group, which regularly holds conferences and issues publications (Smith &Taylor 1986, 1995; Thomson & Smith 1990).
Taxonomy
P. aquilinum is world-wide in its distribution, but it is both genetically and ecologicallyvariable and it is also phytochemically polymorphic (eg with respect to the levels of animal and planttoxins it contains), so that several different schemes of subdivision have been proposed involvingsubspecies (or species) and up to a dozen varieties (Tryon 1941; Cody & Crompton 1975; Page 1997).
Taken at its taxonomic widest, P. aquilinum has had a couple of hundred varieties and formsdescribed (Hultén 1962, Map 131); but that way madness lies! In Britain and Ireland, what previously wasregarded as the solitary species P. aquilinum has been subdivided by Page (1997) into twospecies, P. aquilinum and P. pinetorum, the former being further subdivided into threesubspecies, and the latter (which is confined to scattered sites in Perthshire and Inverness-shire),into two subspecies. The New Flora of the British Isles (1997), on the other hand, takes a verymuch more conservative view of the variation, downgrading Page's species to subspecies, and regardinghis subspecies merely as local ecotypes. We have not differentiated the subspecies of either of theseauthors in Fermanagh, but almost undoubtedly the Bracken in our area is P. aquilinum subsp. aquilinum, by far the most vigorous and abundant weedy formof the plant throughout the whole of the British Isles.
Growth form, size and timing
The thick black rhizome runs deeply buried, between 10 to 50 cm below the surface in well-drained acidsoils (usually less than pH 6.5). The spreading rhizome branches and sends up to the surface thefamiliar annual branched aerial green fronds. The fronds vary greatly in height, but can be up to 2.5 mtall (a maximum of 2.75 m tall has been recorded (Step & Jackson 1945)), when developed over fertiledeep soil. Often fronds are only half that height when the fern is growing on more nutrient-poorsubstrates.
The fronds begin poking above ground in mid-April, and when the hook-like croziers first appear they arevery sensitive to frost and to damage by trampling (Grime et al. 1988). The fronds are slow todevelop, requiring time for the great length of stipe and rachis to harden before the physical strain isput on them of the fully spread-out pinnae. In fact the fronds do not stop growing and using up therhizome energy reserves until the latter half of July. This timing determines the optimum timing forcutting or the use of systemic herbicides for the eradication of Bracken, since the plant's reserves arethen almost non-existent. The energy-containing products of photosynthesis are then being translocateddown into the underground parts of the plant for storage, and the herbicide will be carried down intothe rhizome with this flow of material.
Preferred habitats
Bracken is very frequent on hillsides, sloping, rocky waste ground, woods and roadside embankments. It isespecially common on reasonably deep, well drained, acidic upland pastures, where it can produce frondsup to 2.5 m tall and become locally dominant. The plant does not have a high requirement for soilnutrients and the ideal habitat in Scotland was described as ground providing free-draining slopes ofbrown earth soils on sheltered, lower hill faces of glen sides. It may also occur on similar soils inglen bottoms, but only if the drainage there is unimpeded (MacLeod 1982).
P. aquilinum is much less common on base-rich or shallow substrates where its growth is oftenstunted. In general, Bracken avoids aquatic sites, shallow soils, poorly drained, deep, acid peatlandsand densely shaded conifer plantations. It is also absent from well managed farmland, from frost hollowsand from very exposed situations (Biggin 1982; MacLeod 1982).
While primarily a plant of woodland shade and open moorland pastures, Bracken is also common on roughgrassland on waste ground and roadsides. Although P. aquilinum abhors waterlogging, it frequentlyoccurs near waterways and on the drier margins of lakes and bogs at all altitudes, excepting the mostexposed sites. In general, for reasons that are not at all clear, Bracken seems to frequent shade moreoften in lowland sites than it does in upland areas (Grime et al. 1988). It is also frequentlyfound on cliffs, screes, limestone pavement and other forms of rocky ground. This includes artificialhabitats such as old stone quarries and neglected areas of sand and gravel pits.
Soil preferences
As is the case with other primarily or mainly calcifuge plants, Bracken frequently occurs in limestoneareas of Co Fermanagh. The explanation of this unexpected behaviour is that soils in the west of Irelandhave been so thoroughly leached by prevalent heavy rainfall, acid peat often develops directly on top ofcalcareous rock. The suggestion has been made that P. aquilinum on limestone soils in places suchas Andalucia in southern Spain, may differ to the extent of having only half the normal chromosomenumber (ie 2n= 52) (Molesworth Allen 1968). Studies on the soil preferences of ferns with respect toacidity carried out in W Europe by Koedam et al. (1992), found that P. aquilinum was'acidiphilous' (acid-loving or acid-tolerant). In this study, soil samples taken from the fern root masshad a median pH of 4.0. On the other hand, Jonsell et al. (2000) suggest that in Scandinaviancountries, Bracken is probably quite indifferent to lime.
Bracken litter ecology
Being a very large colonial, deciduous plant, Bracken has the potential to produce an enormous amount ofleaf litter when the aerial shoots die off in late autumn. In communities where P. aquilinumforms virtually pure stands, litter production can be between 8,000 and 14,000 kg/ha/yr (Watt 1976). Therates of both litter accumulation and its decay depend very much on the particular habitat conditions,but in open heathland it can be the actual depth of litter (and the toxic and allelopathic substances itcontains), that allows the species to dominate other plants. In deciduous woodland, on the other hand,litter decay is rapid and very little or none remains after just one year (Watt 1976).
An excessive accumulation of frond litter can prove detrimental to Bracken itself, so that it can fallvictim to its own success. When this happens, the rhizome degenerates, gaps in Bracken cover occur, andthis can allow colonisation by woody species and other plants (Watt 1976). It was his prolonged studiesof Bracken and its competition with heathers which led Watt (1947, 1955), to the influential idea of'cyclical succession' in vegetation, and to his describing the phases of Bracken 'sociology' in terms ofa 'pioneer, building, mature and degenerate' phases that form this cycle (Watt 1964).
Between the extremes of litter persistence mentioned above, there are all degrees of dead frond litteraccumulation. This phenomenon is often affected directly and indirectly by mans' interference throughgrazing, trampling, burning and cutting of the live or dead fronds (Watt 1976). From an ecological pointof view it is probably wisest to consider Bracken as a pioneer species, that will most likely give wayto colonising tree seedlings of species such as birch or oak in succession towards an eventual climaticclimax forest vegetation. Within the process of succession, P. aquilinum occupies a niche as astable, controlled, often very long-lived component species (Page 1986).
Fermanagh occurrence
In Fermanagh, Bracken shows a definite preference for acidic conditions, but it certainly is alsofrequent in limestone areas. This large, deciduous fern has been recorded in 303 tetrads, 57.4% of thosein the VC.
Factors involved in the historic spread of Bracken
Studies in Scotland in particular suggest that colonisation by Bracken has greatly increased from about1750 onwards, and that until at least the first three or four decades of the 20th century, farmers inareas of upland grazing noticed rapid invasion of their pastures. In this manner, Bracken has become areal menace, and one that continues to spread (Rymer 1976; Page 1997). P. aquilinum was probablyoriginally a plant of open woodland and forest margin, occurring mainly on calcifuge terrain where adegree of shade and competition from other species kept its growth in check. Destruction of the woodlandby man's activities, extending from the period of the Neolithic farmers onwards, has removed much ofthese constraints, allowing this vigorous rhizome-possessing species to form the extensive stands itoccupies today on virtually all sorts of sloping rough ground (Rymer 1976; Page 1982(b); 1997, p. 361).
Apart from woodland destruction, numerous other factors are involved in Bracken spread. Probably mostimportant has been the decline in rural (especially upland) human populations, and the consequentabandonment of areas of farmland previously cultivated or more or less intensively grazed. This wouldalso account for ground where Bracken was previously cut, either for its eradication, or morepurposefully for economically significant folk use of the plant (see below).
Subsidiary factors that have encouraged Bracken spread include the general move from cattle to sheepgrazing, with a consequent reduction in trampling pressure; a rising rabbit population with itsselective grazing pattern avoiding Bracken; poor understanding of heather burning regimes leading to aloss of heather cover; plus a general climatic rise in temperatures occurring over the last 250 years(Rymer 1976).
Colonisation of burnt ground
Bracken is an extremely invasive plant, and it is particularly so on burnt ground. The sporophyte rhizomeand the sexual prothallus stage of the species are both very well adapted to rapidly colonise thealmost-virgin ground of recently burnt areas in both woods and heaths, while surviving, competing plantshave yet to recover from the effects of the fire. While the sporophyte plant shows a definite preferencefor calcifuge conditions, the prothallus stage rather unexpectedly is base-demanding. The potash andother minerals released by fire clearly provide particularly suitable conditions for rapid sporegermination, and the released soil minerals promote the growth of both the Bracken gametophyteprothallus and the juvenile sporophyte arising from it after fertilisation (Conway & Stephens 1957;Page 1997, p. 362).
Grazing at either the pioneering or re-colonisation stage of vegetation development, especially closecropping by sheep, is capable of tipping the competitive balance between available species towardsBracken (Page 1982(b)).
The extent of Bracken in Britain
Estimates of the land area covered by Bracken in the United Kingdom of Britain and Northern Ireland rangefrom 3,000-6,000 km2, with the most serious infestations in upland regions of the west andnorth (Fowler 1993). Undoubtedly the scale of the Bracken weed problem in Britain is large and thelandscape and biological conservation implications are quite frightening. The fern causes problems foragriculture, forestry, conservation, shooting interests, recreation, health and water collection(Pakeman et al. 1994). Several surveys made between 1978 and 1990 estimate that Bracken dominatesapproximately 3,600 km2 of Britain's land area, representing around 1.5% of the total landcover. It is present, but not necessarily dominant, in around 17,000 km2, or 7.3% of Britishland cover, and there is a considerable risk of these figures increasing due to changes in both landmanagement and climate (Pakeman et al. 1994).
Bracken continues to spread, and Page (1997) reported this occurring in Britain at a rate of 1-3% peryear, which seems an alarmingly high figure. Very probably the move during the last 100 years away fromcattle to sheep grazing in upland areas has contributed to the extension of Bracken on such pastures,since apart from the differing grazing pressure, the emerging fronds can more readily survive sheeptrampling than that of heavier beasts (Step & Jackson 1945).
Control measures
Efforts to control Bracken largely consist of ploughing it in, regular cutting, crushing, or the use ofherbicides, especially Asulam, which is Bracken specific. However, all of these methods of attack areexpensive, labour-intensive and require safe access to the land by agricultural machinery. Since Brackenoften infests steep or rocky slopes, aerial application of herbicide is often the only current optionfor control, making it both expensive and problematic, since such widely broadcast spray may wellendanger other desirable or protected species. In addition, the problem remains that unless all of therhizome buds are destroyed, the plant will survive and reappear at a later date when control measuresare eased (Taylor 1990).
Rehabilitation of sites is a very important part of conventional Bracken control programmes and thisgreatly adds to the cost, particularly if fencing is required to prevent access by grazing animals(Fowler 1993). A programme of experiments on biological control using two moth species specific toBracken imported from S Africa showed definite promise during testing under semi-natural conditions(Fowler 1993). However, at the eleventh hour funding was refused by government for field trials in GreatBritain (Taylor 1995). Reviewed in a global context, Bracken is encroaching and not retreating where itoccurs, and within the scope of current technology and economics, it is well nigh impossible to reverse,control or eradicate the plant (Taylor 1990).
Reproduction
Under dense woodland canopy Bracken fronds are quite often sterile. However, spore production is enormousin unshaded habitats, where a single frond is capable of producing up to 30 million spores (Conway1957). Having said this, Bracken spore production is sporadic, development being affected by plant andfrond age, degree of shading, exposure, weather conditions, perhaps soil characteristics, and by thegenetic make-up of the individual. In most years spore output is generally poor, at least in Britain andIreland, and some populations appear to be consistently sterile, even when others nearby spore copiously(Dyer 1990).
Despite the potentially enormous spore production, established stands of Bracken often reproduceexclusively by vegetative means due to the amount of frond litter they produce smothering the surface ofthe ground and preventing spore germination. Thus spores are probably only important in the colonisationof new sites on burnt or otherwise disturbed ground (eg, animal burrows, damp hollows and lime-richcavities in rocks, old walls and rubble (Grime et al. 1988; Dyer 1990). More work is urgentlyrequired on the colonising potential of Bracken spores and the significance of Bracken spore banks inthe soil.
Bracken nectaries
As long ago as 1877 Francis Darwin first observed that Bracken plants secrete sugars through numerousfoliar nectaries found all over the plant, but especially on the under-surface at the junctions betweenpinnae and the rachis, and also in smaller amounts at the junctions between pinnules and the pinnamidrib (Page 1982(c)). The size and prominence of the nectaries varies with the particular habitatoccupied, being larger on plants growing in more open sites (Page 1982(c)).
The function of sugar glands like these in Bracken and other plants (including, worldwide, a few otherunrelated ferns), has been a topic of debate since the early years of the 20th century. Some biologistssuppose them to be purely excretory, while others believe them to attract pugnacious ants into amutualistic relationship with the plant, whereby in return for a food reward they protect it fromherbivore attack (Tempel 1983; Page 1997). The real question here is whether or not possession of suchnectaries provides the species with an ecologically significant advantage over plants without suchstructures? Numerous studies carried out have found that it is not easy to answer this apparently simplequestion.
Bracken and ants
Experimental work in New Jersey by Tempel (1983) confirmed that Bracken nectaries are most active in theyoung expanding frond, and that they did attract ants. However, she also showed that mature plantscontinue to secrete small quantities of sugar, even though levels of ant activity sharply fell away onfully expanded fern fronds. Despite indications that Bracken is adapted to some form of mutualisticrelationship with ants, Tempel concluded that no such interaction actually existed in her particulargeographical region, since the ants in her study were non-aggressive and they did not protect the fernfrom herbivore damage. This still leaves a number of open questions relating to the significance offoliar nectaries on Bracken in Britain and Ireland, amongst which must be, is it a topic worthy offurther study?
Toxicity
All parts of the plant, including its airborne spores, contain carcinogens as well as various otherpoisons. Some of these remain toxic after the plant has been cut and dried, so that it can be a dangerto both man and livestock. The carcinogenic and immuno-suppressive effects of Bracken are an active areaof medical research (Cooper & Johnson 1998). As with other plant toxins, the role of poisons inBracken is to deter herbivores and inhibit the growth of neighbouring plants (ie it is allelopathic topotential competitors). The chemical armoury of Bracken is generally extremely effective in theserespects. Allelopathic toxins are contained primarily in Bracken roots, rhizome and litter, and they arereleased into the soil environment to suppress the growth of associated plants (Gliessman 1976).
However, not all Bracken populations possess the full complement of animal deterrent toxins, and theseplants may sometimes be detected by livestock and become heavily grazed. Sheep usually avoid Bracken,but if starving they will graze it and they can then become addicted to it. The same situation applieswith horses. Another Bracken constituent is known to cause thiamine deficiency in non-ruminant animalssuch as horses and pigs (Cooper & Johnson 1998).
Other ill effects of Bracken in pastures include the shelter it provides for the sheep ticks thattransmit Louping ill virus to both grouse chicks and sheep. Sheep ticks are also implicated in thetransmission of Lyme disease to a range of animals including man (Fowler 1993).
Uses
Before the health risks inherent in handling Bracken were known, Bracken was collected and used in farmsfor animal fodder, bedding, kindling, thatch, compost, fertiliser (on account of its potash content) andas packing material for fruit. It was even used as human food: the rhizome contains a lot of starch,although in reality it tastes very astringent (Grieve 1931; Step & Jackson 1945; Rymer 1976). Youngfrond croziers were previously eaten in Japan like asparagus, while in Siberia and Norway expandedfronds were used in the past along with malt to brew some dreadful form of beer (Grieve 1931). Rhizomeswere dried and powdered to make flour from which bread was baked either directly, or after mixing withwheat flour, a practice which was found in native cultures as far apart as New Zealand and Normandy(Rymer 1976).
The astringent properties of the rhizome also saw it being used to dress and prepare kid and chamoisleather, but although this has been reported many times from Lightfoot (1777) onwards, we do not knowwhere or when this was ever the case (Rymer 1976). The ash of Bracken contained enough potash torecommend its use in glass making, and it was also boiled with tallow and used as soap in parts of theEast.
Bracken is a light and quick-burning fuel (making it a severe fire risk when it is abundant in or nearrecreational areas), and it produces a very violent heat (Rymer 1976). In many parts of the BritishIsles, it has been used in the past for burning limestone, for heating ovens used in baking and brewing,and for firing bricks. The cutting of more or less dead Bracken fronds for fuel, thatch, bedding,packing material or other purposes, had the effect of removing some, but not all, of the fern'sfrost-protective litter layer. Since it was gathered late in the season, however, this will have hadrelatively little ecological consequence on the performance or survival of the species.
A most informative, thoroughly researched review of the ethnobotany of Bracken has been published byRymer (1976) and it is highly recommended reading.
Names
The origins of the plant's botanical names are quite fairly described as 'obscure'. The genus name'Pteridium' is derived from the Greek diminutive of 'pteris', fern, from 'pteron', meaning 'a wing','winged', (ie 'little wing'), or 'a feather', an allusion suggesting that some fern fronds resemble abird's wing (Stearn 1992). This notion, except with respect to size, fits the expanded Bracken frondreally rather well. The English term 'fern' is similarly derived from the Anglo-Saxon 'fepern', meaning'a feather' (Grieve 1931).
The Latin specific epithet 'aquilinum' means 'eagle-like', a notion reputedly suggested to the Swedishbotanist, Carl Linnaeus, by the pattern of vascular bundles that is observed when the lower stipe is cutacross obliquely, which supposedly resembles a spread eagle (Grieve 1931; Gilbert-Carter 1964). Forthose interested, this and other similar notions of the name origins are given by Step & Jackson(1945).
Folklore and folk medicine traditions are given full accounts by Grieve (1931, p. 305) and Vickery (1985,pp. 44-45). The most interesting and widespread folklore tradition is that Bracken, or its spores,confers invisiblity (Rymer 1976).
The English name 'Bracken' (sometimes 'Brecken') is the plural of 'Brake', and apparently is derived fromthe Old English 'bracu', possibly referring to something broken. This would be appropriate to the deadfern in winter, forming as it does a dense tangle of broken stems (Grigson 1974). It should be notedthat the name 'Brake' or 'Bracken' was used in pre-scientific days for large ferns generally, and alsomore particularly applied to Pteridium aquilinum (Britten & Holland 1886). Another possibleorigin of 'Brake', 'Brakes' or 'Bracken' is suggested by Prior (1879), who derives it from the German'Brache' or 'Brach-feld', meaning uncultivated land or land that is breakable, or open to tillage aftera term of years, ie land that is not preserved as forest. This particular etymology suggests thatBracken has long been known as an active coloniser of abandoned arable land (Rymer 1976).
Threats
Reviewed in a global context, where Bracken already occurs it is encroaching and not retreating. Withinthe scope of current technology and economics, it is well nigh impossible to reverse the spread in theseareas, or to control or eradicate the plant from them (Taylor 1990).
Native, very rare. Circumpolar temperate.
1806; Scott, Prof R.; Scottsborough lakelet.
June to September.
Growth form and preferred habitats
An erect, medium-sized perennial fern with a creeping rhizome from which arise solitary or clumped annualfronds, T. palustris is a decidedly rare species in Fermanagh, confined to the permanently wet,peaty, but not too acidic, muddy ground dominated by sedges, alder and willow around Upper Lough Erneand several of the smaller lakes in the county. Marshes, sedge fens and wet fen-carr woods by smalllakes are the typical habitats of the species.
Fermanagh occurrence
As the distribution map indicates, T. palustris is represented in ten tetrads, just eight of whichhave post-1975 records. It is mainly concentrated in the south of the county, in wet ground around UpperLough Erne and beside lakes along the SE border of the county. The first Fermanagh record for thisspecies is a recently noted early herbarium specimen in DBN which was collected by Prof Scottfrom near his home in Scottsborough. Marsh Fern has not been re-found at Scott's site, nor at Hart'spre-1887 station on limestone shingle by the River Erne close to Belleek (a record so vague it probablyrefers only to the Co Donegal side of the international border (H34)) (Hart 1898), nor at the Wend of Inver Lough where it was recorded by Meikle and his co-workers in the period 1946-57.
The sites in the River Finn catchment represent the main concentration of the species in NorthernIreland. The sites in S Fermanagh from which the species has been recorded are: Derrymacrow Lough,Abacon Lough, Farmhill Lough, Clonshannagh Lough, Lough Garrow, Killynubber Lough, Inver Lough, thelakelet by the avenue at Crom and the River Finn near Gortnacarrow Bridge. T. palustris is evenrarer in adjacent Co Cavan (H30), where only one of four stations has a recent (1996) record (Reilly2001). Similarly in Co Tyrone (H36), which had two late-19th century sites for the fern, it onlypersists at Enagh Lough, near Caledon (McNeill 2010).
Page (1997) regards Marsh Fern as a plant of essentially Continental climatic conditions. Since Fermanaghis decidedly oceanic (or Atlantic) in its climate, and the local T. palustris sites lie within12-25 km of the west coast of Ireland, the species must be close to the extreme margin of its range,where it is often and extensively replaced by Osmunda regalis (Royal Fern).
Irish and British occurrence
Throughout both Ireland and Britain, T. palustris is a scarce and widely scattered species. TheCensus Catalogue of the Flora of Ireland lists past records from a total of 21VCs, but the map in the 1978 Fern Atlas records only 14 Irish hectads with post-1930 records(Jermy et al. 1978), while the New Atlas map plots 18 hectads with post-1987 records(Preston et al. 2002).
In Britain, while slight concentrations occur on the Isle of Wight (VC 10) and S Hampshire (VC 11)(Brewis et al. 1996), and again in Norfolk (VCs 27 & 28) (Beckett et al. 1999), MarshFern becomes much rarer north of a line between Hull and Liverpool (Jermy et al. 1978; Stewartet al. 1994; Preston et al. 2002).
European and world occurrence
Marsh Fern ranges widely across warm-temperate latitudes of mainland Europe stretching east to Siberia.In the north it reaches N Finland, and it stretches southward to the southern tip of the Peloponnese inGreece, the distribution thinning markedly in both directions (Jalas & Suominen 1972, Map 73).Taking the species in the broadest taxonomic sense, it is disjunctly circumpolar, with gaps in N Asiaand in the eastern half of N America. The NE American and E Asia forms of the plant are now sometimesrecognised as varieties of a separate species, T. thelypteroides (Michaux) Holub. A form of T.palustris s.l. is also found in S Africa, S India and New Zealand, now referred to T.confluens (Thunb.) Morton (Hultén 1962, Map 170), or to T. palustris subsp. squamigera (Schlecht) Hult. (Hultén & Fries 1986, Map38; Page 1997). Without the insight of a trained taxonomist, I am amazed and a little disconcerted bythe fact that Thelypteris palustris has been referred in the past to as many as six other genera,some nowadays totally unfamiliar, so that index searching in older texts often works much better usingthe more stable English common name (Hultén 1962).
Names
The genus name 'Thelypteris' is a Greek compound meaning 'Lady-fern', a name first used by the ancientGreek botanist, Theophrastus, for an unspecified fern. The specific epithet 'palustris' is Latin (givena masculine ending), and means 'of swampy places' (Gilbert-Carter 1964). The standard English commonname of T. palustris is nowadays 'Marsh Fern', a folk name suggested by its habitat. Previouslyit was also called 'Marsh Buckler-fern' and, in the Isle of Wight, 'Ground Fern' (Step & Jackson1945).
Threats
In Fermanagh, as elsewhere in Britain and Ireland, agricultural drainage, cultural eutrophication andscrub encroachment is making inroads on suitable habitats, and Marsh Fern is becoming increasingly rare(Hackney et al. 1992).
Native, very rare. Circumpolar boreo-temperate.
1860; Smith, Rev Prof R.W.; Brookeborough Deerpark.
June to August.
Growth form and preferred habitats
The triangular annual fronds of Beech Fern with their deflexed pair of lower pinnae in a different planefrom the rest of the blade are quite unmistakable, but they are rarely enough seen in Fermanagh. Thisvery misleadingly named fern never occurs under beech trees, the English common name simply being amistranslation of the Greek 'phegos', which means 'oak'. Nevertheless, oak or beech being equallyinappropriate names, this perennial fern is a plant of moist, shady cliffs and damp banks, often nearstreams, in upland ravine woodlands. The creeping rhizome sometimes allows the plant to developextensive colonies in undisturbed sites (Jermy et al. 1978). Stunted fronds of Beech Fern arealso found where water drips through the roots of other plants on cliff ledges, in crevices and amongboulders on rocky slopes.
The species frequents a wide range of rock types and soil pH, but while it appears to prefer soil with areasonable base content (Page 1997), it can also occur under very acidic conditions, eg on the Mournegranites in Co Down (H38). In the latter situation, it must be presumed that there is some slight baseenrichment, however undetected it remains. A degree of inaccessibility tends to assist the survival ofthis fern since P. connectilis is known to be intolerant of grazing (Sinker et al. 1985).
Fermanagh occurrence
This creeping, rhizomatous fern has been recorded in a total of eight Fermanagh tetrads, but only five ofthem have post-1975 records.
P. connectilis was first reported in Fermanagh in the grounds of Brookeborough Deerpark by Smithin 1860. Meikle and his co-workers refound it there in the 1950s, but it has not been seen since then atthis station (Meikle et al. 1975).
The most interesting site in Fermanagh for the Beech Fern was found by the Rev W.B. Steele in 1929 on theS shore of Lower Lough Erne at Carrickreagh. On its first discovery, it was remarkably abundant on arelatively dry, flattish, limestone woodland floor, under mixed oak, birch and hazel. It remainedabundant until 1945 when the site was largely destroyed by the clear felling of the woods and anextension of the nearby quarrying operation (Carrothers et al. 1946). Praeger visited theCarrickreagh site in 1933 and described it in enthusiastic terms, "The fern grows here on limestonerubble thinly covered with humus and mosses, among Primula, Endymion, Hedera,Lysimachia nemorum, Thymus, and Sesleria, forming dense patches up to 20 feet [6 m]across with fronds up to 2 feet [60 cm] high." (Praeger 1934a). In his book The botanist inIreland, he commented on the Beech Fern at the Carrickreagh site as, "the only habitat ofthe kind which I know in Ireland, its characteristic stations being wet chinks or ledges in themountains" (Praeger 1934i). He visited it again around 1938 and reported, "More abundant inthe woods of Carrickreagh than anywhere else I have seen it in Ireland; one dense patch measured 250 ft[76.2 m] by 50 ft. [15.24 m]." (Praeger 1939). At present, just one tiny patch survives in thisarea, consisting of just a couple of fronds on the bank of a stream.
Other current local Fermanagh sites include a strong lowland colony on the Bannagh River near somewaterfalls, and the fern also maintains a precarious existence on high ground as tiny fronds in rockcrevices on the north face of Cuilcagh mountain.
Irish occurrence
Elsewhere in N Ireland, the species is rare and widely scattered, although locally plentiful in theMourne Mountains and the wooded Antrim Glens (Hackney et al. 1992). It has many fewer modernstations in N Ireland than was previously the case and a rather similar situation pertains in theRepublic of Ireland (An Irish Flora 1996; New Atlas).
British occurrence
P. connectilis is widespread and locally frequent in upland parts of the N and W of Britain, thedistribution thinning markedly further south (New Atlas). In these regions, it is most frequentin ancient woodlands dominated by Quercus petraea (Sessile Oak) developed over neutral to acidicsoils. In steeper, less accessible gullies in these woods, it frequently occurs on deeper soilspercolated with base-rich water (R.J. Cooke, in: Preston et al. 2002).
Throughout the British Isles, P. connectilis tends to follow the distribution of a mountain typeof climate, ie cool and with frequent precipitation and high humidity in summer when the fronds arepresent and growing, and fairly cold in winter when they are not (Page 1997). The summer regime clearlyapplies throughout the oceanic area of W Ireland including Fermanagh, while the winters in this area arevery much milder than in mountainous areas of Great Britain.
The creeping rhizome appears to grow slowly (measurement of just how slowly would make an interestingproject), yet the plant is capable of forming very large patches, fully occupying moist, sheltered,generally sloping sites. It therefore appears likely that the plant is long-lived, and thus is anexcellent indicator of long-undisturbed sites, perhaps in some instances, with a timescale measured inthousands of years (Page 1997). The evidence gathered for the BSBI New Atlas survey immediatelyprior to 2000, indicates that the distribution appears stable (Preston et al. 2002).
Reproduction
The nucleus of P. connectilis plant cells contain three sets of chromosomes (ie they aretriploid). While the species does manage to produce the gametophyte generation, cell division isunbalanced when meiosis occurs and thus it cannot form normal gametes (ie male and female sex cells).Nevertheless, the fern shortcuts the sexual process and produces new sporophyte plants withoutfertilisation taking place (ie it reproduces apogamously) (Page 1997).
World occurrence
Beech Fern is widely distributed throughout northern and central temperate parts of Europe and westernAsia (Jalas & Suominen 1972, Map 74). Forms of it, including the diploid and tetraploid parents ofthe British and Irish triploid form, extend it widely around the northern hemisphere making itcircumpolar (Hultén 1962, Map 107; Hultén & Fries 1986, Map 39). The taxonomic uncertainties of thisgroup of ferns can be appreciated when one sees that Phegopteris connectilis has belonged in thepast to no less than six other genera, including Dryopteris, Thelypteris, Lastreaand Polypodium (Hultén 1962; Hultén & Fries 1986).
Names
The genus name 'Phegopteris' was invented by the Swedish taxonomist, Linnaeus, the Greek 'phelos'referring to a species of oak, not the beech, though the word is cognate with the Latin 'fa*gus', thename of the Beech tree. The second part of the genus name is also Greek, 'pteris', meaning 'fern'(Gilbert-Carter 1964).
The Latin specific epithet 'connectilis' means 'well connected' (Hyam & Pankhurst 1995). Analternative English common name for the plant is 'Long Beech Fern' (Hyam & Pankhurst 1995) which,like the much more frequent 'Beech Fern', is a 'book name' rather than derived from folk usage (Britten& Holland 1886). The fern does not appear to have had any uses and there is no folklore associatedwith it.
Threats
The remnant of the Carrickreagh site is threatened by cattle trampling, but the populations at the otherFermanagh sites are kept safe by their remote nature.
Native, very rare, although easily over-looked. European temperate.
1806; Scott, Prof R.; Co Fermanagh.
June to September.
Growth form and recognition
O. limbosperma is a rhizomatous, deciduous species that can be very readily overlooked, beingeasily mistaken for young Dryopteris filix-mas (Male-fern) or D. affinis (ScalyMale-fern), although the shorter, more slender, less scaly stipe, the pinnae tapering nearly to theextreme frond base and the naked, marginally set sori are all useful distinguishing features. The youngfronds of O. limbosperma also give off a distinctive lemon or orange scent when lightly brushed,although as always, appreciation of this depends very much on the individual's sense of smell (Webbet al. 1996).
Preferred habitats
O. limbosperma is a strongly calcifuge plant of open sunlight, but at the same time it oftenoccupies more or less sheltered situations. In terms of soil, it prefers damp to moist, peaty slopeswhich are sufficiently steep to create either surface run-off or sub-surface seepage after typicallyfrequent rainfall. Moving groundwater is characteristic of the specific habitat and Lemon-scented Fernis reckoned to be indifferent to lime, and sensitive to both frost and summer heat (Jonsell etal. 2000). This set of growing conditions creates the cool, constantly moist, well-aerated rootand rhizome environment which O. limbosperma demands, and as a result the species is mostfrequently found on rather steep peaty banks beside mountain or moorland streams, or in open, acidicwoods at somewhat lower altitudes (Jermy & Camus 1991; Webb et al. 1996; Page 1997).
In Flora Nordica (Volume 1), the authors describe the habitat of this fern as oligotrophicforest and heath, ie nutrient poor, unproductive vegetation in terms of growth rate and biomass (Jonsellet al. 2000, p. 50).
Fermanagh occurrence
This is an exceedingly rare and vulnerable fern in Fermanagh, there currently being very few plants inthe county spread over just three widely spaced sites. Previously, there were two other old stations inthe county, but the fern is probably extinct in these now. O. limbosperma has been known inFermanagh from Brookeborough Deerpark since the 1860s, and it is still there, having been refound in1981 by a waterfall on the edge of the forest. Very sparse colonies, each of a few fronds, have alsobeen discovered in two new sites in the last 30 years: at Tullynanny Lough in the SW of the county(Scannell, M.P.H.; June 1974), and in a ravine beside a stream at Stranahone in the NE (RHN & RSF;August 1994). It has not been seen, however, at West & Tetley's 1899 site, simply listed as"Florencecourt", nor at Praeger's similarly vague site, described as, "two stations onthe lower hills W of Church Hill" (Praeger 1904).
Irish occurrence
In N Ireland, Lemon-scented Fern is rare and extremely local, and as with Phegopteris connectilis(Beech Fern), it is very much centred on the Mourne Mountains, Co Down (H38), in a few glens in NEAntrim (H39), and in the Sperrin Mountains and surrounding moorlands of Cos Londonderry and Tyrone (H40& H36) (Hackney et al. 1992; Northern Ireland Flora Website 2014). In the Republic of Irelandlikewise, Lemon-scented Fern is rare and very widely scattered, although it is locally abundant in a fewsites in mountainous regions such as Connemara and Wicklow (Jermy et al. 1978; Page 1997; NewAtlas).
British occurrence
O. limbosperma is much more frequent in suitable habitats in Britain than in Ireland, the speciesincreasing in abundance as one travels both westwards and northwards. This trend culminates in WScotland having the greatest abundance of Lemon-scented fern in Britain and Ireland, and probably alsoon a world scale (Jermy et al. 1978; Page 1997; New Atlas). The New Atlas surveyprior to 2000 found that the distribution in upland areas of Britain remains stable, and concluded thatmany of the losses in lowland areas that occurred before 1930 resulted from the destruction of heathlandhabitat (T.D. Dines, In: Preston et al. 2002).
European and world occurrence
On mainland Europe, the distribution of O. limbosperma most closely resembles that ofDryopteris dilatata (Broad Buckler-fern), occurring predominantly in NW Europe, more or lesscontinuously from the Pyrenees northwards to western Norway and up to just within the Arctic Circle. Italso occurs further south on the Atlantic isles of the Azores and in Madeira. On a world basis, it isregarded as circumpolar, but it only manages this in an extremely disjunct manner (Hultén 1962, Map 144;Jalas & Suominen 1972, Map 72). Despite its very wide but sparse geographic distribution, in termsof abundance W Scotland still very probably represents the world headquarters of what really looks likea very fragmented relict species (Page 1997).
Comparison of Irish and Scottish presence
The very extensive and abundant presence of O. limbosperma in western and central Scotlandcontrasts so sharply with its rare and sparsely spread occurrence in Ireland, and especially so whencompared with N Ireland in particular, that one cannot but wonder at the scale of the difference andponder on the possible reason or reasons. The climate, the rock structure and geological history of thetwo regions are strikingly similar and are intimately linked, the geology possibly most obviously sowith respect to their Tertiary igneous activity (Whittow 1974, 1977). This being so obviously the case,the speculation by Page (1997, p. 275) on the different representation of O. limbosperma in thetwo regions seems unlikely to offer an explanation, since in Ireland he suggests poorer drainage inlowland stations might be responsible, and at higher altitudes he believes that there are more base-richsedimentary rocks than in reality is the case.
As generally happens, however, it is easier to criticise others' explanations than it is to suggest abetter alternative! A careful examination of the habitat requirements and tolerances of the speciesmight generate a fresh hypothesis, but we should not ignore the historical factors either, including forinstance, differing human population pressures and land management regimes stretching across manycenturies. The earlier and almost total deforestation that took place in Ireland, for example, couldwell be significant, as might differing grazing, tillage and burning patterns. It is difficult toimagine anything that would affect O. limbosperma on quite the scale necessary to account for theenormous difference in its presence evidenced on the New Atlas British and Irish hectad map(Preston et al. 2002).
Names
The genus name 'Oreopteris' is a combination of the Greek words, 'oreos' meaning 'mountain' and 'pteris'meaning 'fern' (Gilbert-Carter 1964). The specific epithet 'limbosperma' is derived from the Latin'limbatus' meaning 'a border' or 'margin', and 'sperma' meaning 'seed' (or in this case, 'spores'),referring to the characteristic marginal position of the sori on the underside of the frond (Gledhill1985).
As with Phegopteris connectilis (Beech Fern), Oreopteris limbosperma has moved through sixgenera over the years and, partly as a result of this has also been given a long sequence of Englishcommon names including 'Mountain Fern', 'Mountain Buckler Fern', 'Heath Fern', and in reference to thescented glands, 'Sweet Mountain-fern', 'Scented Fern', 'Lemon-scented Fern', 'Hay-scent Fern' and'Tea-scent Fern', the latter two names being local to Cumberland (Step & Jackson 1945).
There does not appear to be any tradition of use of this fern or folklore associated with it as thespecies was not commonly differentiated from the Buckler-ferns, Dryopteris species.
Threats
The minimal populations in Fermanagh could very easily be eliminated by any disturbance of theirstreamside habitats. This is very clearly a species in urgent need of local habitat management toencourage and support its survival.
Native, common and widespread. European temperate, but also present in E Asia and N America.
1860; Smith, T.O.; Ardunshin.
Throughout the year.
Growth form and preferred habitats
A characteristic leathery, entire-leaved, strap-shaped, wintergreen perennial fern shuttleco*ckappearance, Hart's-tongue is a plant of more or less permanently damp, base-rich habitats. The speciesis very widely distributed and common throughout most of Britain and Ireland. P. scolopendrium isessentially a plant of lowland areas, rarely found above an altitude of 185 m (Jermy & Camus 1991;Page 1997). It is most frequent on limestone and scarce or rare in upland areas where peat cover isextensive. Typical habitats include woods, roadside banks, shady rocky slopes and damp walls.
In parts of Britain and Ireland, where limestone pavements occur, P. scolopendrium is particularlyabundant and luxuriant in the cool, moist, shade, shelter and protection of narrow fissures (calledgrykes or grikes) between the limestone blocks (clints). It is also well developed on damp, shadedlimestone cliffs, where it is often overhung by hazel or ash, the frond blades sometimes in suchhabitats reaching 60 cm or more in length.
The plant is sufficiently shade-tolerant to be a typical species of ash and other mixed, damp, deciduouswoods, especially where these occur on the broken, rock strewn ground of talus slopes below cliffs, orin sheltered, narrow valley woodlands. It is also quite commonly found rooted in artificial habitats,such as the mortar of old, damp walls, especially those of bridges, both those over rivers and of thedisused or neglected railway variety, and around old wells.
Growth rate and reproduction
The entire, long, tongue-like, sterile fronds are produced each spring from the basal rhizome and theyusually survive about a year before browning off and dying. The base of the stipe is thickened andserves as a storage organ (Jonsell et al. 2000). Individual plant rosettes are very slow-growing,and they generally take between 2-5 years to produce their first fertile frond, which in appearance islike the sterile fronds. The fern rosette is long-lived, individuals perhaps surviving for severaldecades, if not longer. The blades of large fertile fronds may bear up to 60 lines of paired sori eitherside of the midrib, and thus have the ability to produce vast numbers of spores (Page 1997).
Fermanagh occurrence
Hart's-tongue is very widely distributed and common throughout Fermanagh being present in 415 tetrads,78.6% of those in the VC. It is most frequently found on the limestone, and is scarcest on uplands withextensive peat.
Irish and British occurrence
Although P. scolopendrium is very widespread in Ireland, England, Wales and lowland Scotland, itis most abundant in W and SW parts of these islands, where conditions of general high humidity and along, mild growing season are conducive to its growth and individual frond survival (Jermy et al.1978; Wardlaw & Leonard 2005).
European occurrence
In Europe, P. scolopendrium has a widespread and more or less continuous southern sub-Atlanticdistribution. It extends in a scattered manner from extreme SW Norway to the Iberian peninsula, NWAfrica, Madeira, Canaries and the Azores, then eastwards to Turkey and the Caspian Sea, but is onlyabundant in the British Isles and on the seaboard of W Europe south to the Pyrenees (Jalas &Suominen 1972, Map 102; Page 1997).
World occurrence
Elsewhere in the world, Hart's-tongue has a very disjunct distribution, with a different chromosome countin eastern N America (diploid whereas the form in Britain and Ireland is tetraploid). What may or maynot be the same diploid taxon (or varieties allied to it) occurs in N Japan and Mexico. It is probablybest to consider these three widely separated taxa as at best subspecies of P. scolopendrium(Hultén 1962, Map 147; Page 1997). In Germany Denmark and Switzerland, and probably in other sites closeto its European distributional limits, the distribution of P. scolopendrium has contracted in thelast 50 years and it has become locally extinct in areas where previously it was considered native(Hultén 1962; Welten & Sutter 1982; Jonsell et al. 2000).
The predominance of P. scolopendrium in mild, western, oceanic areas of Europe, where the climateis ameliorated throughout the year by the warming influence of the Gulf Stream, is undoubtedlyassociated with this slow-growing fern's requirement for a long growing season and constant highhumidity to prevent desiccation. In its more heavily shaded habitats, much of the plant's photosyntheticgain probably takes place during the autumn and winter when higher levels of illumination operate afterdeciduous leaves fall (Page 1997).
Cultivated forms
P. scolopendrium is genetically very variable and easy to cultivate, so it is not surprising thatit has given rise to many cultivated variants. During the Victorian 'Fern Craze', Lowe's FernsBritish and exotic listed 65 cultivars of this species (Lowe 1865). The modern RoyalHorticultural Society Dictionary Index of Garden Plants lists a mere 16 cultivars, most of whichfeature fronds with frilled margins such as var. 'crispum' (Griffiths 1994). Jones (1987) does mentionthat hundreds of cultivars are known, a rough estimate also given by Step & Jackson (1945).
Locally, an unusual form of the plant with fronds that fork at their tips has been recorded on limestonepavement on Heron Island, just off Tully Castle on the shore of Lower Lough Erne.
Names
The name 'Phyllitis' is the ancient name given to this fern by Dioscorides, which it has retained eversince, although occasionally it wanders temporarily into genera such as Asplenium andScolopendrium (Gilbert-Carter 1964). The large number of paired sori either side of the midrib onthe fertile frond gave rise to the Latin species epithet 'scolopendrium', a name given by the classicalherbalist Dioscorides, that compares the plentiful sori to a millipede's leg count (Gilbert-Carter1964).
It is a rather sad fact of life that fern Latinised botanical names have in the last 40 or 50 years beenmuch less stable than their English common names. The most frequent English common name of thisdistinctive, entire-leaved fern is 'Hart's-tongue', which first appeared in print as the Middle English'hertes tongue', in the Grete Herball (Anonymous 1526, see mention of its origin underCeterach officinarum) (Ryden 1984). The name obviously refers to the tongue-like shape ofthe frond blade, and is a translation of the apothecaries' Medieval Latin, 'Lingua cervina' (Gerard1633; Prior 1879). The tongue name also appears in several variants in parts of the British Isles as,'Adder's Tongue', 'Fox-tongue', 'Horse-tongue' and 'Lamb's-tongue'. Additional names suggesting bladelength are 'Long-leaf', 'Finger-fern', 'Seaweed-fern' and 'Snake-leaves'.
Two unusual local names refer to other specific physical features; 'Button-hole' from E. Sussex, ratheraptly refers to the appearance of the young sorus on the back of the frond, while 'Christ's Hair' or'God's Hair', names that originate from Guernsey, Devon and elsewhere. The ‘hair’ in the name is said torefer to the black fibrous bundles (one or two) present in the vascular tissues of the stipe. Both thename and the explanation of its supposed origin are provided by Britten & Holland naming Mr W. G.Piper as the source of the botanical information. The associated story, as quoted in Vickery (1995), ischarming and goes as follows: "The fern was once the pillow for the Son of Man, when He had nowhereto lay His head. In return for this service, He left two hairs of His most blessed and dear head, whichthe plant treasures in her ripe stem, as His legacy – two auburn hairs which children find andshow."
Herbal medicinal uses
The name 'Burntweed' originates from Westmeath, Wales and the Scottish Highlands, and refers to the localuse of the frond to manufacture an ointment for the treatment of burns, scalds and piles (Britten &Holland 1886; Grieve 1931). Other medicinal uses in herbalism were as an astringent treatment fordiarrhoea and dysentery, and as a remedy for removing obstructions from the liver and spleen (Grieve1931).
Threats
None.
Native, frequent. European temperate, but also present as a disjunct rarity in C Asia and N America.
1860; Smith, Rev Prof R.W.; Florencecourt.
Throughout the year.
Growth form and preferred habitats
Black Spleenwort is a very variable evergreen perennial of well-drained, somewhat base-enriched rockyplaces, including limestone screes and dolomitised sandstone scarps, especially in more western, coastaland lowland parts of Britain and Ireland. The plant typically produces a loose tuft of triangular frondsfrom a short, creeping rhizome. It favours sheltered, lightly shaded situations where competition isreduced for a variety of reasons. In very sheltered, more deeply shaded woodlands and hedgebanks, frondlength can occasionally reach 50 cm. Reduced competition often involves fern habitats with little soilsuch as, for instance, tree-shaded rock crevices on cliffs and on old quarry faces. Here, frondssometimes reach up to 30 cm in length, while in more exposed sites with similar strictly limited soilresources they are always very much smaller, often only 10 cm in length or less.
Black Spleenwort also occurs, though not quite so abundantly, on limestone cliffs, and together with themodified sandstone rock mentioned, this behaviour reflects this species known soil nutrient requirementfor at least a trace of base-rich elements, most likely calcium or magnesium (Jermy et al. 1978;Jermy & Camus 1991). This minimal but necessary base requirement excludes A. adiantum-nigrumfrom acidic siliceous rocks, such as quartzite, granite or normal sandstone (Webb & Scannell 1983),but it is not the case, as is sometimes claimed, that this species avoids limestone (Hultén 1962;Sinker et al. 1985), at least not in Fermanagh, nor in the Burren, Co Clare (H9).
Apart from hedgebanks, the fern occasionally occurs in other artificial habitats, such as on oldlime-mortared walls and bridges in lowland areas.
Variation and taxonomy
In recent years, A. adiantum-nigrum has been recognised as having two subspecies: the common andwidespread subsp. adiantum-nigrum, and a second one, to some extent associated with, but perhapsnot totally confined to ultra-basic (especially serpentine) rock, subsp. corrunense Christ (Page1997). The latter was previously confused with the continental European species, A. cuneifolium,and its relationship with this species remains the subject of continuing research. Serpentine and otherultrabasic (or ultramafic) rocks often supply toxic or near-toxic levels of heavy metals such as nickel,cobalt and chromium. The derived soils are extremely infertile, which greatly reduces plant vigour andcompetition (Brooks 1987). Within the Fermanagh western plateau, there is a crescent-shaped outcrop ofintrusive dolerite and basalt which just might support subsp. corrunense, and while it has notyet been discovered, it should certainly be looked out for (Woodland et al. 1977).
Fermanagh occurrence
Black Spleenwort is a characteristic species of the dolomitised, somewhat base-enriched, sandstone scarpsin the more upland SW of Co Fermanagh that is referred to as the Western Plateau. It has been recordedin 73 tetrads in the VC, representing 13.8% of the total area. The occurrence is most unevenly spreadhowever, the fern being predominantly confined to the western half of the county. The most elevated sitein Fermanagh for Black Spleenwort is at Cuilcagh Gap, around 550 m. Otherwise, the generally lowlandpattern that this species displays elsewhere in the British Isles is reflected in Fermanagh (Page 1997).It also occurs, but only very occasionally, on old lime-mortared walls and bridges in the lowlands.Local examples of this occur at Tubbrid churchyard and on the old bridge at Roogagh River.
British and Irish occurrence
A. adiantum-nigrum s.l. rarely occurs in large populations, but is widely distributedthroughout these isles, being most frequent in the S and W, and especially so in mild, coastal districtswhere high levels of humidity and illumination are the norm (Page 1997; Wardlaw & Leonard 2005).
European and world occurrence
A. adiantum-nigrum s.l. is widespread in Europe north to 63o on the W coast of Norway,and south to the Peloponnese, NW Africa, the Caucasus, N Iran, C Asian mountains, SW North America andis also found on some isolated tropical mountains and islands (Jalas & Suominen 1972; Jonsell etal. 2000). The northern hemisphere map of the species (Hultén (1962, Map 142), shows that itabsent from vast tracts of temperate and boreal Asia and N America, to such an extent that to thisauthor’s mind it does not warrant inclusion in an atlas of circumpolar plants. The later publication ofA. adiantum-nigrum s.l. world distribution (Hultén & Fries 1986, Map 44), again highlightshow very far removed the species is from being circumpolar. Preston and Hill (1997) classified the fernas European temperate, at the same time noting that it has an additional very restricted presence in NAmerica, occurs in C Asia and also in widely disjunct parts of the tropics and the Southern Hemisphere,including Australia and Hawaii (Hultén & Fries 1986).
Names
'Asplenium' is derived from the Greek 'a' meaning 'not' and 'splen', 'splene' or 'splenon' referring tothe spleen, alluding to the supposed medicinal properties of the fern genus. The herbal medicinal use isalso invoked by the English common name applied to the genus, ‘Spleenwort’ (Hyam & Pankhurst 1995).The specific epithet is ‘adiantum-nigrum’. ‘Adiantum’ is Greek, ‘a’ meaning ‘not’ and ‘diantos’,moistened, and thus the combination ‘adiantos’ means ‘dry’, ‘not wetting’ or ‘unwetted’, referring tothe fact that fronds of the fern genus ‘Adiantum’ remain unwetted under water. ‘Nigrum’ means ‘black’,so ‘adiantum-nigrum’ translates literally as ‘dry black’ or ‘unwetted black’. However, as the fernAdiantum capillus-veneris has the English common name ‘Maidenhair Fern’, Aspleniumadiantum-nigrum in the past was given the book name ‘Black Maidenhair Spleenwort’. Since there alreadyis a fern with the given (book) name ‘Maidenhair Spleenwort’ (Asplenium trichomanes), to avoidconfusion A. adiantum-nigrum is most usually referred to as ‘Black Spleenwort’ (Step &Jackson 1945).
Although A. adiantum-nigrum is very widely distributed in both Britain and Ireland, it does notappear ever to have had a genuine folk-name or English common name, only the invented, given, book nameslisted above. Lyte (1578), in his ‘Niewe Herball’ mentions this fern under the names ‘BlackOak-fern’ and ‘Petty-fern’, but in reporting this information Step and Jackson (1945, p. 51), expresstheir doubt that these names would have been in use among the people at that early date.
Herbal medicinal uses
Black Spleenwort is said by herbalists to have similar medicinal virtues to other Maidenhairs, “adecoction of it relieving a troublesome cough and proving also a good hair wash. Dosage of infusion: 3tablespoonfuls” (Grieve 1931, p. 303). Allen and Hatfield (2004) reported that in Ireland a cough cureknown as ‘maidenhair’ was once popular among country people in Londonderry (David Moore unpublishedreport 1834-5). These latter authors assumed that this referred to Maidenhair Spleenwort (A.trichomanes), but it might equally well have been A. adiantum-nigrum that was used.Adiantum capillus-veneris does not come into the question on this matter since it was always arare plant of very restricted distribution.
Threats
None.
Native, very rare. Suboceanic southern-temperate.
June 1979; Northridge, R.H.; scarp SW of Lough Achork.
Throughout the year.
Growth form and preferred habitats
Except at the western extremities of its markedly Atlantic distribution, A. marinum is restrictedto a very narrow zone, seldom more than 20-30 m above sea level, where the winter air temperature isameliorated by warm sea spray from the Gulf Stream. The usual habitat requirements of this glossy,evergreen, singly-pinnate, perennial fern are a cool, moist crevice, sheltered from full sun in summerand entirely frost-free in winter (Page 1997).
British and Irish occurrence
Sea Spleenwort is entirely coastal throughout its British and Irish range. In Britain it stretches almostcontinuously from the Isle of Wight on the middle of the south coast of England, westwards to Land's Endand up most of the west coast through Wales to Shetland. On the east coast of Britain it is againrepresented from the far north of Scotland as far south as Scarborough, although here it is lesscontinuous and definitely more thinly scattered.
In Ireland, the New Atlas hectad map shows A. marinum is again very well represented onalmost all of the S & W coasts, but it is absent from a few stretches of the E coast, mainly fromDublin Bay to Wexford (New Atlas).
Fermanagh occurrence
In view of this information, the solitary Fermanagh site in Lough Navar Forest Park is extremelyabnormal, being 17 km inland from the tidal estuary at Ballyshannon and situated at an altitude of 210m. Originally the plant occurred at two places on the same set of N-facing sandstone cliffs, there beingabout 20 plants at one site and about 50 at the other (Northridge et al. 1988). On a visit inJuly 2002, there was a very noticeable decline in the smaller more accessible population, the result ofa*ggressive competition from young Hedera helix (Ivy) stems also present in the crevices.Subsequent visits up to September 2010 showed three plants surviving at the smaller patch; the largerpatch was thriving. In addition, two inaccessible plants growing higher up the cliffs were identifiedthrough binoculars!
Ecology
The Fermanagh sites being so remote from the sea, it cannot be that A. marinum has an absoluterequirement for sodium chloride. The red sandstone of the Lough Navar cliffs has been dolomitized by thevertical percolation of waters rich in calcium and magnesium sulphates, so that mineral replacement hasoccurred and the rock has become base-enriched. At both Fermanagh sites, A. marinum grows wherethe top of the cliff overhangs the base to a considerable degree, providing a measure of frostprotection, although one would think that this would hardly be sufficient to entirely avoid freezingtemperatures at this altitude. Many of the plants grow in a shaded pocket of the cliff while other smallindividuals grow along fault line crevices. A. ruta-muraria (Wall-rue) grows close to some of theA. marinum plants, but many have no nearby competitors.
The winter-green, coarsely divided, leathery, glossy fronds and stiff, dark purple-brown stipes arereadily recognised. It is the opinion of the current author that the thick cuticle of the fleshy frondenables A. marinum to withstand the extreme dryness of this site. A visit to the site in July2002 found that Ivy had invaded some of the cliff crevices and rendered A. marinum locallyextinct.
Fortunately another cliff outcrop further west maintains a vigorous population of the species at itsbase, and the Ivy stems appear to have grown up and beyond the fern, so that they are unlikely tocompete directly for space and light.
Reproduction and possible hybridization
The fronds produce spores abundantly from August onwards and overwinter before decaying the followingspring as new fronds are produced (Page 1997). The breakdown in the Fermanagh site of the usualecological barriers between A. marinum and A. ruta-muraria suggests the possibility of ahybrid, but none has yet been found here. Page (1997) suggests the reason for this failure to hybridizewith any other native northern species of Asplenium is that A. marinum's phylogeneticaffinities appear to lie with an extensive group of species characteristic of wet forests in the tropicsand sub-tropics.
European and world occurrence
The wider distribution of A. marinum is Atlantic-Macaronesian (ie along the W and SW coasts ofEurope, plus the island groups of Madeira, Canaries and Azores (Jalas & Suominen 1972, Map 78; Page1997, p. 72). This, together with its essentially frost-free ecological requirement, tends to supportthe idea that it associates more closely with Asplenium species of warmer climates than thoseencountered at present in the British Isles. In their phytogeographical survey of British and Irishplants, Preston and Hill (1997) classified A. marinum as Suboceanic southern-temperate, althoughthey noted that it also occurs in the Southern hemisphere. Hultén and Fries (1986, Map 40) indicate thatthis refers to presence on the Cape Verde Islands and St. Helena.
Trends in Britain and Ireland
Page (1997) points out that in general A. marinum is much less frequent and less luxuriant in manyof its current British and Irish habitats than it was in the 19th century. Past ravages of ferncollection, together with slowly changing factors, for example, pollution and contamination of shores,and natural changes in climate, that is, global warming, with respect to which this species appears tobe in an extremely delicate balance, are almost certainly involved. Thus this species is under threat,and should on no account be collected.
Names
'Asplenium' is derived from the Greek 'a' meaning 'not' and 'splen', 'splene' or 'splenon' referring tothe spleen, alluding to the supposed medicinal properties of the fern genus. The herbal medicinal use isalso invoked by the English common name applied to the genus, ‘Spleenwort’ (Hyam & Pankhurst 1995).The Latin specific epithet ‘marinum’ means ‘sea’, an obvious reference to the preferred habitat andhence the English common name ‘Sea Spleenwort’.
Like A. adiantum-nigrum, while A. marinum is widely distributed along coastal sites in bothBritain and Ireland, it does not appear ever to have had a genuine folk-name or English common name,only the invented book name, ‘Sea Spleenwort’. Presumably it is too uncommon to have merited folkinterest, and I cannot locate any mention of folk use of the plant.
Threats
Part of one of the Fermanagh sites is being overgrown by Hedera helix (Ivy).
Native, frequent, widespread and locally abundant. Circumpolar southern-temperate.
1860; Smith, Rev Prof R.W.; Florencecourt.
Throughout the year.
Growth form and preferred habitats
Maidenhair Spleenwort is a small, distinctive, evergreen perennial species that colonises crevices onupland limestone outcrops and cliffs. In lowland areas it frequently occurs on the mortar in walls. Itis most abundant and luxuriant when the habitat offers high humidity and it becomes distinctly stuntedin drier, more exposed sites.
Taxonomy
Asplenium trichomanes really consists of a complex polyploid series of forms, within whicha simplified account segregates three ecologically and morphologically recognisable subspecies inBritain and Ireland. All of the Fermanagh records almost certainly refer to by far the most common ofthe three forms, the tetraploid A. trichomanes subsp.quadrivalens D.E. Mey. (but see the separate account below). Essentially this is a calcicolesubspecies, but to a surprising degree it can tolerate habitats with soils having very little calciumpresent (Jermy & Camus 1991; Jonsell et al. 2000).
Fermanagh occurrence
In Fermanagh, this fern species has been recorded in 181 tetrads, 34.3% of those in the VC. Eightscattered tetrads have pre-1976 records only. It is widespread throughout the county, but is especiallyfrequent in crevices on upland limestone outcrops and cliffs. It also occurs less commonly on sandstonescarps in the Lough Navar area which have been dolomitized by seepage of water rich in both calcium andmagnesium carbonates derived from overlying base-rich mica-schists.
In lowland areas of Fermanagh, A. trichomanes is restricted to man-made habitats such as thelime-rich mortar in old walls, bridges, basem*nt areas around older houses, walled gardens and the like.
British and Irish occurrence
A. trichomanes subsp. quadrivalens is common throughout Britainand Ireland, but it is perhaps slightly less frequent in Ireland than in Britain (Jermy & Camus1991). Until the publication of the New Atlas in 2002 there were insufficient data to permit thesubspecies to be mapped separately. The New Atlas maps and those in the New Fern Atlas(Wardlaw & Leonard 2005), both suggest that only subsp.quadrivalens occurs in Ireland. The other two subspecies are very much less common in Britain,and subsp. pachyrachis (H. Christ) Lovis & Reichst. is recorded from just eight hectads inEngland.
European and world occurrence
The collective species, A. trichomanes is common and widespread in W and C Europe, thinning out toboth north and south although reaching well within the Arctic Circle. However, it is currently decliningin Scandinavian countries (Jalas & Suominen 1972, Map 81; Jonsell et al. 2000). In the widestsense, A. trichomanes is a circumpolar southern-temperate species (Hultén 1962, Map 130; Preston& Hill 1997), but is also very well distributed around the southern hemisphere, being present in SAfrica, C and S America, New Guinea, S Australia and New Zealand (Hultén 1962, p. 138; Hultén &Fries 1986, Map 41).
Names
'Asplenium' is derived from the Greek 'a' meaning 'not' and 'splen', 'splene' or 'splenon' referring tothe spleen, alluding to the supposed medicinal properties of the fern genus. The herbal medicinal use isalso invoked by the English common name applied to the genus, ‘Spleenwort’ (Hyam & Pankhurst 1995).
The Latin specific epithet ‘trichomanes’ is the genus name given to Maidenhair Spleenwort byTheophrastus, the ancient Roman doctor. ‘Trichomanes’ is Greek and it refers to a thin hair or a bristle(Johnson & Smith 1946). After the green pinnae die and drop off the black or deep brown stipe of thefrond, the bare remainder adds to the short, dense, wiry tuft of old stipes attached to the plant.Certainly, no female would be flattered to have the hair on her head compared to the wiry tuft of aMaidenhair Spleenwort plant. It follows that ‘Maidenhair’ must refer to hair located elsewhere on thebody, and it is not too difficult to imagine where.
Uses
A tea made with the fronds was described as ‘ sweet, mucilaginous and expectorant’ and was used in herbalmedicine to treat lung disorders. It was also considered a laxative (Grieve 1931, page 303).
Threats
None.
Native, probably frequent but under-recorded at subspecific level. Circumpolar southern-temperate.
1974; Hackney, P.; Boho Caves.
July and August.
Despite the Asplenium trichomanes maps in the New Atlas plotting only subsp.quadrivalens for Ireland, the diploid calcifuge subsp. trichomanes has been recorded fromnine VCs, including in N Ireland, Cos Armagh and Down (H37 & H38). Adjacent VCs to Fermanagh wheresubsp. trichomanes has also been recorded are Co Sligo and both E & W Donegal (H28, H34 andH35). Subsp. pachyrachis (H. Christ) Lovis & Reichst. has not yet been discovered anywhere onthe island of Ireland (Matthew Jebb, pers. comm., 2010).
Having said all that, it is believed that the vast majority of the plants of A. trichomanes inIreland belong to subsp. quadrivalens, although so far there are only eleven records of thissubspecies from ten tetrads in Fermanagh, nine of them the work of Paul Hackney. Vouchers exist forHackney's records in BEL.
While subsp. quadrivalens almost always frequents limestone terrain, the main habitat of subsp.trichomanes is on hard, acidic volcanic or metamorphic rocks, making it unlikely to occur inFermanagh, although it is possible that the third taxon, subsp. pachyrachis could occur with uson steep limestone rock faces. Only time and plenty of further careful recording will tell whether ornot Ireland supports all three A. trichomanes subspecies, and, if it does, their comparativefrequency.
Native, rare. Circumpolar boreo-temperate.
1860; Smith, Rev Prof R.W.; Florencecourt area.
Throughout the year.
Growth form and preferred habitats
This little evergreen perennial fern is generally described as a calcicole species, ie one which islime-tolerant and/or preferring or requiring base-rich conditions. The frond has a delicate texture andthe green rachis is extremely distinctive. A. viride requires much higher levels of humidity thanA. trichomanes (Maidenhair Spleenwort) and will grow in conditions of considerably deeper shade.Unlike A. trichomanes and its subspecies, the rachis of A. viride does not shed itspinnae, but rather when it withers the whole frond shrivels and only a very short persistent brown baseremains (Page 1997; Jonsell et al. 2000).
Normally the species does not tolerate drought or high summer temperatures, and while in natural rockhabitats it regularly associates with A. trichomanes s.l., A. ruta-muraria (Wall-rue) andCystopteris fragilis (Brittle Bladder-fern), unlike these three ferns it is generally absent fromwalls. All Fermanagh records for this species are from naturally occurring rock habitats (ie clifffaces, crevices, ledges and limestone grykes and swallow holes), but in some areas of Britain andIreland A. viride has also been rarely found growing on lime-mortared walls (Jermy & Camus1991).
Fermanagh occurrence
In Fermanagh, Green Spleenwort is rare, never abundant and has been recorded in a total of just eighttetrads. Although a rare species, it is most abundant in several of the deep limestone swallow holesnear Legacurragh above Florencecourt in the south of the county, where it grows as tufts on the verticalfaces of the rock. It also occurs sparingly on both the dolomitized Upper Visean sandstone and shale,and the dolerite and basalt scarps in the Lough Navar area (Woodland et al. 1977). As the tetraddistribution map indicates, in addition to the rather isolated Legacurragh site, A. viride alsohas a second outlier on an isolated scarp near Lough Alaban in Tullyloughan Td, Carrigan Forest. Thefern population here consists of just six plants. The range of substrates mentioned demonstrates thatwhile Green Spleenwort always occurs on soils derived from base-rich rock, it does not necessarilyrequire a high calcium content in the soil it occupies (Jermy & Camus 1991).
British and Irish occurrence
In Britain, Green Spleenwort has a distinctly N, W and upland distribution, with the greatestconcentration of records occurring N of a line from Morecambe to Bridlington. In Ireland, by comparison,it is very much less frequent overall, and it is distinctly western and montane, being confined tomoist, shaded habitats in areas with low summer temperatures (Jermy et al. 1978).
European occurrence
In Europe, A. viride is widespread on calcareous and other base-rich rocks, but further south itis found mainly in the mountains, while in Fenno-Scandinavia it has a north-western and Atlanticconcentration (Jalas & Suominen 1972, Map 83; Jonsell et al. 2000). We could thus describeits European range as being rather similar to that of an Arctic-alpine or Arctic-montane species, butwith an additional more lowland boreal occurrence (Page 1997). In Fenno-Scandinavia, A. viridehas been described as being particularly frequent on ultra-basic rock (eg serpentine), and under suchunusual and generally toxic soil conditions in those latitudes, it can also be found in much drier,sun-exposed sites than is the case elsewhere (Jonsell et al. 2000).
World occurrence
The world map features the species with an uneven distribution in boreal northern and eastern Europe, andthen a rather discontinuous, disjunct circumpolar occurrence through NW Africa, Turkey, the Caucasus,the Urals, mountains of S Siberia and C Asia, W Himalaya, Japan, W & E North America, S Greenlandand S Iceland (Hultén 1962, Map 92; Hultén & Fries 1986, Map 43; Jonsell et al. 2000).
Names
'Asplenium' is derived from the Greek 'a' meaning 'not' and 'splen', 'splene' or 'splenon' referring tothe spleen, alluding to the supposed medicinal properties of the fern genus. The herbal medicinal use isalso invoked by the English common name applied to the genus, ‘Spleenwort’ (Hyam & Pankhurst 1995).The Latin specific epithet ‘viride’ means either ‘youthful’, or rather better, ‘fresh green’. TheEnglish common name is a simple translation of the Scientific name, ‘Green Spleenwort’.
Threats
None.
Native, common and widespread. Circumpolar temperate.
1860; Smith, Rev Prof R.W.; Ardunshin Bridge, the Colebrooke River.
Throughout the year.
Growth form and preferred habitats
This small tufted evergreen perennial is locally abundant on natural habitats such as narrow crevices inrocks and cliff faces, almost exclusively growing on limestones or other forms of base-rich rock.Otherwise this lime-tolerant, strongly calcicolous species is almost exclusively a plant of thelime-rich mortar of old walls and bridges, and it is particularly abundant on rural examples of theseman-made habitats. It develops most luxuriantly in half-shaded, damp sites.
Page (1997) rather tentatively suggests that A. ruta-muraria is, "a fast-growing and perhapsrather short-lived spleenwort. In most situations, frequent re-establishment occurs ...". Thereappears to be very little evidence of dead plants and turnover taking place on walls in Fermanagh, andthe reproductive behaviour of Wall-rue requires and deserves further study.
Fermanagh occurrence
Widespread in Fermanagh, this small, tufted, evergreen fern has been recorded in 153 tetrads, 29.0% ofthose in the VC. Nine of the tetrads have pre-1976 records only. It is very common throughout thecounty, mainly on calcareous rocks, including in shaded crevices in the limestone pavement aboveFlorencecourt and around Knockmore. However, it also grows on the dolomitized sandstone scarps in themore upland, rather exposed and somewhat wetter Western Plateau. Some stretches of the red sandstonescarps here have been chemically altered by long-term percolation of calcium- and magnesium-rich waterderived from overlying rocks (ie., they have become dolmatized).
British and Irish occurrence
Wall-rue is very common throughout Britain and Ireland. However, being calcicole and sensitive toatmospheric pollution, A. ruta-muraria is much less frequent or scarce in more urban andindustrial areas of Britain and Ireland, and in regions where soils are predominantly acidic. Thisincludes substrates derived from siliceous igneous and metamorphic rocks that make up the structure of NScotland and N & W Ireland. Rarity also applies in oceanic areas of very high rainfall, wherestrongly acidic peaty soils develop, irrespective of the geochemistry of the underlying rock (Jermyet al. 1978; Jermy & Camus 1991; New Atlas).
European and world occurrence
This fern is widespread across W & C Europe and extends north of the Arctic Circle (Jalas &Suominen 1972). The species has its headquarters in more central areas of Europe, and geographically isconsidered a continental species, albeit with a particularly extensive range near to the westernAtlantic coast (Page 1988, p. 86).
The circumpolar distribution of A. ruta-muraria is decidedly disjunct, the species being entirelyabsent from W and C regions of N America, and poorly represented with very few, widely spaced sites in EAsia (Hultén 1962, Map 156).
Hultén (1962) described the species as being, "fairly variable, at least 15 varieties having beendescribed from C and S Europe, Morocco and China". Two subspecies of differing chromosome count arerecognised in Europe, but only the tetraploid, subsp. ruta-muraria, is recorded in Britain andIreland (Page 1997). A counterpart, confined to eastern N America, has been described as a distinctspecies, A. cryptolepis, but Hultén (1962) preferred to consider this plant a further subspeciesof A. ruta-muraria.
Extremely rare hybrids
Although Wall-rue commonly occurs immediately adjacent to A. trichomanes subsp. quadrivalens (Maidenhair Spleenwort) in both naturalhabitats and on walls, the hybrid between them, A. × clermontiae(Lady Clermont's Spleenwort), has only once occurred in the British Isles – in 1863 on a mortared wallin Co Louth, Republic of Ireland (H31). It has not been found in Co Down (H38) as erroneously reportedby Stace (1975) and repeated by Page (1997) (Hackney et al. 1992).
A. ruta-muraria does also form another very rare hybrid with A. septentrionale (A. × murbeckii), which has never been recorded anywhere in Ireland, buthas been found three times in Britain, and has also been reported in 13 Scandinavian provinces (Stace1997; Jonsell et al. 2000).
Names and Uses
'Asplenium' is derived from the Greek 'a' meaning 'not' and 'splen', 'splene' or 'splenon' referring tothe spleen, alluding to the supposed medicinal properties of the fern genus. The herbal medicinal use isalso invoked by the English common name applied to the genus, ‘Spleenwort’ (Hyam & Pankhurst 1995).The Latin specific epithet 'ruta-muraria' translates as 'Rue of the wall' or 'Wall-rue'. Folkloresuggested that wearing A. ruta-muraria could protect the individual from witchcraft. Theunrelated flowering plant, Rue (Ruta graveolens) with bluish-green leaves, was a symbol of sorrowand repentance, and as such could be used either to bless or curse, help or harm. In the absence offlowering Rue, the fern, Wall-rue, could be used as a substitute (Vickery 1995).
Wall-rue had the alternative and rather mysterious English common name, 'Tentwort' (Page 1988). Theoriginal version of this name was 'Taintwort', from its use as a remedy for 'the taint', better known tous as rickets (Step & Jackson 1945). Other names applied to the fern include 'White Maidenhair',from the fronds sometimes taking on a blue-green appearance (Gerard 1597), and 'Stone Rue' (Lyte 1578).Herbalists considered it a good remedy for coughs and ruptures in children. It was also thought toprevent hair falling out, and was used for treating shortness of breath, yellow jaundice, diseases ofthe spleen, stopping of the urine, and to help break up kidney stones. Grieve (1931, page 303) detailsthese and many other herbal medicinal uses of this fern.
Threats
None.
Native, occasional or fairly frequent. Submediterranean-subatlantic.
1844; Cole, Hon J.L.; Florencecourt.
Throughout the year.
Growth form and preferred habitats
This distinctive, definitely calcicole, evergreen perennial typically occurs either in crevices in cliffsand screes of limestone or other basic rocks, or in the artificial habitat of old lime-mortared walls.
The prevailing damp and relatively mild climate of Fermanagh allows the evergreen fronds to grow most ofthe year round, and since our high precipitation is rather evenly distributed, checks on C.officinarum growth due to summer drought are few and far between. In the rare event of prolongeddry weather, Rustyback may wilt so severely and curl up so that it appears to have suffered terminally.However, the fronds and their thick backing of overlapping scales have amazing powers of recovery, andthus individuals are very persistent. The fronds can even develop a second flush of sporing sori aftersuch an event (Jermy & Camus 1991). While the fern often appears to be rooted in very dry wall orrock crevices, in reality its roots are frequently embedded in cushions of water-retaining moss or inpockets of damp, black humus derived from dead moss.
Fermanagh occurrence
Only on the limestone cliffs and screes near Boho, at Carrickreagh and in disused quarries at Goladoonear Ederny has Rustyback been found on natural rock surfaces in our survey area. Lime-mortared wallsare not all that frequent in Fermanagh, so the fern's real local stronghold is the mortar on old bridgeparapets. C. officinarum has been recorded in 69 Fermanagh tetrads, 13.1% of those in the VC.More than half the records are from old bridges scattered throughout the survey area. As the tetrad mapshows, it is widely spread across the county, but eleven tetrads have only pre-1975 records, suggestingthere has been some local decline of this species.
British and Irish occurrence
C. officinarum is widely distributed throughout the whole of Ireland, but is probably most commonto the S and W of the island. The latter comment could also be applied to Britain, but here it is evenmore south westerly, being frequent only in SW England and Wales and Cumbria, scattered in the Penninesand SW Scotland, but very rare north of the Grampian Highlands and scarce east of the Pennines inEngland (Jermy et al. 1978; Jermy & Camus 1991).
Europoean and world occurrence
Beyond Britain & Ireland, C. officinarum is widely distributed in SW Europe and theMediterranean basin where it has its main centre of occurrence, but it also extends eastwards to theCrimea, the Caucasus and C Asia. It also stretches southwest to the Cape Verde Islands (Jalas &Suominen 1972, Map 99; Hultén & Fries 1986, Map 47; Page 1997). The species contracted during thelast century and became locally extinct in several areas, mainly along the easterly margins of itsnatural range in Britain, Germany, Holland, Switzerland, and no doubt elsewhere in mainland Europe(Jalas & Suominen 1972).
Names
Conflicting derivations are given for the genus name 'Ceterach' in the literature. Several sources (egGledhill 1985) suggest it is an Arabic name for an unspecified fern, while Gilbert-Carter (1964)believes it is derived from a German word meaning 'itchy', referring to the covering of scales which aresaid to resemble a cutaneous skin eruption! Step & Jackson (1945) refer back to Turner (1548) Thenames of herbes (which is always a good place to start!). Turner refers under the name'Asplenum' or 'Asplenium' to the apothecary's 'Citterache'. Step & Jackson (1945) also regard'Ceterach' (or, alternatively, 'Chetherak'), as possibly being of Arabic origin, apothecaries"using it as a medicine for troubles of the spleen and liver".
The species name 'officinarum' as always is a reference to the apothecary's shop, the 'officina', wheremedicinal plants were kept, and thus infers that the species was used in medicine (Gilbert-Carter 1964).
In his Herbal of 1568, Turner is quoted as saying that he had heard of no English name of thefern, although the ancient name 'Asplenum' and the French(!) 'Ceterache' were familiar to him. Turnerthen suggested several English names himself: "It may well be called in English 'Ceterache' or'Miltwaste', or 'Finger ferne', because it is no longer than a manne's finger; or 'Scale ferne', becauseit is all full of scales on the inner syde." (Turner 1568; quoted in Step & Jackson 1945, p.57). ‘Ceterach’ or ‘Chetherak’ is said to be of Arabic origin, probably handed down by apothecaries whoused it as a medicine for troubles of the spleen and liver (Step & Jackson 1945).
The name 'Miltwaste' refers to the spleen (the 'milt'), the suggestion being that animals eating the fernrootstock (although considering how and where the plant grows, this would in reality seem almostimpossible for them to achieve), were said to suffer wastage of their spleen and liver (Step &Jackson 1945). The origin of other names like 'Rustyback', 'Brown-back' and 'Scale-fern' are obvious toanyone examining the plant. 'Stone-fern' reflects the habitat in which it grows.
The name 'Saxifrage' (ie 'Stone-breaker') has also been applied to this fern. This name first appears in1526 in a work printed by Peter Treveris in Southwark, London entitled The grete herball, ananonymous early English translation of an anonymous French work published in Paris c 1498, knownunder the title Le grant herbier en françoys (Henrey 1975, Volume 1, page 6 & pp 15-22). Inthis work, 'Saxifrage' is simply illustrated as a fern, which was subsequently identified by Britten andHolland (1886) as Ceterach officinarum. This name could refer to any rock or wall growing fern,including several members of the genus Asplenium. C. officinarum was previously calledAsplenium ceterach L., and occasionally it reverts to this name (eg Jermy & Camus 1991). Theidea behind the name 'Saxifrage', whatever plant it is applied to, is that they assist thedisintegration of the rocks on which they grow. Another English name is simply 'Common Spleenwort'(Grieve 1931).
Uses
The appearance of the frond was considered spleen-like, and following the ancient ‘Doctrine ofSignatures’ it could therefore be used to treat ailments of the spleen and other viscera. For instance,an infusion of the fronds was perscribed by herbalists to treat gravel in the liver and spleen (Grieve1931, pages 302-3).
Threats
Re-pointing of bridges, or replacement of old bridges by new ones.
Introduction, neophyte, garden escape or planted, very rare. Circumpolar boreal-montane, but absent as anative from most of W Europe.
9 April 2005; Northridge, R.H. & Northridge, Mrs H.J.; damp trackside on edge of woodland,Knocknabrass Td, Crom.
Fermanagh occurrence
A patch about five metres square of this large, distinctive deciduous fern with its tuft of fronds givingit an attractive shuttleco*ck appearance, was noticed growing, well established in damp ground below anuntarred track at the edge of alder and willow scrub at Knocknabrass Td on the Crom estate. The spot isapproximately 400 m NE of the present occupied castle and quite remote from any garden.
RHN collected a sample portion of the plant, including its stoloniferous base, and grew it on in a gardenpot. Originally it was mistaken for Oreopteris limbosperma which it quite closely resembles, butin 2009 when the potted plant was fully developed RHN recognised it was this species.
Origin, ecology and history in cultivation
M. struthiopteris is a native fern of boreal and montane areas of central and eastern Europe,Asia and N America. It was introduced to garden cultivation in Britain and Ireland as long ago as 1760,and was first recorded in the wild in 1834 (T.D. Dines, in: Preston et al. 2002). As it toleratesshade and waterlogged soils of almost any pH, it has become a popular subject for water gardens inrecent years. Instances of it occurring as a garden escape are quite frequent but widely scattered inBritain from Cornwall (VC 2) to N Ebudes in W Scotland (VC 104) (New Atlas).
Irish occurrence
In Ireland, it is of much rarer occurrence, the Catalogue of Alien Plants in Ireland listing justthree VCs, Cos Leitrim, Antrim and Londonderry (H29, H39 & H40) (Reynolds 2002). Famously, twolargish colonies spread from an abandoned garden at Shane's Castle, Antrim, into damp woodland atMassereene on the NE shore of Lough Neagh. These established patches of the fern, first discovered in1948 by Carrothers, Moon and Davidson of Fermanagh Typescript Flora fame, survive and appearnaturalised, competing with natural vegetation (FNEI 3). We expect the Crom plant will do thesame.
Names
'Matteuccia', is named in honor of Carlo Matteucci (1800-1868), an Italian physicist. The Latinisedepithet 'struthiopteris', from the Greek, 'strouqeios' or 'stroutheios', meaning "of anostrich", and 'pteris', fern. The English common name 'Ostrich Fern' is from a supposed resemblanceof the fronds to the plumes of the large flightless Afican bird. Alternative common names include'Fiddlehead Fern', and the singularly uninspired 'Garden Fern' and 'Hardy Fern' (http://rook.org/earl/bwca/nature/ferns/matteuccia.html)(accessedNov 2014).
Introduction, neophyte, garden escape, very rare.
23 September 2000; Northridge, R.H.; streamside, near Florencecourt House.
Growth form and preferred habitats
In Britain, this deciduous, rhizomatous, perennial garden species of North American and Eastern Asianorigin is a widely scattered, but quite frequent established garden escape or discard. It tends tocolonise wet lake margins and areas where ground water seeps. Other typical habitats include dampwoodland marshy meadows and riversides. The plant, which is frequently grown in gardens, has ahorizontally spreading rhizome which quickly spreads allowing this attractive fern to form large clonalpatches. The fronds are heterophyllous, the plant producing yellow-green pinnatifid fronds up to 90 cmtall and separate, shorter, purplish-brown, fertile fronds that bear bead-like sori tightly clusteredlike grapes.
Since it may quite rapidly outgrow available garden space, Sensitive Fern gets passed around betweengardening acquaintances and also becomes discarded in refuse tips or in areas of waste ground where flytipping takes place (Clement & Foster 1994). The species may also spread naturally by means of itsspores and colonies in Britain have been reported considerable distances from habitation. There is someevidence that this species is currently spreading in Britain at least (T.D. Dines, in: Preston etal. 2002).
Fermanagh occurrence
A recent, solitary record exists in the Fermanagh Flora Database of a large, well-established patch thatwas found growing beside a stream in the outer Pleasure Grounds of Florencecourt House, in an area thatotherwise appeared unplanted.
British occurrence
O. sensibilis is mainly distributed in S & W parts of Britain and according to Jermy &Camus (1991) has also been reported in Ireland. Oddly, in view of this, there is no mention of it in theCatalogue of Alien Plants in Ireland (Reynolds 2002), and while it is mapped in the CD-romdistributed with the New Atlas, the latter plots no hectads for Ireland.
Names
'Onoclea' is from the Greek 'onos', meaning 'vessel', and 'kleio', 'to close', referring to the closelyrolled pinnules of the fertile fronds that enclose the sori. The Latin specific epithet 'sensibilis'means 'sensitive'. The English common name 'Sensitive Fern' originates in N America where the fronds areobserved to be very sensitive to frost, the aerial parts quickly collapsing and dying off when firsttouched by it. An alternative common name, 'Bead Fern' refers to the bead-like sori (http://rook.org/earl/bwca/nature/ferns/onoclea.html)(accessedNov 2014).
Native, very common, widespread and locally abundant. Circumpolar boreo-temperate.
1860; Smith, Rev Prof R.W.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This widespread and abundant usually calcifuge fern is a large, deciduous, perennial with frondsregularly up to 120 cm long, finely-dissected, delicate-looking (and hence lady-like). Althoughextremely variable, it is generally easy to recognise, especially when it is a distinctive light,yellowish-green colour. Some plants of a much darker colour do occur, however, and these need to bechecked more carefully to ensure correct identification. Lady-fern is a very common and widespreadspecies of damp but well-drained, usually (but not obligatorily) shady, acid to neutral habitats.
In earlier centuries, botanists looked for and found their Male- and Female-ferns, their Filix-mas andFilix-femina, the former being coarse and aggressive, the latter contrastingly delicate, finely-cut andlady-like. The female fern originally chosen was our present day Bracken (Pteridium aquilinum),whose lace-like tripinnately-cut fronds suggested feminine grace and delicacy (Step & Jackson 1945).It was the Swedish botanist, Carl von Linné (since he wrote his texts in Latin, most often referred toby the Latinised form of his name, Linnaeus), who transferred the name 'filix-foemina' (nowadays spelt,'filix-femina') to the present species, likewise regarded as characteristically feminine (Grigson 1974).
Fermanagh occurrence
Lady-fern has been recorded in 380 tetrads, 72% of those in the VC. In reality, we would estimate that itprobably occurs in just about every tetrad in Fermanagh except those on: a. the very highest ground,where a combination of altitude and exposure are too great for its tolerances; b. in very heavilydisturbed or waterlogged sites; and c. in county boundary tetrads where only a very small parcel of landlies within our survey area. Typical local habitats include deciduous woods, cliffs, rocky ravines, dampmeadows and by water, including along ditches, paths, shady roadside banks and hedges. Occasionally,Lady-fern is also found growing on damp to wet walls, though plants generally fail to mature in thiscirc*mstance due to the lack of adequate moisture and soil.
Although Lady-fern is a lime-avoiding, calcifuge species, in our damp climate it frequently occurs inshallow, acid, peaty soils overlying calcareous or base-rich rocks. This is especially the case onslopes, which while they are regularly wetted by our skies, permit drainage adequate to support thisfern (Page 1997).
Variation
A. filix-femina is one of the most variable of ferns, so that at least 30 named subspecies,varieties, forms and hybrids have arisen in the wild and under cultivation. Some of these named entitiesare based on the form and cutting of the frond and occurring in widely separated localities and withinseveral races, while others are more geographically limited (Hultén 1962; Hultén & Fries 1986). InFermanagh, as long ago as 1860, Prof Smith described var. convexum Newm., and var. incisumNewm. as, "abundant everywhere" (Smith 1860; Meikle et al. 1975). Some of named formsare difficult to separate on account of the great variability of the species, not to mention theinherent confusion of the taxonomy and nomenclature. An excellent account of this fern in Britain andIreland, its variation, recognition and ecology, is provided by Page (1997).
British and Irish occurrence
A. filix-femina is a very common and widespread species throughout the British Isles, beingespecially common in the wetter and more mountainous western counties. It becomes local only in thesomewhat drier and colder Irish Midlands and in the more continental climate of East Anglia (Jermy etal. 1978; Page 1997; New Atlas).
European and world occurrence
A. filix-femina in the broadest sense is a very widespread circumpolar species of middle latitudesaround the northern hemisphere. The distribution thins southwards towards the southern peninsulas of theEuropean mainland and the Mediterranean isles. It also becomes decidedly more sparsely scattered ineastern parts of continental Europe (Hultén 1962, Maps 168A & 168B; Jalas & Suominen 1972, Map105; Hultén & Fries 1986, Map 49). In the southern hemisphere, Lady-fern has also been reported fromtropical forests and mountains in Zimbabwe, Natal, Java, Peru and Argentina (Hultén 1962; Grime etal. 1988).
Names and Uses
The derivation of the genus name 'Athyrium', is somewhat obscure, but possibly comes from the Greek'anthoros', meaning 'breeding well', perhaps alluding to the varying form of the sori (Gilbert-Carter1964; Hyam & Pankhurst 1995). The Latin specific epithet, 'filix-femina', translates as 'Lady Fern'(Step & Jackson 1945). In herbal medicine, the uses of Lady Fern are as in Male Fern (Dryopterisfilix-mas), but it is considered less powerful in its action (Grieve 1931).
Threats
None.
Native, occasional to locally frequent. Circumpolar wide-boreal.
1860; Smith, Rev Prof R.W.; Florencecourt.
April to December.
Growth form and preferred habitats
Brittle Bladder-fern is a widespread, very variable, small deciduous fern in our survey area. Stronglycalcicole, it is frequent in permanently shaded, rocky ground, Ash woodland, or walls and quarries whereplant competition is limited. It frequently associates with Asplenium trichomanes (MaidenhairSpleenwort), Phyllitis scolopendrium (Hart's-tongue) and Ceterach officinarum (Rustyback).Its thin, delicately cut, deciduous fronds sometimes reach 25 cm or more in length, although usuallythey are much shorter, around 10 cm long. Separate but similar fertile fronds bear numerous sori. Eachyoung sorus is protected by a thin, membranous indusium which is slightly inflated and is pear- orbladder-shaped. The sori turn dark brown or black in colour as the sporangia ripen, and by this stagethe indusium has shrunk and become inconspicuous, facilitating the release of spores on the slightestbreeze.
Fermanagh occurrence
This little fern is a typical plant of the Fermanagh limestones, base-enhanced dolomitized sandstonescarps or other calcium-bearing rock habitats of the west of the county. In Fermanagh, it has beenrecorded in 64 tetrads, 12.1% of those in the VC. Seven tetrads contain pre-1976 records only. Typicallocal habitats include ± permanently shaded, damp crevices on N-facing, sometimes wooded cliffs andravines, rocky slopes, screes, swallow-holes, caves and narrow fissures (ie grykes) in limestonepavement. As the tetrad distribution map indicates, apart from the calcareous or base-rich natural rockoutcrops of W Fermanagh, Brittle Bladder-fern is only occasional elsewhere in the county and here it isconfined to scattered man-made habitats, such as quarries and the weathered lime-mortar of shaded oldwalls and bridges.
British and Irish occurrence
In Britain & Ireland, the distribution of C. fragilis is markedly northern and western, beingstrongly associated with the wetter and higher rocky ground in these regions. In other parts of theseisles, C. fragilis ascends well over 915 m in sheltered, moist cliff crevices, but in the veryoceanic climate and with the relatively low relief base-rich rocks in Fermanagh, the highest it reachesis the very modest 400-420 m of Trien Mountain. The complete inability of the species to tolerate summerdrought means that it is more or less absent or very rare in the S & E of both Britain and Ireland(Page 1997).
In Ireland, the distribution and frequency of Brittle Bladder-fern most closely matches that ofHymenophyllum wilsonii (Wilson's Filmy-fern), another very delicate, even thinner-textured fernspecies, the distribution of which is also very much governed by constantly high atmospheric humiditylevels (Jermy et al. 1978; New Atlas).
Variation
C. fragilis is a very variable polymorphic fern with a number of named varieties, some of whichare very probably linked to the different chromosome numbers recorded for the species which form apolyploid series. Tetraploid and hexaploid plants, plus the pentaploid hybrid between them, have beenfound in the British Isles. The degree of frond dissection appears to be correlated with chromosomenumber: the more dissected it is, the more likely the plant is hexaploid. An octoploid form has beenfound in Europe, which might also crop up if searched for (Jermy & Camus 1991; Page 1997).
European and world occurrence
In Europe, the species is very widely distributed throughout moister northern and middle latitudes,becoming somewhat less evident in the Mediterranean basin, yet reaching the Azores (Jalas & Suominen1972, Map 110; Page 1997). A closely related species, previously regarded as C. fragilis subsp. diaphana (Bory) Litard., occurs throughout Madeira (Presset al. 1994), and another related form also occurs in the Canary Islands (Page 1997).
Taken in the broadest sense, C. fragilis s. lat. is an extremely widespread circumpolar species,its natural range stretching right across N America as well as throughout Europe and Asia. Taxonomicvarieties of it also occur in large parts of Africa down to the Cape, in Australia, Tasmania, NewZealand, in Kerguelen and South Georgia, as well as in S America and the Falkland Isles (Hultén 1962,Map 55; Hultén & Fries 1986, Map 52).
Names
The genus name 'Cystopteris' is Greek, combining 'kustis', 'kystis' or 'cystis', meaning "abladder", and 'ptěris', "fern", referring to the bladder- or pear-shaped outline of theindusium (Gilbert-Carter 1964). The shape of the latter always reminds me of the large flasks ofcoloured water that in my Londonderry childhood of the 1950s often graced pharmacists' premises, andwhich still act as icons of their profession. The Latin specific epithet, 'fragilis' usually translatesas 'brittle', 'fragile' or 'easily broken', which in this case applies to the brittle stipe or stalk ofthe frond. Translating the Scientific name thus, we derive the English common name 'BrittleBladder-fern'. However, an alternative translation of 'fragilis' is 'wilting quickly' (Stearn 1992),which is also appropriate for this fern and can readily be applied to its delicate fronds.
Threats
Upgrading and 'tidying' of old walls and bridges.
Native, common, very widespread and locally abundant. Submediterranean-subatlantic.
1860; Smith, Rev Prof R.W.; Co Fermanagh.
Throughout the year.
Growth form and recognition
This large, common rhizomatous fern develops dense shuttleco*cks of many fronds. In the middle part of thefrond the base of every pinnule has a larger lobe, ie the ultimate segments are highly asymmetrical attheir base (Webb et al. 1996; Rich & Jermy 1998, p. 29). This feature, plus the soft texture(always best appreciated by touching the plant), and the grass-green colour of mature fronds make thisan easily recognised species, although some individual plants whose texture is less soft will need to beexamined more closely to distinguish them from P. aculeatum (Hard Shield-fern) (Page 1997).
The fronds expand in late April or early May and are generally wintergreen with us in Ireland, the oldfronds only dying off as the new fronds begin to unfurl. (In colder parts of Britain the fern isregarded as semi-evergreen, fronds dying off after the first hard frost.) Sporing is copious and beginsin July or early August. It may continue until the fronds wilt and collapse in the following spring.
Preferred habitats
Soft Shield-fern occurs on a variety of moderately acidic to neutral, damp but not wet soils of mediumbase-status. The species is sometimes described as a moderate calcicole (T.D. Dines, In: Preston etal. 2002). It is very common and widespread, and under suitable conditions it can dominate theground layer in humid woods at low altitude. P. setiferum is generally absent from upland areasabove c 200 m, and also from strongly acidic peaty ground. Apart from damp mixed deciduous woods, themost frequent habitats of P. setiferum in Fermanagh as elsewhere in Britain and Ireland are onslopes in scrub, along hedgerows, riverbanks and in similar sheltered, shady, damp, but well-drainedplaces.
Fermanagh occurrence
In Fermanagh, this large, rhizomatous fern is common and very widespread having been recorded in 344tetrads, 64.6% of those in the VC. In terms of tetrad frequency this makes it the eighth most widespreadfern in our survey area.
Irish occurrence
Soft Shield-fern is very common throughout most of Ireland except in the more exposed, strongly acidic,constantly wet peatlands of the W & C where it becomes decidedly rare or absent (Jermy et al.1978; An Irish Flora 1996; New Atlas).
British occurrence
Soft Shield-fern is abundant in suitable sites in SW England and SW Wales, but as one moves either eastor northwards it becomes steadily rarer, until in Scotland it is virtually confined to western coasts.The most northerly point P. setiferum reaches anywhere in the world is near Ullapool on the westcoast of Scotland (Jermy et al. 1978; New Atlas).
European and world occurrence
Overall P. setiferum is very definitely an Atlantic or oceanic species, most abundant along theAtlantic coastline, but occurring inland throughout W and S Europe and along the northern shore of theMediterranean, becoming more discontinuous further east but reaching the Caspian Sea area. It occurs onall the islands in the Azores, and further south reaches Madeira and the Canary Islands (Jalas &Suominen 1972, Map 121; Hultén & Fries 1986, Map 62; Page 1997; Vertag 2002).
Hybrids
P. setiferum hybridizes with P. aculeatum to form P. ×bicknelli, which has been recorded once in Fermanagh (see our account of this hybrid).
Names
The genus name 'Polystichum' is derived from two Greek words 'polus', 'many', and 'sticos' or 'stichos','row' or 'file'. The genus name thus translates as 'many rows', a reference to the regular rows of sorion the fertile frond (Gilbert-Carter 1964). The specific epithet 'setiferum' means 'bristle-bearing' andis derived from the Latin words 'seta' or 'saeta' meaning 'a bristle' or 'a stiff hair', and 'fero'meaning 'to bear' or 'to carry' (Gilbert-Carter 1964).
In past years, the general population did not distinguish any species of Polystichum other thanP. lonchitis (Holly-fern), so there are no folk uses, nor any English common names other thanobvious, invented 'book names' (Step & Jackson 1945). The name 'Shield fern' is given because thecircular, centrally stalked (peltate) indusium that protects the young sorus is considered reminiscentof the Medieval circular Buckler shield.
Threats
None.
Native, very rare but probably overlooked and under-recorded.
December 1980; Northridge, R.H.; shaded riverbank, Ballyvelin Bridge, Colebrooke River nearMaguiresbridge.
Throughout the year.
Growth form and appearance
P. × bicknellii is evergreen and calcicole like its P.setiferum (Soft Shield-fern) parent and it can be distinguished by the presence of mostlyabortive spores. The plant closely resembles a large, leathery, robust, dark green form of P.setiferum.
Fermanagh occurrence
This hybrid has been found only once in Fermanagh as detailed above, but it should be looked for inshaded, mainly lowland areas where both parent ferns occur (Northridge et al. 1988). Apart fromthe limestones of the Western Plateau, the Florencecourt and Maguiresbridge areas and the ground lyingnorth of Kesh are the main localities in Fermanagh where both parents commonly meet and thissporadically occurring hybrid is most likely to occur.
British and Irish occurrence
Throughout Britain and Ireland, this hybrid occurs thinly scattered as isolated individuals, mainly inecologically open, calcareous or base-rich soils. Typical habitats include rocky woodlands, limestonegorges, stream banks and old disused quarries. It seems particularly associated with ground where bothparents are involved in recolonisation after disturbance (Jermy et al. 1978). The hybrid is veryeasily overlooked, and Page (1997) therefore believes it is under-recorded.
Threats
None.
Native, frequent. Eurasian temperate.
1860; Smith, Rev Prof R.W.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
While P. setiferum (Soft Shield-fern) is quite gregarious and forms large 'shuttleco*cks', eachcomposed of numerous fronds, in comparison P. aculeatum rarely occurs as anything more thanscattered individual plants. Also, individuals of Hard Shield-fern are typically small to medium-sized,often with just a few fronds per shuttleco*ck crown.
P. aculeatum usually grows in shade, and it is very often associated with exposed calcareous orother forms of base-rich rock. Of the two shield-ferns, the evergreen P. aculeatum tends to bethe more upland in character, growing in damp pockets of soil in a wide range of shaded places whereoutcrops of limestone rock occur, including stabilized scree, around swallowholes, in deep grykes inlimestone pavement, on steep wooded slopes, river banks and by streams and waterfalls.
The hard leathery texture of the fronds of P. aculeatum and their glossy, dark-green colour whenmature makes most plants easy to distinguish from Soft Shield-fern. If in any doubt, then the plant isfar more likely to be P. setiferum. A useful and reliable characteristic of P. aculeatumis that the ultimate segments of pinnae in the middle of the frond are more or less symmetricalat their base and the innermost pinnule on the upper side of each pinna is usually very much larger thanall the rest and it is more deeply divided (An Irish Flora 1996; Page 1997).
Fronds of P. aculeatum expand from the beginning of May onwards and they begin to spore inmid-July. Like Soft Shield-fern, the fronds are wintergreen and they persist longer in their second yearthan those of P. setiferum (Page 1992).
Fermanagh occurrence
Although, as the tetrad distribution map indicates, this is a rather widely scattered species throughoutmuch of Fermanagh, having been recorded in 130 tetrads, 24.6% of those in the VC, there really are onlytwo areas in Fermanagh where Hard Shield-fern occurs with any notable frequency. These are along damp,shaded riverbanks and roadsides in the Maguiresbridge district and to the north of Kesh. In these twolocalities, it is more frequent than the generally much more common and widespread P. setiferum.
In addition to its shaded and rocky natural or semi-natural habitats, in Fermanagh Hard Shield-fern isalso recorded in urban situations around Enniskillen, and in more rural areas it is often associatedwith man-made structures such as old walls, bridges, weirs and even along old disused railway lines.
Similar behaviour is described in England, where additional man-made base-rich sites colonised includecanal-sides, locks and bridges (Jermy et al. 1978). In SW Scandinavia, Hard Shield-fern occurs onmountain cairns, 'stone-fences' (presumably this refers to our 'dry stone walls'), and along ditches(Jonsell et al. 2000). The leathery fronds of P. aculeatum are considerably more hardythan those of P. setiferum, enabling it to survive in more easterly areas of the British Islesand Europe which suffer heavier winter frost than western areas like Fermanagh.
Irish occurrence
In the past, Hard Shield-fern has been recorded at least once in every Irish vice-countyexcept Mid-Cork (H4) (Scannell & Synnott 1987). While this is a fact, in reality over most ofthe island the fern is only thinly scattered, seldom frequent, and occurs chiefly in the N & W ofIreland (Jermy et al. 1978; Webb et al. 1996; New Atlas).
British occurrence
In Britain, P. aculeatum is much more widespread than P. setiferum, extending from theEnglish Channel coast to Orkney in the far north (but not Shetland, where it is considered a rareintrodution). It is absent, however, from considerable areas of the English East Midlands. It is mostcommon in N England and Scotland, but less common than P. setiferum in SW England and S Wales(Jermy et al. 1978; Jermy & Camus 1991; Stace 1997; Preston et al. 2002). In areas ofthe British Isles where their distributions overlap, P. aculeatum and P. setiferum meet insuitable habitats intermediate in altitude, and their hybrid, P. ×bicknellii (recorded once in Fermanagh), is not uncommon and should be looked for (Preston etal. 2002).
European and world occurrence
In Europe, Hard Shield-fern is widespread but uncommon, predominantly occurring in the W and C temperateregions, extending to 64°N up the coast of SW Norway, and stretching south to the Azores, Majorca,Corsica, the S Peloponnese and the coast of N Africa (Jalas & Suominen 1972, Map 120). Eastwards thefern spreads ever more discontinuously, reaching the Caspian Sea and possibly beyond. Probable relatedtaxa occur in the Himalaya and in Japan (Stace 1997). Hultén (1962, Map 141) and Hultén & Fries(1986, Map 61) treat P. aculeatum in an extremely broad sense, merging it at least in part withP. setiferum and related species, so that their maps and accounts are for once singularlyunhelpful.
P. aculeatum is sometimes grown for ornament, and Jones (1987) lists four garden varieties, ofwhich 'Pulcherrimum Gracillimum' is described as "the most beautiful British fern" [incultivation].
Name
The genus name 'Polystichum' is derived from two Greek words 'polus', 'many', and 'sticos' or 'stichos','row' or 'file'. The genus name thus translates as 'many rows', a reference to the regular rows of sorion the fertile frond (Gilbert-Carter 1964). The Latin specific epithet 'aculeatum' means 'havingprickles, thorny or prickly' (Gledhill 1985), which in the present author’s opinion, rather overstatesthe case as the frond does not feel prickly when handled.
Threats
Excessive clearance or tidying of riverbanks.
Native, very rare. Circumpolar boreo-temperate.
July 1979; Northridge, Mrs H.J. & Northridge, R.H.; cliff 1 km SE of Lough Achork, in the Lough NavarForest Park.
Throughout the year.
Growth form and preferred habitats
Holly-fern produces its glossy, evergreen, rather leathery fronds in shuttleco*ck-like rosettes from ashort, stout rhizome, often embedded in tight crevices on exposed rock faces. The English common name'Holly-fern' is not really all that appropriate, since it is not 'holly-like' at all, except in that thesimply pinnate fronds are definitely evergreen (each persisting two or three years). The fronds areshiny, and the individual pinnae bear teeth that look rather spiny, but are really quite flexible. Boththe growth of fronds and the establishment of new plants are slow, but compensating for this fact,individual plants and fronds are clearly long-lived.
P. lonchitis is a definite calcicole, being confined to calcareous or base-rich rocks, whichincludes the dolomitised sandstone it frequents in Fermanagh. The species typically grows in cool,moist, well-drained positions at the base of cliff faces, or in crevices on or near ledges. In numeroussites, it frequents stabilised boulder scree and, in England, occasional plants of P. lonchitisare found in moist, deep grykes in limestone pavement and around the entrances of sinkholes.
Fermanagh occurrence
One plant of this species is known to have survived for 30 years at its site in a crevice on a N-facingdolomitized sandstone cliff in Lough Navar Forest (Northridge et al. 1988). In 2000, the originalplant was joined by a small plant 20 cm higher on the cliff, which we regard as most probably anoffspring of the established plant. By 2010, there were five plants at the site, the original plus twosmall daughter rosettes slightly higher up on the cliff face, plus two nearly mature rosettes on theground at the base of the rock face, which previously had been overlooked.
In June 1999 R.D. Porley, an English bryologist, surveying mosses and liverworts on the Lough Navarscarps for the Environment and Heritage Service, found another small plant which he identified as P.lonchitis on Bolusty Beg, almost exactly 2 km due north of the earlier known station. Thepresent author and his Botanical Society joint-Vice County Recorder, Robert Northridge, only discoveredthis claim when the bryological results were published in September 2002 (the Irish Naturalists’ Journalwas a year late in its appearance). In 2003 Robert Northridge visited the site and saw an immaturePolystichum plant near a red marker stick. The plant was too immature to determine to specieslevel. On 1 February 2004, RHN revisited the site and the plant had matured enough for it to bedetermined as P. aculeatum (Hard Shield-fern). It is difficult to distinguish juvenile specimensof these closely related ferns and the mistaken identification is a perfectly understandable one.
P. lonchitis is a circumpolar arctic-alpine species, and at its Fermanagh site it grows beside aplant of Asplenium viride (= A. trichomanes-ramosum) (Green Spleenwort), another northernor arctic-alpine species similarly confined to calcareous or other base-rich rock habitats. Normallyboth these species occur in upland areas in Britain and Ireland, which are typically cool in summer(summer maximum around 27C), and where base-rich pockets of soil are kept permanently moist by waterseepage (Page 1997).
Sori are produced on the top portion of the frond only, and spores are released from mid-summer until thefollowing spring. Clearly conditions at our Fermanagh site are suitable for reproduction, since newyoung plants have become established near the original specimen in recent years. Apart from thewell-colonised stable block scree stations in Glenade, Co Leitrim (H29) (the undoubted Irishheadquarters of the plant, where the species grows large, luxuriant clumps), Holly-fern occurs in mostof its Irish sites as small, widely scattered, individual cliff crevice plants. It is quite possiblethat somewhere in the Lough Navar area another plant might occur on nearby cliffs or screes of suitablerock chemistry.
Irish occurrence
The plants at the solitary Fermanagh site are the only representatives of this species known fromNorthern Ireland, although elsewhere in the Republic of Ireland there are a few scattered localitiesdown the W coast in Co Donegal (H34 & H35), Co Leitrim (H29) (definitely its Irish headquarters), CoSligo (H28), Mayo (H27), Co Galway (H16) and Kerry (H1 & H2) (Jermy et al. 1978; CensusCatalogue of the Flora of Ireland 2).
In limestone pavement in N England, occasional plants of P. lonchitis are found in moist, deepgrykes and around sinkhole mouths, making it all the more odd that the species is absent from theidentical habitats which are so vastly more abundant in the Burren district of Co Clare. Having saidthat, A. viride (Green Spleenwort) is another inexplicable Burren fern absentee.
British occurrence
In Britain, P. lonchitis is similarly rare and local from N Wales, N England and S Scotland, butit becomes more widespread in the mountains of C & NW Scotland (Jermy et al. 1978; Page1997). The records in the 1978 Fern Atlas suggest that there may be a contraction in the numberof sites for Holly-fern in parts of Ireland, England and Wales, but this does not appear to apply in themain area of the species in N & W Scotland (Jermy et al. 1978). In limestone pavement in NEngland, occasional plants of P. lonchitis are found in moist, deep grykes and around sinkholemouths.
European and world occurrence
In Europe, P. lonchitis is widespread in cooler areas, ie montane and high latitude regions. Thedistribution extends northwards along W Scandinavia reaching well inside the Arctic Circle. The speciesis also present in Iceland and the Faroes. In continental Europe, it is mainly associated with the Alps,Pyrenees and other outlying mountain areas, reaching its southern extremities in SE Spain, Italy,Corsica, the Peloponnese and W Crete (Jalas & Suominen 1972, Map 119). The overall Europeandistribution pattern is remarkably similar to that of Asplenium viride (= A.trichomanes-ramosum), Green Spleenwort (Jalas & Suominen 1972, Map 83). As in the case inDryopteris carthusiana (Narrow Buckler-fern), P. lonchitis in Scandinavia occupies a muchwider range of habitats than it does in the British Isles, including forest (deciduous forest as well asmixed and spruce forest), together with crevices in lava-fields, and even occasionally on man-madestructures e.g. stone walls ('stone-fences') (Jonsell et al. 2000).
Beyond Europe, Holly-fern ranges around cooler parts of the northern hemisphere in a circumpolar mannerfrom the Caucasus, the N Urals, mountains of C Asia, the W Himalaya, Japan, W and NE North America andGreenland (Hultén 1958, Map 219; Jonsell et al. 2000).
Names
The genus name 'Polystichum' is derived from two Greek words 'polus', 'many', and 'sticos' or 'stichos',meaning 'row' or 'file'. Taken together 'many rows' is a reference to the regularly arranged rows ofsori on the fertile frond (Gilbert-Carter 1964). The specific epithet 'lonchitis' is a Latinised form ofthe Greek 'loncho' meaning 'spear-shaped' or 'lance-shaped', obviously referring to the outline of thefrond. It was also a name given to an unknown fern by Dioscorides (Gilbert-Carter 1964; Gledhill 1985).
The English common name, 'Holly-fern', is a rather curious one, the only similarities between the twoplants being the evergreen, thick textured leaves. The numerous teeth on the fern pinnae only lookprickly or spiny, but are in fact quite soft, unlike the real Holly's leaf prickles. In common with the'Oak Fern' and 'Beech Fern', there is no ecological linkage between the fern appellation and the treefor which it is named, and one is left to wonder at the philosophy behind the obscure names humans giveto the things about them!
Threats
The Fermanagh Holly-fern station is vulnerable having survived one extensive fire on its cliff in recentyears; another such event could easily destroy it.
Native, common, widespread and locally abundant. Circumpolar temperate.
1860; Smith, Rev Prof R.W.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Male-ferns in Britain and Ireland were originally conceived to be a single species and the history of thesubsequent splits and their naming is a complicated one. In Fermanagh, we have records for a total offive species of Dryopteris plus two hybrids. Among the male-ferns, we recognise and distinguishD. filix-mas and D. affinis (Scaly Male-fern), the latter an apomictic species, severalforms of which have in the past been considered and recorded as separate species. D. filix-mas isa large, common deciduous fern that is distinguished from D. affinis by the lack of a black markat the point where the pinna midribs meet the rachis, and by its sparser, paler, straw-coloured scaleson the stipe.
D. filix-mas is a large, vigorous, shuttleco*ck-forming perennial, the vertical rhizome of which islong-lived and can become quite massive. It is common in woods, hedgerows, streamside banks, ditches,roadside verges, rocky slopes, screes and cliff ledges. The species has an unusually wide tolerance ofsoil pH (from pH 3-8), nutrient status and water supply, which is reflected in the huge variety ofhabitats it occupies. Experimental measurements made in W Europe to determine the role of rootcation-exchange properties of fern species found that D. filix-mas was indifferent to both soilpH and calcium carbonate content (Koedam et al. 1992).
D. filix-mas performs best in lowland sites with relatively fertile soil and little disturbance.On well-drained slopes, within mixed deciduous woodland, it can dominate stretches of the floorvegetation. Often in this type of seasonally shaded habitat, its main competitors are other large ferns.Most frequently these are D. dilatata (Broad Buckler-fern) and Athyrium filix-femina(Lady-fern), but sometimes it also overlaps with the closely related D. affinis.
Unusually for a fern, D. filix-mas is very tolerant of atmospheric pollution and it colonisesshady, damp, urban areas and industrial sites including rubbish dumps, old brickwork and other lesssalubrious situations. It is probably true to say that its occurrence is limited only by extremesof exposure, wetness (ie permanent water-logging) and shade, and by heavy disturbance including grazingpressure.
Fermanagh occurrence
This species and D. dilatata (Broad Buckler-fern) are the two most common ferns in Fermanagh, thelatter perhaps being slightly more widespread. The Fermanagh Flora Database contains records of D.filix-mas from 462 tetrads, 87.5% of those in the VC, and it is clearly widespread and abundantalmost everywhere except in aquatic, heavily disturbed, or very exposed situations. It dominatesstretches of the woodland floor at the base of the cliffs of Poulaphouca, on Bilberry Island on LowerLough Erne, and in parts of the Cladagh River Glen.
Phenology
Fresh annual fronds of D. filix-mas unfurl from early May and are fully expanded by the end ofthat month. D. filix-mas is one of the hardiest ferns in the British Isles and its fronds arewintergreen to semi-persistent in Fermanagh, depending on degree of exposure, although in truth underour dull, grey skies they probably are semi-senescent for much of the winter (Grime et al. 1988).By late February or earlier, the old fronds are often broken down and they are certainly dying off, soonto be replaced by the growth of fresh new croziers on the upright rhizome.
Reproduction
Male-fern plants produce colossal amounts of spores which are released from August to November (Page1997). Despite this fecundity, it is not at all clear with what degree of success it achievesestablishment of the sporophyte generation. That this does happen is attested by the presence of young,small, sporophyte plants embedded in cushions of moss on little rises on the floor of woodlands, andoccasional plants developing in crevices at various heights on walls (Willmot 1985). A preliminarypopulation study in Derbyshire woodland suggested that recruitment of small D. filix-mas plantswas a relatively rare event when compared with D. dilatata, but that once plants of the formerestablished, they might live for a long time (Willmot 1985). A study in Russian woods suggested thatplants mature and spore only when over six years of age, and they may survive for 30 to 40 years orlonger (Pogorelova & Rabotnov 1978).
On walls, observation indicates that small plants frequently persist for many years, but they seldomachieve maturity and sporing ability. Possibly reasons for such failure are excessive dryness, lack ofnutrients, or perhaps because of eventual, inevitable disturbance (Grime et al. 1988).
British and Irish occurrence
D. filix-mas is extremely common throughout the whole of the British Isles, and in fact in termsof spread it is second only to Pteridium aquilinum (Bracken) in the number of hectads in which itoccurs (Jermy et al. 1978; New Atlas). Page (1982, 1997) suggests that the presence ofMale-fern in Britain and Ireland has diminished in the last two thousand years with the gradual removalof much of the forest vegetation, the natural habitat of the species. Considering just how common andwidespread a plant it remains, and its almost unrivalled ability to colonise artificial, man-madehabitats, it is hard to see that there is any cause for concern about the species yet, since it clearlymanages extremely well in the substitute habitats it now so fully occupies!
European and world occurrence
In Europe, Male-fern is widespread at middle and northern latitudes, thinning northwards but reaching theArctic circle in Scandinavia, and while present in Iceland it is absent further north in Svalbard(Spitsbergen, Bear Island and Jan Mayen) (Tutin et al. 1993). The distribution also thins towardsthe south in the Iberian Peninsula, and Male-fern is absent from the Azores, Madeira and the Canaries.Of the Mediterranean islands, it is present only on Corsica and Sicily (Jalas & Suominen 1972, Map123).
On the world scale, taking D. filix-mas in a broad sense (and recognising that the taxonomic andnomenclature confusion that exists within this group of ferns creates problems when it comes toestimating distribution), beyond its main European base it displays a disjunct circumpolar distribution.In the past, several forms were given separate taxonomic recognition, especially in the far east ofAsia, S Africa, S America and a number of island groups in the southern hemisphere, includingMadagascar, Hawaii, the Falklands and the Galapagos (Hultén 1962, Map 110; Hultén & Fries 1986, Map64).
Names
The genus name 'Dryopteris' was first given by the Greek physician, Pedanius Dioscorides (c. 40-90 AD),to a fern growing on oak trees, and is a compound of the Greek 'dryas' = 'oak', and 'pteris' = 'fern'(Gilbert-Carter 1964). The specific epithet 'filix-mas' was first given by the German botanist, LeonhartFuchs (1501-66), and is derived from Latin 'filix' = 'fern', and 'mas' = 'male', from the supposedlybig, bold 'masculine' appearance of the species in comparison with the much more delicate, finelydivided fronds of the 'Lady-fern' (Gilbert-Carter 1964).
Uses
The above mention of these ancient and medieval herbalists gives an indication of the fact that Male-fernhas a very long history of medicinal use. An oil extracted from the rhizome was used from ancient timesas a vermifuge (ie used to kill and expel flatworms, tapeworms and liverfluke). It has also been appliedfor worming in veterinary medicine (Vickery 1995), although the rhizome also contains a dangerous toxin,thiaminase. The latter is known to cause thiamine deficiency in animals such as horses, cattle and pigswhich have eaten the fern. When oil of Male-fern was used for worming, a single sufficient dose wasreputed to produce a cure at once, but too much of the toxic drug is poisonous and can cause coma andblindness.
D. filix-mas also contains filixic acid (filicin), and the main toxic activity of the species isdue to a phloroglucinol derivative of this substance (Cooper & Johnson 1998). As with all herballore quoted here, BE WARNED, and do not attempt to administer any such drug without the guidance of aqualified, licensed medical or veterinary practitioner. Another past herbal use for powdered root ofMale-fern was for treating rickets in children, and an ointment was also made and used for healingwounds (Grieve 1931).
Male-fern had other uses similar to those of Bracken, the ash of the burnt fern having applications inglass-making and soap. Young croziers were boiled and eaten like Asparagus. In Norway, at times ofhardship, it was used to make beer, the dried fronds being said to make an excellent bitter. Frondsinfused in hot water have also been regarded in the past as good fodder for sheep and goats (Grieve1931).
Threats
None.
Very rare, but easily overlooked and very probably under-recorded.
1989; Tickner, M.; Stony Islands, Lower Lough Erne.
This deciduous rhizomatous hybrid is under-recorded throughout Britain and Ireland, but itusually occurs on damp, acidic soils in sheltered woodland, ditches, hedgerows and coniferous foresttracks, usually in lowland areas, and especially where the habitat has been disturbed. Plants may occureither as scattered individuals or in small clumps, with or without the presence of one or both of theparent species. Unfortunately, the intermediate hybrid is fully fertile and back-crossing is prevalentproducing a swarm of variation that very greatly complicates identification of all three taxa. SinceD. affinis (Scaly Male-fern) in particular is highly variable in any case, this makes a badsituation even worse. As Botanical Society of Britain and Ireland appointed joint-Vice-county Recordersfor Fermanagh, Robert Northridge and myself (Ralph Forbes) have been present when even the greatest fernexperts in Britain could not agree on the identity of these hybrid plants, for instance on the floor ofthe Cladagh River Glen!
Fermanagh occurrence
There are just eleven records, therefore, in the Fermanagh Flora Database, occurring in nine tetrads. Themajority of them were made by Matthew Tickner of the RSPB when he was surveying the flora of smallislands in Lower Lough Erne in summer 1989. The seven additional islands on which he claimed D. × complexa grew were as follows: Bingham's Rock, Coghran's, Gravelly,Inishmakill, Inishturk, Sam's and 'Stone Park West'. During a BSBI field meeting on 24 August 2004, thevisiting English botanist Ken Trewen added a further three records with sites listed as follows:woodland in Correl Glen NR, determined by KT; scrub covered scarp on S side of Glencreawan Lough,determined by KT; woods beside Lower Lough Erne below the Cliffs of Magho, determined by KT.
Native, common, widespread and locally abundant. European temperate.
1858; Smith, T.O.; Ardunshin.
Throughout the year.
Growth form and variation
This is an extremely variable apomictic species of complex ancestry which is divided into four subspeciesby Page (1982) and into five morphotypes by Jermy & Camus (1991). Page (1997) reverts to threesubspecies, and in the second edition of the New Flora of the British Isles 1997, Stace takes thesame approach. Like other entirely asexually reproducing species complexes, D. affinis does notcarry out meiosis or reduction division during spore formation, so that its spores are of the samechromosome number as the parent plant. When the spores germinate and produce prothalli, no sexual fusiontakes place, so the new sporophyte arises by apogamy (ie an unreduced female gamete, or a cellassociated with it, forms the embryo of the next sporophyte generation). It is therefore geneticallyidentical to the original parent plant and the sexual mechanism has been by-passed. Over time, geneticmutations occur, however, and, if the offspring are viable, the mutations are maintained by this methodof non-sexual spore reproduction. Eventually, given sufficient time, this produces a multitude ofself-perpetuating varieties or 'micro-species' as in this particular case (Jermy & Camus 1991).
These varieties, morphotypes, subspecies or micro-species – whatever we decide to call them, can probablyall act as male parents in crosses with other species of the genus Dryopteris, and we do have alimited number of records in Fermanagh of the hybrid that D. affinis forms with D.filix-mas (D. × complexa).
Identification
Despite the above, D. affinis is usually easily separated from D. filix-mas (Male-fern) bythe presence of a dark lead-grey or blackish spot at the point at which each pinna meets the rachis,which itself is densely clothed in masses of orange-brown or golden-brown, chaffy scales. The new annualfronds of D. affinis are also produced two to five weeks later than those of D. filix-mas,and their stipes are more densely clothed with chaffy golden-brown or light orange-brown scales (Page1997).
The three subspecies recognised in New Flora of the BI (1997) are really very difficult todifferentiate in a consistent manner, and Robert Northridge and the current author (Ralph Forbes) asjoint Vice-county Recorders of Fermanagh, have not tried to distinguish them here. There is certainlymore than one subspecies or 'morphotype' of D. affinis present in Fermanagh, although withlimited manpower we have not yet been able to differentiate and properly record them. As Page (1997)points out, "satisfactory identification of the variants of this taxonomically very complexapogamous species usually requires a symphony of characters to be taken into simultaneous account."
Preferred habitats
In general, D. affinis occurs in the same wide range of natural and artificial (man-made) habitatsas both D. filix-mas and D. dilatata with which it frequently overlaps (Page 1997). Theseinclude deciduous woods, along open rides or fire breaks in conifer plantations, hedgerows, streamsides,well-drained places on open hillsides and in mountain glens, plus on or in crevices in urban brickwalls.
Toxicity
As with other Dryopteris species, D. affinis contains toxins and is intolerant of grazingpressure.
Fermanagh occurrence
In Fermanagh, D. affinis has been recorded in 345 tetrads, 65.3% of those in the VC. This makes itcommon and more or less widespread throughout, although it is still scarce in some parts of thelowlands, particularly those with less acid or more base-rich soils. Typical habitats are woods, shadedbanks and mountain screes. At higher altitudes, D. affinis certainly appears more common thanD. filix-mas which it appears to replace in these circ*mstances. In Fermanagh, it is very likelythat the triploid subsp. borreri (Newman) Fraser-Jenk. will become recognised as the common formof the species, a pattern of occurrence that is perhaps just beginning to appear elsewhere in Britainand Ireland.
British and Irish occurrence
Like D. filix-mas and D. dilatata (Broad Buckler-fern), D. affinis is a long-livedplant, most frequent in the wetter, more ceanic climate of western parts of the British Isles, frequentbut more local in the east (New Atlas). It ascends in Fermanagh to moderately high levels, around300 m on mountain cliffs and screes. Being difficult to identify, the three subspecies recognised byStace (1997) were poorly recorded during the BSBI Atlas 2000 survey, and they were not mapped in the2002 New Atlas.
Elsewhere in Ireland, Hackney et al. (1992) listed three subspecies occurring in the three VCscovered by The Flora of the NE of Ireland; subsp. affinis in open situations at allaltitudes; subsp. borreri in similar situations to the previous, but also in woodland and morefrequent than subsp. affinis at lower levels; and subsp. cambrensis, which is confined toopen, upland situations, for example, on mountain cliff ledges and in crevices. D. affinis wasnot subdivided in The Flora of County Dublin (Doogue et al. 1998), and there are just twoindividual records by Clive Jermy of subsp. affinis and subsp. borreri dating from 1984listed in The Flora of Cavan (Reilly 2001). In the Flora of County Waterford, Green (2008)lists nine records of subsp. affinis and two of subsp. borreri made by four different recorders.
European and world occurrence
On account of the taxonomic and nomenclature changes this taxon has gone through in the last 40years it is difficult to assess the real distribution of D. affinis on a European, let alone aworld basis. Nevertheless, it is mapped by Hultén & Fries (1986, Map 65), and regarded by them andmost other botanists as a mainly European temperate species.
Names
The genus name 'Dryopteris' was first given by the Greek physician, Pedanius Dioscorides (c. 40-90 AD),to a fern growing on oak trees, and is a compound of the Greek 'dryas' = 'oak', and 'pteris' = 'fern'(Gilbert-Carter 1964). The specific epithet 'affinis' is Latin meaning 'related' or 'similar to',presumably referring to D. filix-mas, the other scaly Male-fern (Gledhill 1985).
Threats
None.
Native, locally frequent to occasional. Oceanic temperate.
1858; Smith, T.O.; Tempo.
Throughout the year.
Growth form and preferred habitats
The somewhat crimped appearance of its distinctive light-green fronds, and the long, purple stipe of theplant make this wintergreen plant a distinctive and easily recognised fern. The fronds when lightlybruised in the field give off a slight, sweet smell, but when collected and dried for the herbarium,they at first give off a much more distinct coumarin odour reminiscent of new-mown hay, and hence theEnglish common name (Page 1997). D. aemula is a fern of permanently moist, but essentiallywell-drained acidic to neutral, often peaty soils of low base content. It typically occupies woodedslopes, shaded banks and sea-cliffs (Wardlaw & Leonard 2005).
Fermanagh occurrence
In upland mixed deciduous woodlands in Fermanagh, such as the Correl Glen NR, D. aemula can formthe dominant ground cover over quite large areas of shaded, rocky ground on damp, acidic soils. Thespecies is also a characteristic plant of the sheltered, N-facing, more acidic scarps of the WesternPlateau. It is also found on some of the wooded islands of Lower Lough Erne, and on steep, woodedstreamsides elsewhere in the county. Altogether, D. aemula has been often recorded in 80Fermanagh tetrads, 15.2% of those in the VC. The fern is frequent in the western half of the county,particularly in moist woods and shady banks, but it is only occasional and very scattered elsewhere.
Irish occurrence
In Northern Ireland, D. aemula is noticeably more widespread in the wetter, more oceanic westernparts of Cos Fermanagh, Tyrone and Londonderry (H33, H36 & H40), while in the east of the provinceit is largely but not entirely confined to the wooded coastal glens of Co Antrim (H39), and to moreupland woods and stream-sides of south Down (H38) (Hackney et al. 1992). In the Republic ofIreland, D. aemula is quite frequent and widespread in counties along the western and southernAtlantic coasts, but is encountered much more rarely or completely absent along coastal counties in theeast adjacent to the Irish Sea, and similarly rare in inland situations (New Atlas).
British occurrence
In Britain, as in Ireland, the distribution of this distinctive fern is predominantly western, extendingright from the SW tip of Cornwall to Orkney (but not reaching Shetland). There are a few easternoutlying populations in damp, acid, mainly coastal ground in both Britain and Ireland, but neverthelessthe predominant distribution is markedly western (Jermy et al. 1978; Wardlaw & Leonard 2005).
The slow growth rate of this species and the gradual, unhurried deployment of additional fronds after arelatively rapid spring flush of growth, means that the immature fronds of D. aemula are rathersusceptible to frost both in late spring and in the autumn (ie an early winter or cold snap). The lengthand reliability of the frost-free period is the most likely factor restricting the distribution of thefern to markedly oceanic areas, and at the same time helps explains its absence elsewhere (Page 1997).
The inference immediately drawn from the distinctive distribution pattern of D. aemula is that itis sensitive to winter cold and late frosts, and according to Page (1997), its habitats are low-lying,"most being within about 30 m [100 ft] of sea-level, although it occasionally ascends higher,especially in Ireland". The mild influence of the Atlantic Gulf Stream has a more pronounced effecton winter, late spring and early summer temperatures in western Ireland than is the case in Britain. Ineastern Ireland, D. aemula is known to ascend mountains to 370 m (1200 ft) in Co Wicklow (H20),and 440 m (1430 ft) in Co Down (H38), while in south Co Kerry in the extreme SW of the island (H1), thefern reaches an altitude of 646 m (2100 ft) on the high Reeks (Hart 1891; Brunker 1950; Hackney etal. 1992). Page (1997) therefore appears to be somewhat overstating the difference in the fern'sbehaviour between Britain and Ireland, since by comparison D. aemula reaches 220 m (715 ft) onthe Furness Peninsula in Cumbria (VC 69), 770 m (2500 ft) in E Perthshire (VC 89), and an incredible1015 m (3300 ft) on Braeriach in the Cairngorms (VC 96) (Wilson 1956; Halliday 1997).
European and world occurrence
This distinctive species has a pronounced western distribution in Europe as a whole, and Ireland is oneof its strongholds (Page 1997; NI Vascular Plant Database 2014). It was listed as vulnerable in theCouncil of Europe report on the Rare, Threatened and Endemic Plants of Europe (Anon. 1977). InFrance, D. aemula is strictly confined to western parts of Normandy and Brittany. The remainderof its disjunct mainland continental distribution is thinly scattered along the Cantabrian coast of NSpain and Portugal. The only other known world locations for this fern are on the higher mountains ofthe Azores, the Canary Islands and Madeira (Jalas & Suominen 1972, Map 133; Jermy et al.1978). In Madeira it is described as "frequent amongst rocks, in woods and along levadasthroughout" (ie beside artificial open water channels) (Press et al. 1994).
The fact that Hay-scented Buckler-fern has its world distribution centred and concentrated in the BritishIsles, means that although it is not overall a rare or even a scarce species here in these islands, wedo have a special duty to conserve and manage its sites and study its requirements on the grounds of ourInternational Biodiversity responsibilities.
An interesting species comparision
Examination of both the British Isles and European species distribution maps indicates that there is adefinite similarity between the pattern of D. aemula occurrence and that of Hymenophyllumtunbrigense (Tunbridge Filmy-fern). The match of these two physically very different species isparticularly close within Britain and Ireland, but on the continental mainland H. tunbrigense hasa number of additional stations in E France, SW Germany and NW Italy that are not shared withD. aemula (Jalas & Suominen 1972, Map 69; Tutin et al. 1993). D. aemula andH. tunbrigense both occur on the Sussex Weald and in a number of other disjunct sites in thecooler, more eastern areas of southern England, a fact apparently associated with local pockets of highhumidity in these parts of the country (Jermy et al. 1978; Jermy & Camus 1991).
The habitat requirements of these two very different looking ferns are startlingly similar. Both requirefree-draining yet permanently moist soils, year-round high atmospheric humidity, plus shelter from fullsun and desiccating winds. Both ferns grow in sheltered shade in a similar manner, rooted in peaty, acidsoils on mossy boulders or on mossy rock slopes, forming carpets or curtains of cascading pendulousfronds. They grow rather slowly, and both can also be epiphytic on mossy tree trunks, as they are inFermanagh, eg in the Correl Glen Nature Reserve.
Names
The genus name 'Dryopteris' was first given by the Greek physician, Pedanius Dioscorides (c. 40-90 AD),to a fern growing on oak trees, and is a compound of the Greek 'dryas' = 'oak', and 'pteris' = 'fern'(Gilbert-Carter 1964). The specific epithet 'aemula' is Latin meaning 'striving', 'rivalling' or'imitating' (hence our familiar word, 'emulate'), and presumably this refers to D. dilatata orD. carthusiana which species D. aemula rivals in beauty and competes with in Nature (Step& Jackson 1945; Gilbert-Carter 1964; Gledhill 1985).
Threats
None.
Native in N England, a definite mis-identification here.
1860; Smith, Rev Prof R.W.; "From the vicinity of Brookeborough".
Growth form and preferred habitats
A medium-sized, finely-divided, stiff, upright-fronded, deciduous Buckler-fern with a distinctive dull,greyish-green, mealy surface, this is a rare plant of base-rich rocks, including deep crevices (grikes)in limestone pavement, coarse limestone screes (block screes) and rock crevices where moist, humus-rich,peaty soils develop. D. submontana prefers a degree of shelter from weather and adequateprotection from grazing is essential to its survival, but is intolerant of all but light shade. Ittherefore tends to occur in relatively inaccessible places, such as rock ledges, deeper, wider grikes inlimestone pavement and amid thorny or evergreen scrub that provides shelter and protection.
British Isles occurrence
This is a rare plant confined to a limited area of limestone terrain in the northern English Pennines,although there are also rare outlying stations in N Wales and the NW Midlands.
Fermanagh occurrence
This fern, of which there is just the solitary record listed above, was recorded at the time asLastrea rigidum (= L. rigida (Sw.) C. Presl). However, it must certainly be wronglyidentified, since this very rare, deciduous, calcicole species which demands sheltered, moist,humus-rich soils, has never been found anywhere else in Ireland. Neither Meikle and hisco-workers (who in 1957 and 1975 referred to the plant as Dryopteris villarii (Bell.) Woynar),nor we, can identify what fern might have been taken for this in error by an expert pteridologist likeRev Prof Smith.
Very rare, but a definite error here.
1860; Smith, Rev Prof R.W.; "From the vicinity of Brookeborough".
This very rare hybrid was again recorded from the same site as the species above, the station beingvaguely described and reported by Smith in his 1860 paper in the Natural History Review7(2): 40. Again, as in the case of Dryopteris submontana (Rigid Buckler-fern), withhindsight, Smith was certainly mistaken. Rev Prof Smith was a foremost British fern expert and the firstdiscoverer of many of Fermanagh's ferns, and while he did make a few errors, the fern taxonomy of hisday was very different from ours. We must not lose sight of that significant fact and make unjustcriticism of his mistakes.
The plant would much more likely to have been D. × deweveri (ie D.carthusiana × D. dilatata), since both of these parent species occur in CoFermanagh, and the hybrid between them rarely occurs or is rarely reported elsewhere in Ireland. D.cristata (Crested Buckler-fern), on the other hand, is totally unknown anywhere on the wholeisland of Ireland (Meikle et al. 1975 Revised Typescript Flora). Regrettably we do nothave any other records of D. × deweveri in the Fermanagh Flora Database tosupport this suggestion. The New Atlas and New Fern Atlas hectad map of D. ×deweveri plots a total of just ten symbols of any date for Ireland, so this hybrid is also veryclearly seriously under-recorded on the island (T.D. Dines, in: Preston et al. 2002; Wardlaw& Leonard 2005).
Native, occasional. Eurosiberian boreo-temperate.
1860; Smith, Rev Prof R.W.; Tempo.
Throughout the year.
Growth form, identification and preferred habitats
This deciduous, bipinnate fern is very much a species of wet, peaty, cut-over lowland raised bogs andlakeshore marshes and fens, especially those overlying rich alluvial soils. The lightish-green uprightfronds with long basal stipes are usually produced in sparse, irregular groups (never in tightshuttleco*cks), which makes it easy to distinguish the species even at some distance from the much morerobust and very much more common D. dilatata (Broad Buckler-fern).
The plant has either a short, decumbent rhizome crown (ie reclining but rising at the tip), or a moreslender creeping rhizome, the latter type spreading through wet peat and mossy cushions and branching toproduce new crowns which send up groups of aerial fronds at intervals.
In our Fermanagh experience, there often may be only one or two individual fronds sprouting in 30 cm highvegetation on a large expanse of bog, so that this is a species that must be actively searched out. Onceone has developed an eye for its particular habitat and manner of growth, however, it can be found quitefrequently.
The literature suggests that D. carthusiana occurs as a plant of wet woodlands, usually with analder-willow-birch canopy and with a floor dominated by Sphagnum bog mosses (Page 1982, 1997).However, we do not find it under these conditions anywhere in Fermanagh.
Fermanagh occurrence
In Fermanagh, D. carthusiana has been recorded in 62 tetrads, 11.7% of those in the VC. Seven ofthese tetrads have pre-1976 records only, a proportion that suggests that the habitats this fernoccupies are under threat (see below). As the distribution map indicates, Narrow Buckler-fern is verywidely but rather thinly scattered, mainly across wetter areas of the Fermanagh lowlands.
Reproduction
In addition to vegetative spread, mature fronds produce numerous asexual sori, which spore freely fromJuly to September (Hyde et al. 1969). The fronds are summer-green only, dying and disappearingquickly after the first winter frost (Jermy & Camus 1991; Page 1997). Interestingly, in Scandinavia,Jonsell et al. (2000) suggest that it is the fertile fronds which die off, while most sterileones persist overwinter.
British and Irish occurrence
D. carthusiana is widespread in lowland area of both Britain and N Ireland, especially in Britainsouth of a line between Stranraer and Berwick-upon-Tweed, and in Ireland, north of the Internationalborder with the Republic. However, its frequency has quite rapidly declined throughout these islandssince the 1930s. The decline is perhaps most obvious in Ireland where the species was never all thatfrequently reported to begin with, although there remains an outside and unlikely possibility that thespecies may not be discriminated by sufficient Irish field workers to give an accurate picture (Jermyet al. 1978; Webb et al. 1996; Page 1997). Irrespective of this possibility, there can beno doubt that Narrow Buckler-fern was much more familiar and frequently found by Victorian fieldbotanists than it is today, since it is now a locally frequent to occasional, or even a rare species inparts of the British Isles (Webb et al. 1996; Page 1997).
Being a plant of wet, peaty habitats which naturally follow a transitional pattern of dynamic successiongradually moving towards drier seral stages as organic matter accumulates, it is not surprising thatD. carthusiana populations are eventually eclipsed by these environmental and vegetationalchanges. They are also vulnerable to the much more drastic and rapidly operating effects of artificialdrainage for farming, peat-cutting or other land-development processes which have increasingly affectedlowland wetlands in Britain and Ireland during the last 50 or more years (Jermy & Camus 1991).
European and world occurrence
In continental Europe and W Asia, D. carthusiana is widespread in mid-temperate latitudes of N andC Europe thinning somewhat northwards (although reaching within the Arctic Circle in Scandinavia), andsouthwards to the Mediterranean (Jalas & Suominen 1972, Map 129). Related forms or species occur ineastern N America allowing Hultén (1958, Map 155) to include D. carthusiana (as D.spinulosa) in his amphi-Atlantic group of species. In eastern and central N. America there areclosely related taxa that Hultén and Fries (1986, Map 67), plot as var. intermedia and var.fructosa.
In Scandinavia, D. carthusiana appears to occupy a much greater range of habitats than in Britainand Ireland, including much drier sites such as rock crevices, screes, tall-herb meadows and dunes, aswell as on stone walls and urban situations (Jonsell et al. 2000).
Name
The genus name 'Dryopteris' was first given by the Greek physician, Pedanius Dioscorides (c. 40-90 AD),to a fern growing on oak trees, and is a compound of the Greek 'dryas' = 'oak', and 'pteris' = 'fern'(Gilbert-Carter 1964). The Latin specific epithet 'carthusiana' refers for some unknown reason to theGrande Chartreuse Monastery of Carthusian Monks, near Grenoble, in France (Gledhill 1985).
Threats
Drainage of fens and bogs, and mechanical peat cutting.
Native, common, widespread and locally abundant. European temperate.
1806; Scott, Prof R.; Cuilcagh Mountain.
Throughout the year.
Growth form and preferred habitats
The dark-centred scales on the stipe, the down-turned margins of the pinnules and the dark-green colourof the frond readily distinguish D. dilatata from two much rarer Dryopteris species, D.carthusiana (Narrow Buckler-fern) and D. aemula (Hay-scented Buckler-fern).
The typical habitats of this very common deciduous fern are woods, hedgerows and shaded banks, but italso appears in upland areas on open rocky slopes and in rock crevices and, as a weedy species, in moreurban and waste ground situations. Thus, like D. filix-mas (Male-fern), it occurs in a largevariety of damp, lowland shade or, in upland, more open habitats throughout almost the whole range ofaltitude. The biology, ecology and distribution of these two extremely common ferns in Britain andIreland are very similar and their ecological niches clearly overlap considerably at many shared sites.D. dilatata is more frequently found and is the more abundant of the two species on permanentlywet, but not waterlogged soils, at pH levels below 5.0, on bogs, or on acidic, moderately fertile,organic substrates in woods, scrub, hedgerows and on the banks of rivers and streams.
In mixed deciduous acid woodland generally dominated by oak, D. dilatata can carpet the damp,shady floor vegetation, and developing from massive, old, more or less upright rhizomes it forms adense, mid-green sward of fronds up to 1.5 m tall. In old, less disturbed woods of this type, D.dilatata sometimes also grows as an epiphyte in mosses on the rugged bark of the larger trees inthe same way that Polypodium species very often, and Blechnum spicant (Hard-fern),occasionally does (Page 1997). Like Male-fern, D. dilatata also invades conifer plantations, mostfrequently being found along tracks and fire-breaks, and particularly along the sides of ditches anddrains associated with these less shaded, better drained conditions.
D. dilatata is perhaps slightly less frequent than D. filix-mas in limestone areas ofFermanagh, although in our wet western oceanic climate, an insulating layer of peat regularly forms overbase-rich rocks, and thus plants that are widely regarded as calcifuge can frequently be found alsogrowing in limestone districts.
Of these two common fern species, D. dilatata is also more often found than D. filix-mas inmore open sites on higher ground, such as on more or less steep, rocky slopes, stabilised screes and inrock crevices. Relatively dwarfed plants of D. dilatata are abundant in the clefts between rocksfor instance on the summit of Cuilcagh, our highest mountain.
D. dilatata is such a rapidly growing and maturing fern that in less natural, urban and disturbedhabitats it can also behave like a weed species, colonising crevices in damp brickwork in the mannerD. filix-mas sometimes does, but doing so even more effectively than the latter. It is also quitecommonly found in less well-tended gardens, growing out of soil on steps, competing with decorativespecies in tubs and in greenhouse pots. The pronounced reproductive ability and wide range of variationwithin the species, suggests that D. dilatata is possibly still capable of further increasing itsdistribution and range of habitats within Britain and Ireland (Grime et al. 1988).
Fermanagh occurrence
D. dilatata is almost ubiquitous throughout the county. It is both the most frequent and the mostwidespread fern in Fermanagh being present in 462 tetrads, 87.5% of those in the VC. In the wetterWestern Plateau uplands of Fermanagh, D. aemula tends to replace D. dilatata both in wet,acidic, shallow rocky ground in shade and also as an epiphyte in oak woods, eg in the Correl Glen NR.
Reproduction
Broad Buckler-fern produces sporing sori on all but the smallest plants, a feature rather different fromMale-fern, which instead takes up to six years growth to achieve sporing fertility (Page 1997). Frondsare less frequently wintergreen than those of D. filix-mas, but D. dilatata produces itsfresh annual fronds and sporing sori much earlier in the season than Male-fern. Spores are clearlyproduced in massive quantities, and from the wide range of habitats and geographical spread in theBritish Isles, dispersal is very efficient. Probably it is only the essential requirement for freemoisture to enable the functioning of the delicate prothallial stage which limits the plant and preventsthe even more common occurrence of the species.
A field study by Willmot (1985) found that small sporophyte plants of Dryopteris dilatata andD. filix-mas in woodland, developed in cushions of moss. In the several woods he studied, allpopulations of D. dilatata produced an excess of small, sterile plants over larger, older fertileones, while the age structure in D. filix-mas populations did the opposite. Severalinterpretations of this observation are possible, but a likely one is that Broad Buckler-fern eitherproduces more new sporophytes each year, or that members of this vulnerable stage survive better thanthose of Male-fern under the site conditions studied and are recruited into the mature population moresuccessfully. There appears to be very few field studies of the population behaviour of any fern in theBritish Isles, and Willmot's work urgently requires to be followed up and emulated with other species.
Vegetative reproduction
D. dilatata also has a greater tendency to carry out vegetative reproduction than D.filix-mas; the rhizome of some plants, typically when they are growing on shallow soil overlyingrock, very occasionally produce long, slender, creeping, offset branches which bear a sequence of smallcrowns each producing new fronds (Page 1982; Grime et al. 1988). Page (1997) believes that thisvariant is an environmentally induced form, a suggestion which appears very likely the case.
Toxins
As with Male-fern and other Dryopteris species, D. dilatata is intolerant of grazingpressure and contains the toxic substances thiaminase and filixic acid which can cause blindness and,very rarely, the death of cattle which have eaten the rhizome through lack of other more suitablegrazing material (Cooper & Johnson 1998).
European occurrence
In Europe, the distribution of D. dilatata is very much more limited than D. filix-mas,being rather confined to the western region of middle temperate latitudes, at the same time thinningconsiderably towards the Mediterranean. It does however extend northwards along the Atlantic coast ofNorway, and just reaches the Arctic Circle near Bodo (Jonsell et al. 2000).
World occurrence
Beyond Europe (in the Florae Europaea sense), D. dilatata only occurs (presumably rarely)in Asia Minor and the Caucasus (Jalas & Suominen 1972, Map 130). Related forms occur in N America,and possibly also in the Far East (Hultén 1958). Forms in Greenland and Iceland were previously recordedas D. dilatata (Böcher et al. 1968; Löve 1983), but these have been reassigned to D.expansa (Northern Buckler-fern), an amphi-Atlantic species which is now regarded as one of thediploid parent species of tetraploid D. dilatata (Kristinsson 1987; Grime et al. 1988;Jonsell et al. 2000).
Name
The genus name 'Dryopteris' was first given by the Greek physician, Pedanius Dioscorides (c. 40-90 AD),to a fern growing on oak trees, and is a compound of the Greek 'dryas' = 'oak', and 'pteris' = 'fern'(Gilbert-Carter 1964). The specific epithet 'dilatata' is Latin meaning 'broad' or 'spread out', and isderived from the past participle of 'dilato', itself from 'latus' meaning 'broad' (Gilbert-Carter 1964).As the species in the past was not differentiated from ferns in general, it has no local English commonnames nor any folklore (Step & Jackson 1945).
Threats
None.
Native, common, widespread and locally abundant. European temperate and widely disjunct circumpolar.
1860; Smith, Rev Prof R.W.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
A very characteristic, easily recognised wintergreen, rosette-forming, heterophyllous, strongly calcifugefern of wet, acidic, generally peaty conditions in a wide variety of habitats, B. spicant isespecially common in damp woodlands, both deciduous and coniferous. The species is also commonly foundon more open upland heathy moorland and montane blanket bog, especially along stream and ditch banks insuch sites. Hultén (1962) comments that, "in most parts of its area [ie he is referring here to itswhole range], it is a calcifuge, but this is not always so in Scandinavia." Jonsell et al.(2000) instead regard it as a plant of, "mostly oligotrophic ground [ie nutrient-poor,unproductive]; apparently indifferent to lime."
Ecology
Hard-fern does not tolerate drought, but rather it requires permanently damp, humid, yet relativelyfreely drained, sheltered and usually somewhat shaded conditions for optimum growth and competitiveability (Page 1997). Frequently it becomes locally dominant, forming extensive patches of overlappingrosettes in damp hollows in woodland, or along steep, damp, acid riverbanks, where the depth of shadevaries from light to moderate. Apart from woods, cliffs, stream and roadside banks and old, acidic-rockquarries, the fern is much sparser or absent in lowland habitats, particularly in the areas of betteragricultural soils.
Hard-fern is well named, the fronds being rather leathery, not to say rigid, so that they appearunattractive to all but extremely hungry grazing animals. Putting this another way, the rosettes aretolerant of moderate but not heavy grazing pressure. Management of upland grazing involving cyclicalburning to create a mosaic of young and older vegetation is undoubtedly detrimental to this fern, whichcan be locally eliminated by such practices. (Sinker et al. 1985; Page 1997).
Reproduction
The plant produces separate, quite dissimilar sterile and fertile fronds: the latter are longer and bearmuch narrower pinnae that on the underside bear paired linear sori covered by two long indusia eitherside of the midrib.
Fermanagh occurrence
B. spicant occurs on moors and upland bogs in sheltered spots right up to near the highest levelson mountains in Fermanagh. Very dwarf specimens of Hard-fern grow in sheltered damp hollows, along peatbanks and between rocks close to the summit of Cuilcagh, the highest mountain. B. spicant findsvery many suitable sites in Fermanagh and has been recorded from 366 tetrads, 69.3% of the tetrads inthe VC, making it considerably more frequent and widespread than even Pteridium aquilinum(Bracken) (301 tetrads, 57.0%). It is found almost throughout the VC, but avoids lime and base-richconditions and therefore is absent particularly from some fertile, intensively cultivated lowland areas.
In the very wet oceanic climate of Fermanagh, peat can develop directly over limestone and calcareoussandstone so that B. spicant can occur, but only very locally, in small pockets or on widerstretches of damp, acid, organic soil, even in what appears from the map or general appearance to begeologically unsuitable limestone terrain.
British and Irish occurrence
Hard-fern is very common and widespread throughout most of the British Isles, especially in the wetter Nand W areas. It is much less prevalent, or indeed absent, in parts of the east and midlands on bothislands. It is most markedly absent on the clay, chalk and limestones of S England. B. spicanthas contracted to an unknown extent in the Irish and English Midlands, probably due to a combination offactors causing habitat loss, including woodland clearances, destruction of lowland heathland to createimproved pasture, general intensification of farming practices, building development, and industrial anddomestic air pollution; it is sensitive to all of these (Jermy et al. 1978; Jermy & Camus1991; Page 1997).
European occurrence
B. spicant is widespread in W and C Europe, oceanic conditions allowing it to spread northwards upthe coast of Norway to within the Arctic Circle. It is also found on the Atlantic islands (Iceland, theFaeroes, Azores, Madeira and the Canaries). Towards the Mediterranean it becomes more dispersed andlocal, but it is found (however rarely), on the southern tip of the Iberian Peninsula, Corsica,Sardinia, Sicily and Crete. There appears to be considerable doubt about its presence on the BalearicIsles (Tutin et al. 1993) and Jalas & Suominen (1972) do not map it there. In SE Europe, itreaches, but is scarce and local, in Turkey and in Asia Minor (Jalas & Suominen 1972, Map 139; Page1997).
World occurrence
Beyond Europe, B. spicant has a very widely disjunct discontinuous range in middle temperatelatitudes around the northern hemisphere, occurring locally in N Asia, Japan (where a var.nipponicum (Kuntze) Miyabe & Kudo is recognised), Alaska and the Eastern Pacific states of NAmerica (Hultén 1962, Map 143; Hultén & Fries 1986, Map 73). In the opinion of the current author(Ralph Forbes), the distribution is so extremely disjunct it really is stretching the concept tobreaking point to refer to it as circumpolar, but in some heavily qualified context it might fit thisdescription.
Names
The genus name 'Blechnum' is derived from a Classical Greek fern name 'blechnon', which in view of therarity of the modern species of this name in the Mediterranean area, probably was not applied to thesame plant at all (Gilbert-Carter 1964; Stearn 1992). The Latin specific epithet 'spicant' means'tufted' or 'spiked', probably referring to the often shuttleco*ck manner of growth of the whole plant,or to the relatively rigid, spike-like appearance of the pinnae (Johnson & Smith 1946; Step &Jackson 1945; Gledhill 1985).
Alternative local English common names include 'Deer Fern', 'Foxes Fern', 'Herrin'-bone Fern' or'Fishbone Fern' (the latter two both being fitting names, especially when applied to the shape of thedistinctive fertile frond), 'Rough Spleenwort' and 'Snake Fern' (Britten & Holland 1886; Step &Jackson 1945).
The species is not very variable, but a small number of varieties of the fern, including a crested form,are quite commonly grown in gardens.
Threats
B. spicant is susceptible to the more intensive agricultural practices and associated habitatchanges, eg heather burning, bog drainage, removal of hedges and woodland felling. On the other hand, itinvades coniferous plantations, where it thrives along the sides of firebreaks and drainage channels. Onbalance it is holding its own and is much too common to be under any immediate threat. However, changesin the management of upland areas could affect this species either way.
Introduction, neophyte, deliberately planted, rare but certainly often over-looked or ignored andtherefore under-recorded.
24 July 1986; Northridge, R.H. & Forbes, R.S.; Gubbaroe Point, shore of Lower Lough Erne.
June and July.
Self-sown and naturalised or deliberately planted, this is certainly under-recorded in Fermanagh since wetend to walk along with crooked necks inspecting the ground rather than looking up at the trees!Recording introduced, deliberately planted alien trees was certainly not regarded as a worthwhileexercise by earlier field workers including Meikle and his co- workers, and indeed many such treesremain ignored to this day, even when they might be capable of occasional naturalisation. A. albais a case in point, it being able to regenerate freely from self-sown seed in mixed woodland on fertilesoils. Saplings can develop even under a woodland canopy (M.E. Braithwaite, in: Preston et al.2002). A native of the mountains of C & S Europe (Jalas & Suominen 1973, Map 152) A. albawas first introduced to the British Isles in 1603. Formerly it was widely planted as a specimen tree ingardens and in timber plantations, especially in the N & W of these islands (Mitchell 1974; Stace1997). Interestingly, it was first recorded from the wild in the British Isles as late as 1914.
Fermanagh recorded occurrences
In Fermanagh, we have only four records of A. alba in semi-wild situations on wood margins andthickets. Even with these few records, we have no definite evidence and cannot be absolutely certainthat they are self-sown rather than planted. There is evidence of other alien trees being planted atCladagh River Glen for instance. The details of the other three stations are: two records by D.M. Smith,W. McKenna & Ms E. Kennedy in July 1990, in Glen Wood, Florencecourt and Corry Point Wood FNR, LowerLough Macnean; and Cladagh River Glen NR, 6 June 1991, Ms B. Hamill & M. Bradley.
Other Silver Firs
The other Abies species most likely to be found self-sown and naturalised in parts of Britain andIreland is A. procera (Noble Fir), although we do not as yet have any records for it inFermanagh. It is a very handsome tree with blue-grey foliage and it is frequently planted as a singlerow around blocks of forestry plots (including some in the Lough Navar Forest Park), to make theplantation more attractive in appearance.
A. alba is now very seldom planted because its foliage is more susceptible to rust fungusand woolly aphids than other Silver Firs, eg A. grandis (Giant Fir) and A. procera. Inamenity plantations, A. alba has been largely replaced by A. nordmanniana (Caucasian Fir),of which Forestry Service timber trials are also under way. In the last 20 years or so, A.nordmanniana has also begun to oust Picea abies (Norway Spruce), as the most popular'Christmas tree', since unlike the latter it does not drop its leaves indoors over the holiday period(Stace 1997). The gaunt 'stag-head' tops of surviving old trees of A. alba are often rathernoticeable from a distance in estate parks and woodlands.
Like the other species of Abies and indeed all exotic conifers, A. alba is inconsistentlyrecorded in the flora of Britain and Ireland, making it very probably seriously under-recorded (M.E.Braithwaite, in: Preston et al. 2002).
Names
The genus name 'Abies' is from the Latin 'abire' meaning 'to rise', which is considered by some to referto the great height some species of the genus can attain under good growing conditions (Hyam andPankhurst 1995). The specific epithet is Latin meaning 'white', possibly referring to the distinctivewhitish stomatal bands on the undersurfaces of the leaves.
Threats
None.
Introduced, neophyte, rare but much ignored and under-recorded.
1986; Waterman, T. & Brain, P.J.T.; lakeshore woods at Knocknabrass Td to Doohat Td, Upper LoughErne.
June to August.
Self-sown and naturalised frequently throughout B & I (Stace 1997), or deliberately planted, butagain as with other introduced conifers, very often ignored by field botanists including those inFermanagh even though in these islands it is the tree planted in numbers greater than all the otherscombined! In forestry plantations, it outgrows and out-yields every other tree species on vast tracts ofpoor, degraded wet mountain sites. It is also one of only three tree species to exceed 60 m in height inthe British Isles (so far, we might add, since some introduced species are still too young to have madethis sort of growth) (Mitchell 1996).
Fermanagh recorded occurrences
In Fermanagh, it has been recorded so far in just 21 tetrads in woods and lakeshores on islands in UpperLough Erne and thinly scattered in the near vicinity of obvious plantation woods from which seed musthave originated. As the distribution map shows, most of these plantations lie close to the countyboundary. The other record details are: Inishleague Island (West), 17 June 1987, R.S. Weyl & Mrs J.Whatmough; Inishlught Island, 1987, R.J. Bleakley; Glen Wood, Florencecourt, 1990, D.M. Smith, W.McKenna & Ms J. McConnell; Cornagague Lough, 1991, W. McKenna & Ms J. McConnell; Knocks Td, ENEof Lisnaskea, 1995, RHN & RSF; 1 km E of Eshcarcoge Td, August 1995, RHN & RSF; shore atplantation, 1.5 km NW of Garrison, July 1996, EHS Habitat Survey Team. RHN & HJN added six morerecords in autumn 2010 as follows: Cashel Crossroads; N of Rotten Mountain bridge; Meenawanick NE ofBrickagh; SW of Tonymore; Corraleek SE of Garrane; and Tullykeeran Td, Pettigo Plateau.
Names
'Picea’ is the classical Latin name of the genus and is derived from the Latin 'pix' meaning 'pitch', areference to the resin obtained from the tree
(Hyam & Pankhurst 1995). The specific epithet 'sitchensis', means 'of Sitka', geographical referenceto a borough and city in Alaska, NW America.
Uses
Although popularly denigrated and even vilified by "the undiscerning public" and "greenconservationists", as creating, "huge sterile blocks of alien conifers blanketing thebeautiful moors in ugly rows of uniform dark green", Mitchell (1996, p. 91), from whom these arequotes are taken, puts up a strong case for the virtues of Sitka Spruce, especially in terms of the manyspecies of birds it supports.
Threats
Sitka Spruce is another species susceptible to the cankerous killer disease caused by the readilytransmitted fungal pathogen Phytophthora ramosum which is currently spreading in Britainand Ireland.
Introduced, neophyte, usually or always deliberately planted, occasional, but ignored and thereforeseriously under-recorded.
1975; Faulkner, Dr J.S.; Marble Arch/Cladagh River Glen NR.
April to August.
Norway Spruce, the very familiar Christmas tree, is not much planted today as it was formerly in NI,being very much overtaken as a timber tree by Picea sitchensis (Sitka Spruce). This isparticularly the case on poorer, wet and more exposed upland soils. It is also unsuited to dryconditions or deep peat. Having said that, it is still not as frequently recorded in Fermanagh as itprobably should be since farmers carrying out shelter-belt or private forestry plantation often plantsurplus trees into their hedges. It seeds freely and can colonise open, disturbed ground and adjacentheath or moorland, provided grazing animals allow its survival (Stace 1997).
In Fermanagh, we have records of it from just eleven tetrads, mainly around the Upper Lough Erne basinand, as the distribution map shows, scattered very thinly elsewhere. It has been recorded in Fermanaghon lakeshore and riverside woods, plus along roadside hedgerows, but botanical recorders are oftenunfamiliar with exotic conifers and tend to ignore the trees, considering them obviously planted and ofnegligible conservation interest. Norway Spruce is not a long-lived species and very often specimens inthe wild are stunted and horribly deformed by poor growing conditions.
One of the most interesting things about P. abies is its rather peculiar natural distribution. Itis a widespread Eurasian boreal-montane species, yet with the exception of a tiny outlier on the HarzMountains in N Germany it is entirely absent from SW Europe (Jalas & Suominen 1973, Map 157). Thisdistribution makes it clear that it could not and did not migrate back into the British Isles after thelast Ice Age, although fossil evidence proves it had done so in previous warm Interglacial periods(Mitchell 1996; M.E. Braithwaite in: Preston et al. 2002).
Names
'Picea’ is the classical Latin name of the genus and is derived from the Latin 'pix' meaning 'pitch', areference to the resin obtained from the tree. The specific epithet 'abies' is derived from the Latin'abire' meaning 'to rise', which is considered a reference to the considerable height some species ofconiferous trees can attain under good growing conditions (Hyam and Pankhurst 1995).
Threats
None.
Introduced, neophyte, rarely recorded but often ignored and therefore under-recorded.
1968; unnamed recorder; Marble Arch/Cladagh River Glen NR.
July to November.
As with other introduced conifers, this deciduous conifer is often ignored by field recorders as being oflittle or no interest, it being generally assumed, often perfectly correctly, to be deliberatelyplanted. The four records that belong here are identified only as being a form of larch, but veryprobably they are the hybrid larch, Larix decidua × L. kaempferi (seebelow).
Introduced, neophyte, rarely recorded but often ignored and therefore under-recorded.
15 August 1986; Waterman, T. & Brain, P.J.T.; Kilturk Lough, Killalahard Td, Upper Lough Erne.
May to October.
Although this deciduous conifer, which is endemic to the Alps and the Carpathian Mountains (Jalas &Suominen 1973, Map 161), can readily self-seed in the British Isles, in the few sites where we know ofthe species in Fermanagh, it is almost certainly planted. Even more definitely, however, it isunder-recorded in the county, there being records from only nine tetrads. The Fermanagh records are fromlakeshore and riverside woods, mainly sited around the larger, lowland lakes, most of them undoubtedlyestate plantations (details below).
The late Alan Mitchell, a forester and major authority on the identification and measurement of Britishtrees described European Larch as being, "among the most valuable and decorative of all the treeswe grow" (Mitchell 1996). L. decidua was introduced to B & I probably just before 1629.John Parkinson knew of it as a rarity, but he had never seen it cone (Parkinson 1629). The tree wasfirst recorded from the wild in Britain in 1886, although before that date botanical field recordersvery probably had simply ignored it (M.E. Braithwaite, in: Preston et al. 2002).
Though L. decidua is still occasionally planted for its excellent timber, it is unfortunately verysusceptible to canker, so that more disease resistant larches, and especially the hybrid L. ×marschlinsii (Hybrid Larch), which arose around 1897 as a cross between European Larch and theintroduced L. kaempferi (Japanese Larch), are now very much more preferred (Mitchell 1996).
Fermanagh recorded occurrences
The details of the remaining eight records are: W end of Inishleague Island, Upper Lough Erne, 17 June1987, R.S. Weyl & Mrs J. Whatmough; shore of Lower Lough Erne, N of Strahenny Point to Temple Hill,Rossfad Td, 13 July 1987, RHN; wood on Doocharn Island, Upper Lough Erne, 1 October 1987, B. Nelson;Glen Wood, Florencecourt, July 1990, D.M. Smith, W. McKenna & Ms E. Kennedy; Correl Glen woodland, 7May 1992, J. Farren & T. Waterman; cutover bog 1 km NE of Dresternan Lough, towards Rosslea, 4August 1995, RHN & RSF; Bilberry Island, Lough Melvin, 17 July 1996, EHS Habitat Survey Team; foresttrack, Tullyvocady Td, N of Derrin Mountain, 26 October 2010, RHN & HJN.
Names
'Larix' is the classical Latin name for this tree, Larix decidua, and the specific epithet'decidua' denotes that it is deciduous, somewhat unusual for a coniferous tree.
Threats
A leaf blight and canker-causing fungal disease organism, Phytophthora ramorum, potentially fatalto a range of woody species including both Japanese and European Larches, is actively spreadingnorthwards in Britain and Ireland at present. Between 2002-2009, the disease was found at 34 sites in NIreland, mostly on Rhododendron and other ornamental species at sites which included plantproduction/retail premises, private gardens, private estates and public parks. All outbreaks weresuccessfully controlled. The disease first attacked Japanese Larch in eastern N Ireland in 2010. It hasnot yet been found on Hybrid Larch. A second fungal species, P. kernoviae, has also appeared,although so far it is attacking mainly Rhododendron. These pathogens can be spread on footwear, vehiclewheels, tools and machinery, by the movement of infected plants and in rain, mists and air currents.Felling and burning of infected trees is the only effective containment measure known.
Introduction, neophyte, rare but often ignored and therefore under-recorded.
2 August 1989; Forbes, Dr R.S.; roadsides around Gortaree district.
April to September.
A F1 hybrid between L. decidua (European Larch) and L. kaempferi (Lamb) Carrière(Japanese Larch), L. × marschlinsii Coaz first arose in Scotland in1897. Nowadays, it is much more widely planted for timber than the European Larch since it is bothfaster growing and much more resistant to Larch canker. The hybrid is fully fertile and regularlyregenerates from seed. It also back-crosses readily with either parent, although the European species isbecoming increasingly rare.
While the F1 plants are greatly preferred for forestry and are widely planted in the BritishIsles, the hybrid was not recorded from the wild until 1983. The distribution of the hybrid is verysimilar to that of L. kaempferi, both plants being very widespread but quite definitelyunder-recorded (Mitchell 1996; M.E. Braithwaite, in: Preston et al. 2002).
Fermanagh recorded occurrences
There are only three records of this hybrid in the Fermanagh Flora Database. The details of the other twoare: roadside near Scarfield Bridge, Colebrooke River, 12 April 1996, RHN & RSF; and DerrysteatonTd, Galloon Island, Upper Lough Erne, 1 September 2001, RHN & RSF.
Names
The Latin specific epithet 'kaempferi' is given in honour of the late 17th century Germannaturalist, physician and traveller Engelbert Kaempfer, who explored regions from Russia to Japanbetween 1683 and 1693 and wrote both a flora and a history of Japan. The specific epithet 'marschlinsii'is another genitive case meaning 'of Marschlins', possibly or apparently referring to an area inSwitzerland that seems most famous for its castle.
Threats
This hybrid faces the same threat from fungal attack as L. decidua.
Introduction, neophyte, occasional, but usually obviously planted. As a native, Eurasian boreal-montane.
1882; Barrington, R.M.; Co Fermanagh.
Throughout the year.
British and Irish occurrence
A very commonly grown, extremely familiar conifer Scots Pine is almost always deliberately planted inFermanagh as is generally the case elsewhere in most of Britain and Ireland. The fossil record provesthat P. sylvestris was present throughout both islands early in the Post-glacial period, havingrapidly spread north as glaciers retreated. Nowadays, although remaining very widespead, it is regardedas native only in the Caledonian forests of the eastern Scottish Highlands (mainly Deeside andSpeyside)(Proctor 2013).
Fermanagh occurrence
In Fermanagh, very occasionally we suspect it of seeding itself spontaneously, and when this occurs it isalmost always close to habitation or to plantation woods. Scots Pine is still by far the most commonlyfound gymnosperm in Fermanagh, having been recorded in 155 tetrads, 29.4% of those in the VC. While nolonger widely planted by the Northern Ireland Forest Service for commercial timber, it is still activelyplanted by landowners in plantation woods, on boundaries and for wind-breaks. It is widespreadthroughout Co Fermanagh, in woods, hedges and rocky glens, but is especially frequent in the east wheremore intensive farming takes place. Such trees are capable of regeneration and their seed may disperse,self-sow and become naturalised.
Status in Ireland
Although P. sylvestris was present in Ireland after the last Ice Age and was thus originally anative species, it was over-exploited and died out around the Sub-Boreal to Sub-Atlantic periods (ie3000 BC to around 1500 AD). It was subsequently re-introduced in the early 18th century using Scottishseed (McCracken 1971), so that nowadays it is generally considered native nowhere in Ireland (Carlisle& Brown 1968; Mitchell 1986).
Fossil record and Uses
There has to remain some slight doubt on this interesting matter, however, since the fossil record showsdifferences in the decline of pine in separate parts of the island. It had disappeared from NE Irelandby around 2000 BC, but seems to have survived in the SW in marginal, exposed sites with poor soils untilaround 200 AD. The fossil record shows that whenever bog surfaces became relatively dry, pine couldinvade them, and we know that some (probably stunted), trees were still growing on Midland Irish raisedbogs at around 300 AD (Mitchell 1986). It probably also hung on into historic times in other areas ofIreland with exposed conditions and dry limestone soils, such as the Burren, Co Clare (H9), and onheaths. An early Irish Law text from the eighth century AD survives and it lists the penalties forunlawfully interfering with trees and bushes. Evidence form this text indicates that pine, which wasgreatly valued for its resin and was used for making pitch to caulk boats, was still fairly common inIreland in the eighth and ninth centuries AD (Kelly 1997, p. 383). The question of pine survival is anacademic one, however, and to all intents and purposes, the botanical and associated entomologicalevidence points to P. sylvestris either becoming extinct or surviving in such minute numbers thatit can no longer be represented in the gene pool of present-day pines in Ireland (Webb & Scannell1983; Speight 1985).
World occurrence
Scots Pine is a polymorphic Eurasian species and it is the first or second most widely distributedconifer in the world: it’s rival for this distinction is Larix sibirica (Siberian or RussianLarch). The natural range of P. sylvestris stretches from beyond the Arctic Circle in Scandinaviato southern Spain, and from western Scotland to the Okhotsk Sea in eastern Siberia (Hultén & Fries1986, Map 80). Within this range it grows at elevations from sea level to 2,400 metres (8,000 feet), theelevation generally increasing from north to south.
A forester’s verdict: In his last and postumously published book, Alan Mitchell's Trees ofBritain, the author explains that at forestry school Scots Pine was known as "the facilesnare", because it caught out the uninitiated tree planter. The point being made is that Scots Pineis easy to raise and it establishes well on a wide range of soils and sites, including some of the mostdifficult, but in almost all circ*mstances it will be outgrown and out-yielded by other timber species."It is unable to exploit better sites and even at its best is a smaller and slower growing treethan any other used in forestry." (Mitchell 1996, p. 118).
Threats
None.
Introduction, neophyte, very rare but very probably under-recorded.
25 May 1988; Northridge, R.H.; Carn Hill near Drumcullion.
Fermanagh occurrence
This evergreen conifer is widely used in forestry plantations in Fermanagh, but it has only beenrecorded in the county on seven occasions as a naturalised escape from cultivation. The first case wasgrowing on the margin of a cut-over bog and on the adjacent roadside at the site listed above. Twofurther examples of self-sown plants alongside forest tracks were noted by RHN & HJN at TullykeeranTd, Pettigo Plateau, 4 October 2010 and Tullyvocady Td N of Derrin Mountain, 26 October 2010. Othersites include near Moysnaght, and in two places around the shores of Lough Vearty where RHN recorded inDecember 2010.
British and Irish occurrence
Elsewhere in these islands, Stace (1997) mentions P. contorta being self-sown from only two areas,Cardiganshire (VC 46) and the Scilly Isles (VC 1b), but clearly it does so also in Fermanagh. The NewAtlas hectad map now shows that P. contorta has been quite frequently and widely recordedacross both Britain and Ireland, but especially so in the N & W regions of both islands.
Between 1945 and 1980, the two forms of this species were the second most abundant trees planted inBritain and Ireland, being used in re-afforestation to raise the planting limit above 100 m, and toextend planting to deep peat soils. The Shore Pine (var. contorta) was first introduced toforestry by A.C. Forbes of the Irish Forest Service, who discovered it as a rogue growing among a batchof seed of Pseudotsuga menziesii (Douglas Fir) imported from western N America in about 1920. Henoticed that this form made good strong plants, but later it was found that when grown in the veryexposed sites to which they appeared to be ideally adapted, Shore Pine soon became bowed at the base,the effect of strong wind on their very rapidly growing young stems.
Subsequently this led to growth trials of the inland form of the species, var. latifolia(Engelmann) Critchfield (Lodgepole Pine), and it is this particular variety which today forms much ofthe maturing pine forest in plantations, especially in the wetter, cooler N & W parts of Britain andIreland (Mitchell 1996).
Introduction, neophyte, very rare, but possibly over-looked.
15 October 1987; Waterman T.; lakeshore, Inish Rath Island, Upper Lough Erne.
There is just a solitary record of this commonly planted conifer in the Fermanagh Flora Database, made ona DOE field survey of Upper Lough Erne. It was probably self-sown on the lakeshore, but might have beendeliberately planted to provide shelter for other saplings. This dark, funereal, shrubby evergreen treeis all too commonly planted in parks, gardens, churchyards and shelter-belts (Mitchell 1996). It growsvery vigorously and in Britain and Ireland frequently regenerates from seed, plants quickly becomingnaturalised on open habitats such as along banks, by walls, on woodland margins and lakeshores.
A native of NW America (eg California and Oregon), C. lawsoniana was first introduced to theBritish Isles in 1854 and is now represented in horticulture by a huge range of cultivars (Griffiths1994). It is much less frequently planted in rural than in urban areas, but it is sometimes recommendedfor shelter-belts and for under-planting as a nurse species in conifer plantations.
Nowadays, Lawson's Cypress is increasingly widespread in the wild, but was not recorded as such until aslate as 1958. It appears to be much less frequently recorded in unplanted sites in Ireland than it is inBritain (M.E. Braithwaite, in: Preston et al. 2002).
Native, rare. Circumpolar boreo-temperate.
1904; Praeger, R.Ll.; SW portion of the hill above Doagh Lough.
Throughout the year.
Growth form and preferred habitats
This evergreen, dioecious conifer varies from a prostrate, dwarf in exposed upland and coastalsituations, to an upright shrub up to 10 m tall usually growing in damp, lakeshore or riverbank sitesbut it can also occur over limestone, chalk and slate soils on heaths, moors and in pine and birchwoodlands.
The prickly, spreading needle-like leaves borne in whorls of three represent so-called 'juvenilefoliage', and the subsequent adult foliage is comprised of tightly appressed overlapping scale leaves inopposite pairs, resembling the genus Cupressus (Cypress). All, or almost all forms of juniperoccurring naturally in Britain and Ireland produce only more or less prickly, linear, juvenile foliage.Fertilised female cones ripen in their second or third year, the overlapping scales swelling andbecoming fleshy to form the familiar 5 mm, blackish globular edible berry-like fruit.
Fermanagh occurrence
J. communis has been recorded in a total of 16 Fermanagh tetrads, 14 of which contain post-1975records. They probably constitute juniper's main stronghold in Northern Ireland following its decline inCo Antrim in the 1980s and 1990s (N Ireland Vascular Plant Database 2002).
Response to grazing pressure
The juniper population decline in Co Antrim is very probably a direct result of the drastically increasedgrazing pressure on the Antrim plateau due to European Community sheep headage payments in the 1970swhich encouraged overstocking of upland pastures.
In the Republic of Ireland, a similar outcome has been found in upland grasslands in Connemara, causingpeat erosion and a loss of species diversity (Bleasdale & Sheehy Skeffington 1995). Studies on J.communis conservation and regeneration in the English Chiltern downlands, the Lake District andin the Scottish highlands, all indicate that the species is sensitive to grazing pressure, particularlyby sheep (Fitter & Jennings 1975; Ward & Lakhani 1977; Miles & Kinnard 1979a & b;Dearnley & Duckett 1999).
British occurrence
Juniper is a locally common and very variable shrub in various areas of Britain over a wide range ofsoils and habitats, but it is most widespread in the north and west of the country and is especiallyfrequent in Scotland. It occurs in NW Wales and is also rather thinly scattered in southern England,where there have been many lowland extinctions over the last century or so (Ward 1981; Preston etal. 2002).
Reproduction
In common with most other species, juniper requires open, bare ground or very short turf for regenerationfrom seed to occur, and while the necessary vegetation gaps are usually present on steep slopes, on morelevel terrain openings that might permit colonisation are generally provided by heavy grazing pressure,or more rarely by fire. After germination takes place it is vital for the establishment and longer-termsurvival of juniper seedlings that they are allowed to grow on under managed conditions providing aprotected, considerably lightened grazing regime.
With respect to seedling survival, the seasonal timing of sheep browsing, as well as the extent ordegree of grazing pressure, has been shown to be significant in chalk downlands in Oxfordshire. Autumnand winter grazing increases juniper seedling mortality and stunting, whereas summer grazing of thegrassland is apparently beneficial: the sheep having sufficient browsing material, simply leave theyoung juniper plantlets alone, and their grazing curtails competition from grasses, tall herbs andhawthorn scrub without killing the juniper (Fitter & Jennings 1975).
Variation
In Fermanagh, all juniper shrubs are confined to the limestone areas of the county, on scarps, scree,pavement and rocky grassland. It is thus restricted to the Monawilkin, Hanging Rock, Marlbank andFlorencecourt areas of the county. Although the growth form in Fermanagh is invariably prostrate, theplants are almost certainly exposed ecotypes of subsp. communis. In truth, however, this form ofthe plant is very difficult to distinguish from subsp. nana (Hook.) Syme in many parts ofIreland, and probably the same is true elsewhere. It is therefore possible that the latter may alsooccur in some of our most exposed Fermanagh sites (Webb & Scannell 1983; Dearnley & Duckett1999).
In many places in Fermanagh, juniper grows on scarps and rocky ground closely associated with Taxusbaccata (Yew). Some individual juniper plants cover several square metres, and since prostrategrowth forms of the species are very slow-growing, this suggests they might possibly be ancient clones.Studies elsewhere in Britain, however, have shown that juniper is a notoriously difficult species toaccurately age without cutting live samples of the stem – a destructive process and something we couldnot justify under any circ*mstances. The difficulty in measuring the age of individual specimens arisesbecause, firstly, the stem diameter of J. communis is not closely related to age, and secondly,the stems are usually eccentric in shape. This makes sampling for girth measurements or taking cores toring count for age an inherently inaccurate process (Dearnley & Duckett 1999).
Seed viability
The maintenance of high seed viability has been shown in several studies in England to be important forthe conservation of J. communis (Ward 1989). In the English Lake District, a study found thatsites with large populations of juniper (ie more than 1,000 bushes) had significantly higher seedviability than those in small populations, and a reference site which had for 70 years been protectedfrom sheep grazing, produced the greatest juniper seed viability index of all (Dearnley & Duckett1999).
As far as we are aware, no study has yet been made of the ability of juniper in N Ireland to regenerate,and in the light of the obvious contraction of the species on the Garron Plateau in Co Antrim, not tomention the question of the effect on this northern-montane species of Global warming (or as we havecome to experience it over the last decade - Global wetting and winding!), conservation backgroundresearch work needs to be carried out to ascertain the threats to the species, and its ability tosurvive at all its northern Irish sites under current management practices and levels of disturbance.
European occurrence
J. communis is very widespread over most of northern and western Europe, while to the south itbecomes increasingly rare and mainly a mountain plant (Jalas & Suominen 1973, Map 181).
World occurrence
A very variable, polymorphic species, Hultén & Fries (1986, Map 82) recognised six subspecific taxa(either varieties or subspecies), and they plotted the total species distribution as almost completelycircumpolar.
Toxicity and uses
Juniper 'berries' are the ripe female cones of the plant, and they contain high levels of resin (10%), anunnamed essential oil, plus terpene derivatives and a bitter substance (probably an alkaloid), which hasbeen given the name 'juniperine' (Cooper & Johnson 1998). The berries have a long history ofculinary use, as a flavouring of both meat dishes and gin. The plant is poisonous if eaten in quantity,being particularly dangerous to pregnant animals since it can cause severe cramps and even abortion.Fortunately, however, no specific cases of such poisoning have been reported in Britain or Ireland(Cooper & Johnston 1998).
Names
The genus name 'Juniperus' is Latin not Greek, being a name first given by Virgil to the plant (ChicheleyPlowden 1972). The Latin specific epithet 'communis' means either 'common' or 'clumped', ie 'growing incompany' (Gledhill 1985).
Threats
Apart from excessive grazing pressure throughout its range in Northern Ireland, a couple of Fermanaghsites on the Marlbank could be threatened by improvements for agriculture. Elsewhere in parts of bothBritain and Ireland there has been loss of scrub habitat suitable for juniper due to burning of moorsand heaths, succession to woodland and afforestation (M.E. Braithwaite, in: Preston et al. 2002).
Native, occasional. European temperate.
1739; Henry, Rev W.; Hanging Rock NR.
Throughout the year.
Growth form and preferred habitats
A widespread European native and widely cultivated dioecious evergreen tree casting a very dense shade,Taxus baccata L. s.s. (Yew) is one of eight species that formerly comprised T. baccata L.s.l. (Hultén & Fries 1986). The linear leaves are arranged in two lateral ranks and the seed issurrounded by a dull red, fleshy, cup-like edible aril. The Yew tree is unusual for a native conifer, orrather, a 'Taxad', in Britain and Ireland, in being both very slow growing, and also very long-lived(see below). Most particularly, it is unusual through being capable of both vegetative reproduction bylayering, in addition to sexual increase and dispersal by seed production (Milner 1992).
It is a small tree in stature when compared with many of its relatives, reaching only a maximum of around20 m in height, and very often achieving only half this measurement under natural or semi-naturalgrowing conditions. While it is of restricted stature, the Yew can spread its canopy extremely wide inrelation to its vertical measurement. On limestone pavement, especially if grazed by goats, Yew can alsorarely appear as a prostrate, mat-like shrub.
Sexual reproduction
T. baccata is usually dioecious, having separate male and female trees. Very occasionally,however, monoecious plants are recorded. For instance, an otherwise female tree may bear one or a fewbranches with small, yellow male cones, rendering the tree monoecious (Nelson 1981; Nelson & Walsh1993). The female reproductive structure is produced on a short side bud and each consists of aninsignificant fleshy disc with a single central ovule (Ross-Craig 1967-70, Part 27, Plate 45; Milner1992). Pollen release takes place between February and April, wind pollination being the rule, althoughhoney bees do frequently visit male cones to collect the early season pollen that they need to feedtheir developing brood. Although native Yew trees are hardly all that common, thanks to the frequency ofplanted specimens the amount of pollen the male trees release is so great that the species is in the topten plants for pollen abundance in Britain (Milner 1992).
From August to October, the ripe red fruit of the Yew is unmistakable (except possibly for aninsect-induced gall that attacks some other alien taxads), consisting of a single, smooth,brownish-purple seed surrounded or embedded in a fleshy, sweet, edible red or pink translucent aril thatattracts birds and other animals. (The aril is an outgrowth of the seed coat, which actually is an extraintegument layer of the ovule) (Holmes 1979; Lang 1987).
Fermanagh occurrence
T. baccata is one of the very few Fermanagh plant species for which we have records dating before1800. William Henry recorded it from both the Hanging Rock area and from around Upper Lough Macneanprior to 1739 (Henry et al. 1987). Rutty (1772) also recorded it, "from the islands of LowerLough Erne". Yew trees, which are certainly native rather than planted specimens, still do occurboth on the cliffs and rocky slopes of Hanging Rock Nature Reserve and on the shores and islands ofLower Lough Erne, though there are no recent records from Upper Lough Macnean. Having said that,probably there are more planted Yew trees in Fermanagh than naturally arising specimens, a situationthat is now common throughout these islands (Nelson & Walsh 1993).
T. baccata has been recorded from 64 Fermanagh tetrads, 12.1% of those in the VC. As a nativespecies it occurs on the old, pre-drainage shores of Lough Erne and on limestone cliffs and steep scarpsin the Monawilkin, Knockmore and Florencecourt areas. Elsewhere, the species has either been planted oris bird-sown in hedges. As the tetrad distribution map shows, it is quite widespread. The FermanaghFlora Database records include some planted trees in demesnes, gardens and graveyards, together withtrees in hedgerows which undoubtedly are bird-sown.
The tetrad map indicates that the main area for the species in Fermanagh lies within the region withlimestone soils, particularly around Lower Lough Erne and the upland limestone plateau lying to thesouth of it. However, the species is not confined to well-drained, lime- or base-rich soils, although itmay prefer them, rather it also grows on more acidic terrain. It occurs in mixed deciduous woods overlimestone rock, mainly but by no means exclusively associated with ash and hazel.
British and Irish occurrence
While the BSBI's 1962 Atlas attempted to record and display the predominantly native occurrence ofYew in Cumbria, S England and Wales, together with a very thin and mainly coastal occurrence in Ireland(Walters & Perring 1962), the editors of the four year (1996-1999) survey for the New Atlasfound that it proved impossible to distinguish native from introduced trees, and consequently theypublished a map which treats all Yew records as native (Preston et al. 2002). Even with thislimitation, while the New Atlas hexad map shows the species widespread throughout the BritishIsles, it remains most prevalent south of a line between Carlisle and Newcastle, while in Ireland it isthinly scattered throughout (M.E. Braithwaite and M.J. Wigginton, in: Preston et al. 2002).
European occurrence
The Yew tree is sensitive to frost, which limits its northern distribution both in Scotland and elsewherein W Europe, since it is confined in Scandinavia to southern coastal districts of Norway and Sweden(Jonsell et al. 2000). On the continent it is quite widespread in C and S regions, but strangelyabsent from most of France, N Germany and Denmark (Jalas & Suominen 1973, Map 194).
Historic timber uses
Yew is the tree most closely linked with history and legends throughout the British Isles. In particular,probably on account of large dimensions and supposed great age, the tree has been associated withnotions of immortality and with religious and/or revered burial sites dating back well into Druidical orindeed Neolithic pre-history. Some of the notions associated with the tree continue up to the presentday (Mitchell 1996). In early Christian Ireland, Yew was rated in an eighth century law text along withjust six other trees as a 'Noble of the wood'. Apparently it rated so highly because Yew wood was thepreferred material for domestic vessels, such as eating utensils (Kelly 1997, pp. 380-3). In addition,the timber being extremely hard and both water and insect resistant, it had very many other uses. Yewtimber was greatly valued before the use of iron became general, both for its durability and itselasticity. The latter property, for instance, recommended its use for the manufacture of longbowweapons for hunting and in war for well over 300 years (Grieve 1931, p. 866; Milner 1992, pp. 40-3).
In a wide ranging essay extolling the Yew tree, Grigson (1952) discussed how valued the trees were forsheltering dwellings from the prevailing winds. When used in this manner in Fermanagh, they are almostalways planted on the SW side and close to the house.
Famous Fermanagh specimens
The story of the 'Florencecourt Yew', T. baccata var. fastigiata, is covered separatelybelow. The other most famous Yew tree in Fermanagh is 'The Crom Yew', or rather 'Yews' in the plural,which probably are the oldest trees of any species in Northern Ireland. Having said this, they areestimated to be only 800 years old at most (Browne & Hartwell 1999). The Crom Yew stands on theeastern bank of Upper Lough Erne near the ruins of Old Crom Castle. The late Alan Mitchell (1996, p.157), a very well-known tree expert of the second half of the 20th century, gives a very amusing accountof 'The Crom Yew', of which he read several descriptions in Irish books and magazines, and examinedseveral drawings of the tree. Accounts of the tree mentioned its extraordinary crown, spread in a lowcanopy supported by 16 oak posts (and previously by 34 brick pillars), and under whose shelter 200guests of Lord Erne were once served tea. In 1895, the said Lord wrote that it had a 6 foot [1.83 m]bole, girthing 12 feet [3.66 m] at ground-level, and a spread 77 feet [23.47 m] in diameter. Elwes &Henry (1902) quoted Lord Erne and his "60 supports", and described the tree as,"resembling an enormous green mushroom".
Alan Mitchell visited Crom in 1983, and to his great surprise found not one but two very similar trees,brother and sister, planted about 20 ft [6 m] apart! The trees have had their branches interwoven sothat they share the same crown and its supports. The second tree is clearly a twin of its partner, andmust have been there for between 400 and 800 years (depending upon which account of its planting youbelieve). The unsolved mystery must remain why these biological details had been ignored or overlookedin the various accounts of The Crom Yew and by the artists who depicted it? (Mitchell 1996, p. 157). Thetrees have been 'tidied' in more recent times, and as a result they have lost some of their mystery. Itis now easy to get under and through the combined canopy of the two siblings. Browne & Hartwell(1999) provide three photographs in their booklet, but Packenham (1996) has captured their appearanceeven better in his book Meetings with remarkable trees.
Yew toxicity and seed dispersal
Yew is very poisonous. Indeed, it is claimed that every part of the species except the fruit aril istoxic, but even it may be slightly so (Cooper & Johnson 1998). The fleshy, mucilaginous arilsurrounding the seed is eaten by a wide variety of birds, the poisonous seed passing undigested throughtheir intestines and voided with their faeces (Lang 1987). Uneaten fruits remain in good condition formany weeks, but eventually they begin to ferment on the branch. Birds that eat these old, decomposingarils may become intoxicated or ill.
Fruit eating normally reaches a peak in November, and by January only a few, more concealed arils remainon the Yew branches. Members of the thrush family (especially Song Thrushes, Blackbirds and MistleThrushes) are the main feeders, but Robins and Starlings are also important. Greenfinches aresignificant seed-predators, and Great tit* are to a much lesser extent. The Greenfinch has a techniqueof removing the aril and seedcoat first, before consuming the remainder of the seed, which stronglysuggests that the toxins are contained in the testa (ie the seedcoat) (Snow & Snow 1988).
A great deal of study has been carried out on the toxins, a number of which have names based on the genusname. One of the many is 'taxine', which is present in all parts of the tree, and is a complex mixtureof at least 11 poisonous alkaloids. Taxine is rapidly absorbed from the digestive tract and interfereswith the action of the heart. Another poison is a cyanogenic glycoside called 'taxiphyllin', and thereis also an irritant volatile oil. In the last 30 years, a chemically altered alkaloid derivative oftaxine extracted from T. baccata leaves, named 'taxol', has proven useful for treating ovarianand breast cancers. Due to a vast effort by biochemists worldwide, taxol can now be efficientlysemi-synthesised from Yew leaf hedge clippings.
The toxicity of Yew is not diminished by wilting or drying, so that clippings or even fallen leaves arestill highly poisonous (Cooper & Johnson 1998). Yew is considered by some to be the most toxic plantin these islands, yet there are conflicting reports of its toxicity to grazing animals. In general, thetrees should always be considered highly toxic, but if eaten regularly or often in small quantities,there may be no adverse effects. Cooper & Johnson (1998) report deer regularly grazing on Yew on theNorth Downs in Surrey without being poisoned, and we have observed the same thing happening with feralgoats in the Burren, Co Clare. Despite these two exceptions, a long list of stock and wild animals areknown to have been fatally poisoned, often collapsing and dying within a couple of hours of ingesting alethal dose, which may be as low as 0.5-2.0 g per kg body weight for animals such as horses (Cooper& Johnson 1998).
Medicinal uses
The fact that Yew is potentially lethal to humans has been known since ancient times, and as with manyother poisonous plants, this has somehow led to its medicinal use. It was given to 'steady the heart',and as an antidote for adder bites and against rabies (Cooper & Johnson 1998). As usual, it shouldbe emphasised that NOBODY SHOULD EXPERIMENT WITH POTENTIALLY LETHAL POISONOUS PLANTS.
Most human poisoning with Yew involves children eating quantities of the red arils and the seeds theysurround. Provided the poisonous seeds are not chewed, they should pass through the gut harmlessly, orwith only minor digestive disturbances being noticed (Cooper & Johnson 1998).
The question of Yew longevity
The longevity of Yew trees and shrubs has been appreciated for hundreds of years, and huge trees arevenerated or have even been worshipped in ancient times. However, it has only quite recently beenrealised that some specimens may be thousands of years old (Milner 1992; Thomas 2000). Unfortunately,the yew trees of greatest girth, and all those over about 400 years age, are invariably hollow (Mabey1996). For this reason, accurate tree-ring counts or radio-carbon dating of specimen trees areimpossible, the early growth wood being absent (Mitchell 1996). Indeed, we do not even know at what agethe central heartwood begins to rot, or when and why the trees go into slow-growth mode (Mabey 1996).
Consequently, researchers have turned to documentary evidence, comparing girth measurements andcalculated growth rates and annual ring counts of presumed very old and younger trees (Milner 1992).This method of estimating age applied to 70 trees over 300 years old scattered throughout England andWales, allowed Allen Meredith to calculate that during the first 500 years of growth, a churchyard yewincreases in girth an average of 1.1 cm per year until it reaches approximately 5.5 m (A. Meredith,quoted in Milner 1992, p. 82).
However, it is clear from documented measurements made of particular trees, that the accuracy of thismethod is very suspect, or that many old specimen trees grow much more slowly than this calculationsuggests, and in some cases their girth may cease increasing for periods of 300 years or more!Furthermore, Yew is known to be capable of growing without the formation of tree-rings (Thomas 2000, p.45), and thus, even when accurate ring counts exist, they may provide an underestimate of the real ageof the specimen.
The growth of Yew is so irregular, variable and anomalous, that unfortunately no generalised growth curvecan be fitted to the data (Mitchell 1996). Dated trees however do exist, for instance that at DryburghAbbey, Selkirk, which the monks are known from documentary evidence to have planted in 1136. In 1894,John Lowe measured the girth of this yew to be 11ft 4in [3.45 m], and 90 years later it had added teninches [25.4 cm] (Mitchell 1996). Thus a tree 858 years old had a girth of 12ft 2in [3.71 m]. Reallylarge old yews may have girths of up to 30ft [9.14 m], but with the known possibility of slowing of thegrowth rate, and the decay of central wood, we can only guess at their age. As far as we can tell, theyare probably several thousand years old. The best account of this interesting but problematic topicappears in Alan Mitchell's Trees of Britain (Mitchell 1996, pp. 153-6).
Variation
The extent to which T. baccata varies is demonstrated by the fact that including the fastigate'Irish Yew', or 'Florencecourt Yew' (dealt with separately), there are as many as 49 cultivars listed inDallimore & Jackson (1966) Handbook of Coniferae and Ginkgoaceae. Even allowing forinevitable duplications among cultivated varieties, this still represents a striking array of geneticvariation within the species.
Folklore
A considerable body of folklore attaches to the Yew tree, which is conveniently summarised in Grigson(1987), Milner (1992) and Vickery (1995). Several of the folk notions associated with the Yew recountedby these authors might not be all that ancient, but may really have originated from the speculations ofthe famous late 18th century cleric, Gilbert White, who pondered at length on a specimen in Selbornechurchyard (Grigson 1952, p. 12). The Selborne tree was a much measured individual, which unfortunatelybecame uprooted by a gale on 25 January 1990 (Mabey 1996).
Names
The genus name 'Taxus' is probably derived from the Greek 'taxon' meaning 'a bow', the strong, flexiblewood having been used for making longbow weapons for centuries. The Latin specific epithet 'baccata'means 'berried', although a better spelling would be 'bacatus' as it is derived from 'baca', meaning'berry' (Melderis & Bangerter 1955; Gilbert-Carter 1964).
The English common name 'Yew', or 'Yeugh' has various spellings in old authors, for example, 'Ewgh','Ewe', 'Ife', 'Ugh', 'Uhe' or 'Uhe tre', 'Vew' and even 'View'. In Anglo-Saxon, it was 'iw', MedievalLatin, 'ivus', 'iva', or 'iua', and there are cognate names in other Germanic languages and in theCeltic languages (Grigson 1974). However, it appears that this Anglo-Saxon name was applied to severaldifferent plants (Prior 1879). Some authorities derive the name from Gaelic 'iw', meaning 'green' (ieevergreen), but according to Prior (1879) there does not appear to be any such word. As the currentauthor has no skill in Gaelic, he awaits knowledgeable opinion on this matter. Prior links the MedievalLatin version of the name 'iva', to that of Ivy, and also to 'chamaepitys' (Mediterranean BlackCypress), "through a train of blunders" (Prior 1879), which sums up the confused situationrather aptly.
Yew 'berries' are variously referred to as, 'Snat-berries', 'Snottle-berries', 'Snottergall','Snotty-gogs' and 'Snoder-gills', an elegant reference to their sliminess (Prior 1879; Britten &Holland 1886).
Threats
Nowadays, most truly native trees grow on inaccessible cliffs, though some other rare possibly indigenousspecimens are browsed by sheep and goats, despite being poisonous.
Originally a very rare native, but now always deliberately planted; occasional.
1825; Mackay, J.T.; "First observed at Florencecourt".
Growth form and origin
This is the famous upright or fastigate 'Florencecourt Yew', planted world wide and known to all,although probably hated by some on account of its very dark, almost blackish-green, often dusty, or evensooty, fastigate foliage and its rather gloomy funereal associations with church and other graveyards.Although it originates here in Fermanagh, we have not bothered to record its local distribution in ourvice county botanical survey as it is always planted.
The original one or two specimens of this unusual growth form were collected as juveniles some timebefore 1767, or perhaps as early as 1740 (Nelson 1981). It was discovered at a place calledCarraig-na-madadh or 'The Rock of the Dog', on the NE slopes of Cuilcagh mountain above Florencecourt bya Mr George Willis, a farmer who lived at Aghtenroark (actually, Aghatirourke), in the parish ofKillesher (see pseudonymous account by Norval, The Gardeners' Chronicle 1873, p. 1336). Willisplanted one specimen in his own garden which eventually died around 1865, but fortunately he presented asecond to his landlord, Lord Mountflorence (later created the Earl of Enniskillen), who lived inFlorencecourt house. This plant has survived to this day and is the mother tree of all 'FlorencecourtYews' everywhere.
The original station
Charles Nelson and John Phillips investigated the original find area on Cuilcagh thoroughly in 1980 withthe help of W. Forde, a former gamekeeper of Lord Enniskillen whose family had handed down knowledge ofthe spot through the generations. No trees whatsoever now grow in the rock strewn area of blanket bognear the boundary of Aghatirourke and Beihy townlands, below an exposed sandstone outcrop about 2 mhigh, which has been renamed locally 'Willis's Rock', grid reference: H141297 (Nelson 1981; Morton1998).
Reproduction
The Florencecourt Yew is a female 'berried' tree, so that all its descendants raised vegetatively mustalso be female. The fastigate form does not breed true from seed, so the gene producing the uprightgrowth form appears to be recessive (Milner 1992, p. 42). Very occasionally a mutant branch with malecones is produced. This also happens rarely with the common, or 'English', yew (Morton 1998, pp. 196-7).
Garden introduction
The fastigate form of Yew is very easily propagated from cuttings, and when around 1780 the Florencecourtspecimen was admired by George Cunningham, a Liverpool nurseryman who obviously could see its commercialpotential, Lord Enniskillen was persuaded to give him some slips. Probably it was Cunningham who firstintroduced the tree to the horticultural trade (Nelson 1981), and possibly not Lee and Kennedy ofHammersmith (Bean 1970-80), although they too, along with other private Irish gardeners, may have beengiven cuttings sometime around 1780 (Nelson 1981).
Irrespective of exactly who introduced the tree to commerce, by 1838 the 'Irish Yew', or better, the'Florencecourt Yew', was available to the public at a low price (Bean 1980, 4, p. 566), and waswidely disseminated as we can easily see by today’s many large specimens and avenues of the tree aroundthe country in gardens, demesnes and church associated sites.
Thomas Packenham featured and photographed the mother tree with himself standing under it in hisexcellent book, Meetings with remarkable trees (Packenham 1996), although he comments ratherdisparagingly of its present day appearance. It is also illustrated in the late Dinah Browne'sattractive booklet of Northern Ireland's special trees, Our remarkable trees, photographed byMike Hartwell (Browne & Hartwell 1999).
Condition of the original tree
The original fastigate tree at Florencecourt grows beside a small stream in a glade surrounded andsheltered by laurel and other taller trees, on land owned and managed by the N Ireland Forest Service.The tree survives rather than thrives, being rather looser in habit than normal, scraggy and ragged dueto its uncongenial position, which was heavily shaded in the past by laurels, and in a soil too damp forgood growth. It is misshapen by the repeated taking of cuttings, and while some clearing of trees aroundit and careful pruning of the specimen carried out in 1980 has helped rejuvenate it, the tree is stillsomewhat overgrown with lichens. Nevertheless, it remains recognisable and while it lives is ofbotanical interest.
Threats
None.
Introduction, archaeophyte, extremely rare and almost certainly extinct.
Eurosiberian boreo-temperate.
1864; Dickie, Prof. G.; Pettigoe.
Growth form and preferred habitats
Asarum europaeum is a patch-forming perennial of shady places, which was widely grown in medievaltimes by herbalists from at least 1200 AD onwards, according to Harvey (1990).
Fermanagh occurrence
This unique record originates in Dickie's (1864) Flora of Ulster under the heading, "Specieswhich may be considered not strictly indigenous". It is unique since there does not appear to beany other record for this species anywhere in Ireland at any time (Preston et al. 2002). Dickierecorded the plant from, "waste places at the village of Pettigoe". Being a village on theborder with the Republic of Ireland, the record might equally apply to Co Donegal (H34) as to Fermanagh.
George Dickie
The finder of the plant in Pettigoe, George Dickie (1812-1882), was a native of Aberdeen and a medicallyqualified graduate of both Aberdeen and Edinburgh. He came to Belfast in 1849 as Professor of NaturalHistory at the then new University of Queen's College (now The Queen's University of Belfast). Hereturned to Aberdeen as Professor of Botany in 1860. While in Ireland, Dickie collected the material forhis Flora of Ulster (1864), which Praeger (1949) described as, "an excellent little bookwhich embraced not only that province" (ie Ulster), "but included the interesting area ofSligo and Leitrim." Dickie produced two further Floras dealing with parts of eastern Scotland, andhe became an FRS in 1881 (Praeger 1949).
Regarding the accuracy of Dickie's Asarabacca record, Robert Northridge has suggested the possibilitythat the professor could have made an error: the two dark green and rather glossy, evergreen,kidney-shaped leaves of A. europaeum are smaller, but similar in shape to those of Petasitesfragrans (Winter Heliotrope), a plant which today is abundant on roadsides around Pettigo.However, I reckon that a man of Dickie's medical and botanical learning and experience would certainlyknow a Birthwort from a Butterbur! This is not to imply that even the most eminent professor cannot makemistakes. I recall David Webb detailing some of his own errors at a BSBI AGM held in the BotanicGardens, Glasnevin in Dublin. The odds are in Dickie's favour when it comes to a medicinal plant likeAsarabacca.
British status and occurrence
This species is also a declining and rare plant in Britain, with 60 of the 77 hectad squares plotted inthe New Atlas having pre-1970 records only. Although previously claimed as being native, it is nolonger regarded as anything but an ancient introduction (ie an archaeophyte) (Coombe 1956; G.M. Kay, in:Preston et al. 2002).
Medicinal uses
The root and leaves of A. europaeum are acrid and contain a volatile oil, a bitter matter and asubstance with properties like camphor (Grieve 1931, p. 64). It was used amongst other things as a curefor hangovers, as a purgative and to promote sneezing, although Grieve indicates that even in the 1930sit had been replaced, "by safer and more certain remedies". It is clear from a quotation inGrigson (1987, p. 225), taken from a book by John Pechey (1694) The Compleat Herbal of PhysicalPlants, that Asarabacca was also used as an abortifacient, "Tis diuretick also, and forcesthe Courses: wherefore Wenches use the Decoction of it too frequently, when they think they are withChild."
European occurrence
A. europaeum is a European temperate species, widespread in middle latitudes and towardsthe east, but absent as a native from most of W Europe (Jalas & Suominen 1976, Map 368).
Names
Of the curious name, 'Asarabacca', Grigson (1987) says, "Dioscorides described the plantvery precisely in his Di Materia Medica, under the name 'Asaron'. He also wrote of abacchareis, which some herbalists took to be Asarum europaeum as well, though the twodescriptions do not tally. Virgil in the Eclogues wrote of a baccar which grew with ivy,in the way of A. europaeum. As if to compromise and resolve the matter, apothecaries squashed thetwo names into one, to give the strange word 'Asarabacca'".
Threats
None.
Native, frequent. European temperate.
1860; Smith, T.O.; Lough Eyes.
May to September.
Growth form and preferred habitats
This conspicuous rhizomatous perennial is frequently found in open water floating leaf plant communitiesin still, shallow-water lakes of all sizes in our area, but it also extends into adjacent reedswampshallows. In other parts of Britain and Ireland, White Water-lily is also found to a much lesser extentin slow flowing ditches and in river backwaters, but while we have over 280 records for N. albain Fermanagh, only once has it been listed from this type of habitat, from a drain at Cornaleck, UpperLough Erne in 2007.
In the type of still water bodies that N. alba frequents, the bottom is typically mud, silt orpeat, and it is only rarely of shingle or rock. N. alba tolerates a wide range of water chemistryand it is found in calcareous turloughs (ie vanishing lakes) and marl lakes, as well as in decidedlyacidic upland lakes, often over peaty bottoms. In the Britain and Ireland, it is normally regarded as aplant of relatively shallow, still, lowland waters, but it can survive in waters up to 5 m deep(Heslop-Harrison 1955b). It reaches an altitude of 405 m at Tarn Fell in Cumberland (C.D. Preston, in:Preston et al. 2002). Colonisation of deeper waters must be entirely by means of vegetativereproduction involving rhizome fragments since seedlings cannot grow at the low light levels thatprevail at depth. N. alba also occurs in waters of all levels of productivity ranging fromeutrophic to oligotrophic (Preston & Croft 1997).
Although generally found in open, full sun conditions, N. alba is occasionally found in the shadeof overhanging trees, or more frequently among tall emergent stems, such as those of Phragmitesaustralis (Common Reed), where light levels may be reduced by up to 50% of full sun(Heslop-Harrison 1955b).
Under favourable growing conditions, N. alba can dominate open water in smaller lakes, the largeleaves of the species completely covering the surface. Such large clonal individuals may be of verygreat, or indeed, indefinite age, and can measure 10 m or more in diameter. Large colonies are generallyring-like in form, the central older parts of the clone having eventually died off (Heslop-Harrison1955b).
Recognition
In the absence of flowers, the leaves can be distinguished from those of Nuphar lutea (YellowWater-lily), by being almost circular in outline, having the lateral veins branching at wide angles andforming a rather inconspicuous net-like web near the margin. The leaf stalks are also rounded or oval incross section, not angular as in Nuphar (Haslam et al. 1975; N.F. Stewart, in: Rich& Jermy 1998).
Alternative growth form
The species is also known to possess a terrestrial growth form with tufts of erect leaves with rolledmargins, occasionally reported in marshy habitats in several areas of Europe, eg in W Ireland on AchillIsland (Praeger 1934i, paragraphs 24 & 408), by a pool south of Lough Gill near Castle Gregory (H28)and around the shores of Lough Carra (H26), where it grows on wet, calcareous marl (Heslop-Harrison1955b). Although we have similar marl lakes in Fermanagh, to date we have not observed this ratherunusual form of the species anywhere in the VC.
Fermanagh occurrence
N. alba has been recorded in 98 Fermanagh tetrads, 18.6% of those in the VC. As the tetraddistribution map indicates, it is frequent in the W, C and SE of the county, but absent from theopen-water areas of our largest lakes, Upper and Lower Lough Erne. The species lacks submerged leavesand is, therefore, much less tolerant of disturbed water than Nuphar lutea (YellowWater-lily). Thus in Fermanagh's larger lakes, and especially in the navigable channels betweentheir many islands where water can readily become turbid through wind and wave action or as a result ofboating activity, White Water-lily is absent. In these lakes, it is confined to sheltered bays andbackwater shallows protected from excessive disturbance by reedbeds.
Phenology and sexual reproduction
The floating leaves are produced annually, first making their appearance at the water surface around May.The flowers, which also float, are produced from early June to August. In comparison with Nupharlutea, the blooms open much wider, and at up to 20 cm in diameter, they are the largestindividual wild flower in these islands by a wide margin. The white flowers, which are very primitive(probably dating back to the Jurassic Period), have large numbers of separate, spirally arranged parts(tepals). The flowers close at night, or when it rains, during the 4-7 days each one lasts. The femaleorgans ripen first (ie the flowers are protogynous), with a subsequent prolonged male phase. Pollinationis effected either by bees or by other insect visitors, but should this fail, selfing occurs(Heslop-Harrison 1955b; Velde 1986).
After pollination the flower is pulled about 30 cm below the water surface by contraction of the stalk.The flower rots and the fruit ripens and eventually sometime between September and early November itbursts irregularly, releasing a floating mass of mucilage and embedded seeds which resembles frog-spawnand thus attracts both water birds and fish.
Seed production and dispersal
Typical seed production averages around 500 per fruit, although the number of flowers per plant and theseed set per flower, both vary considerably with growing conditions. The seed mass mucilage consists ofthe transparent arils around each seed aggregated together. It entraps air, floats and enables dispersalacross the water body. According to Dutch studies, this flotation may be of short duration, since anyrainfall releases the entrapped air and the seeds then immediately sink (Velde 1986; Smits et al.1989). Duck, Coot and other water birds actively eat the seed, and fish such as Common Carp will also doso, but only if they are starving. Unfortunately, at least in the case of these particular animals,ingestion does not appear to aid the plant's dispersal since, unlike Potamogeton species(Pondweeds), the seed coat is weak, allowing it to be either completely digested or killed by passagethrough the animal's gut. This fact does not rule out the possibility that other birds and herbivorousfish with less efficient digestion, might internally transport and excrete viable seeds (a mechanismtechnically known as 'endozoochorous transport'), but as far as I know there is no positive evidence ofthis occurring at present (Smits et al. 1989).
Transport of species between water bodies
Due to this inability of seeds to survive passage through the animal gut, aquatic plants like N.alba having a weak testa, can transfer between water systems (ie long-distance orjump-dispersal), only through external adherence of seed on the body of an animal vector. This might beachieved either through chemical stickiness, or physical projections involving some kind of burr-likeseed or fruit with a spiny or a hooked surface. Our two Water-lily species, Nymphaea alba andNuphar lutea, have no such fruit or seed properties, and thus they are not specifically adaptedfor external transport on animal bodies. They might still, however, adhere for a short time in mud onthe feet, coat or down of an animal moving between adjacent lakes or streams.
Thus it is not completely impossible to imagine the occasional or rare stochastic longer range dispersalevent occurring. Studies of very long distance bird transport of seed to remote oceanic islands hasshown that the down on young fledglings may well prove significant, since seed attaches very much morereadily to this type of surface than to feathers, particularly pre-flight, preened ones (Falla 1960;Carlquist 1974). In the case of Water-lily species, it is likely that external animal attachment in mudwould persist only for a short space of time, perhaps a few minutes, since it has been shown that theseed is susceptible to dehydration. However, I believe that the experimental test used to examine thisfactor was unrealistically extreme, as it involved air drying seed in a dissector for a period of 28days before testing their viability! (Smits et al. 1989).
Long-distance dispersal must be a repeatable, recurrent, even if random event; it is not sufficient forit to be fortuitous. Thus animals which seasonally migrate along regular routes, such as birds, arefavoured vectors to carry the plant propagules (Cruden 1966).
Seed properties
Seed germination is usually successful and seedlings can sometimes be found in large numbers, but few ofthem survive overwinter. In water, seed may survive dormant for up to three years (Jonsell et al.2001), although other studies suggest the seed bank is only ephemeral (Thompson et al. 1997).
Irish occurrence
In Northern Ireland, N. alba, like Nuphar lutea, is very much more frequent in the southernhalf of the six counties, but overall it is the less common of the two Water-lilies (NI Flora Web Page(accessed 2015) http://www.habitas.org.uk/flora/species.asp?item=2757). The overall Irish distributionshows N. alba concentrated in the N and W of the island, with a thinly scattered occurrence inthe centre of the island, and a remarkable almost total absence from the SE, where it is introduced inWexford (H12), Carlow (H13), Laois (H14) and Dublin (H21) (Praeger 1934i, paragraphs 270 & 495;Scannell & Synnott 1987).
British occurrence
The British distribution in the New Atlas fails to distinguish native populations from the manyintroductions that are now known to occur as a result of the fashion for garden ponds. 'Escapes' and'deliberate releases' from such ponds are especially frequent in SE England. The New Atlas maptherefore shows N. alba occurring widespread in the lowlands throughout the whole territory, butbecoming much less frequent in the E and the NE as one travels northwards (C.D. Preston, in: Prestonet al. 2002).
Allowing as best one can for the introduced populations, it clearly remains the case that the somewhaterratic pattern of N. alba distribution in Britain and Ireland must depend both on the currentpresence of suitable lowland, nearly-still-water habitats, and the long-term effectiveness of dispersalby water, and perhaps by other means, between discrete water bodies and separate catchment areas.
Clinal variation in Europe
In Europe, the genus Nymphaea is generally recognised as consisting of four or five taxa atspecies or subspecies level. Of these, N. alba and N. candida are the most similar, to theextent that they are regularly considered as subspecies of N. alba. While these latter two formsusually remain geographically separate, N. candida being more NE European and Asian than N.alba, intermediate forms and small areas of territorial overlap do occur between them, so thatin northern Europe a 'cline' may exist, ie a gradient of gradually changing variation running betweenthe extremes of the two forms (Heslop-Harrison 1955b).
European occurrence
Beyond Britain and Ireland, the form of the species that occurs with us (N. alba) is common andwidespread in W and C Europe, and to the NW it stretches beyond 68°N in coastal Norway, although absentnorth of 60°N in Sweden and present, but thinning northwards in Finland and the Baltic States (Jonsellet al. 2001). In southern Europe, the distribution thins, both in the Iberian peninsula andtowards the Mediterranean, where there are a number of extinctions, eg on Sicily and the southern end ofSardinia. This thinning and local extinction pattern is also repeated towards E Europe and SW Asia(Jalas & Suominen 1989, Map 1509).
World occurrence
The southern limit of N. alba is reached in Algeria, and the SE extremity in Kishmir and theHimalaya (Heslop-Harrison 1955b; Hultén & Fries 1986, Map 812).
Fossil history
The fossil seed and pollen record of N. alba extends back to the Cromer Forest Bed interglacialseries and it appears in all subsequent warm periods including the present Flandrian/Littletonian inBritain and Ireland. Indeed, the species is capable of surviving considerable frost, being presentwithin the Arctic Circle at present, and a continuous fossil record runs back to the middle and latestages of the last major Ice Age, called the 'Weichselian' in Britain and the 'Midlandian' in Ireland.It is possible, therefore, that N. alba might have survived in situ the British Islesduring much colder phases than the present, perhaps living quite close to the edge of the ice sheetsunder periglacial conditions (Godwin 1975).
Toxicity and uses
All parts of the plant, except the seeds, contain the alkaloid nupharine, the amount present varying withthe season (Heslop-Harrison 1955b). The rhizome and leaves have a history of use in herbal medicine, asin addition they contain tannin, gallic acid, resin and mucilage. Grieve (1931, p. 484) details theplant's use in cases of dysentery, diarrhoea, gonorrhoea and leucorrhoea. The leaves and 'roots' werealso used to poultice boils, tumours, ulcers and inflamed skin, and an infusion was gargled for mouthand throat ulcers.
Other folk uses include the starchy rhizome as a food in parts of Finland and Russia, and the same plantorgan was a source of purple-black dye for wool and yarn in the Inner Hebrides (Heslop-Harrison 1955b;Vickery 1995).
Names
The genus name 'Nymphaea' is from the Greek name 'Nymphe' given by Theophrastus to an unknown water plantafter one of the three half-divine water nymphs who in mythology inhabited seas, streams and woods(Gilbert-Carter 1964; Chicheley Plowden 1972). The Latin specific epithet 'alba', means white.
There are at least 16 local English common names listed in total between Britten & Holland (1886) andGrigson (1987). Some names are poetic and very evocative of the beautiful floating flower, eg 'Lady ofthe lake' and 'Swan amongst the flowers'. In his famous evocation of Cotswold village life, Ciderwith Rosie, the author Laurie Lee (1959) spoke of the white flowers, "... they poured fromtheir leaves like candle-fat, ran molten and then cooled on the water."
A high proportion of the English common names refer either to 'water', eg 'Water Bells', 'Water Blob','Water Socks', 'Water Rose', or to 'floating', for instance, 'Floating Dock', of which 'Flatter Dock' isprobably just a variant. References to the conspicuous broad leaves of the plant are only to beexpected, and comparison with 'dock', as already mentioned, eg 'Can-dock', 'Can-leaves'. 'Can' is areference to the carafe or jug-like shape of the fruit capsule (Grigson 1987). 'Bobbins' is a nameapplied to both Nymphaea alba and Nuphar lutea, being a further shape comparison of theglobular fruit to a lace or weaving bobbin. The name 'Cambie-leaf' is a curious one of northern Scottishorigin, applied there again to both of the common water lilies. I would welcome any explanation of thederivation. Yet another name applied to both species in early herbals is 'Nenuphar', which Grigson(1987) reports came down via Mediaeval Latin from the Sanskrit 'nilotpala', the name for the Blue Lotusof India, Nymphaea stellata. By comparison, Caltha palustris (Marsh Marigold) is 'Petienenufar' (ie 'Petty nenufar'), in Turner's The Names of Herbes, 1548 (Watts 2000).
Threats
None, except perhaps undue water turbidity resulting from disturbance.
Native, very rare. European temperate.
July 1946; MCM & D; Carrick Lough, Dresternan Td, NW of Derrygonnelly.
Stace (1997) recognises this form with its smaller flowers and leaves as a subspecies, which occurs inunproductive lakes in W Ireland and N & W Scotland. The authors of the Typescript Flora (andthe Revised Typescript Flora) listed this subspecies separately but in the latter addedthe comment, "This is merely a starved form of N. alba growing in a habitat deficient innutrients."
The status of subsp. occidentalis requires further investigation, particularly since Stace goes onto point out that intermediates occur between it and subsp. alba and that the intermediates arenot confined to the areas where the two forms overlap (Stace 1997).
There are just two records of the supposed subspecies in the Fermanagh Flora Database, both made byMeikle and co-workers. The second site was at Carricknagower Lake, also on the Western Plateau, recordedin July 1947.
Native, common and widespread. Eurosiberian boreo-temperate.
1860; Smith, T.O; Lough Eyes.
May to January.
Growth form and preferred habitats
A familiar floating-leaved water-lily of lowland lakes, lakelets and slowly flowing streams and rivers,N. lutea is a perennial with a creeping rhizome. It is usually found in both lake and river waterbetween 0.5-2.5 m deep, over mud and silt bottoms, and it typically bears annually renewed leaves of twokinds: the familiar floating, rounded leathery plates, plus large, crumpled, inconspicuous ones whichare translucent and are kept permanently submerged. Possession of these submerged leaves allows N.lutea to dampen the physical effects of water movement, and it tolerates much more mechanicaldisturbance and associated water turbidity than the related Nymphaea alba (White Water-lily)can manage. However, N. lutea is less tolerant of base-poor sites than the latter and ittends to occur in basic to only moderately acidic waters, seemingly restricted to pH 6.0 and above(Heslop-Harrison 1955a; Preston & Croft 1997). In sheltered, still water sites where the twowater-lily species cohabit, which they quite often do, N. lutea typically occupies the deeper,more nutrient-rich water.
The floating leaves of N. lutea are clearly oval rather than circular in outline and the lateralveins divide repeatedly and forking regularly in a herring-bone pattern until they reach the margin,while in cross section the leaf stalks are triangular or semi-circular (ie definitely angular) (Haslamet al. 1975; N.F. Stewart, in: Rich & Jermy 1998).
Again, like Nymphaea alba, N. lutea occurs in both the lowland floating leaf open waterplant community and occasionally along the margins of reedswamp, particularly where the water depthshelves steeply, or where there is a faster current beyond. Colonies of Yellow Water-lily can varyenormously in size, from isolated plants in less favourable sites, to situations where it dominateshectares of water at a stretch, spreading both by vegetative growth and branching of the horizontalrhizome, and by seed if water depth, flow and turbidity permits germination and establishment(Heslop-Harrison 1955a; Jonsell et al. 2001).
Flowering reproduction
Individual plants of N. lutea in cultivation are known to survive for at least a century and it islikely that under favourable growing conditions in the wild, plants may persist absolutely indefinitely.In a suitable environment, young plants flower annually and they do so quite freely from their thirdyear of growth onwards. It is not unusual to find a mature plant bearing 15 or more flowers at somestage between early June and late August. The cup-shaped solitary yellow flowers, 4-6 cm across, areborne a few cm above the water surface on very long rigid peduncles. Unlike White Water-lilies, once theflower buds open they do not close again during their 4-8 day flowering period.
The flowers are protogynous (ie female first), the stigmas ripening slightly earlier than the stamens(Velde 1986). Pollination is carried out by a variety of flies, bees and beetles, probably in that orderof importance, and there may even be Nuphar specialist pollinating flies (Lippok & Renner1997). The insects are attracted by a strong flower scent, reminiscent to many humans of plum brandy ora combination of fruit and alcohol. The active ingredient of the perfume is actually ethyl acetate,which is an organic chemical combination of acetic acid and ethyl alcohol (Genders 1971, pp. 27 &42). The alcohol component is produced in the plant's roots, which appear to tolerate and cope withanaerobic conditions in the muddy bottom exceptionally well. Alcohol is a by-product of the partialbreakdown of starch and sugars in the absence of oxygen and, of course, in high concentration it is alethal toxin that can kill cells. N. lutea transports the alcohol aloft from the submerged rootsto the floating leaves and to the flowers, from which it evaporates harmlessly away. Effectively, thisis a form of plant excretion, getting rid of a toxic waste-product into the atmosphere (Fitter 1987, p.218). Consequently, one of the better known English common names of the plant is 'Brandy bottle',although in part this might also be due to the globular shape of the ripe fruit.
The flower visitors are rewarded by a copious and freely available supply of both pollen and nectarfoods. The stamens are unusual in that while the anthers are introrse (ie their slit-like openings aredirected towards the centre of the flower), as they split to release their pollen they arch overbackwards and thus they present their pollen outwards towards the petals. Consequently, automaticself-pollination is avoided and out-breeding is strongly favoured, although an insect walking orstaggering around on the flat and relatively broad stigmatic disk, can still occasionally manage toself-fertilise the flower (Heslop-Harrison 1955a).
Fruit and seed production
Again, in further contrast to Nymphaea alba, the fruit develops and ripens just at the watersurface, rather than submerged (Velde 1986). The number of seed per fruit is very variable and appearsto be under both genetic and environmental control. In cultivation in Sweden, for instance, a mean of361 seeds per fruit was measured for a sample of 70 fruits, the range of seed production being 45-651per fruit (Heslop-Harrison 1955a). When ripe, the whole fruit may detach and float around for two orthree days before the contained mucilage swells and the fruit bursts or irregularly disintegrates,releasing the ten to 20 fruiting carpels, each containing numerous seeds.
Seed dispersal
In contrast with Nymphaea, seeds of Nuphar do not possess an aril to assist their flotationin water, but the slimy and spongy pericarp tissue which contains small air or gas bubbles serves thesame function in N. lutea. Smits et al (1989) concluded that Yellow Water-lily carpels hadvery poor buoyancy, and that after a day or so (or even less if it rains and releases the air from thebubbles), this spongy mucilage disintegrates and the seeds then immediately sink to the bottom. Sincewater transport appears to be the most obvious means of dispersal for aquatic macrophytes, the plantpropagules of N. lutea appear to be strangely ill-equipped and poorly adapted by evolution forflotation over anything other than short distances. It is known that young seedlings of some aquaticmacrophytes also possess powers of flotation and thus a secondary, additional phase of dissemination mayoccur at a later stage, as is the case with some emergents, eg Baldellia ranunculoides (LesserWater-plantain) and also some submerged plants, eg Hottonia palustris (Water-violet) (Sculthorpe1967, p. 329), but as far as we know this possibility has never been observed in N. lutea.
The findings of Mason (1975) with regard to the behaviour of long-established alien aquatic plants in NewZealand underline the poor dispersal abilities of many of them, including N. lutea. Mason showedthat although these introduced species were locally abundant, they colonised other isolated water bodiesonly slowly, if at all, when they were spread deliberately, or accidentally, by man. Without this humanintervention they were static and completely confined to existing sites.
Animals that live in, or frequent water, are often considered important dispersal agents, capable ofcarrying seeds or other transferable vegetative plant parts (Ridley 1930; Sculthorpe 1967). In the caseof N. lutea, however, Smits et al. (1989) found no evidence of any adaptation to enableany imaginable form of such animal transport, either attached externally, or ingested and voided in aviable condition through the alimentary canal of fish (Carp), or birds (Mallard duck and Coot). However,viable seeds of N. lutea were once discovered in the excreta of a Heron. The seeds were presumedto have been eaten by a fish, which was subsequently eaten by the bird! Heslop-Harrison (1955a) alsoquotes the work of a Finnish researcher, who found that seeds removed from the gut of Ridd fish(Scardinius erythrophthalmus), germinated more readily than seeds sown directly from the plant.
More recent work in a sheltered muddy site on the River Rhone in France found that while seed productionof N. lutea was at least 600 per m2, not a single seedling of the species was recordedin numerous sample bare mud quadrats examined over an entire five year study period. The onlyexplanation which could be offered was that the muddy sediment in question was very loose and easilyre-suspended in water, so that the seeds could sink into the mud to a depth from which seedlingemergence simply became impossible. The dynamics of the Nuphar population at this particular sitetherefore rested entirely upon vegetative extension of the rhizome system (Barrat-Segretain 1996). WaterHens and Grebes might assist in vegetative dispersal, since aquatic birds such as these have beenobserved carrying uprooted portions of Nuphar rhizome during their nest building (Heslop-Harrison1955a).
Seed germination
The findings and conclusion of the Rhone work described above is rather surprising in the light of Dutchstudies which indicate that germination in this species, while often erratic, is actually better underanaerobic conditions when compared with aerobic, provided the seed has been cold-treatment stratified.The same Dutch study also found that the presence of ethanol and ethylene helped stimulate germinationin both Nuphar lutea and Nymphaea alba (Smits et al. 1995). Both seedling andmature N. lutea plants are also known to produce allelopathic compounds, eg resorcinol, plus anumber of
alkaloids such as nupharolutine, which appear capable of inhibiting the growth of competing species(Elakovich & Yang 1996; Sutfeld et al. 1996).
Fossil record
The fossil pollen and seed record of N. lutea extends back through all the interglacials to thePastonian. Both pollen and seed are also frequently recorded in all zones of the current Flandrianinterglacial (also known as the Littletonian in Ireland), and pollen appears in low frequency throughoutthe Late Weichselian glacial stage (the Midlandian in Ireland). Godwin (1975) concludes, therefore, thatthere is a strong case for regarding the species as persisting in Britain and Ireland right through thePleistocene, although as one would expect, it must have been restricted during the glacial stages.
Fermanagh occurrence
Nuphar lutea is four times more commonly recorded and at least twice as widespread in Fermanagh asNymphaea alba (White Water-lily). There are records of N. lutea in 188 Fermanagh tetrads,35.6% of the total in the VC. Yellow Water-lily is particularly abundant and ubiquitousthroughout the whole of the Upper Lough Erne catchment, but is very much less frequent in Lower LoughErne, where indeed it is absent from most of the shore.
Irish occurrence
In Ireland, N. lutea is much more common and widespread than Nymphaea alba, especially inthe Midlands, although like the latter species, it also becomes rare or absent from most of the SE ofthe island.
British occurrence
Having smaller flowers that are much less decorative than White Water-lily, N. lutea is not nearlyso likely to be introduced in English sites and the New Atlas therefore attempts to map thenative British distribution. This shows frequent throughout C and S England, but absent from the extremeSE, the Channel Islands, and also from much of the N of Scotland including the northern isles. Thereappears to be little change in the distribution when it is compared with that in the 1962 Altas(Walters & Perring 1962; Preston et al. 2002).
European and World occurrence
In Europe, it is widespread throughout, extending to 67oN, but thinning markedly southwards inthe Iberian and Italian peninsulas, and is scarce throughout the Mediterranean basin, although presentand rare in E Algeria (Jalas & Suominen 1989, Map 1514). Eastwards, it stretches through the MiddleEast, the Caucasus to C Asia, Siberia and Manchuria and there are closely related forms in N America(Heslop-Harrison 1955a; Hultén 1971, Map 158). As with Nymphaea alba, the wide range of N.lutea in Europe and Asia rules out broad regional climatic factors as agents creating theobserved uneven British Isles distribution. This must instead reflect the availability of suitable waterbodies. The fact that it is more or less a lowland plant again probably reflects its minimum pH (ie pH6.0), rather than lower temperatures, although higher exposure and more frequent wind on heights mightcreate wave and turbidity problems which would also limit the species (Heslop-Harrison 1955a).
Names
The genus name 'Nuphar' is derived from the Iranian 'nufar' or 'naufar' meaning or referring to awater-lily (Gilbert-Carter 1964). The Latin specific epithet 'lutea' meaning 'yellow' is derived fromthe name 'Lutum', of the Dyer's Wintergreen or Weld, Reseda luteola, which yields a yellow dye(Gilbert-Carter 1964; Stearn 1992).
Being a common and conspicuous water plant it is not surprising that N. lutea has at least 25local English common names, of which five are shared with Nymphaea alba (Britten & Holland1886; Grigson 1987). The name 'Blob' or 'Water-blob', is one of six names shared with Calthapalustris (Marsh-marigold). The word 'blob' or 'bleb' is Anglo-Saxon and means a bladder, bubbleor blister (sometimes, as in the Ranunculaceae, indicating the capacity to raise a blister on skin), andthe adjective 'blub' refers to something swollen, plump or round, all descriptive terms which could beapplied equally well to flowers of Nuphar and Caltha (Prior 1879; Britten & Holland1886; Watts 2000). Several names suggest the yellow flower in referring to butter and other allusions,for instance 'Butter Churn' and 'Butter Pumps' both refer secondarily to the shape of the fruit. Thename 'Golland', 'Water Golland', or in numerous variant dialect spellings as, for example, 'Gowan','Gowlan' and 'Gowland', is shared not only with Caltha palustris (Marsh-marigold), butalso with Trollius europaeus (Globeflower), the yellow buttercups and, indeed, with almost anyyellow flower, or even with one which has a yellow centre, like Bellis perennis (Daisy).'Golland' is thought to be derived from the Anglo-Saxon 'gold', or if we prefer a remote ancestry, theSuio-gothic 'gul', 'gol', meaning 'yellow' (Britten & Holland 1886). It could also be related orcompared to the Norwegian 'gal' or 'gaul', again meaning simply 'yellow' (Grigson 1974). 'Clot' and'Clote-leaf' are names applied to both N. lutea and Verbascum thapsus (Great Mullein).'Clote' is Old English and can mean 'wedge', which is probably a reference to the large, broad leafshape (Watts 2000). Other names and their allusions are discussed under Nymphaea alba, as are theherbal medicinal uses that Nuphar lutea shares with that species.
Threats
None.
Native, but also very probably introduced: very rare, a recent arrival. Circumpolar southern-temperate,but almost cosmopolitan.
25 July 2006; ENSIS New Lake Survey; Killymackan Lough ASSI.
Growth form and history of discovery in Fermanagh
This submerged, floating, truly aquatic species typical of eutrophic, still or slow-flowing water, wasfound in quantity on 6 October 2010 by Robert Northridge at a jetty near the National Trust's CromCastle estate Visitor Centre on the shore of Upper Lough Erne. The Rigid Hornwort was floating in atangled mat of aquatic species including Elodea nuttallii (Nuttall's Waterweed). A specimen ofthe hornwort was collected and sent to DBN for confirmation. Dr Matthew Jebb confirmed theidentification and we were confident that this was a new County Record.
On checking this supposition with the CEDaR Database for N Ireland, Robert Northridge and the currentauthor were very surprised to discover a previous Fermanagh record had been listed. The record detailswere given as: 8/9 August 1968 at Mill Lough near Bellanaleck, by staff of the Department of Agriculture& Rural Development NI, Fisheries Division. We presume the record was made during survey work onwater quality or fish stocks. In checking this Mill Lough record, we found that no voucher exists, norany note of the recorder's name (or names). CEDaR supplied three other plant records for the same dateand lake: Littorella uniflora (Shoreweed), Phragmites australis (Common Reed) andSchoenoplectus lacustris (Common Club-rush). All three are common widespread species in Fermanaghand, together with five other common wetland species on a total list of 18 records across several lakes,they give us no real indication of the recorder's identification skill.
This particular Mill Lough (there are three others in Fermanagh) has been visited by RHN on threeoccasions (1979, 1985 and 2000) and also by the NI Lakes Survey 1988-90, without occurrence of C.demersum. RHN revisited Mill Lough on 13 October 2010 and made a thorough search of floatingplants at seven jetties around the lake. C. demersum was not found, but masses of Water Crowfootwere present, either Ranunculus circinatus (Fan-leaved Water-crowfoot) or R. trichophyllus(Thread-leaved Water-crowfoot), with very short submerged leaves that might easily be mistaken for RigidHornwort by inexperienced botanists or non-botanical field workers. In view of all this, we havediscounted the 1968 Fisheries record entirely and have asked for it to be removed from the CEDaRDatabase.
On mentioning the above error to CEDaR staff, Robert and I were disconcerted to be then given additionallake data for Fermanagh collected at two sites for NIEA by the English consultant company, ENSIS in 2006and 2007, and from another four sites on Upper Lough Erne discovered by the NIEA's own Lake Ecology Teamin August 2010. Since the timing of these October 2010 revelations coincided with evidence given to thecurrent authors regarding the recent arrival of Elodea nuttallii in Upper Lough Erne, RHN carriedout a speedy late season survey of Upper Lough Erne shores (3-5 November 2010) to search for these twospecies. This led to the discovery of C. demersum at a further eight sites in the southern partof Upper Lough Erne. Thus Rigid Hornwort is now known to have first appeared in the VC in 2006, andduring the last four years it has spread to a total of 11 Fermanagh tetrads as shown in the map.
Irish occurrence
Although occasionally abundant in lakes, pools and canals, C. demersum remains a relatively rarespecies in Ireland. The 1987 Cen Cat Ir Fl 2 listed 17 VCs where it had been recorded at leastonce. Inspection of the New Atlas hectad map using a transparent overlay of the 40 Irish VCs (notas accurate or as straight forward an operation as it sounds) increases the number of VCs where it hasoccurred up to the year 2000 to 23. The VCs featured in these two sources do not completely agree,however, so while a wider distribution is indicated, we cannot conclude that the species has spread tosix additional VCs. The major change in C. demersum distribution in NI has been its arrival inLough Neagh in 1988 and its subsequent spread around the lake basin. Nevertheless, this was preceded inCo Tyrone (H36) by a find in 1982 at Ballagh Lough, 5.5 km NE of Fivemiletown (McNeill 2010). This siteis only 4 km from the boundary of Fermanagh.
Although it flowers regularly and freely in B & I, Rigid Hornwort very rarely sets seed, reproducingalmost entirely by vegetative fragmentation. While it is considered native in some parts of Ireland, thespecies is commonly used to oxygenate garden pools and indoor aquaria. We therefore suspect that theCrom colony at least is an introduction originating from recently discarded cultivated material. Cromjetty is a heavily visited public amenity.
British occurrence
C. demersum is widespread in lowland aquatic habitats across the length and breadth of GreatBritain. However, it is very unevenly spread, being much more frequently recorded in the EnglishMidlands and the South East of the country (Preston et al. 2002). This pattern of distribution,taken together with the fact that fruiting in Britain and Ireland is rare at present, suggests thattemperature may be limiting both the occurrence of the species and its sexual reproduction.
Rigid Hornwort is not confined to lakes, but occupies a wide variety of still and slow-moving eutrophichabitats including ponds, rivers, canals and ditches. In smaller water bodies such as ponds and ditches,it often forms large, dense mats of growth that can rise above the water surface. Comparison of theNew Atlas hectad map with the 1962 BSBI Atlas shows it is now more frequently recorded,but this is probably a reflection of better co-ordinated plant recording, rather than a real increase inpresence. In any event, at least the distribution in Britain appears stable, if not definitelyincreasing (C.D. Preston, in Preston et al. 2002).
World occurrence
Rigid Hornwort is an extremely widespread circumpolar species, not only in Eurasia and N America, butalso in C and S America, C and S Africa, S India and S Australia (Hultén & Fries 1986, Map 817).
Fossil record
The long-spined fruits of C. demersum are very easily recognised fossils and the sedimentaryrecord shows that this species has persisted in Britain as far back as the Cromer Forest Bed series andthrough all the subsequent interglacial periods including the present Flandrian. The record shows thatit was also present in the less severe terminal parts of the glacial stages. Godwin (1975) quotesSamuelsson (1934) regarding the post-glacial immigration of C. demersum back into northernFinland in the early Post-glacial period as glaciers retreated, where fossil records show the speciesestablished a considerable distribution. However, the record in Finland also demonstrated a subsequentcontraction and readjustment of the species distribution, possibly due either to changes in waterchemistry or to subsequent climate deterioration.
A study of lake deposits on the Scottish isle of Skye by Birks (1969) suggests that C. demersumsuffered a restriction of range southwards since the Late Weichselian glacial period, a notion whichsupports the case for a present day temperature limitation on the species.
Names
The genus name 'Ceratophyllum' is from the Greek 'keras', meaning a horn, and 'phyllon', a leaf, the manydivisions of the leaves suggesting the shape of horns. The Latin specific epithet 'demersum' means 'downunder the water' or 'submerged'. The English common name 'Rigid Hornwort', is a so-called book name andthe plant does not appear to be associated with any folk traditions. Despite having 'wort' as part ofthe name, the species does not have any use in herbal medicine that the present author can detect.
Native, very rare, almost certainly an identification error. Eurosiberian temperate.
August 2010; EHS Lake Ecology Team Survey; shore of Cornaleck Td, Upper Lough Erne.
Growth form and history of possible Fermanagh discovery
Although this free floating or lightly anchored aquatic has a similar ecology and reproductive capacityto C. demersum (Rigid Hornwort), in Ireland it is extremely rare, having been mapped in theNew Atlas in a total of just three hectads in two widely separated regions near the east coast inCos Wexford and Down (H12 and H38). A water quality survey carried out by the Water Management Unit(WMU) of the Northern Ireland Environment Agency (NIEA) at 15 sites around Upper Lough Erne in August2010 produced two unexpected records that were communicated to the current authors in late October,2010. The two sites were the one listed above and another at Knockninny Quay. Plant material was locatedat depths of 0.7-1.6 m at Cornaleck, and a small quantity only was found at a depth of 1.6 m atKnockninny Quay.
C. submersum was first recorded in Ireland in August 1989 when the NI Lakes Survey discovered itin two lakes on the Lecale Peninsula near Downpatrick (Smith & Wolfe-Murphy 1991). To date, theseare the only known sites for Soft Hornwort in Northern Ireland. We therefore made enquiries to ascertainif vouchers had been collected at the Upper Lough Erne sites, and whether or not the identification ofthe species material had been checked by a known botanical authority. Plant material had indeed beencollected and was identified in the WMU laboratory in Lisburn. Unfortunately, specimen samples were notretained. The determination of C. submersum from C. demersum was based entirely on thedividing of the leaves, "into threes, rather than dichotomously" (B. Walker, pers. comm. 18February 2011).
The significant leaf distinction between these two Ceratophyllum species refers to the number oftimes the leaf segments divide dichotomously: once or twice forked for C. demersum, threeor four times forked for C. submersum. The leaf segments are obviously toothed in C.demersum and much less so in C. submersum, and the leaves are more rigid and darker incolour in C. demersum in comparison with C. submersum (Stace 1997).
Without vouchers these two C. submersum records are very doubtful and cannot stand as First andSecond County Records. We believe the plants were incorrectly identified. However, the possibilityremains that Soft Hornwort just might be present in eutrophic, base-rich Fermanagh waters growing alongwith C. demersum and it should certainly be looked for. The point has been well made by therecently retired Curator of the BEL herbarium, that, "it is a regrettable tendency thesedays for agencies and consultants not to collect or retain vouchers. It undermines all the time andexpense spent in doing surveys." (P. Hackney, pers. comm. 23 February 2011).
C. submersum is reputed to flower more freely in Britain and Ireland than C. demersum, andalthough vegetative reproduction by fragmentation is certainly the norm in both species, the small,hard, nut-like fruits may well be dispersed by waterfowl, thus enabling jump dispersal of the species(Preston & Croft 1997). On account of wider geographical distribution and biodiversity aims, C.submersum is included on the NIEA list of Priority Species of special concern requiring localconservation action.
Names
The genus name 'Ceratophyllum' is from the Greek 'keras', meaning a horn, and 'phyllon', a leaf, the manydivisions of the leaves suggesting the shape of horns. The Latin specific epithet 'submersum' means'submerged'. The English common name 'Soft Hornwort', is a so-called book name, and the plant does notappear to be associated with any folk traditions. Despite having 'wort' as part of the name, the plantis rare and does not have any use in herbal medicine that the present author can detect.
Native, common and very widespread, locally abundant. Circumpolar wide-boreal.
1881; Stewart, S.A.; Co Fermanagh.
February to November.
Growth form and preferred habitats
A perennial with a short, thick, rather tuber-like horizontal rhizome, C. palustris occurs in avery wide variety of wet habitats, including the margins of rivers, streams and ditches, in wet woodsand fen carr around lakes and ponds where it grows well in partial shade in low-lying ground. Vegetativegrowth begins early in the year and, like Ranunculus ficaria (Lesser Celandine) and Anemonenemorosa (Wood Anemone), it is pre-vernal in its flowering when growing in deciduous woodland.It also occurs more locally in seasonally flooded meadows and pastures where it can sometimes form largedominant patches, but it does not tolerate permanently flooded ground (Grime et al. 1988).
Typically C. palustris prefers neutral to base-rich mineral or fen-peat soils, rather than veryacidic ones and it is, therefore, rare or absent on boglands and on soils with a pH below about 4.5(Grime et al. 1988). Like several other members of the Ranunculaceae, it tends to frequent areasof disturbed ground with relatively high exposures of bare earth and it often grows on banks of streamsand lakes close to the water's edge where fluctuating water levels help minimise competition fromtaller, tufted, potentially dominant emergent species (Grime et al. 1988). As with most wetlandspecies, it is chiefly found in lowland situations, but it is known to ascend to 1,100 m in Scotland(Wilson 1949). With increasing altitude the species occurs more frequently in open, completely unshadedsituations.
Reproduction
In the normal upright growth form of the plant there does not appear to be any form of vegetativepropagation, rather reproduction and dispersal relies entirely on seed. However, Grime et al.(1988) reported that shoots detached during disturbance are capable of regeneration, a feature whichdeserves to be further investigated, since, for example, trampling by grazing livestock and draincleaning operations must often result in plant fragmentation.
The showy and very familiar flowers of Marsh-marigold are produced early in the year from March onwardsand, depending on the season, they continue flowering through to July or even August. The flowers arepetal-less but have five to eight sepals, glossy yellow above, often greenish beneath, which take on theprotective and insect-attracting, advertising roles of petals. Flowers of C. palustris areself-incompatible and are visited and pollinated by a variety of insects, including flies, bees andbeetles, which collect both nectar (secreted by the several carpels at their base and, therefore, partlyconcealed, though generally copiously produced) and plentifully produced pollen as food sources(Hutchinson 1948, p. 139; Proctor & Yeo 1973; Fitter 1987; Jonsell et al. 2001).
Fruit and seed production
The distinctive star- or crown-like fruit (hence the English common name 'Kingcup'), usually containsfive or six compartments (ie follicles), which split open along their upper sides to reveal a double rowof seeds (Melderis & Bangerter 1955). Salisbury (1942) found that a considerable proportion of theovules formed normally abort, so that while the usual number of ovules per follicle is 17 or 18, thenumber of seeds is generally much fewer, the calculated mean being nine per follicle. The average plantexamined by Salisbury (1964) produced 2,700 seeds, though as Butcher (1961) earlier pointed out, thespecies is very variable in both reproductive and vegetative respects, and therefore such estimatesshould be regarded as merely a guide.
Seed dispersal
There is no specialised seed dispersal mechanism: seeds simply drop out of the boat-shaped open follicleand since they possess a corky swollen attachment, they may float for between one and four weeks (Ridley1930, p. 198; Jonsell et al. 2001). After flowering the leaves of the plant have been observed toincrease very considerably in size (Step & Blakelock 1963).
Seed germination and longevity
Caltha seeds have a well-defined chilling requirement which ensures that they remain dormantduring winter and germinate in the spring (Grime et al. 1988). A survey of soil seed bank data inNW Europe has provided information from 18 sources, 12 of which regarded the seed as transient, two asshort-term persistent (ie surviving for 1-5 years), and two as long-term persistent (ie surviving for atleast five years). The two remaining studies were undecided as to which category the species fitted(Thompson et al. 1997). Clearly there is no consensus on this important matter and further studyis required.
Genetic variation and the 'creeping' form: In the Typescript Flora ofFermanagh, which details finds up until the early 1950s, Meikle remarked on variants of C.palustris with slender, hairless, creeping stems rooting at the nodes and bearing sparseinflorescences of small flowers that are frequent on lake shores and in wet woods in Castlecoole andelsewhere in Co Fermanagh. The extremes of variation in C. palustris are very distinct and thedifferent forms are said to remain distinct (though rather less so) in cultivation. However, it is noteasy to distinguish them satisfactorily from typical C. palustris at a taxonomic level, becausenumerous intermediate forms exist. Stace (1991, 1997) believes that these few-flowered, procumbentplants are best recognised as var. radicans. In the critical Flora Nordica volume 2(Jonsell et al. 2001), Piirainen elevates this variety to subspecific level as subsp.radicans (T.F. Forst.) Syme. Whatever we decide to name it, this creeping form was first recordedin Fermanagh by Barrington (1884) as C. radicans (Forst.), and although Meikle states that thevariant has been recorded by him and his co-workers in all four of their district subdivisions of thecounty (Meikle et al. 1975), subsequent local recorders have tended to ignore the form, and infact there are just three records of this form of the plant in the Fermanagh Flora database. Theprostrate creeping form of the plant spreads both by seed and by rooting shoots, so it definitelydemonstrates vegetative reproduction. The seeds of this form of the plant lack the corky appendage anddo not float, which must restrict its dispersal considerably (Jonsell et al. 2001, p. 331).
Genetically, in terms of chromosome numbers, C. palustris contains a polyploid series ranging fromdiploids to octoploids, plus a number of aneuploid
forms. However, the chromosome variation is not correlated with variation in form, nor with geography(for details see Jonsell et al. 2001, p. 329).
It is interesting to note that argument regarding the taxonomy of C. palustris and the status ofits varieties has rumbled on since 1768, when Philip Miller produced the 8th edition of his GardenersDictionary. [NB There is no apostrophe in Gardeners in the original work]. In this work heproposed distinguishing Caltha minor for montane (ie upland) plants having round, crenate,heart-shaped leaves and smaller flowers than normal C. palustris. This suggestion was made only15 years after the Swedish taxonomist, Carl Linnaeus, described and named C. palustris (Panigrahi1955). Despite its age, the author of this latter paper has conveniently summarised the history of thedebate on the several forms of C. palustris, their morphology, cytology and geography.Panigrahi gives full weight to several significant contributions based on Irish material by Robert LloydPraeger. Anyone requiring a more modern treatment of the subject should consult Woodell &Kootin-Sanwu (1971) or Jonsell et al. (2001).
Grazing and toxins
The fact that it can become so prevalent in wet grassland is unfortunate for the farmer and his animals,since like other members of the Ranunculaceae, Caltha contains a significant quantity of thedangerous toxin protoanemonin (for details of this poison see the Ranunculus acris synopsis).This makes plants acrid tasting and they are avoided by stock animals to such an extent that Cooper& Johnson (1998) have not reported any recent cases of poisoning in the British Isles. Salisbury(1964) comments that C. palustris acts as a purgative and that it leads to diminished milkproduction in cattle which graze it. He also reports it being known to cause death in both cattle andhorses.
Fermanagh occurrence
Marsh-marigold is the tenth most frequently recorded vascular plant in Fermanagh in terms of records andit is the ninth most widespread species in the VC, being found in 376 tetrads, 71.2% of our totaltetrads. It is absent only from the highest altitudes, permanently flooded, or excessively acidic soilsbelow pH 4.5.
British and Irish occurrence
C. palustris is common and very widespread throughout Great Britain, but is absent from areas ofintensive farming such as Cambridge. It is also common and widespread in Ireland, but more or lessabsent from the more acid boglands of the S & W. Rather remarkably, it is a rare and introducedspecies in the Channel Isles (Garrard & Streeter 1983; Preston et al. 2002). Despite thedestruction of many wetland areas in Britain and Ireland during the past 40 years, the index of changecalculated and provided in the New Atlas is -0.26. At least at the hectad scale, this suggeststhere is little evidence of any change in the species distribution in these islands (R.A. Fitzgerald,in: Preston et al. 2002).
Fossil record
Seed and pollen of C. palustris are well represented in the fossil record although Calthatype pollen also includes Aquilegia vulgaris (Columbine). Godwin (1975, p. 119) suggests that theabundance of site records in the Late Weichselian ice age, strengthens the possibility that the species,"survived the glaciations in this country". One has to presume that as an English writerGodwin was referring to his native island (Great Britain) alone, but perhaps he did not mean to be soinsular, and really meant to specify periglacial survival of C. palustris in all unglaciatedareas within Britain and Ireland. An opinion on this matter from a suitably qualified authority would beappreciated.
World distribution
In Europe, C. palustris is widespread in both temperate and Arctic areas, but in Europe it becomesscarcer towards the south and around the Mediterranean, reaching only N Portugal. It is absent from theGreek Peloponnese and from all the Mediterranean islands (Jalas & Suominen 1989). Due to differingviews of the taxonomy of Caltha palustris, the species has been mapped in different ways; forexample, Hultén recognises and maps several different subspecies so that it becomes necessary tomentally combine these variants to achieve an overall picture (Hultén 1971, Maps 75 & 76). Once thisis done, C. palustris in this broad sense is shown to be circumpolar in Arctic and boreal,northern temperate regions.
Uses
Considering the fact that the plant is not a Near Eastern nor a southern species, and thus it neverattracted the attention of the older Classical civilisations who handed on a knowledge of plants andtheir uses to Mediaeval and Renaissance Europe, the long list of English common names that Grigsonprovides must derive almost entirely from the colourful show the plant makes in the countryside and inthe water garden. Its prominence, even in habitats disturbed by man and his animals, and the religiousand folk beliefs associated with it, rather than any widespread use in folk or herbal medicine, musthave made Caltha palustris very much a favourite plant (Grigson 1987).
Being a severe irritant and causing blisters, it was widely known to have caused serious effects whenattempts were made to use it in herbal medicine. Nevertheless, it did have some medicinal use associatedwith mythical beliefs on the treatment of fits (Grieve 1931, p. 519). The leaves were also boiled andeaten like spinach and flower buds were occasionally used like capers − although even Grieve regardingthis type of usage as "rather inadvised" [my italics].
Names
The genus name 'Caltha' is derived from the Greek 'kalathos' meaning 'a cup', a reference to thebowl-like shape of the flowers. The Latin specific epithet 'palustris' means 'of marshes' (Melderis& Bangerter 1955). C. palustris has a huge number of English common names. Grigson (1987)lists no fewer than 93. The most widespread English name, 'Marsh-marigold', refers to its use in churchfestivals in the Middle Ages, as a flower devoted to the Virgin Mary. Several other names also refer toMary and to May-Day festivals (Grieve 1931; Grigson 1987).
Threats
Many of the plant's meadow sites are vulnerable to drainage operations and could easily be destroyed.However the plant is so widespread and abundant in Co Fermanagh that the threat level to the species isnil.
Native, very rare. European boreal-montane.
1896; Mr Pike, of London; Gorminish Island, Lough Melvin.
January to September.
Growth form and preferred habitats
In Britain and Ireland, this rather lovely perennial typically occurs in cool, damp,often shaded habitats, including lake margins, stream and river banks, open woodland or their margins,traditionally managed hay-meadows and rarely on upland rock ledges where the plants may never flower. Itneeds protection from grazing by sheep and goats to survive. Even when it is not flowering, the largebasal palmate leaves of the species are perfectly recognisable. The plant prefers basic soils and isgenerally associated with limestone districts, and especially in sites where there is some degree ofnutrient enrichment from mesotrophic flushing groundwater.
Fermanagh occurrence
T. europaeus occurs, sometimes in considerable quantity, in just one very specialised Fermanaghhabitat. It occupies a narrow zone about 2 or 3 m wide, just below the winter high water-mark, on therocky limestone shores of two of the larger lakes in the county under the light shade and shelter ofalder, willow and rose scrub woodland. The species has been recorded from a total of five Fermanaghtetrads on the shores of Rosskit, Gorminish Island and Bilberry Island on Lough Melvin, and from RushinPoint and a couple of nearby sites on Upper Lough Macnean, where it also occurs on the Co Cavan side ofthe lake (Northridge 1995). RHN has visited the Upper Lough Macnean shore when only the flowers andupper leaves were emergent from the water surface, presumably after a period of heavy May rainfall!
Irish occurrence
Apart from the sites just mentioned, the only other extant Irish stations for T. europaeusare in Co Donegal (H34 or H35 or both?) (three or four sites),although previously it also occurred in Co Leitrim (H29). It might well survive on another, differentBilberry Island, which occurs in Co Leitrim, the one that lies just south of Patrick's Island, but thisstill needs to be investigated (Northridge 1995).
British occurrence
T. europaeus is a decidedly northern species in the British Isles, being much more common inScotland and Cumbria than elsewhere. The British distribution is totally confined to the area north of aline between Cardiff and Whitby (R.A. Fitzgerald, in: Preston et al. 2002). This distributionpattern is very similar to that of Cirsium heterophyllum (Melancholy Thistle) (another Irishrarity present in Fermanagh), and in Britain the two species are often associated in damp calcareousgrassland (Clapham 1978).
European occurrence
In Europe, T. europaeus is native and widespread throughout N and C regions, but it is absent frommost of the westernmost parts of the continent, although widespread and frequent again in the mountainsfurther south from Spain eastwards to Greece (Jalas & Suominen 1989, Map 1557). In Scandinavia, itextends beyond 70°N and it also stretches eastwards to W Siberia (Jonsell et al. 2001), and onthe margins of its indigenous distribution, it is occasionally recorded as an established introduction(Jalas & Suominen 1989).
It was been shown by Conolly & Dahl (1970) that the distribution pattern of T. europaeus liesnorth of the current 27°C maximum summer temperature isotherm, quantifying the basis on which it occursin Britain and indeed on most of the European Continent. In these areas, it is considered a species ofnorthern or montane to sub-alpine damp, calcareous meadows and pastures, stream banks, the margins ofwoods and rarely on cliff ledges (Halliday 1997).
Indigenous populations in Scandinavia occupy a rather wider range of habitats than in Britain andIreland: there it occurs in varying conditions of light, open to moderately shady habitats, on moist tomesic, nutrient-rich soils that are usually refreshed and enriched with moving groundwater. Typicalhabitats include tall-herb vegetation, scree, fen margins, Willow and Alder thickets, and the banks ofwater courses. Also in Scandinavia, T. europaeus is closely associated with man-managed'apophytic' situations, including wooded pasture, forest glades, former hay-meadows, field margins androadsides (Jonsell et al. 2001). In many of these latter more regularly disturbed stations, T.europaeus, in Scandinavia at least, is recognised as an established introduction, its seed veryprobably being transported around the countryside by cutting machinery and the hay fodder market.
Decline in range and environmental pressures
In both Britain and Scandinavia, T. europaeus appears to have declined in range over a longperiod, probably for at least a century. As usual in such situations, decline and retreat isparticularly obvious at the margins of the previous distribution. (Halliday 1997; Jonsell et al.2001; R.A. Fitzgerald, in: Preston et al. 2002). In part, T. europaeus appears to beretreating under pressure from grazing animals, yet like other members of the Ranunculaceae, it containsthe blister-inducing bitter toxin protoanemonin, which in theory ought to deter browsers and give thespecies a measure of protection (Cooper & Johnson 1998). However, further north, Arctic Reindeer areknown to graze T. europaeus and other northern representatives of the plant family, andGlobeflower seeds are internally transported and occur in reindeer excreta (Ridley 1930, p. 373). Thelevels of toxin in Trollius may be insufficient to deter deer and other large herbivore species,so it might be possible for cattle, sheep and goats to browse on it without suffering ill effects, againassisting the plant in its seed dispersal. Work is required to investigate these possibilities inBritain and Ireland.
In Britain, the main cause of the obvious decline in the species appears to be the agriculturalimprovement of land (especially in hilly areas), by drainage, the earlier cutting of herbage for silagethan previously for hay, and the widespread application of fertilisers stimulating the growth of morevigorous species with which T. europaeus simply cannot compete (Halliday 1997; R.A. Fitzgerald,in: Preston et al. 2002). In Ireland, populations on or near river and stream banks have alsobeen destroyed by flash floods (eg in Co Donegal), a weather related phenomenon which seems to beincreasing in frequency and may well be related to global climate change (Curtis & McGough 1988, p.100; Sheppard 1991).
Garden escapes and relict status
Globeflower is a longstanding, popular garden border perennial that is undemanding and easy to cultivate.While it must occasionally 'jump the garden fence', as it is claimed it does in Scandinavia (Jonsellet al. 2001), in Britain and Ireland there are only a few stations on the New Atlas mapwhich suggest or indicate such a happening (Preston et al. 2002).
In Ireland, T. europaeus and the other members of the Northern Montane phytogeographical group areregarded as relict plants of past climate stages and much changed environments that are approaching theend of their local occupation (Matthews 1955, p. 117). This understanding fits very well with therestriction in habitat found in Fermanagh, the contraction in range elsewhere, especially in Ireland,and with the developing picture of Global warming (or in our corner of the British Isles, moreaccurately Global wetting and blowing) (Plantlife Report: Death Knell for Bluebells? GlobalWarming & British Plants, Anon. 1991).
In view of the large number of apparently suitable sites for T. europaeus on Lower Lough Erne, itis sad to think that the species nowadays appears to show no capacity whatsoever for natural dispersaland colonisation over the relatively short distances involved (8-13 km).
Fossil record
The fossil record is sparse but is sufficient to confirm the native status of Globeflower (Webb 1985). Itconsists of a tentative Flandrian post-glacial seed record from Ireland of Mesolithic age, and pollenrecords exclusively from Scotland throughout the entire post-glacial (Godwin 1975).
Sexual reproduction
T. europaeus has a short, stout, erect, fibrous rootstock and it has no powers of vegetativereproduction, instead relying entirely on seed production for increase and dispersal of the species(Clapham et al. 1962).
Insect pollination
Globeflower has a most interesting relationship with three species of small flies belonging to the genusChiastochaeta. The globular yellow flowers produced from June to August never open theiroverlapping petal-like sepals, yet despite this (or maybe because of it and the concealment andprotection this floral behaviour affords the flower's pollen and nectar food supplies), they are visitedby adults of these species of fly which meet, mate and feed on the hidden food sources. IndividualTrollius flowers last for only about five or six days and they are self-incompatible. Aftermating inside one of the closed flowers, the female insects lay their eggs (usually just one perflower).
The Chiastochaeta fly species differ slightly in their sexual behaviour, for example, the stage offlowering at which they lay their eggs, the exact positioning of the eggs, and the paths along which thelarvae bore inside the carpels during their development, and in this manner the fly species manage toavoid competition to a remarkable degree, ie strict resource partitioning is present (Pellmyr 1989). Theinsect species which lays its eggs earliest during the short life of the flower is the one which is mosteffective in achieving pollination, but this is entirely incidental as far as the insects' movements areconcerned.
Flower-insect mutualism
Each flower is reputed to produces a rather large number of ovules (around 400 or so) and only a fewyoung seeds of each flower are eaten by the fly larvae, so the cost to the plant is fairly minimal. Thisis reckoned to be a fine example of an obligate mutualistic relationship occurring in the temperatezone. Mutualism is a phenomenon that is associated much more frequently with biology in the tropics(Pellmyr 1989). Although there is a cost to the plant, the relationship with these small fliesessentially benefits both partners, and the trade-off by the plant of a few of its many seeds enablescross-pollination of its un-opening, self-incompatible flowers.
When fertilised, the numerous carpels of T. europaeus develop into many-seeded follicles, whichwhen ripe split to release the seed.
Seed dormancy and germination
The seed is dormant at dispersal and requires cold-stratification during the following winter to breakdormancy (Milberg 1994). Experimental germination was equally effective in both light and darktreatments, which suggests that seeds might germinate even when they are too deep in the soil forseedlings to emerge. In turn this suggests that Trollius does not form a persistent soil seedbank and indeed this appears to be the case. In an experimental study comparing Primula veris(Cowslip) with T. europaeus, after 16 months burial, 85% of the Primula seeds, but only 8%of the Trollius seeds remained viable (Milberg 1994).
Competitive ability
T. europaeus is a poor competitor and in Finnish meadows has been known to suffer a prolongeddelay in maturity, successful flowering and fruiting being curtailed for up to eight years. In theabsence of competition, the species can fruit in its first season (Linkola 1936, quoted in Salisbury1942, p. 54). The species is polycarpic and probably long-lived, continuing to fruit for a number ofyears after attaining maturity, so that the loss of between two and seven years' seed output in Finlandmay represent only a small percentage of the total production throughout the plant's life. However, theprolongation of the juvenile phase represents a greatly increased risk of mortality prior to anyreproduction and this could be a significant factor determining the survival ability of the species in aparticular plant community (Salisbury 1942). This delay in plant maturity is sometimes referred to as'the demographic penalty', affecting the overall 'fitness' of a plant population (Silvertown &Lovett-Doust 1993, p. 158).
Plant and flower size plasticity
The flowers of T. europaeus at the Fermanagh lakeshore sites are much smaller, approximately athird the size of those the author has often observed elsewhere, including those in Teesdale, Co Durham(VC 66) (Clapham et al. 1978) and regularly found in the Swiss and Italian Alps and the Pyreneesat very much higher altitudes. This suggests a particular genetic clone with smaller flower occurs inFermanagh, perhaps a response to a challenging waterside environment.
All plants allocate their limited photosynthetic energy and mineral nutrient resources under constraintsand limits imposed by their local environment which ultimately acts upon the plant genotype, but isexpressed in the readily observed and more easily measured variation of the population phenotype(Silvertown & Lovett-Doust (1993). Conflicting demands inevitably lead to trade-offs betweendifferent activities. Two classes of trade-off are virtually universal: (i.) a trade-off betweenreproduction and other activities which is manifest as 'the cost of reproduction', or in proportionalterms as the 'reproductive effort' (RE); and, (ii.) a trade-off between the size and the number ofoffspring produced. For perennial plants, absolute allocation to reproduction usually increases withplant size. However, depending on the species under examination, the proportional reproductiveallocation, ie the reproductive effort (RE), can either decrease, increase, or remain independent ofplant size.
Comparative reproductive effort
A study of reproductive effort of populations of T. europaeus and Ranunculus acris (MeadowButtercup), in subarctic Swedish Lapland compared the species at differing altitudes up to and above thetree-line (at 870 m). This showed that for both species plant size, measured in terms of 'mean plantbiomass' (ie mean dry weight), was approximately two or three times larger below the tree line comparedwith above it (Hemborg & Karlsson 1998). For Trollius, plants at the high altitude sitesshowed no relationship between RE and size, while below the tree line, RE varied inversely with plantsize, ie reproductive effort generally decreased as the Trollius plants became larger.This might be due to a limitation imposed by the number of flowering meristems the Swedish populationcould support, since all the plants in the study, irrespective of size only bore one flower per stem.Moreover, floral structures, eg flower size and ovule number, may be limited by low plasticity, featureswhich are very likely both genetically and environmentally regulated (Schmid & Bazzaz 1992).
At low altitudes in Britain and Ireland, and in the French Alps, most plants of T. europaeus havetwo or three flowers per stem, the lateral flowers usually being smaller than the terminal. This factalone would produce different size-effects on reproductive effort in comparison with this Swedish study(Hemborg & Karlsson 1998).
Toxins and herbal use
Globeflower contains protoanemonin, the same toxic principal present in other members of the family.Being a decidedly local plant, T. europaeus has never had the herbal medicinal reputation anduses of its more common and readily available relatives. The exception to this is perhaps found inRussia and parts of Sweden where the species is very prevalent (Grieve 1931).
Names
The genus name 'Trollius' is a Latinised version of the older Swiss-German name 'Trollblume', firstcoined by Gesner in the 16th century, which translates as 'rounded flower' (Gilbert-Carter 1964;Gledhill 1985). 'Trollblume' may well be a contraction of 'die rolle Blume', referring to the rolled in,or closed in, petals of the flower (Grigson 1987, p. 31). The Latin specific epithet is geographical andobvious.
There are 16 local English common names listed by Grigson (1987), of which various forms and spellings of'Locken Gowan' are the most frequent. 'Gowan', 'Gowlan' and so on are derived from the Anglo-Saxon'gold' meaning 'yellow', while 'Locken', 'Lockety', 'Locker', 'Lapper' and so on, refer to the locked orclosed petaloid flower parts (Britten & Holland 1886; Grigson 1987). 'Locken Gowan' has also beensometimes used to refer to Caltha palustris (Marsh-marigold) and there is a fair degree ofoverlap with some of the other English common names between these two species. Other local names aremore predictable not to say prosaic, eg 'Goldilocks', 'Golden Balls' and, of course, 'Globeflower'. Thecurrent author particularly likes the imagery generated by the name 'Bull-Jumpling', which hails fromKinross-shire: the 'bull' part is most probably a corruption of the Old English, 'boll', meaning, 'anyglobular body' (Prior 1879). The origin of 'Jumpling' we can only guess at. Another name mentioned byGerard (1597) in his Herball, he being the first English writer to mention the whereabouts of thespecies in northern England, was 'Troll floures', or 'Troll flower'. Grigson (1987) is dismissive ofthis name, emphasising his belief that Trollius is not a flower in any way linked to the evilNordic trolls. Prior (1879) disagrees though, comparing the name with the Scottish 'Witches Gowan'.According to him both these names were given on account of the plant's acrid poisonous properties.
Threats
In Fermanagh, reclamation of shore lines and, at least on Bilberry Island, grazing by goats. Potentialshoreline building development could also prove deleterious.
Introduction, neophyte, garden escape, very rare, or a possible mis-identification, but very probably nowextinct.
21 August 1986; Waterman, T. & Brain, P.J.T.; Derrymacrow Lough, near Crom, Upper Lough Erne.
There is just a solitary record for this often short-lived, but occasionally well-established perennialin Fermanagh. The site for this particular plant record is given as, "grassland and woods to thewest and south of the lake". It could be a correct identification of a garden plant naturalised onthe Crom Estate, or might possibly be a mis-identification of another naturalised colony of H.viridis (Green Hellebore). It has only been recorded once and therefore very probably has diedout.
Both these Helleborus species occupy rather similar semi-shaded wood or scrub habitats, on shallowcalcareous soils, and their biology and poisonous properties also closely parallel one another. The mostnotable differences are the wintergreen leaves of H. foetidus and its very much higher andpotentially lethal content of the toxic protoanemonin (Cooper & Johnson 1998).
Elsewhere in Ireland, H. foetidus is a rare, winter-flowering garden escape. Omitting Fermanagh,the Cat Alien Pl Ir lists just seven other Irish VCs with records, including in the North, CosTyrone and Armagh (H36 and H37).
Names
Some authorities derive the genus name 'Helleborus' from the Greek 'helein' or 'elein', meaning 'to kill'or 'to injure', and 'bora', meaning 'food', indicating the poisonous properties of the plants (Melderis& Bangerter 1955; Cricheley Plowden 1972). Other writers suggest the name 'helleboros' was theancient classical Greek name for H. orientale (Gilbert-Carter 1964; Stearn 1972).
The Latin specific epithet 'foetidus' translates as 'stinking' or 'bad smelling' (Gledhill 1985).
The English common name 'Stinking Hellebore', listed above, is a straight translation of the botanicalname and as such is a mere book name (Britton & Holland 1886). Other, much more interesting commonnames with genuine, local folklore connections do exist (Prior 1879; Britten & Holland 1886; Grigson1955, 1987). These include 'Bear's-foot' and 'He-barfoot' (presumably referring to the digitate leafshape), and 'Setter' and 'Setterwort', the latter a name used in his early English herbal by John Gerard(1597). The derivation of the 'Setter' names, which are associated with healing cattle, is given in thiswork under Helleborus viridis and will not be repeated here. Other local English names for H.foetidus with cattle connections are 'Gargut root' (originating in parts of Norfolk), 'gargut'being "a disease incident to calves" (Britten & Holland 1886, p. 199), and 'Ox-heal', fromthe Anglo-Saxon 'oxnalib', again referring to settering cattle with the plant root (Prior 1879, p. 174).
Introduction, neophyte, garden escape, rare.
12 July 1946; Carrothers, E.N., Meikle, R.D. & Moon, J.McK.; Rossclare Bay, near Killadeas, LowerLough Erne.
February to July.
H. viridis, H. foetidus (Stinking Hellebore), H. niger L. (Christmas-rose) and H.argutifolius Viv. (Corsican Hellebore), plus numerous other varieties, subspecies, species andhybrids are common and popular tuberous garden perennials grown for decoration throughout these islands(Mathew 1989; Griffiths 1994). Green Hellebore regularly escapes from cultivation and it appears to havebecome well established and naturalised in at least some Irish sites (An Irish Flora 1996; CatAlien Pl Ir).
Irish and British occurrence and status
In Ireland, H. viridis has always been accepted as being a persistent alien introduction, but inBritain small, permanent, supposedly or traditionally native populations are widely distributed. Theplant usually occurs on chalk or limestone in shady lowland habitats such as wooded glades, rockystream-sides and in old hedgerow banks. However, H. viridis has been grown in gardens throughoutBritain and Ireland since medieval times (Harvey 1990; Landsberg 1996) and it was first recorded 'in thewild' around 1562. It is therefore difficult or virtually impossible to distinguish native fromintroduced populations and, after so long a period in cultivation, it appears rather foolish to even tryto do so.
Fermanagh occurrence
H. viridis has been rarely recorded in ten tetrads, seven of them with post-1975 records. In thelate 1940s and early 1950s, when Meikle and his companions were exploring the flora of Fermanagh, theyrecorded five widely spaced stations of the plant. Nowadays we know of six scattered populations,including two that they originally discovered − at Rossclare Bay and near Monea Castle. The typicalhabitats include woods, scrub, shaded river banks, and on scree below cliffs, usually on limestone.Meikle recalled seeing the plant in scrub on the limestone shore of Lower Lough Erne at Rossclare onfamily holidays in the 1930s (Carrothers et al. 1947), and a large patch of about 18 plantssurvives to this day in the same area. At Coffey's Ford, SW of Kinawley, there are 20 large patchesgrowing beside a stream, clearly indicating that the plant is reproducing successfully.
In the Revised Typescript Flora of Fermanagh, Meikle remarked that the plant was often found farremoved from gardens and that it appeared, "perfectly spontaneous". He also suggested thatsome of the existing populations might have been derived from stocks of the plant deliberatelycultivated for cattle medicine (see below) (Meikle et al. 1975).
Growth form and reproduction
The species has a short, stout, ascending, blackish rootstock as its perennating organ, but the attachedpair (or more) of leathery radical leaves and the aerial stem do not overwinter. Being a long-livedperennial, the current rootstock of the plant tends to be surrounded by a substantial woody cluster ofold decaying stem-bases left from previous years (Ross-Craig 1948, Part 1, plate 38), but the plant hasno real means of vegetative reproduction.
Growth begins early in midwinter, and the individual plant produces its two to four yellowish-greenflowers around the second half of February or early March. Flowering generally continues on into April.Each flower contains 9-12 pocket-like green nectaries that attract early flying bees as pollinators. Theinsects collect both nectar and pollen from the flowers, both of which are urgently required to feed thedeveloping brood of the bee colony in the spring.
Seed production and snail dispersal
The flower usually has three carpels and the fruit is a many-seeded follicle. The individual seeds arerather large, 4.5 × 3 mm, and dark brown in colour. Salisbury (1942) examined a small sample of 25plants and calculated the mean annual seed output as 191, ± 48, per plant. The seeds possess an obviousappendage growing out from the seed coat. This is an edible elaiosome or oil-body that attracts antswhich carry off the seeds when they are shaken out of the open fruit onto the soil or other surface,thus assisting the species' dispersal (Beattie 1985).
It has also be reported that the elaisome of the related species H. foetidus attracts snails whichdevour the oil-body and in so doing get some of the seed adhering to their slimy body. Dymes (1916)observed that the snail resented the presence of seeds on its head or tail and actively sloughed themoff. However, the snail was quite unconcerned if the seed stuck to its body near the shell on the headside. A snail was observed to carry a seed in this manner for a distance of 35 cm. The Garden Snail(Cornu asperum = Helix aspersa), on average travels 5 cm per minute and Dymes measuredthem travelling a distance of 5.4 m for food. While molluscs move slowly, they do get around and theyare abundant and widespread in many types of plant community across Britain and Ireland. If snailscommonly carry seeds in the manner described, they would certainly play a significant role in seeddispersal (Ridley 1930, p. 150).
This is yet another plea for a return to careful natural history observation, something for which peoplein the British Isles were once famous, and could be again. No laboratory required; just eyes,imagination, time, patience and a notebook, although a hand-lens and a stereo-zoom low-power microscopewould certainly also be useful!
The solitary determination of the soil seed bank in the NW European survey indicates that the GreenHellebore seed is transient (persisting in the soil for less than a year) (Thompson et al. 1997).A correspondent of Salisbury's indicated that the rate of germination success is low (Salisbury 1942, p.178).
Reasons for population decline
The waning of H. viridis populations in Britain and Ireland that is apparent from the NewAtlas hectad map may partly be attributable to records of introductions which failed to persist.Additionally, some long-established populations are known to have been lost as a result of changes inagricultural and land management practices taking place during the last 50 years, including theclearance of hedges and copses, and the gradual cessation of woodland coppicing (R.A. Fitzgerald, in:Preston et al. 2002).
Toxins
The principal toxin hellebore plants contain is protoanemonin, as found in most other members of thefamily Ranunculaceae. The content of this irritant varies widely with the species. The scale of thevariation is demonstrated by the fact that measurements showed H. foetidus (Stinking Hellebore)contained 672 µg/g, while H. viridis had just 28 µg/g of the toxin present (Cooper & Johnson1998). Numerous other references (including Clapham et al. 1962) mention the presence of twofurther toxic glycosides named helleborin and helleborein, and the fact that the plant has a burningtaste.
Use in herbal medicine and related plant names
Decoctions of hellebores (both species covered here) were used in former years as purgatives, localanaesthetics, abortifacients, or to clear parasitic infestations of the skin or animal coat (Cooper& Johnson 1998). The association of garlic with veterinary application of Green Hellebore root incattle follows an old belief, possibly dating from the ancient Classical Period. This suggested thatsince hellebore was such a powerful herb, a certain amount of prayer and ritual should be observed whenlifting its rootstocks. "The person digging them up had either to chew on, or shortly before haveeaten, several cloves of garlic, simply to ward off the poisonous effluvia of its roots." (LeStrange 1977, p. 136).
In the Revised Typescript Flora of Fermanagh, Meikle records being told by a local hotelier whenhe was on an outing in the Lough Melvin area during 1949 that, "the plant is (or was) used for adisorder of cattle, being pounded with butter and garlic and rubbed into an incision in the animal'stail, 'until you could smell the garlic on its breath.'" We are not told what the cow was sufferingfrom, but other sources indicate it was used either as a purgative for worms, or to clear the skin andcoat of lice (Grieve 1931; Le Strange 1977).
While the hotelier in Fermanagh described the application of the remedy to the tail of the animal, otheraccounts tell of an incision being made in the cow's dewlap (the loose fold of skin hanging under theanimal's throat), and the Green Hellebore rootstock, or a preparation made from it, inserted into thewound. One of the less well known English common names of the species is 'Setterwort', which is derivedfrom the term used to describe the aforementioned process, which is 'settering' or 'pegging' the dewlap.Prior (1879, p. 213) who details this plant name and the medicinal term, mentions that 'setter' is acorruption of 'seton', derived from the Italian 'setone', meaning a large thread of silk. Possibly thethread was used to sew up the wound in the dewlap. An alternative name for Green Hellebore is'Pegroots', from the operation of 'pegging the dewlap'. Prior (1879) also indicates that settering wasused to treat lung problems in cattle, such as coughs or wheezes (Prior 1879; Grigson 1987).
Cases of cattle poisoning have occurred as a result of this process, as one could easily imagine, thesymptoms of which included prostration, loss of appetite, swelling of the neck, loss of condition of thecoat, muscular tremors and difficult breathing (Cooper & Johnson 1998). Clearly the plant is not oneto be handled more than necessary!
European occurrence
H. viridis has a strictly western discontinuous, native distribution on continental Europe,centred on France and stretching south to N Spain and N Italy, northwards to C Germany and east toPoland. The species has established alien status both within this range and to the north of it (Jalas& Suominen 1989, Map 1524). In plant geography, its distribution is summarised as suboceanictemperate (Preston & Hill 1997). The plant was introduced to New England for its somewhat dubiousmedicinal properties and it has become naturalised in N America (Grigson 1987).
Names
Some authorities derive the genus name 'Helleborus' from the Greek 'helein' or 'elein', meaning 'to kill'or 'to injure', and 'bora', meaning 'food', indicating the poisonous properties of the plants (Melderis& Bangerter 1955; Cricheley Plowden 1972). Other writers suggest the name 'helleboros' was theancient classical Greek name for H. orientale (Gilbert-Carter 1964; Stearn 1992). The Latinspecific epithet 'viridis' simply translates as 'green'.
English common names additional to those already mentioned above include 'Bear's-foot' and 'Boar's-foot'− a bear is called a 'boar' in Scotland, especially in northern Scotland, according to a source quotedby Britten & Holland (1886, p. 55); the allusion is to the digitately lobed leaf of the plant (Prior1879). Another interesting name is 'Fellon-grass' which was applied to several quite different plants ofwhich Hellebores were just one. In Westmorland, the name was applied to H. viridis. A 'fellon'was a boil or swelling, most commonly encountered in children. Housewives grew the plant to treat thesechildhood skin problems and partly also to treat against worms. It was a dangerous treatment, however,and it sometimes killed both the worms and the patient (Grigson 1987).
Threats in Fermanagh
The old, well-established Rossclare and Monea Castle sites could be threatened by building development.
Introduction, neophyte, very rare garden escape.
1884; Barrington, R.M.; Castle Hume estate, Lower Lough Erne.
May to July.
Irish status
When it is found in Ireland, which is only rarely the case, A. napellus is always regarded as anaturalised garden escape. The plant generally occurs on roadsides and open areas on field margins.
Fermanagh occurrence
A. napellus s.l. has been recorded in Fermanagh on a total of eight occasions and at only fourscattered sites in recent decades. A possible reason for the near disappearance in the 'wild' of thisconspicuous, tall, beautiful, blue-flowered garden escape could be its well-known extremely poisonousnature, which may have led to its eradication by landowners when found in order to protect grazingstock.
The details of the other seven Fermanagh records are: Galloon Td, Upper Lough Erne, 1951, MCM & D;Arney village, 1952, MCM & D; Clonelly, NW of Kesh, 25 July 1976, Miss N. Dawson; Colebrooke Church,1 July 1997 & 24 June 2003, RHN; roadside at Killadeas, near hotel entrance, 24 May 2002, RHN; androadside at Cornamucklagh Td, NE of Brookeborough, 29 May 2004, RHN.
Growth form and reproduction
Monk's-hood has a blackish tuberous taproot or rootstock as its perennating (ie overwintering) organ,from which arises a usually unbranched flowering stem up to 1.5 m tall, but generally less. The planthas no means of vegetative reproduction and relies entirely on seed for its increase and dispersal. Thecowl-hooded or helmet-shaped deep reddish-violet or purplish-blue flowers are produced in a longterminal raceme that may consist of around 30 flowers if the stem is unbranched and up to 100 whenbranched. The irregular flower conceals two long nectar-secreting spurs inside the hood, which areinterpreted either as petals or as staminodes (modified sterile stamens) (Blamey & Grey-Wilson1989). Each flower contains 3-5 carpels which, after the stigma has been pollinated by long-tonguedbumblebees, ripen to form upright, many-seeded follicles.
The seeds are rather large (6 × 3 mm) and they bear three wings, one of which is slightly wider than theothers (Butcher 1961; Clapham et al. 1962). The 'wings' are not very large, but they undoubtedlyassist the seed to travel slightly further through the air when shaken out of the censer-like fruit. Aswith Aquilegia vulgaris (Columbine), one would not expect the plant to have great powers ofdispersal, yet its occurrence in the wild indicates it is well able to escape from gardens on a regularbasis. Apart from this, the reproductive ecology of A. napellus appears a completely closed book.I have not located any information on seed dormancy, longevity or germination.
A supposed native British variant
A form of the plant referred to as A. napellus subsp. napellus (orsubsp. anglicum) has traditionally been considered indigenous in S Wales and parts of SW England (Watson1883; Druce 1932; New Atlas). The semi-native habitats it frequents are characterised bycalcareous to slightly acidic soils along stream banks that are often shaded, in damp, open woods ormeadows (R.A. Fitzgerald, in: Preston et al. 2002). This supposedly native British population ismapped by Jalas & Suominen (1989, Map 1576), showing in addition to its British distribution asolitary record of a plant of similar form in the Pyrenees.
Since A. napellus has been grown in gardens for centuries and first made a 'wild' appearance inthe British flora as late as 1821 (R.A. Fitzgerald, in: Preston et al. 2002), I find it quiteamazing that anyone today would simply assume and assert that a species or subspecies is native (and/orendemic) to a region without first making a rigorous objective examination of all the circ*mstantialevidence that might be assembled to support such a status (Webb 1985; Forbes 2000). The 'endemic native'plant is reputedly found flowering in early summer in shady riverside sites in SW England and S Wales(A.J. Silverside, in: Rich & Jermy 1998).
Aconitum napellus subsp. napellus is distinguished from many ofthe more widespread garden forms of the plant (which are sometimes of hybrid origin), by having lessdeeply cut leaves, but with more finely pointed ultimate leaf segments. It also has a slightly earlierflowering period and the helmet of the flower is hemispherical, not elongated (A.J. Silverside, in: Rich& Jermy 1998).
Apart from its taxonomy and alkaloid content, the plant appears little studied and I cannot locate anyrecent references on its biology or ecology, let alone on its status, apart from the brief treatment byJohn Akeroyd in Scarce plants in Britain (Stewart et al. 1994).
Irish occurrence
A. napellus s.l. is slightly less rare than elsewhere in Ireland in the six county province ofNorthern Ireland, where it has records in five counties (the exception is Co Armagh (H37). In theRepublic of Ireland, the species has only five widely scattered records (Preston et al. 2002).
British occurrence
In Britain, A. napellus s.l. is widespread throughout as a garden escape, but it has a greaterpresence in western districts of England and Wales, while north of the Scottish border it becomessomewhat more frequent in eastern areas. It is very possible that some of the mapped plants really arethe garden hybrid A. × cammarum L., a cross between A.napellus and A. variegatum (R.A. Fitzgerald, in: Preston et al. 2002).
European occurrence
A. napellus s.l. is a very variable species endemic to W and C Europe, its distribution on thecontinental mainland extending south to C Spain and stretching eastwards to the Carpathian mountains. Itis absent, however, from most of the Mediterranean basin (Jalas & Suominen 1989, Map 1575). Thetaxonomy of the species (or polymorphic aggregate of forms) is greatly confused by the recognition bysome of a range of subspecies (which others elevate to species rank), plus a history of very many namechanges. As is the case with Aquilegia vulgaris (Columbine), which sometimes occurs in similarshady and damp habitats to A. napellus, the comparative scarcity of the species and its insectpollination syndrome would hardly lead us to expect the appearance of either its pollen or its seed inthe fossil record, and indeed none exists (Godwin 1975).
Toxins
The plant contains a co*cktail of at least four alkaloids including aconitine, which even on its own ishighly toxic, so that Monk's-hood has the reputation of being the most poisonous plant in the BritishIsles (Cooper & Johnson 1998, p. 188). Perhaps because its poisonous nature is so well known andhence its subsequent removal from sites where grazing animals might find it, there are very few reportsin recent years of animal or human poisoning by the species in these islands.
Names
The genus name 'Aconitum' is Latin and is thought by some to be derived from the Greek name 'Akoniton'(although Gilbert-Carter (1964), for instance, regards the etymology as doubtful). 'Aconitum' is aclassical name first given to an unknown poisonous plant by Theophrastus (Gledhill 1985; Stearn 1992)and later reused by the Swedish botanist, Linnaeus, for the current genus. The Latin specific epithet,'napellus', is a diminutive of 'napus', which means 'little turnip', an obvious allusion to the tuber ofthe plant (Stearn 1992).
The English common name 'Monk's-hood' was first given by Lyte (1578), who described "The flowers beas little hoodes", translating the name directly from Dutch and German (Grigson 1974). In hisexcellent The Englishman's Flora, which deserves to be on every Celt's bookshelves too, Grigson(1955, reprinted 1987) remarks that the local names of the garden form of this very poisonous plant areall charmingly innocent. Most of them relate to the odd form of the flowers, "and especially to thefluttering, dove-like nectaries". They include 'Doves in the Ark', 'Lady Lavinia's Dove Carriage'.Many alternative names refer to bonnets, caps, helmets or hoods, for instance, 'Old Woman's Nightcap'and 'Face in Hood' (Britten & Holland 1886).
Fermanagh threats
None, as it is much too rare to be a threat to natural vegetation, or to grazing stock. With itsrecognised status as a rare, neophyte, garden escape, we are not concerned about threats of any kind tothe survival of Aconitum napellus.
Native, common, widespread and locally abundant. Eurosiberian temperate.
8 April 1862; Smith, T.O.; near Ardunshin.
March to November.
Growth form and preferred habitats
One of the prettiest if not the earliest of the welcome harbingers of spring in its typical habitat ofdeciduous woods and hedges, A. nemorosa is a perennial geophyte with a shallow, brittle, slender,brown rhizome which branches and forms clonal patches up to 5 m in diameter (Shirreffs 1985). As withother vernal species, A. nemorosa is really a shade-avoiding rather than a shade-tolerant speciessince it exploits the light phase in woods, scrub and hedges before the leaf canopy develops. While ittolerates a wide variety of soils, as a rule A. nemorosa is best developed on soils that aremoist to wet in spring. Such soils are often of relatively heavy texture or rich in humus, ie between 7%and 20% organic matter (Grime et al. 1988), or even more than this in Fermanagh.
The very superficial root system of A. nemorosa runs at a depth of just 5-10 cm, a feature whichexposes the plant to early season drought but which allows it to survive in much wetter soils than itscommon competitor, Hyacinthoides non-scripta (Bluebell), the bulbs of which are drawn downyear-by-year by contractile roots, often reaching depths of 15-25 cm where they may easily sufferwaterlogging (Grabham & Packham 1983). While this is the case, A. nemorosa is not really awetland species, it can merely tolerate moist soils on the fringes of marsh, fen and bog, although insome of these situations it may have to endure periods of submergence.
While Wood Anemone appears to prefer less fertile limestone woodland soils, it is not in any way confinedto them, but rather it may be expected in any shady situation including low growing ericaceous heath. Itdoes not, however, penetrate peat bogs where the soil pH falls below 3.5, which appears to totallyexclude the species (Shirreffs 1985; Grime et al. 1988).
In hedges, limestone pavement and long-established grassland, the presence of A. nemorosa canfrequently be associated with previous scrub or woodland cover, so that the plants are seen as remnantsof previous vegetation. When found on cliffs, however, or in sheltered spots above the tree-line, therecan be no question of relict status, and one wonders exactly how the species was transported to theseelevated sites. Then the plant typically occurs in rock clefts, or on sheltered, often north-facingledges, or shaded by overhanging sub-shrubs or trailing vegetation, all conditions providing theshelter, shade and high humidity the species requires.
Fermanagh occurrence
Locally, the species is very common and widespread, especially in lowland Fermanagh, having being foundin 290 tetrads, 54.9% of those in the VC. The most typical habitats it occupies are deciduous woods,hedgerows and river banks but, in the prevailing wet conditions of Fermanagh, A. nemorosa iscapable of extending into grassland and other open habitats, including more rarely, cliff ledges andscree on the talus slopes beneath cliffs.
British and Irish occurrence
Wood Anemone is common and widespread throughout Britain and Ireland, although absent from Orkney andShetland and rare in areas like the English Fens and other exposed situations where woodland (and indeedanything approaching dry land) are sometimes scarce. The previous dearth of records from the Republic ofIreland, which was regarded as probably under-recording (Shirreffs 1985), was remedied to a considerableextent in the New Atlas survey. The distribution remains fairly patchy in the Republic, except inparts of the far south and in the Dublin and Wicklow area, where there are more resident plant recorders(New Atlas). However, one must never overlook or underestimate the likely limiting ecologicalfactor(s), and local excesses of soils, exposure and wetness must certainly also restrict distribution.
Phenology and growth
The shoot emerges from below ground in March, pushing up through the leaf litter crozier-like, with threefolded leaf-like bracts surrounding and protecting the solitary flower bud. The shoot soon straightens,the palmately cut bracts unfurl and expand, and the flower stalk elongates carrying the flower wellabove the ring of three involucral bracts. The flower bud then loses its green tinge and, since theperianth in this species consists of just one set of leaf-like segments, the tepals or petal-like sepalsexpand (plant anatomists tell us they are not true petals), and the first anemone flower opens 'forbusiness'. This usually occurs around the end of March or the beginning of April depending on season,habitat and geographical location (Shirreffs 1985). The true leaves are very similar in appearance tothe bracts, but they are produced a short distance further along the rhizome from the flower-stalk andthey do not appear until after the flowers have opened (Step & Blakelock 1963).
Growth rates in A. nemorosa are extremely low: seedlings take at least five years to form a viablerhizome, plus perhaps another five to ten years for the plant to become capable of flowering. Theaverage annual extension in adult plant rhizomes is only 2.5 cm, so vegetative spread is also extremelyslow (Ernst 1983; Shirreffs 1985). By June, the sexual reproductive cycle has been completed, and in theshaded floor vegetation the plant rapidly dies down and disappears completely below ground by around themiddle to end of July.
Flower variation and pollination
All parts of the anemone plant are very variable (ie phenotypically plastic), and this is particularlytrue of the flower. Flower stems vary in height between 10-30 cm above ground (with a mean of 14 cm),each bearing a solitary blossom 10-40 mm in diameter. The perianth is composed of from 4-11 ellipticsepals (most frequently six or seven). The sepals are usually white inside, purple tinged or streakedoutside, but purple, blue and pink forms also occur, the former sometimes quite frequently (Shirreffs1985). Stamens are numerous, usually about 45 in three ranks of differing filament length, and as WoodAnemone flowers offer no nectar, unspecialised insects visit them to collect the openly presentedprotein-rich pollen as food for themselves and their brood. The insects attracted by the perianth'sappearance, movement and food reward range from honey-, bumble- and solitary-bees, to beetles, flies,thrips and bugs (Proctor & Yeo 1973).
When the flower first opens the maturing stamens are crowded over the stigmas and prevent them from beingpollinated, though pollen is already being shed from the ripe, outer anthers at this stage. After abouta week, the rest of stamens ripen and diverge, and during the second week the white, translucent stigmascan be pollinated. Self-fertilisation is prevented by an incompatibility mechanism, makingcross-pollination obligatory (Proctor & Yeo 1973). The flowers are held erect during the day andmove in the slightest breeze, but they droop and fold at night, or in dull or wet weather. The stigmasshrivel and blacken after pollination and a crowded head of single-seeded achene fruits then develops.
Reproductive biology
The number of carpels in the flower varies from 9-42, with a mean of 22 (Salisbury 1942). Shirreffs(1985) found the mean number of carpels ranged from 16 to 31 at different sites, the lower figures beingassociated with non-woodland sites, such as open grassland. Flowering density is greatest in woodlandssince this is where the species forms dominant carpets. The mean number of fruiting flowers in a typicalA. nemorosa carpet is around 152/m² (Salisbury 1942), with higher figures in coppice (380flowers/m²), but much lower figures than this are found in densely shaded areas.
The cluster of achenes breaks up and the individual fruits are shed from May to June, depending on thelocal climatic and micro-climatic conditions. When shed, the achenes of A. nemorosa contain animmature embryo that requires a moist, cold, after-ripening period lasting from 4 to 6 months beforethey ripen sufficiently and become capable of germination (Vegis 1961). Germination occurs in thefollowing spring, but the typical rates that occur in the wild are poor. Ernst (1983), however, foundsomewhat improved figures of between 5% and 35% germination occurred after long, cold winters inGermany, with lesser figures being obtained after mild winters. This indicates that seedling recruitmentinto the existing mature population is irregular. Ernst concluded that recruitment of seedlings andyoung plants (ie 2nd to 5th year classes) in the study area was inadequate to maintain a viablelong-term population. The mortality of the young plants was very high in the first and second years(88.2% ± 13.5%), which is comparable to that shown by other, bulbous vernal species, namely Alliumursinum (Ramsons) (Ernst 1979) and Narcissus pseudonarcissus (Daffodil) (Barkham 1980).
In his study, Ernst (1983) calculated that A. nemorosa generally does not invest more than 5% ofits biomass resources into sexual reproduction, while most of the fixed energy (production) is used tomaintain the rhizome at 40 to 50 % of the total plant biomass. The annual increase of the rhizome can beas much as 150 mg. Rhizomes older than 15 to 25 years are brittle and readily separate from the parentplant. This fragmentation constitutes a rather unspecialised form of vegetative reproduction and, in thesites studied, it appears to be the main mechanism maintaining the population.
Population persistence
Wood Anemone showed a remarkable degree of persistence in neglected coppice uncut for 30 to 40 years in EEngland, being present in 70% of plots in five such woods. This figure was surpassed only by Rubusfruticosa (Bramble) which had a 100% occurrence (Brown & Oosterhuis 1981). It is interestingthat during the same study, germination tests carried on for two years with soil samples taken from theupper 15 cm of the profile (after litter removal), found no seedlings of A. nemorosa present, norindeed any seedlings of Hyacinthoides non-scripta (Bluebell) or Mercurialis perennis(Dog's Mercury), although all three of these often dominant carpet-forming woodland species had survivedin at least 50% of the neglected, overgrown woodland coppice investigated. This agrees with the generalfinding that species of shaded habitats tend to lack mechanisms for widespread and rapid seed dispersal(Webb 1966; Brown & Oosterhuis 1981): the seeds of such species are heavy, seed production isrelatively low (Salisbury 1942) and they are in the main dispersed by rainwash, in clinging mud, or byants (Ridley 1930).
Possible dispersal mechanisms
Anemone nemorosa has no specialised means of achene dispersal, although it has been suggested thatants may be involved (Ridley 1930; Oberdorfer 1970). The achene has no attached food body, so that antsand other animals are unlikely to show any interest in them. However, if ant dispersal (ie myrmechory)really is the sole, or even the principal method of seed dispersal, then the efficiency and efficacy ofthe process must be rather severely limited since Ernst (1983) found the distance between parent plantsand established seedlings was never more than 13 cm. Brown & Oosterhuis (1981) observed that even arelatively narrow strip of non-woodland habitat around 50-100 m wide, would create an ecological hurdlethat most plant species of shaded habitats could hardly ever cross. The apparent lack of an effectiveseed dispersal mechanism in A. nemorosa, together with the exceedingly slow diffusive spread ofthe rather long-lived, creeping rhizome, results in the observed single species carpet of interminglingclones of the plant which we see mainly in the relative stability of woodland or undisturbed scrubvegetation.
Competing woodland species
The carpet growth of A. nemorosa enables it to shade out many smaller competing species, allowingit to become dominant at sites which are wet in the spring. However, it is usually unable to competewith taller growing vernal species, including for instance Hyacinthoides non-scripta (Bluebell),Allium ursinum (Ramsons) and, in Britain, but not in Ireland, Mercurialis perennis (Dog'sMercury). In these instances, A. nemorosa is either completely ousted by the shade of the tallerplants and their competition for other limited environmental resources, or it survives in smallernumbers only as a subsidiary companion species (Shirreffs 1985). However, where there is a considerabledegree of woodland disturbance (eg grazing, trampling or coppicing), A. nemorosa is often betterable to withstand these external pressures than can Bluebells, Ramsons or Dog's Mercury, especiallywhere such disturbance is combined with seasonally wet soils (Grime et al. 1988).
Toxins
A. nemorosa has some degree of protection from grazing animals since it contains the volatile,oily, irritant substance, protoanemonin, the concentration of which reaches its peak when the plant isflowering and most conspicuous. The toxin has an acrid taste and causes burning in the mouth and throat,effectively deterring animals from eating much of it (Cooper & Johnston 1998). This said, a study inWarwickshire woods made 50 years ago, when rabbit populations were very much more active than now, founda number of widespread woodland herbs, including A. nemorosa, suffered heavy grazing pressurenear warrens, sometimes almost to vanishing point (Knight 1964). Several fungi, both Ascomycete andBasidiomycete, attack A. nemorosa leaves and rhizomes and can suppress flowering partially orcompletely (Ernst 1983; Shirreffs 1985).
Fossil record
Fossil pollen of Anemone-type has been found in Scotland from the late-Glacial period(13,000-10,000 BP), but it is not specifically that of Anemone nemorosa, but could also come fromthe related species Actaea spicata (Baneberry) or Pulsatilla vulgaris (Pasqueflower)(Shirreffs 1985).
World occurrence
Beyond the British Isles, A. nemorosa occurs throughout the suboceanic northern temperate zone ofboth Europe and W Asia and reaches 67° N, just within the Arctic Circle in Norway (Shirreffs 1985;Hultén & Fries 1986, Map 827; Jalas & Suominen 1989, Map 1630; Jonsell et al. 2001).Forms of A. nemorosa are widely grown in gardens within and beyond the natural range of thespecies. Griffiths (1994) lists 19 garden cultivars of the species and Jonsell et al. (2001)mention an additional yellow form cultivated in Sweden.
Herbal use
A. nemorosa has entirely fallen out of use in herbal medicine today, although the older herbalistssuch as Gerard and Culpepper listed numerous ailments it was supposed to alleviate, eg headache,rheumatic gout, lethargy and for cleansing ulcers (Grieve 1931). As the plant is decidedly poisonous,the modern advice is to BEWARE of any such remedies.
Names
The name 'Anemone' is often said to be derived from Greek 'anemos', wind, plus the feminine patronymicsuffix, making it 'daughter of the wind' (Gilbert-Carter 1964). The connection with the wind is somewhatobscure in this particular species, however, although the flower does dangle and flutter in the breezeif it is strong enough, so perhaps the alternative explanation may fit better. This suggests the name isa corrupted Greek loan word of Semitic origin, referring to the lament for slain Adonis, or Naaman,whose shed blood produced the blood-red flowers of Anemone coronaria (Crown Anemone), orAdonis annua (Pheasant's Eye), both common spring species in the Mediterranean region(Gilbert-Carter 1964; Stearn 1992). The Latin specific epithet 'nemorosa' means, 'growing in woods' or'in shady groves' (Gledhill 1985).
There are dozens of English common names listed in Grigson (1987), the two most frequently used being'Nemony' or 'Neminies', a simple contraction of Anemones, and 'Wind-flower'. Both of these names areborrowed from Anemone coronaria, famous in Greek legend as mentioned above. Other names include'Wood Crowfoot', 'Moonflower', 'Cowslip' (the latter rather odd), and two names that refer to an odour,'Smell Foxes' and 'Smell Smock', both of which might be derived from the sharp, unpleasant taste and thefaint smell of A. nemorosa (Grigson 1987).
Threats
Anemone nemerosa regenerates mainly by rhizome growth and while large clones do develop and genetscan persist for a long number of years, seed persistence is low, dispersal is very poor and seedlingestablishment is extremely slow. Thus the plant is a poor colonist of new sites and Grime et al.(1988) believe it is decreasing in England in grassland habitats at least, and perhaps also in somewoodlands. Certainly it is not equipped for jump-dispersal and the colonisation of new habitats, so itmay well be a relict in many of its existing sites.
Introduction, neophyte, naturalised garden escape, rare. European temperate, widely naturalised beyondits native range.
1951; MCM & D; railway crossing at Aghalurcher Old Church.
Throughout the year.
Growth form and preferred habitats
This vigorous deciduous perennial with its climbing, scrambling and trailing stems belongs to the onlygenus in the family Ranunculaceae that contains woody members. Typically it clambers over shrubs andtrees, or clings on walls or rocks, holding onto its support by the twining stalks (petioles) of itscompound, once-pinnate, opposite leaves that act like tendrils. The grip of the petiole-tendrils is verytight, and as they age they harden and become wire-like, so that C. vitalba sometimes stranglesand kills the stems of the plants supporting it (Melderis & Bangerter 1955; Step & Blakelock1963a).
Being deciduous, the leaves of C. vitalba drop off in the late autumn, although their twistedtendril-like petioles persist for a while after the leaf blades have disappeared. During the winter, theplant relies for its entire support on the entanglement of its woody stems with those of the tree orshrub on which it is climbing and despite winter storms this always seems to suffice (Fitter 1987).
Soil preferences
The species performs best on base-rich or calcareous soils, of which at least in certain parts of Britainand Ireland, it is a useful and reliable indicator species (Lousley 1969, p. 15). In central Europe,however, Ellenberg (1974) found that C. vitalba grows on a wide range of soils from weakly acidto weakly basic. However, to really thrive it requires a soil with moderate to high fertility and mediumto good drainage. In another English study, low calcium levels in soil appeared to retard the growth ofthe species (Buxton 1985).
A detailed experimental investigation of the species soil nutritional requirements in New Zealand foundthat growth of C. vitalba increased with increasing levels of lime; this was especially so whenthis was accompanied with increasing rates of applied phosphate. Maximum growth occurred at pH 4.7,while plants were killed by a pH as low as 3.7, presumably due to the toxicity of available aluminium atthis acidity (Hume et al. 1995). This study also showed that the plant's response was to high pHand/or low aluminium concentrations, rather than to high concentrations of calcium, indicating that atleast in New Zealand, C. vitalba is not a true calcicole species (ie lime-loving) (Hume etal. 1995).
While showing a definite preference for calcareous or base-rich soils, C. vitalba is notcompletely confined to them, but may also occur on other dry, stony sites and on disturbed, enrichedsoils, including on waste ground and in rock quarries (Sinker et al. 1985).
Origin of most Clematis plants
The garden source of the C. vitalba plants found in the wild in Britain and Ireland is often notimmediately obvious. The genus Clematis is a very popular, indeed at present a very fashionablehorticultural subject, with many very beautiful species and legions of novel varieties widely availablein the nursery trade. However, C. vitalba itself is much too rampant and weedy a plant to begrown by many gardeners on its own account. The answer to this apparent garden usage, yet lack ofdecorative worthiness, has to do with the horticultural production of rapidly flowering newClematis varieties and especially of sterile hybrids such as the familiar C. × jackmanii. Hybrids and other varieties can be multiplied byinternode cuttings, but a flowering plant of the desired hybrid or variety is more quickly and morecertainly obtained by grafting the variety on to a seedling rootstock of C. vitalba, oralternatively on another cultivated form, C. viticella (Purple Clematis).
If the grafted individual is planted with the region of the tissue union below the soil surface, after afew years the grafted variety will have developed its own root system (Salisbury 1935, p. 151). On theother hand, if an inexperienced gardener plants the graft with the union exposed, there is nothing toprevent the C. vitalba rootstock developing its own competing stems. In addition, if the graftedplant is incorrectly pruned the scion may be damaged, killed or entirely removed, again allowing theC. vitalba rootstock to take over. Such plants invariably prove too dominant or rampant in thegarden setting and they eventually end up being discarded on refuse tips, manure heaps, or on wasteground, where they may survive and continue to grow, reproduce and release their wind-borne seeds intosurrounding wild habitats.
Fermanagh occurrence
C. vitalba is either deliberately planted, or much more probably, a naturalised wind-dispersedgarden escape in Fermanagh, growing in just a few old, rather neglected hedgerows and thickets. Thefirst record of this climber in Fermanagh was made as late as 1951, incidentally providing a very goodexample of the lack of previous recording of alien and introduced species in the county. It wasdiscovered in what was then a very typical habitat of the species − by a railway crossing. C.vitalba is very much associated with railways throughout the British Isles (Hackney etal. 1992). Previously, when the Fermanagh railway was in operation, C. vitalba grewalongside the permanent way, and having plumed achenes for fruit, it is very easy to imagine it readilyspreading along open areas on embankments, cuttings and road crossings. After the closure of the railwayin 1957, although it could equally disperse along roadsides, the species had fewer opportunities forcolonisation and it has almost certainly dwindled to arrive at the current level of rarity. We cannotknow this for certain, the plant having only ever been recorded in seven Fermanagh tetrads, six of themwith post-1975 records. Occasionally, however, as at Gubbaroe on the limestone shore of Lower LoughErne, when it grows in full sun, colonies of C. vitalba become so vigorous the plant can coverand dominate very large sections of hedgerow and scrub thicket. In such situations, it may totallyobscure and overwhelm young trees and shrubs and even threaten older, established plants.
The other Fermanagh record details are: Carrickreagh Bay, Lower Lough Erne, 1983, RHN; Scottsboroughlakelet, 28 August 1988, RHN & RSF; Cloughmore, 2 km SE of Rosslea, 28 August 1988, RHN & RSF;ride in conifer plantation, Gubbaroe Point, Lower Lough Erne, 1 January 1990, RHN; Cleenishgarve Island,Lower Lough Erne, 17 June 1990, RHN; Killadeas Td, Lower Lough Erne, July 1993, I. McNeill; near oldhouse, E of Gubbaroe Point, Lower Lough Erne, 18 April 1998, RHN; near old house, Mullaghfad Td, E of
Brookeborough, 21 September 1998, I. McNeill; Gublusk Bay, Lower Lough Erne, 8 August 1999, RHN; samesite, 12 October 2002, I. McNeill; fence on Irvinestown Road, Enniskillen, 19 September 2010, RHN &HJN; Carrickreagh Bay, Lower Lough Erne, 29 September 2010, RHN.
Flowering reproduction
C. vitalba flowers rather late in the year from July to September and the branched clusters ofcreamy or greenish-white, star-like 2 cm diameter flowers, like those of Caltha andAnemone, are petal-less. The usually four, but occasionally five or six sepals, which are hairyon their outer surface, again take on the role of the missing flower whorl (Webb et al. 1996).The Clematis flower has a faint but pleasant vanilla fragrance (Genders 1971), but it does notproduce any nectar for pollinating visitors. The flowers are visited by flies and by bees that collectthe abundant protein-rich pollen as a food reward and carry out cross-pollination. Wind-pollination mayalso be involved, but the extent of this in unknown.
Fruiting and dispersal
The characteristic fruits are produced in the autumn in large numbers, and usually sufficient silveryachenes are retained on the receptacle to keep the plant conspicuous in the hedgerow throughout thewinter months. It has been estimated that around 17,000 seeds (achenes) are produced for every 0.5m2 of C. vitalba canopy and they are dispersed by wind, water, people and othervertebrates (Cronk & Fuller 2001, p. 70). In linear habitats, eg along roadsides and beside railwaylines, the dispersal of the many achenes of C. vitalba with their firmly attached white orsilvery grey, long-plumed, feathery styles is obviously enormously facilitated by the sucking linearslipstream of swiftly moving traffic.
Germination and seed survival
Study of C. vitalba germination ecology in New Zealand has shown that achenes (ie single seededdry fruits), retained on the vine over the winter, have a high degree of dormancy and viability, and thesporadic release of the seed from the parent plant effectively acts as a form of aerial seed bank(Bungard et al. 1997). The published survey of soil seed banks in NW Europe contains very littleinformation on C. vitalba, but one report did suggest that a short-term persistent buried soilseed bank may exist (ie seed surviving between one and five years) (Thompson et al. 1997). Inaddition to a minimum light requirement (equivalent to 5% of full sun), the achenes require a period ofchilling to break dormancy, conditions that effectively time germination to the spring following theirproduction.
Behaviour as a serious weed in New Zealand
Although over most of its European range C. vitalba is usually an innocuous climber, the speciescan become a serious weed in young forestry plantations causing losses due to overgrowth of saplings. InNew Zealand, it is a particularly vigorous and harmful invasive alien in forestry plantations and alsoin conserved remnants of native podocarp forest and it is responsible for losses of both foreststructure and indigenous species biodiversity (Ogle et al. 2000; Hill et al. 2001). C.vitalba was first recorded as a weed in New Zealand in 1940 (Webb et al. 1988), althoughit was known much earlier in gardens and as a local garden plant escapee. It is thought to have arrivedas a garden plant from Europe, and the first herbarium specimen of a wild plant of the species wascollected in 1936 (West 1992). C. vitalba now occurs as an adventive species almost throughoutthe lowlands of New Zealand, except for regions north of latitude 37˚S (Webb et al. 1988; West1992). It is probably the most publicised environmental weed in New Zealand, and community groups,government departments, local authorities, schools and paid contractors have tackled infestations overlarge and small areas, either mechanically or chemically (Timmins 1995). It has come to public noticemostly because it invades and smothers indigenous forest. In 1998 it was the subject of 37% of thecomplaints about plant pests made to the Regional Council which oversees the indigenous forest areaaround Taihape in central North Island, New Zealand, which was more than any other species, includingagricultural weeds (Rowatt 1998). The seriousness of its weed status is illustrated by the fact thatresearch is underway to identify an insect suitable for biological control, and the introduction of theEuropean leaf-miner Phytomyza vitalbae Kaltenbach which attacks the species, is being activelyconsidered (Hill et al. 2001).
In the warm temperate and moist to wet conditions prevailing in New Zealand, C. vitalba can regrowfrom fragments after cutting and this is recognised as important in its invasive spread in this part ofthe world. Regeneration of fragments is related to age, since older stem sections have better waterretention and larger nutrient resources available than softer, young tissues (Kennedy 1984). Under theprevailing growing conditions the species has a high growth rate, with young plants and new shootsextending up to 2 m per year. If given full sun, plants also reach reproductive maturity early in life,producing seed when one to three years old and reproducing vegetatively after just one year's growth(Cronk & Fuller 2001, p. 71).
Irish occurrence
C. vitalba is an established alien in Ireland, where although widely scattered throughout, it isvery much more frequent south of a line between Limerick and Dundalk.
Fossil record and species status in Britain
Despite an extremely skimpy fossil record in Britain from the Atlantic Period onwards (7,500-5,200BP)(Godwin 1975, p. 119; Rackham 1980, p. 108), C. vitalba is at least traditionally regarded asnative in most of the area of England and Wales south of a line between Anglesey and The Wash. North ofthis juncture it is considered an alien introduction and its presence diminishes rapidly towards SScotland (Preston et al. 2002). It is difficult to envisage exactly what criteria determine thedistinction between native and alien in these circ*mstances (Webb 1985). It is sensible to take acautious approach when attempting to interpret the published map (Preston et al. 2002) and,indeed, for any species great care is needed when distinguishing native from alien status for regions onthe same land mass.
European and world occurrence
Beyond Britain and Ireland, C. vitalba is considered native in S, W and C Europe and has alienstatus in a narrow zone north of its native range in Holland, Denmark and Germany (Jalas & Suominen1989, Map 1679). It is also regarded as native in N Africa, W Turkey, the Lebannon, the Caucasus, N Iranand Afghanistan (Griffiths 1994; Jonsell et al. 2001). As already mentioned, it is a veryinvasive naturalised introduction in New Zealand and is also naturalised in parts of both S Australiaand N America (Cronk & Fuller 2001).
Knowing the extent of the major weed problem this alien climber has created in New Zealand over the past50 years and the threat it poses to the survival of remnants of indigenous vegetation and rare specieson those islands, it is rather surprising that at least one horticultural supplier is still offeringC. vitalba for sale to gardeners in N America, although it does warn of its vigour and potentialspread (http://www.botanical-journeys-plant-guides.com/clematis-vitalba.html,accessed 19 January 2016). As Cronk & Fuller (2001) point out, "It is important that theproblems associated with it are made known so that future introductions to potentially 'invasible' [ahorrid, invented word] areas are prevented."
Toxins
Like other members of the family, C. vitalba contains an appreciable amount of the poisonoussubstance, protoanemonin and, although animals rarely browse the plant because of the acrid taste andirritant effect on the mouth, it is known to have killed cattle. Contact with the plant sap can alsoblister the skin (Cooper & Johnson 1998).
Names and uses
'Clematis' is derived from the Greek 'klema', meaning 'a vine branch', alluding to the vine-like twiningclimbing habit of the plant. The Latin specific epithet 'vitalba' translates as 'the white vine' andrefers to the wild species (Johnson & Smith 1946).
There are a large number of English common names in existence: Grigson (1955, 1987) lists 36.'Travellers’ Joy' was a name coined by John Gerard for his English herbal of 1597, presumably because heknew that the plant grows on waysides and hedges. The most widespread English common name is 'Old Man'sBeard', a reference to the silvery-white twist of long, feathery styles that adorn the fruit achenes. Itshould be remembered that very often in folklore the 'old man' referred to is the Devil. Similarallusions include 'Bushy Beard', 'Daddy's Whiskers', 'Grandfather's Whiskers' and 'Father Time'.
Other names refer to the rope-like nature of the older climbing stems, eg 'Bullbine', 'Hag-rope' and'Devil's Guts'. The woody branches with their characteristic flaking stringy bark have been used to makelightweight baskets in the past (Hutchinson 1972). In Devon, Clematis stem was woven to make thebottoms of pots for catching crabs (Vickery 1995, p. 375).
Another group of names indicates the use of stems as a cheap tobacco substitute, eg 'Boy's Bacca','Gipsy's Bacca', 'Tom's Bacca', 'Smoking Cane' and 'Poor Man's Friend'. According to Grigson, the youngand the poor used to smoke cigar lengths of the dry stem, "as they draw well and do not burst intoflame". Personally, I cannot imagine the desperation and the taste! However, there is evidence thatthis type of cigarettes was also smoked elsewhere, as there are equivalent names for the plant, eg'Rauchholz' ('smoke wood') in German, 'Smookhout' in Dutch, and 'Fumailles' and 'Bois à fumer' in French(Grigson 1955, 1987). If anyone is tempted to try this, beware; unless the stem is dead and completelydry the irritant toxin is present and can cause ulceration of the lips and mouth (Mabey 1977, p. 159).
There are also a number of other peculiar, unexplained ideas contained in further quite widespreadalternative English names, for instance, 'Honesty', and some connection with the virtuous Virgin Mary,for example, 'Lady's Bower' and 'Maiden's Hair' (Grigson 1987). Prior (1879) reckoned the name,'Virgin's Bower', which was also used by Gerard (1597), alluded not to the Virgin Mary, but rather tothe virgin Queen Elizabeth. In names, as in other matters, truly it is often easier to pose a questionthan to answer it!
In herbal and homeopathic medicine, several European species of Clematis, being diuretic anddiaphoretic, have been used in treating ulcerous diseases such as syphilis, gonorrhoea, cancer and otherinflammatory conditions including those of the eye (Grieve 1931). According to this herbal source theroots and stems of C. vitalba bruised and boiled for a few minutes in water and then digested fora while in sweet oil, made a cure for itch. There is folklore indicating that a C. vitalba stemtwisted into a ring and worn round the neck was used in herbal medicine to cure convulsions in children(Vickery 1995, p. 375).
Considering the poisonous nature of the species, an interesting usage of it as food has been recentlypublished. In a remote and isolated valley in NW Tuscany called 'Garfa*gnana', traditional gathering anduse of a wide range of herbaceous plants has survived, so that for example, the inhabitants make avegetable broth containing at least 20, but often more than 40 wild plants (Pieroni 1999). The souprecipe does not contain C. vitalba, but it is the main ingredient in a very popular vegetableomelette. Young shoots are boiled before they are incorporated with eggs, and sometimes with cheese, andthe mixture fried so that the protoanemonine is inactivated (Pieroni 1999).
Threats
There are no conservation problems locally, but rampant C. vitalba can smother other plants.
Native, very common and widespread, locally abundant. Eurasian, but widely naturalised in N America, sonow circumpolar wide-boreal.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form
This variable, wintergreen perennial is a widespread and abundant medium-tall herb (up to 50 cm), ofmoist to seasonally wet (but not waterlogged), pastures, mown meadows and roadside verge grasslands. Itis also present in unmanaged grasslands, eg in woodland clearings and on all forms of rock outcrop.Although it possesses a small wintergreen leaf rosette the species produces very little growth untilFebruary or March and for its energy requirements the overwintering plant relies on starch stored in itsshort, stout rootstock (Harper 1957). Coles (1971) refers to the compact rootstock as 'premorse', asopposed to a longer, more spreading, rhizomatous one. The term is derived from the Latin 'praemorsus',meaning 'bitten off', which is a rather appropriate description of the rootstock (Holmes 1979).
Possible native habitats
As with Ranunculus repens (Creeping Buttercup), while fossil evidence proves R. acris isundoubtedly a native species in Britain and Ireland (see below), it is very definitely a 'follower ofman'. Nowadays, more often than not it occupies habitats managed, opened up, or created – intentionallyor not – by human activity (Harper 1957). It is not really possible to be certain what the naturalhabitats of R. acris were in these isles prior to human arrival, although Harper (1958) hassuggested that it most likely frequented a variety of damp ground communities, ranging from marshes andCarex elata (Tufted-sedge) dominated fens, to mountain grasslands above the climatic forestlimit.
Variation
R. acris is a polymorphic species that in Europe can be subdivided into four subspecies (seeFlora Europaea 1, Tutin et al. 1993, p. 274), two of which occur in in the BritishIsles, the common subsp. acris and the much rarer northern subsp. pumilus (Wahlenb.) A.& D. Löve, that only appears in Scotland. R. acris can also be subdivided into threevarieties, a larger, very widespread var. acris, a considerably smaller (up to 20 cm tall) var.pumilus Wahlenb. that is restricted to the Scottish Cairngorm mountains, and a hairier var.villosus (Drabble) S.M. Coles, which is common in undisturbed areas of N Scotland (including theisles) and W and C Ireland (Stace 1997, p. 88). Var. villosus was not recorded in Co Fermanaghand certainly is under-recorded across its range.
Soil moisture preferences
The soil moisture preferences of R. acris (although it is not strictly confined to them)are intermediate between those of two other, very common, closely related buttercups, R. bulbosus(Bulbous Buttercup) on drier forms of rocky ground or on shallow soils and R. repens in wetterhollows, or poorly drained soils. Certainly, R. acris is always absent from areas which sufferserious midsummer drought, but thanks to its geographical west Atlantic situation, prolonged droughtvery rarely, if ever, occurs in Fermanagh (Harper & Sagar 1953; Harper 1957). Meadow Buttercup istaller than these other two mentioned buttercups and thus is better able to compete and survive in themore closed, tightly knit turf of low to moderately fertile, fairly productive, herb-rich water meadowsthat are such a significant conservation feature of lakeland Fermanagh. These damp, waterside meadowsthat flood regularly or from time-to-time, harbour some rather aggressive competitive plant species,including the grasses Lolium perenne (Perennial Rye-grass) and Agrostis stolonifera(Creeping Bent) and two even more locally common and dominant herbs than Meadow Buttercup, Juncuseffusus (Soft-rush) and Filipendula ulmaria (Meadowsweet).
Fermanagh occurrence
Throughout Fermanagh, R. acris is common everywhere except at the highest, most exposed ground, orin permanently wet, or very acidic habitats. Despite these substrate limitations it is still present in487 tetrads, 92.2% of those in the VC, making it one of our most widespread species. In terms of tetradnumbers, it ranks fourth equal with Angelica sylvestris (Wild Angelica) and it is the eighth mostfrequently recorded species in the Fermanagh Flora Database.
Even at this high level of frequency of records, it is possible that our statistics might understate thepresence of this species. One of the acknowledged errors in any botanical field survey is the likelihoodthat very common and familiar plants tend be overlooked, simply because they have become so unremarkablein the recorder's mind, and regularly it will be assumed that they have already been entered on anotherfield recording card and consequently they are overlooked or ignored!
Land management and plant species competition
The farmland grasslands where Meadow Buttercup populations persist to the greatest extent are regularlygrazed or mown. This form of ecological pressure is important in maintaining the herb's population size,since without regular disturbance opening up the vegetation and offering periodic temporary release fromcompetitive stress, the species would survive only at much lower frequency. If the grazing or cuttingregime becomes relaxed, R. acris cannot thrive and maintain its abundance in the longer termamong more vigorous, gregarious and aggressive plant species.
Writing prior to the latest (and greatest) intensification of farming in the last half-century, Harper's(1957) research indicated that the abundance of R. acris in pastures and meadows provides anindex of the age of the grassland and that its frequency increased with overgrazing or with regularcropping for hay. Meadow Buttercup has an acrid taste that makes herbivores avoid it and thus it tendsto spread and increase in heavily grazed communities (see below for details of its poisonousproperties).
The widespread introduction and development of intensively managed 'improved' rye-grass based sowngrasslands, replacing old permanent meadows and pastures, together with regular or frequent applicationof chemical fertilizer or slurry manure, has led to a considerable decline in the abundance of R.acris and, indeed, of most other lowland grassland herbs throughout the British Isles. Theseforms of government subsidized agricultural 'progress' are aimed at greatly increasing farm productivityand together with other changes in farmland management, eg changing over from arable to grasslandfarming, improved land drainage operations, swapping from sheep to cattle husbandry and from hay tosilage-making, are all practices which have resulted in depletion of plant species diversity on managedland.
Fermanagh has suffered the accumulated effects of these changes along with almost everywhere else inthese islands from the second half of the 20th century onwards. The major transformations have been inthe more intensively managed productive farmland, which in Fermanagh is most frequently situated in theeastern lowlands. However, where farming has become intensified, drainage generally leads to increasedpollution of both surrounding semi-natural ground and adjacent water bodies. This results frominevitable agricultural chemical runoff, both nutrient and toxic. As a result, permanent 'unimproved'pasture has become restricted to less productive ground, either on less accessible steep or rockyslopes, very shallow soils, or in more upland areas featuring small pockets of grazing that areinaccessible to tractors and spraying machinery. Thus, apart from on lightly or occasionally managedlowland wayside grasslands, R. acris is now most frequent and abundant on pastures at higheraltitudes, except where such ground becomes excessively acidic (ie below pH 4.0) and nutrient leached,or where grassland gives way altogether to either ericaceous heath or bracken.
British and Irish occurrence
The New Atlas hectad distribution map shows R. acris having almost total cover acrossBritain and Ireland, the few absences being on exposed western coasts and in a few scattered squares inIrish and Scottish VCs which probably have no resident botanist. In these latter areas, recording evenof such a common species inevitably tends to be incomplete, despite the efforts by non-residents. TheRelative Change Index of R. acris, measured in Britain (but unfortunately not surveyed inIreland), between the recording periods of the two BSBI plant Atlases, is calculated as +0.30.The index is interpreted as indicating that R. acris is a stable species in the area studied(Preston et al. 2002).
Buttercup population studies
Detailed demographic studies of field populations of three common buttercup species, R. acris,R. bulbosus and R. repens, were made in the early 1970s at Bangor in N Wales by Dr J.Sarukhan, supervised by Prof. John Harper. Despite its age this pioneering study remains highlyrecommended reading since the authors thought deeply about plant population processes and used theirdata, and that of earlier workers, to deduce important generalisations regarding plant behaviour.
Plant ecology has few general concepts that scientists totally agree about, and absolutely no 'Laws'exist as they do in subjects like Physics. However, it is a useful and illuminating exercise to try andgenerate such general principles whenever possible. Harper and Sarukhan give us a clear picture of therecruitment rate of each buttercup species from seed and also from vegetatively produced daughterplantlets (ie ramets). This fecundity is then compared with the species population mortality and thusthe flux or rate of turnover can be calculated (Sarukhan & Harper 1973; Sarukhan 1974, 1976).
The concept of 'a plant population' is an extremely dynamic one, even more so than the concept of'vegetation', and in the case of the former it involves the gains and losses of individuals happening atthe same time. Thus a census approach is essential to population studies, marking and following the fateof individuals, including some that are actively growing and others that may be going through a dormantphase (ie buried viable seeds have also to be considered as part of the total plant species populationpicture).
Comparative patterns of population turnover in three buttercups
In his research studies, Sarukhan found that individuals of the three common buttercup species had quitedifferent levels of longevity. This reflected their methods of reproduction and the balance of strategybetween seed and vegetative ramet reproduction that each employed. Reproduction in R. bulbosus isexclusively by seed, that of R. repens is predominantly vegetative with some seed, and R.acris is mainly seed, but very occasionally it reproduces vegetatively by branching of itscharacteristic short rhizome.
During the two and a half year period of Sarukhan's study of the three buttercup species, the populationflux was greatest in R. acris. Indeed, he found that in some study plots there was no permanentpopulation, only a series of temporary but overlapping short-term cohorts, establishing rapidly fromseed and then soon dying. R. acris seed germinated in April and May, with a swift and highmortality peak in May and June, and mortality continued gradually at very low levels until the followingspring. A similar picture of high seedling production and mortality occurred in R. bulbosus,although germination in this species took place in the autumn rather than the spring. In both thesebuttercup species, the high initial seedling mortality is balanced by a long life span for thoseminority of individuals that manage to survive and become mature established plants.
The high risk of death at the early seedling stage probably reflects not only the very obvious innaterisks involved in establishing a viable root and shoot system, but additionally, and probably even moreso, the genetic load of unfit genotypes being carried by the species. The new gene combinations that areproduced by sexual reproduction are, by their nature, experimental and elimination of the plants withthe most unfit genotypes can be expected to occur early in life, as was the case with all threebuttercups (Sarukhan & Harper 1973; Sarukhan 1976).
In comparison with the other two buttercups, R. repens (Creeping Buttercup), the only one of thethree with appreciable vegetative multiplication, suffers little loss of vegetative individuals (ieramets) at the very beginning of their life, but it replaces a large number of them more frequently thanthe other two species do. The life expectancy of a R. repens ramet was short, ranging onlybetween 1.2 and 2.1 years. Very few plants of R. repens derived from seedlings lived for morethan 1.5 years, whereas mature plants of both the other buttercups could survive for much longerperiods; in the case of R. acris, individuals were shown to live up to eight or more years(Sarukhan & Harper 1973).
R. repens, with its dependable, high level of vegetative reproduction, comparatively weakflowering and seed production, but great seed longevity, contrasted strikingly with the other twobuttercup species which had little or no vegetative reproduction, high seed output, rapid germination,high seedling mortality and a rather short half-life of the buried dormant seed (ie just 5 months in thecase of R. acris and 8 months in R. bulbosus) (Sarukhan 1974).
Buried seed longevity
Other estimates of buried seed longevity are given in the detailed European survey of this topicpublished by Thompson et al. (1997). The survey lists a total of 40 estimates for R. acrisdivided as follows: transient (less than one year) = 19; short-term persistent (between 1 and 5 years) =10; long-term persistent (at least 5 years) = 3, and present in soil but unassigned to any of thesethree categories = 8. The equivalent figures for R. bulbosus seed are: 8, 4, 4 and 5 – a total of21 estimates. It is not unusual for seed longevity estimates to vary, but for both R. acris andR. bulbosus the predominant impression remains that buried seed is relatively short-lived.
Flowering reproduction
Flowering in R. acris stretches from May through to August, peaking in mid-June. Seeds are shedfrom July onwards. The female flower parts ripen first (ie the flower is protogynous) and a largevariety of short-tongued insects visit them to collect nectar, including Honey Bees (Apismellifera) (Harper 1957). Spatial fragrance patterns within the bowl-shaped flower guide theinsect visitor to the nectary, which is partially concealed by a flap at the base of each petal(Bergstrom et al. 1995). Self-incompatibility in the species is described as, "often verymarked, but not [occurring] in all populations" (James & Clapham 1935). Althoughcross-pollination is frequently achieved, some degree of apomixis (ie agamospermy − seed productionwithout any fertilisation or pollen involvement), does occur, perhaps in as many as 1% of flowers(Marsden-Jones & Turrill 1952).
Flowering in R. acris is so very variable in response to its environment that it has proveddifficult to measure its flowering capacity. Even Salisbury (1942, 1964), who was very keen to computesuch statistics, did not attempt to do so for this species. In his comparative study, Sarukhan (1974)was more daring than Salisbury and in his sample he found that the number of flowers per plant couldrange from 1-20. Sarukhan also found a high proportion of plants of both R. acris and R.bulbosus each produced between 40 and 140 seeds in total. The maximum seed number produced by anindividual plant in his populations was 281 for R acris and 287 for R. bulbosus.
Due to grazing and other forms of disturbance, Sarukhan's study showed that not all plants that floweredmanaged to set seed, but in R. acris about 40% of flowers did. Over a two year period, similarproportions of R. acris plants flowered in Sarukhan's study, but the ratio of seeds per floweringplant fluctuated wildly, from 26 in 1969, to a little over 1.0 in 1970 (Sarukhan 1974). Since Harper(1957) reported the number of seeds per flower varying between 0 and 40, and the number of flowers perplant was also very variable in his study, the variability helps explain Salisbury's reluctance tomeasure the reproductive capacity of these buttercup species.
Seed dispersal
As with other buttercups, R. acris has no specialized seed dispersal mechanism, but viable seedhas been found in the droppings of birds including the House sparrow (Collinge 1913). Furthermore, Dore& Raymond (1942) calculated from analysis of the seed content of farmyard manure, that a single cowmight disperse around 22,000 R. acris seed per ha during a 165 day grazing period. However, asSarukhan (1974) pointed out, while the seeds are 'dispersed' in this manner, they are also concentratedin 'local droppings'! Voles and Field mice are very probably the main rodent seed predators ofRanunculus species in grasslands in Britain and Ireland, but birds such as Pigeons and Pheasantsmust also consume and destroy huge quantities of buttercup seed.
Absence of Ranunculus acris hybrids: Hybrids betweenR. acris and other buttercup species have never been recorded anywhere in Britiain and Ireland,and while crosses with both R. repens and R. bulbosus have been reported from severalEuropean countries, Stace (1975, p. 124) regarded them as doubtful and requiring confirmation, and thenew Hybrid Flora of the British Isles (Stace et al. 2015) makes no mention of themwhatsoever.
Fossil history
Fossil seed (ie achenes) have been found in sediments of the four most recent interglacial periods (fromthe Comerian onwards) and also from the two most recent glacial stages. This evidence conclusivelyproves that although R. acris populations increase when human populations disturb natural orsemi-natural forms of vegetation, the species very definitely is native in these islands (Godwin 1975).
Toxins
Meadow Buttercup contains the acrid, irritant, poisonous principle protoanemonin, an unstable compoundderived from the glycoside ranunculin. The concentration of protoanemonin increases during growth of theplant and reaches a maximum during the flowering period. Being an unstable chemical, however, the dryinginvolved in hay-making readily converts protoanemonin into an inert, non-toxic substance called anemoninand thus dried fodder containing buttercups is perfectly safe to give to animals. Also, R. acriscontains much lower toxin concentrations than is found in Bulbous Buttercup (Cooper & Johnson 1998).Due to the presence of this acrid, bitter-tasting toxin, both of these buttercup species are unpalatablewhen fresh and they are avoided by grazing animals unless the beasts are actually starving (Harper1957).
R. acris has been known to poison both grazing cattle and sheep, but few cases have been reportedin recent years, probably because pastures nowadays contain much smaller proportions of these speciesthan previously was the case for the reasons discussed above (Cooper & Johnson 1998). In Norway in1988, however, five cows in late pregnancy were turned out to graze on a field of poor pasturecontaining abundant R. acris. They developed severe diarrhoea, a rapid pulse and noisyrespiration and all of them died or had to be destroyed because of their deteriorating condition(Heggstad 1989). Other experimental studies in Canada found that cattle gradually fed increasing amountsof R. acris in the flowering stage could cope very well with between 7 and 25 kg of the plant perday for two weeks towards the end of the experimental trial (Therrein et al. 1962; Hidiroglou& Knutti 1963, both references quoted in Cooper & Johnston 1998).
European and world occurrence
Beyond Britain and Ireland, R. acris s.l. (ie the polymorphic species we are considering here) iscommon over the whole of C and N Europe including the Faeroes and Iceland. There is some dispute as towhether or not it is native in Greenland, since material from there was indistinguishable from plantsknown to have been definitely introduced to Spitzbergen and N America (Coles 1971). R. acris hasa more limited distribution in S Europe; it is absent from Portugal and areas S of latitude40oN (Jalas & Suominen 1989, Map 1714). The coverage of the species in Italy wasunder-recorded by the latter map, since Pignatti (1997, 1, p. 306) maps a complete cover of thepeninsula. R. acris is also found in Morocco, and while it is very rare on Madeira, it is absentfrom all of the Canaries (Hultén 1971, Map 288; Press & Short 1994).
R. acris is very widely naturalised across the world and as a result has become circumpolar in theN Hemisphere (Hultén & Fries 1986, Map 844; Preston & Hill 1997). It is also introduced in a fewtemperate areas of the S Hemisphere, eg in S Africa and New Zealand (Hultén 1971; Hultén & Fries1986).
Names
'Ranunculus' is derived from the Latin 'rana' meaning 'a frog', an allusion to the fact that so manymembers of the plant genus live in or near water, the habitat of frogs (Johnson & Smith 1946). TheLatin specific epithet 'acris' means 'acrid', ie with a sharp, burning, peppery taste (Gilbert-Carter1964). This is very descriptive of all parts of the plant. Contact with skin can cause severe blistering(Grieve 1931; Cooper & Johnston 1997). Despite this, the caustic sap has been used in herbalmedicine to remove warts and the plant has also been used to treat headache, gout and even cancer(Grieve 1931). As ever, a health warning should here be attached to these comments and no recommendationwhatsoever is intended or implied by the inclusion here of this information.
The English common names include a recognition of the danger of handling the plant, eg 'Blister cup' and'Blister plant' are both listed by Grigson (1987). Britten & Holland (1886) provide a list of 38varied common names including 'Clovewort', 'Crowflower' and 'Bassinet'. The latter name means 'a smallbasin', apparently a reference to the bowl shape of the flower and, therefore, quite widely applied to awhole range of flowers, including all species of buttercup, Caltha palustris(Marsh-marigold) and many Geranium species. 'Crowflower', 'Crowfoot' and other versions of thisname, refer to the deeply cut leaf shape of many buttercup species and their relatives plus, again, somespecies of Geranium, eg G. pratense (Meadow Crane's-bill) (Grigson 1974). Other commonnames refer to the gold or yellow flower colour, but the strangest name of all appears to be 'Crazy'(see below in the species synopsis of Ranunculus repens) (Prior 1879).
Threats
None. R. acris is readily exterminated by modern systemic herbicides and the species has only anegligible, short-lived, soil seed bank. It has survived for thousands of years despite theselimitations.
Native, common, very widespread and locally abundant. Eurasian boreo-temperate, but almost cosmopolitanas a weed across both hemispheres.
1861; Smith, T.O.; vicinity of Ardunshin.
Throughout the year.
Growth form, ecology and Fermanagh occurrence
This vigorous, rosette-forming, wintergreen, short-lived perennial is very variable in form (especiallywith respect to leaf shape and degree of hairiness), and it can grow in almost any habitat provided itis damp (except very acidic peat bog), meaning virtually everywhere in Fermanagh! Creeping Buttercup isin fact the second most frequently recorded and widespread plant in the Fermanagh Flora database, beingfound in 515 tetrads, 97.5% of those in the VC. Creeping Buttercup thrives and, indeed, is most abundantin heavy mineral or clayey soil where drainage is naturally impeded (Harper 1957). In a survey of theSheffield area, Grime et al. (1988) found R. repens almost entirely absent from infertileacidic soils with a pH below 4.5 and from permanently flooded sites. Creeping Buttercup is also commonon wooded, open marsh or fen-fringed lakeshores, riversides and stream banks, on ditches and byroadsides, especially on wet, heavy soils. In these situations, its long creeping stoloniferous stems,rooting at their nodes and their deep, stout, tenacious roots emanating from a short erect rhizome, makethe plant extremely difficult to eradicate or control.
R. repens can regenerate from very small root fragments, plus its seeds show dormancy andsustained viability in the soil seed bank (see below for details) and it is also resistant to manyherbicides. These factors combine to make it particularly difficult to devise an effective weed controlstrategy for it in cultivated ground (Lovett-Doust et al. 1990).
In lakeshore grasslands, Creeping Buttercup occupies a zone between drier ground, where it is forced tocompete with R. acris (Meadow Buttercup), and wetter soil lower on the shore where Calthapalustris (Marsh-marigold) becomes dominant. R. repens is also a common weed of disturbedsoil and gravel and in these more open habitats it can tolerate very much drier conditions, rapidlyestablishing and spreading vegetatively, its numerous stolons quickly forming large clonal patches(Harper 1957). Salisbury (1964, p. 203) reported that, under favourable soil conditions, an individualplant could spread vegetatively over 4 m2 in a single year. However, the species isphenotypically very plastic, and particularly so with respect to stolon production. Stoloniferous growthclosely reflects both soil fertility and the intensity of plant competition.
Effect of trampling on populations
Creeping Buttercup can tolerate frequent disturbance and a considerable degree of soil compaction and asa result it is very common around field gates, along paths and on forest and woodland tracks andclearings. Damp, heavy soils frequently become 'poached' or puddled by the hooves of cattle or otherstock animals (especially around gates or feeding troughs) and grasses are often killed under theseconditions. R. repens is an efficient pioneer species colonising this type of disturbed, bareground. It invades rapidly through germination of re-exposed buried dormant seed and from nearby plantsby the extension of their stolons and the attached rooting plantlets.
The number of daughter plantlets (ie ramets) has been shown to increase with trampling of the vegetation(Diemer & Schmid 2001). The leaves of trampled plants spreading vegetatively in this manner areespecially large, which enables R. repens to cover and hold on to previously open patches in theturf or soil and prevent invasion by competing pioneer species (Harper 1957).
Since R. repens possesses an impressive ability to rapidly colonise disturbed ground, to a largeextent it has become a follower of man. The plant is a significant weed of gardens, waste disposalareas, building sites, dredgings of river banks, hedgerows, roadsides and, indeed, it occurs on any formof disturbed ground, including in the depressions made by animal hooves which tend to be damper thansurrounding ground through holding rain, dew, or water from other sources (Harper 1957).
Turloughs ('vanishing' lakes) − an unusual and specialised habitat
Other open habitats which R. repens occupies in Fermanagh include limestone pavement and scree. Insharp contrast to these latter relatively dry conditions, it also occurs on the shores of a special typeof limestone lakes called turloughs (ie so-called 'vanishing lakes' that drain vertically intounderground cave systems).
Recent studies in W Ireland have shown that turlough populations of R. repens differ from moretypical broad-leaved ruderal plants in both their leaf form (they have more highly-dissected and moreglabrous leaves) and in their physiology (the turlough plants have a higher rate of aerial and submergedphotosynthesis) (Lynn & Waldren 2001, 2002).
Life expectancy in comparative Buttercup population studies
During detailed population census studies of R. repens and two other buttercup species (see theR. acris species account above), the life expectancy of individuals of R. repens decreasedsignificantly with increasing density of the plant population (Sarukhan 1976). The study also clearlyshowed that the highest mortality rates per week were obtained, not in the unfavourable phases of thephysical environment (ie during the winter), but rather they coincided with active growth phases of theplant (Sarukhan & Harper 1973).
Flowering reproduction
R. repens flowers are quite variable in size and in the number of shiny yellow petals theypossess. They contain nectar and are pollinated by honeybees over a 4-9 day lifespan. The curve of thehoneybee's body closely mirrors and 'fits' that of the stamen cone of the buttercup to a remarkabledegree but while Percival (1955) noted bees actively collecting nectar from R. repens flowers, heremarked that pollen collection has very seldom been seen. Having said this, all buttercupflowers are primitive and unspecialised, meaning that their nectar and pollen flower foods are availableto all types of insect visitors. Thus they attract a great many different insect species and areprobably pollinated by many of them. The flowers are so unspecialised, they could possibly also beself-pollinated by raindrops (Van den Berg et al. 1985; Proctor et al. 1996; Jonsell etal. 2001). Having said this, cross-pollination is very much the norm, but a low level of selfingis also possible. There is no evidence of apomixis (ie the asexual formation of seed withoutfertilization taking place) (Coles 1977).
Fruiting performance
Under favourable conditions of slight or negligible competition, the average R. repens plantindividual produces five fruit heads although the number ranges from 0-38. However, the frequencydistribution is very heavily skewed, the most frequent class having just three fruiting heads per plant(25%). Each fruiting head contains a mean of 30 achenes, giving a mean total output of 150 ± 10 achenesper flowering plant. When subjected to marked competition, however, R. repens flowering becomessuppressed and, if the plant persists, as it does in some wet habitats, the reproductive balance is evenmore completely directed towards stolon development, with seed production then becoming meagre orcompletely absent (Salisbury 1942, p. 226).
Individual plants normally die off after they have successfully fruited, being replaced by a daughterplantlet produced vegetatively on a very short stolon, ie the plant is usually (but not always)monocarpic. This is another example of a biological concept (ie monocarpic versus polycarpicreproduction), which is not absolute, but rather, it is somewhat 'leaky' or facultative in its mode ofoperation (Forbes 2000, p. 187). The connecting stolons generally (but not always) die off in theautumn from September onwards, leaving the daughter rosettes produced at their nodes as independentplantlets (Van den Berg et al. 1985; Jonsell et al. 2001).
Population studies and plant plasticity and variation
R. repens is one of the best studied weedy plants from a population biology or demographicperspective, having been the subject of major studies by Prof. John Harper and several of hisco-workers, including amongst others, Lovett-Doust (1981). The latter showed that Creeping Buttercuppopulations studied in grassland and open woodland habitats follow similar seasonal patterns. Thepopulations examined remained remarkably stable from year-to-year, but the density of plants in thewoodland was significantly greater than that in the particular grassland examined. This suggests thatsome form of density related self-regulation of population size occurs at a figure referred to as 'thecarrying capacity'. The latter varies according to a number of environmental factors. The average timefor complete turnover of rosette populations was calculated as 2.17 years for woodland and notsignificantly different at 2.27 years for grassland rosettes (Lovett-Doust 1981). The creeping habit ofthe species is a response to the pressure of close grazing (or mowing) and if this is removed the plantwill grow more upright (Harper 1957).
The population dynamics of R. repens in pastures in N Wales were examined over a four year periodand computer modelled by Soane & Wilkinson (1979). These workers found little evidence of selectionamong families of clonal rosettes, or against new seedling recruits within populations. Theirmeasurements showed that the number of original genotypes present in a population of R. repensdeclines continuously at an approximately exponential rate. Local dominance by a few clones is thereforeto be expected unless new genotypes are recruited into the population, eg by seedling establishment.There was no evidence that selection was maintaining a diversity of genetic individuals (ie genets)within the R. repens population, but although recruitment of new seedlings was low enough to bedescribed by them as 'occasional', nevertheless it clearly plays a very significant role in the longerterm, through determining the number of genets represented in the population, and thus maintaininggenetic variation within these populations (Soane & Watkinson 1979).
Variation
Creeping Buttercup is continuously variable in so many of its characters that Coles (1977) found nojustification for the distinction of any infraspecific units within the species (ie forms, varieties orsubspecies).
Possible allelopathic effect on competing plants
There is a suggestion that R. repens may exert an allelopathic effect (ie a chemical inhibition)when competing with the roots of other plants, possibly involving phenolic compounds (Whitehead etal. 1982). Hatfield (1970) regarded R. repens as responsible for serious depletion ofpotassium and other elements from soil and he proposed that the roots secrete a toxin causingneighbouring plants to suffer from a nitrogen deficiency (Lovett-Doust et al. 1990). More work isrequired to clarify the real position on this topic, but as yet nobody has proven that any allelopathiceffect exists, although it might be more likely occur in soils rich in lime, or after lime has beenapplied (Whitehead et al. 1982).
In upland leached acidic soils and in other situations of low fertility, for instance in peat bogs and inwetter, marshy areas, R. repens is much less competitive than in drier, lowland situations andhere it tends to be replaced by R. flammula (Lesser Spearwort).
Colonising ability
The immediate opportunistic response of R. repens to disturbed environmental conditions permittingcolonisation of new territory tends to be increased stolon development, rather than seedling production.This is thought to be due to stolon production and flowering being largely coincident in May and June,plus the fact that vegetatively produced offspring often do not flower in their first year of growth.However, juvenile plants may do so if the habitat is very open and in these circ*mstances they generallyflower rather late in the season, up until about October (Harper 1957).
On the other hand, seed germination is greatest in late spring, with just a few seedlings appearing inautumn, and then only if there is a combination of high soil temperature and abundant moisture.Germination is very rapid (almost immediate) when seeds are exposed by soil disturbance. Seedlingsestablish readily where ground is open and particularly when the water table is high but the soil is notcompletely flooded.
Very rapid colonisation of bare ground may be achieved in the year of germination (Harper 1957).Salisbury (1942, p. 225) illustrates a case where a plant in open garden soil occupied more than 0.5m2 in its first growing season, producing 35 rooting nodes, of which 23 bore inflorescencesand twelve remained vegetative. Clearly, this is merely an isolated instance, and growth rate will bedependent upon habitat conditions, but it provides a helpful indication of the level of colonisingability the species is capable of achieving.
Further detail of R. repens population behaviour is present in my R. acris species accountunder the heading, 'Comparative patterns of population turnover in three buttercups'.
Seed dispersal
No specialised dispersal mechanism exists in R. repens, most of the dry, smooth seed simply beingdropped beneath the parent plant. However, cattle and horses are known to help disperse R. repensby transporting the seed in their gut, it having been found in their droppings. Various birds dolikewise, notably the House sparrow, but while Partridges, Pheasants and Pigeons are found frequentlywith a high crop content of R. repens seed, it is unlikely to pass through their guts in a viablecondition. Plants growing beside water may disperse seed in flowing streams and occasionally wholeplants will dislodge and migrate downstream in the adjacent flow. These processes may possibly beassisted by the disturbing activity of various ducks and water hens (Harper 1957; Salisbury 1964).
Other animals may also act as agents of dispersal, including those which transport propagules externallyon their surfaces. This form of dispersal agency includes man and his vehicles. Darlington (1969) foundthat washings of mud from motor tyres in the month of June contained a considerable number of R.repens achenes. In another novel and unusual, not to say quaint study of the trouser turn-upfluff of schoolboys who walked across fields to school, he showed that of the 70 plant species the boysunwittingly transported, 11% of the propagules were of R. repens (Darlington & Brown 1975).These authors pointed out that with the exception of adherent burrs like those of Galium aparine(Cleavers), the majority of the fruits and seeds in the turn-up are carried loose in the contained dustand fluff, "so that the wearer becomes a sort of peripatetic censer mechanism for scatteringpropagules, notably the smoother kinds (R. repens and others), as he walks about"(Darlington & Brown 1975, p. 34).
Seed dormancy and survival in soil
Dormancy is enforced by burying the seed and large populations of buried viable seed havebeen reported. Seed survival ability varies enormously, presumably dependent upon soil moisture,nutrient levels, stability and disturbance.
The survey of soil seed bank of NW Europe tabulates results of no less than 98 records of buried seedsurvival. Of the four seed bank categories listed, the representation of R. repens appears asfollows: transient (surviving less than 1 year) 21; short-term persistent (between 1 and 5 years) 26;long-term persistent (at least 5 years) 30; and present in soil (but not assigned to one of theforegoing) 21 studies (Thompson et al. 1997).
British and Irish occurrence
R. repens is extremely common and widespread over almost all of Britain and Ireland, becomingslightly less frequent in the NW and highlands areas of Scotland (Preston et al. 2002).
World occurrence
It has an almost continuous Eurasian boreo-temperate native range (Jalas & Suominen 1989, Map 1712).R. repens is also introduced and partly naturalised in both N & C America and has thus becomecircumpolar in its Northern hemisphere distribution. It is also an introduction in South America, SouthGeorgia, New Zealand, Tasmania and Great Barrier Island (Hultén 1971, Map 225; Hultén & Fries 1986,Map 842; Preston & Hill 1997).
Toxicity
Unlike other common buttercup species, R. acris, R. bulbosus and the more scarce R.scleratus (Celery-leaved Buttercup), it appears that Creeping Buttercup in the British Islesnormally contains only a low concentration of the Ranunculus poisonous principle protoanemonin (atoxic cardiac glycoside). Consequently, R. repens does not harm stock animals − including horses− and they frequently browse upon it (Cooper & Johnson 1998).
In the wider geographical range of the species however, there are instances where the levels of toxins inpopulations of R. repens are sufficiently high to make the plant distasteful, or even seriouslypoisonous, making it capable of causing diarrhoea and abdominal pain in cattle and sheep with symptomsthat persist for up to 14 days (Lovett-Doust et al. 1990). In Chile, for instance, in recenttimes R. repens poisoning was held responsible for causing abortions in a herd of dairy cattle(Morales 1989).
Herbicidal control
Although resistant to a wide range of herbicides, R. repens is very sensitive to translocatedselective herbicides such as 2,4-D, MCPA-salt, MCPB-salt, paraquat and aminotriazole (Lovett-Doust etal. 1990). Selective herbicides containing aminopyralid, such as Milestone and VM, can be usedto kill Creeping Buttercup. Aminopyralid products such as these are available at farm supply stores andshould only be used in areas listed on the label, ie pastures, hayfields and other agriculturalsettings. Fortunately, aminopyralid products do not harm livestock, provided all precautionary advice isfollowed. The Nature Conservancy Wildland Invasive Species Team publishes an online Weed ControlMethods Handboook that is regularly updated (Tu, M. et al. 2001: http://invasive.org/gist/products/handbook/methods-handbook.pdf
Accessed 25 January 2016). This tabulates herbicide advice and makes recommendations on other ecologicalmanipulations which help limit the weed population. To eradicate Creeping Buttercup from grassland itwill probably be necessary to apply herbicide up to three times, since mature plants can often recover,and seed in the soil seed bank will germinate and may re-establish the plant. Sprayed ground will needto be monitored and seedlings removed before they develop runners.
Uses
As R. repens tissues normally contain only a small percentage of protoanemonin, the species hasnot been used in herbal medicine in the same way as its close relatives, R. bulbosus (BulbousButtercup) and R. acris (Meadow Buttercup). In fact, Grieve (1931) does not mention it at all inher comprehensive book, A Modern Herbal.
Names
'Ranunculus' is derived from the Latin 'rana' meaning 'a frog', an allusion to the fact that so manymembers of the plant genus live in or near water, the habitat of frogs (Johnson & Smith 1946). TheLatin specific epithet 'repens' means 'creeping'. As one might expect, R. repens shares manyEnglish folk or common names with R. acris. Additional ones include 'Devil's Guts', 'Gold-balls','Granny-threads', 'Hod-the-Rake', 'Lantern Leaves', 'Meg-many-feet', 'Ram's Claws', 'Sitfast','Sitsicker', 'Tether-Toad' and 'Toad-tether' (Britten & Holland 1886). Many of these names refer tothe spreading stolons and/or the tenacity with which its roots cling to the ground, making the plantdifficult to eradicate.
Threats
None.
Native, frequent. European southern-temperate, also naturalised in N America and New Zealand.
1882; Stewart, S.A.; Knockmore Hill.
April to September.
Growth form and habitat preferences
Early in the growth season, when plants of this perennial are not yet in flower, the lower leaves of eachindividual's basal rosette lying flat to the ground are a distinctive identification feature of BulbousButtercup and, in addition, the corm-like, swollen stem-base can often be felt underneath the leaves.The lowest leaves being close-pressed to the ground (a feature described by Harper (1957, p. 332) as,'strongly epinastic' − an interesting term), appears to confer a distinct competitive advantage overother plants in the sward, since it allows the R. bulbous individual to form a 5-7 cmsaucer-shaped hollow in the turf, from which it manages to exclude other species. When Bulbous Buttercupis in flower, the down-turned (ie reflexed) sepals are very obvious and they provide another ready meansof recognition.
While Bulbous Buttercup is a characteristic perennial of Fermanagh's limestone pastures, it is not astrict calcicole, ie it may prefer and occur most abundantly on well-drained, lime-rich soils,but it is not entirely confined to them. Rather, the species also occurs on dry roadside banksand verges as well as on well-drained unimproved meadows and pastures, including some over more neutralto moderately acidic soils (ie around pH 5.0 and above). This can also involve the shallow peaty soilsthat form over leached Carboniferous limestone in Fermanagh and other areas in western Ireland.
Essentially a lowland plant of open, full-sun situations, R. bulbosus can withstand moderateamounts of disturbance such as trampling and since it contains protoanemonin and is unpalatable, stocknaturally avoid grazing it (Cooper & Johnston 1998). Severe trampling and the associated degree ofsoil disturbance and compaction will, however, obliterate the plant and its complete absence fromfrequently used pathways in grassland is often very obvious. Bulbous Buttercup also avoids stronglyacid, wet, shaded or indeed overly fertile, productive conditions. In the latter circ*mstance, tallerand more vigorous plants out compete it (Harper 1957; Grime et al. 1988).
Fermanagh occurrence
R. bulbosus is much less common in Fermanagh than either R. acris (Meadow Buttercup) orR. repens (Creeping Buttercup) and has only been recorded in 66 tetrads, 12.5% of those in theVC. Nine of these tetrads contain only pre-1976 records, indicating a loss of suitable habitat which iseasily identified with agricultural improvements made to lowland limestone grasslands in recent decades.As the distribution map shows, Bulbous Buttercup is widely scattered across Fermanagh, but it isdefinitely most frequent in the Monawilkin, Knockmore and Marlbank limestone areas.
Flowering
R. bulbosus flowers earlier than its closest buttercup relatives, R. acris and R.repens, the peak of its flowering normally occurring in mid- to late-May. Fruit has normallyripened by the end of June and the aerial parts may brown and die off soon afterwards, so that thesummer-dormant plant survives as the corm just below the soil surface − a drought-avoiding mechanism notvery necessary in the wet climate of western Ireland! The species can thus be very inconspicuous for afew weeks in mid-summer, particularly if we have a dry spell of weather. However, in our damp, mildclimate, with rainfall typically occurring regularly throughout the summer, Bulbous Buttercup plantsproduce fresh basal leaves soon after flowering (ie by the end of July or in early August), and theymaintain their growth until winter cold eventually stops them in October or November. The leaf rosetteis maintained throughout the winter and it recommences vegetative growth early in the year oncetemperatures begin to rise. Although the corm-like stem base and all the other parts of the plant arerenewed annually (Harper 1957, p. 333), this method of perennation means that the plant tissues neverbecome old and senescent and thus established individuals of R. bulbosus can be exceedinglylong-lived and, indeed, in stable habitats they may persist indefinitely. There is nothing againstsaying that, in certain circ*mstances, some individuals could be thousands of years old!
As with other common buttercup species, R. bulbosus shows considerable variation in form inrelation to prevailing environmental conditions (ie phenotypic plasticity), especially with respect tothe size of corms and leaves and the number of flowering stems per corm (Harper 1957; Coles 1973). Thetypical plant has a corm 1.5 cm in diameter which produces just one flowering stem, but larger corms cancarry up to eight or more flower stems and Harper (1957) reports a record breaking plant bearing 42!
The flowers open daily for about 4-7 days and are visited for both pollen and nectar by honey bees andmany other short-tongued insects. Cross-pollination is the norm since a high degree ofself-incompatibility exists, but while some evidence of selfing and a low degree of agamospermy, ie seedformation without fertilization, had been reported in the past (Harper 1957). However, Coles (1973)concluded after he had carried out greenhouse tests that R. bulbosus is totallyself-incompatible, cross-pollinated and sexual, ie the flowers are fully outcrossing (xenogamous), andthe breeding system is ‘panmictic’ (ie it involves random matings)(Richards 1997, page 6). Each flowerproduces between 20 and 30 achenes or seed (ie the achenes are single-seeded dry fruits).
Variation in seed production
The scale of seed production, or the reproductive capacity of the species, is another characteristic thatvaries enormously with the environment and particularly with the competitive situation of the plant. Itis hard to measure or summarise reproductive capacity, but Salisbury (1942) provided experimentalevidence by comparing the growth and productivity of plants in a meadow under three regimes: nocompetitors, and slight and severe levels of plant competition. As Harper (1957) pointed out, thecompetitive measurements Salisbury made were carried out without any experimental control, yet theystill give a clear impression of the overall scale of the effect of competition on the reproductivecapacity of the species. In the case of R. bulbosus, the sample under severe competition produced69 fertile carpels per plant, while that with no competitors produced 687, a tenfold difference insexual productivity.
Seed dispersal
The ripe seed drop off the receptacle and there is no specialised means of dispersal. Internal transferin the gut of animals through being eaten by birds or stock animals, plus external carriage in mud byanimals, including man and his vehicles, are probably significant. Wind and rain-wash may also play somerole in dispersal of R. bulbosus achenes (Harper 1957). Other bird species, such as Pigeons, areknown to be responsible for a considerable level of seed predation, and Voles and Field mice also makedepredations.
Phenology
Bulbous Buttercup germinates mainly in the late summer and early autumn on bare ground in gaps created inturf by disturbance such as trampling and overgrazing (Sarukhan & Harper 1973; Sarukhan 1974, 1976).In Britain, molehills may be significant in this respect, providing fresh, bare soil, but fortunatelythere are no moles in Ireland. Although tremendous rates of seedling loss are involved, effectiveestablishment from seed is the only reproductive mechanism that Bulbous Buttercup displays, cormdivision being extremely rare.
Seed survival in the soil seed bank
Buried seed longevity frequently appears to be brief: eight out of 15 studies in a major survey in NWEurope which gave survival estimates in years, indicated that the seed bank of R. bulbosus insoil is transient (surviving less than 1 year), and four indicated that it is short-term only (survivingmore than 1 year but less than 5) (Thompson et al. 1997). The remaining three studies gaveestimates of R. bulbosus seed longevity that ranged from 5 years to over 30! Clearly there isconsiderable variation in estimates of seed longevity, but the predominant mode remains transient toshort-term only.
Fossil record
The fossil record of R. bulbosus is more slight than that of other species in the genus, but thepattern is similar through numerous glacial and interglacial stages and thus provides conclusive proofthat the species is native in Britain and Ireland (Godwin 1975).
British and Irish occurrence
R. bulbosus is widespread in Britain and Ireland, but less so than either R. acris or R.repens. In Ireland, it is widely distributed but is less frequently recorded in western andsouthern areas of the island. The pattern in Britain shows the species is less frequent in the N & Wof Scotland. While this may in part be due to the upland, acidic nature of the terrain in the N & Wof both islands, the fact that the species is only conspicuous and readily identified in the earlysummer, probably means that it is under-recorded in areas where botanical recorders are themselvesscarce, or indeed, rare (New Atlas).
The calculated Change Index value measuring change between the two BSBI plant Atlases (publishedin 1962 and 2002) is -0.48, which is taken to indicate very little loss over the 40 year period (R.A.Fitzgerald, in: Preston et al. 2002). The results of the 2003-04 BSBI Local Changere-survey in Britain of the 1987-88 BSBI Monitoring Scheme survey squares suggest that R.bulbosus is actually on the increase in Britain. Unfortunately, this re-survey was not extendedto the island of Ireland. In comparison with other widespread plant species of predominantly calcareousor neutral grasslands, Bulbous Buttercup can cope relatively well with factors which lead to rankerswards, including eutrophication (essentially, nitrogen enrichment) (Braithwaite et al. 2006).
European occurrence
In continental Europe, depending upon which source is consulted, either two or three subspecies of R.bulbosus are now recognised: subsp. bulbosus which occurs in the British Isles, andsubsp. adscendens which is confined to the Mediterranean region, presumably including N Africa(Coles 1973). Subsp. adscendens may be subdivided into subsp. castellanus in NW Spain andsubsp. aleae in S Europe, extending eastwards to Hungary (Tutin & Akeroyd, in: Tutin etal. 1993). Plants of R. bulbosus in N Africa, N Turkey, Georgia and Azerbaijan also liewithin subsp. aleae (Hultén & Fries 1986, Map 845).
Subspecies bulbosus is widespread over W and C Europe, but it is absent from much of the N and Eof the mainland. Having said this, it is frequent in Denmark and southern Sweden, rarer on the southcoast of Norway, and extends east to Belarus and the Balkans (Hultén & Fries 1986, Map 845; Jalas& Suominen 1989, Map 1743; Jonsell et al. 2001, p. 289). Some outlying occurrences in CFinland and C Russia are considered occasional only (Hultén & Fries 1986).
World occurrence
Bulbous buttercup is a naturalised alien in N America, apparently invading from both E and W coasts,although much more frequent in eastern states. It is also a commonly naturalised weed in New Zealand(Harper 1957; Hultén & Fries 1986, Map 845; Jonsell et al. 2001).
Names
'Ranunculus' is derived from the Latin 'rana' meaning 'a frog', an allusion to the fact that so manymembers of the plant genus live in or near water, the habitat of frogs (Johnson & Smith 1946). TheLatin specific epithet 'bulbosus' means 'having a bulb', but of course technically it is not a leafystorage organ, that is, a bulb, but rather stem tissue (Gilbert-Carter 1964). Thirty-eight Englishcommon names are listed by Britten & Holland (1886), many of which are widely applied and some ofwhich are merely mis-spellings, for example, 'Bolt', for 'Bout', which is derived from the French'Bouton d'or', referring to yellow flower-buds of this and other species (Prior 1879). One of the moreinteresting names is 'Lodewort', said to be an Anglo-Saxon name for the species, but also applied bysome to R. aquatilis (Common Water-crowfoot) (Britten & Holland 1886). Another name ofinterest is 'St Anthony's Rape' or 'St Anthony's Turnip', from its corm being a favourite food of pigs,and he being the patron saint of pigs (Prior 1879, p. 204).
Uses
Although in Britain and Ireland it is not nearly as common a species as several other buttercups, R.bulbous is very conspicuous early in the growing season and, perhaps for this reason, it haslong been used in herbal medicine for its blistering properties. Grieve (1931) summarises the many uses,some of which like the cure for a headache which involves applying the acrid juice to the nostrils,makes the present author's mind boggle at the very thought. THIS DEFINITELY IS NOT ADVICE TO BEFOLLOWED.
Threats
Improvement of grasslands involving ploughing, reseeding and application of fertiliser.
Native, frequent, widespread and locally abundant. European boreo-temperate.
April 1854; Smith, T.O.; vicinity of Ardunshin.
March to July.
Growth form and habitat preferences
Goldilocks Buttercup is a perennial with a short, stout rootstock and in Fermanagh it mainly occurs (orperhaps is most noticed) beneath hedges, especially on roadsides where its distinctive pale yellowflowers can be spotted even from a car. Less frequently it is found in quite deep shade in hazel woodson limestone, eg in the Screenagh River Glen. These are moist to fairly dry sites and R.auricomus appears to require moderately fertile, base-rich, generally calcareous soils. Itreally thrives when it is supplied with a good depth of rich leaf-mould in undisturbed corners of oldwoodland or under scrub in light to half-shade conditions.
R. auricomus avoids both very acid and very dry sites (Garrard & Streeter 1983) and it appearsto be both a weak competitor and intolerant of grazing or cutting, tending to shun unshaded meadowgrassland for these reasons (Salisbury 1942, p. 54; Sinker et al. 1985). R. auricomus alsotends to be a lowland species in Britain and Ireland, although in some parts of Scotland at least, itcan rarely be found on open moorland when it is protected from grazing by boulders, and it may alsooccur rarely on mountain ledges (R.A. Fitzgerald, in: Preston et al. 2002).
Further north, in the Nordic region of continental Europe, R. auricomus shows a very much widerhabitat range than is observed in Britain and Ireland, appearing in much more open situations, eg inmeadows and grazed pastures, in wetter littoral and riparian habitats with bare soil, in mountainsnowbeds and on scree. It is also a weed in cultivated and disturbed ground in the more northern part ofits species distribution. Another difference is that it appears indifferent to lime in these northernterritories (Jonsell et al. 2001).
Overwintering (perennation) and phenology
The rootstock of R. auricomus overwinters with its bud at or just beneath the soil surface (ie itis a rosette-forming hemicryptophyte, or a buried geophyte). It begins growth in the very early spring,enabling it to flower a couple of weeks before R. bulbosus (Bulbous Buttercup) and several weeksearlier than all the other buttercups in Britain and Ireland. In a comparative study of the floweringbehaviour of five buttercup species in central Germany, R. auricomus was first to flower and ithad a five week flowering period (Steinbach & Gottsberger 1994).
British and Irish occurrence
R. auricomus is almost certainly an under-recorded species in Britain and Ireland and particularlyso in the less-frequented corners of these islands, due to its early flowering season. This runs fromMarch to May and peaks in April, so that it may be missed by 'summer botanists'. Despite their awarenessof this, Webb & Scannell (Flora of Connemara and the Burren, 1983) regarded GoldilocksButtercup as a rare species in W Ireland and it is generally considered that R. auricomusdeclines in frequency in Britain and Ireland as one goes northwards and westwards, a belief reinforcedby the distribution shown in the New Atlas (Preston et al. 2002).
R. auricomus is frequent and widespread in England and to some extent at least it mirrors thedistribution of chalk and limestone. In Wales, Scotland and Ireland, however, R. auricomus isvery much more scattered and the mapped distribution in these areas does not reflect the calcareousgeology. This is especially the case in Ireland where the distribution of the species is much betterrepresented than was the case in the previous 1955-60 BSBI Atlas survey (Perring & Walters1962, 1976; Preston et al. 2002, page 3).
Fermanagh occurrence
In Fermanagh, there were just 18 pre-1975 records for R. auricomus, but when RHN started lookingfor it around 1986, he found that it was very widespread, quite frequent and locally abundant. We nowhave over 250 records and the distribution map shows Goldilocks Buttercup present in 121 tetrads, 22.9%of those in the VC. Harron in his Flora of Lough Neagh (1986) made the very same discovery aroundLough Neagh, where he considered R. auricomus present in greater abundance than anywhere else inUlster (ie in the nine county Irish province).
Apomixis, an unusual breeding system producing an exceptional range of variation
R. auricomus is a facultative apomictic, ie in addition to the normal sexual process it displayspseudogamous agamospermy − which is a shorthand technical way of saying that it can set seed asexually,but only after pollination takes place (either by crossing or selfing). Despite the requirement forpollination, in the case of apomixis, no actual fertilisation takes place, yet seed isproduced. For a simple introduction to this complex matter see Proctor et al. (1996), pp.348-349, and for a more detailed explanation see Richards (1997a), p. 405 and pp. 411-420).
As a further complication, the apomictic microspecies created are all at least tetraploid (Jonsell etal. 2001). As a result of its dual reproductive methods, R. auricomus is extremelyvariable in form, especially with regard to petal development. A very full treatment of the microspecieshas recently appeared in Flora Nordica, 2 (in English), where a total of no less than 605microspecies in the Nordic countries are described (Jonsell et al. 2001). The agamospecies havenot yet been formally described within the British Isles, but Stace (1997) reckons at least 100 R.auricomus microspecies exist in these islands.
Flower variation
In Fermanagh, as elsewhere, plants of R. auricomus are sometimes found with five perfect petalsas, eg in the Teemore district, but more frequently the petals are reduced in number, distorted in theirdevelopment, or even in some cases totally absent. When the petals are much reduced or are completelyabsent, the sepals, which are usually greenish, may develop yellow colour and become shiny and petaloid,thus taking on the advertising role as insect attractants (Hutchinson 1972).
Pollination compared with related species
R. auricomus has either a cup-like nectary without a covering scale, or it has a small orabortive scale (Butcher 1961; Clapham et al. 1962). The flowers attract insect visitors, but insmaller numbers than most other terrestrial buttercup species. A study of five common buttercupspecies in central Germany found that while in its natural habitat R. flammula (LesserSpearwort) attracted up to a mean of 35.7 insects per hour, R. auricomus achieved a mean of just2.3 per hour. When cultivated in a garden bed R. auricomus was visited by ten species of insects(evenly divided between Diptera, Hymenoptera and Coleoptera), whereas R. acris (MeadowButtercup), R. flammula, R. bulbosus (Bulbous Buttercup) and R. repens (CreepingButtercup) had visits from 54, 41, 37 and 28 species, respectively. Thrips were the only insectsobserved collecting nectar from the Goldilocks Buttercup, while the other nine visiting species tookonly pollen. In its natural habitat, R. auricomus was visited only by Coleoptera at the very lowrate quoted above (Steinbach & Gottsberger 1994).
Seed production
The present author does not know of any study of the reproductive capacity of R. auricomus(certainly nothing published in English).
Seed dispersal
Nordic studies of seed dispersal in their much wider range of habitats emphasised the role of humanactivities (eg transport of hay), for both local and long-distance microspecies movements (Jonsell etal. 2001). However, this is obviously irrelevant, or much less relevant, when the plant lives inwoods and hedges rather than in meadows, as is the situation in most of Britain and Ireland. How doesGoldilocks Buttercup get about? Does it have a dispersal mechanism at all? The seed (actually achenes)are described as, "very shortly pubescent" (Stace 1997), perhaps suggesting they might adhereto animal coats. Another project beckons!
Buried seed longevity
The survey of soil seed banks in W Europe found a total of 15 estimates , ten of which regarded R.auricomus seed as transient, two considered it long-term (ie persisting for at least 5 years),and three did not specify any duration (Thompson et al. 1997).
A call for basic ecological information
A careful but necessarily incomplete internet search failed to unearth any other information on the lifehistory or ecology of this interesting species group. Perhaps if less research emphasis were placed onits genetics and reproductive strategy and a preliminary study initiated on the population biology,life-table and natural history of the species group, it might prove worth the effort.
European and World occurrence
In W & N Europe, R. auricomus s.l. is widespread everywhere except Spain and the Mediterraneanmainland, where it becomes rare and scattered towards the south. It is absent from all Mediterraneanislands except Corsica (Jalas & Suominen 1989, Map 1809). R. auricomus s.l. also occurs in NAsia, Alaska, NE Canada and Greenland (Jonsell et al. 2001).
Uses
Despite its attractive-sounding English common name, the species does not appear to have any folk-lore oruse specifically associated with it. This is probably because the plant is too rare, or is seldomrecognised.
Names
'Ranunculus' is derived from the Latin 'rana' meaning 'a frog', an allusion to the fact that so manymembers of the plant genus live in or near water, the habitat of frogs (Johnson & Smith 1946). TheLatin specific epithet 'auricomus' is a combination of 'aurum' meaning 'gold' and 'coma', meaning 'hairof the head' or 'locks', and thus translates as 'with golden hair', presumably a poetic likening of thespring carpet of yellow flowers to a blonde head of hair (Gilbert-Carter 1964). The English common name'Goldilocks' or 'Goldylocks' is a straightforward translation of its Latin specific name, first used byWilliam How in his Phytologia britannica of 1650. An alternative name is 'Wood Crow-foot'(Britten & Holland 1886).
Threats
Removal of both hedges and small patches of woodland.
Native, scarce. Circumpolar boreo-temperate, disjunct in E Asia and widely naturalised in the southernhemisphere.
1934; Praeger, R.Ll.; around Enniskillen.
May to September.
Growth form and habitat preferences
R. sceleratus is quite a tall (30-70 cm), conspicuous but rather scarce, much branched, yellowishgreen plant producing a host of rather small yellow flowers. This many-seeded winter or summer annual isa pioneer coloniser of shallow water, or wet, disturbed, nutrient-rich (especially nitrogen-rich), baremud after it has been thoroughly disturbed, eg heavily trampled and poached by drinking livestock. Theanimals also provide, of course, the required nitrogen in their excretion (van der Toorn 1980; R.A.Fitzgerald, in: Preston et al. 2002). The wet, open, almost always lowland habitats it frequentsare generally flooded and deeply submerged for part of the winter months and often indeed remain so intothe late spring or even the summer in our wet Oceanic (or Atlantic) climate. R. sceleratus istotally absent from soils below about pH 4.0 and it never occurs in permanently flooded aquaticsites (Grime et al. 1988).
With regard to its ecological status, R. sceleratus is always a pioneer species colonising baremud. Among its many associates are Myosotis scorpioides (Water Forget-me-not), Rorippapalustris (Marsh Yellow-cress), Persicaria hydropiper (Water-pepper), Alismaplantago-aquatica (Water-plantain), Veronica beccabunga (Brooklime), Lemna minor(Common Duckweed), L. trisulca (Ivy-leaved Duckweed), Callitriche stagnalis (CommonWater-starwort) and Bidens cernua (Nodding Bur-marigold). In terms of plant communities, R.sceleratus belongs chiefly to the NVC OV32 Myosotis scorpioides-Ranunculussceleratus open, nitrogen-rich, often muddy and disturbed, intermittently wet ground community(Rodwell et al. 2000, page 434), an Association of the Bidention Alliance which goes undervarious names in different parts of W Europe (White & Doyle 1982). It does also occur however, as aregular associate in seven other aquatic and swamp communities listed in Rodwell et al. (1995).
Like most pioneer colonisers of bare ground habitats, the presence of R. sceleratus tends to beephemeral. Where disturbance occurs more rarely or irregularly, it is gradually crowded out by thearrival of taller, more permanent colonising vegetation dominated by species such as Phragmitesaustralis (Common Reed), Typha latifolia (Bulrush), Schoenoplectus lacustris(Common Club-rush), Equisetum fluviatile (Water Horsetail), Cicuta virosa (Cowbane) andIris pseudacorus (Yellow Flag). The latter, together with various sedges and a collection ofother invading and carpeting species, can very quickly cover and occupy previously bare mud,out-competing and excluding R. sceleratus. In addition, should the inhabited site dry out duringa prolonged drought, R. sceleratus quickly succumbs. It is too fleshy and succulent to survivedry conditions for long (Grime et al. 1988).
Fermanagh occurrence
This is a rather scarce annual which had only been seen twice in Fermanagh before 1980, but since then ithas been recorded at 15 new sites covering 21 tetrads, 4% of the total in the VC. As the distributionmap shows, it is thinly scattered in seasonally flooded water meadows around the Upper Lough Erne basin,mainly in the south of the county, with one outlying station at Derryclawan near Enniskillen. The latteris the only station where it was found in any real quantity and here it grows on cattle poached, wet,well-dunged, anaerobic or very poorly-aerated mud on the bed of an old lake exposed in summer after aspell of dry weather.
Flowering
R. sceleratus flowers are remarkable for their outsized, elongated, pineapple-shaped receptacle,which sits quite incongruously amongst the encircling small petals and tends to dwarf them. Flowers areproduced mainly from May to September, but chiefly from June to August. They attract flies and bees withfreely presented nectar and they may be cross-pollinated by wind or by their winged insect visitors(Clapham et al. 1962; van der Toorn 1980). In addition, if this fails to occur, the travels ofthrips and aphids crawling around individual plants enables self-pollination (Baker & Cruden 1991).
Seed production
The many seeds of the plant (ie the achenes − single seeded dry fruits), are produced at rates of between70-100 per receptacle and up to 45,000 per plant (with a mean of 26,000, however) (Salisbury 1942).Another estimate given by van der Toorn (1980) indicated that a large plant in very good growingconditions and with little or no competition can produce up to 50,000 achenes. The individualseeds/achenes are smaller than those of R. flammula (Lesser Spearwort) and many times smallerthan those of R. lingua (Greater Spearwort) (Clapham et al. 1962).
Seed dispersal
Seed dispersal involves wind and water. Seeds float for at least an hour, but generally somewhat longer,the distance travelled obviously dependent upon rate of water flow and density of waterside vegetation(van der Toorn 1980).
Germination and life-cycle/growth strategy
Celery-leaved Buttercup seed which has been stratified by winter cold for 4 to 6 months and subsequentlyexposed will germinate in the spring or early summer (van den Toorn & ten Hove 1982). The speciescan complete its life-cycle in two months (ie behaving as a summer annual). If it germinates later inthe summer, for instance in August or September, it is frost resistant and may persist through thewinter as a submerged dormant leaf rosette. It then recommences growth when the mud it occupies isexposed the following year − ie it behaves as a winter annual, thus giving the species a dual lifestrategy (Bakker 1966; van der Toorn 1980). During the second year, these winter annuals will probablyseed early in the summer and, since a proportion of the fresh seed can germinate immediately, there maybe sufficient time for a second generation to grow and complete their life-cycle in the same season(Bakker 1966; Grime et al. 1988).
In the first year or so after germination, R. scleratus can build up its population very quicklydue to its enormous seed output and their easily achieved dispersal by wind and water.
Buried seed survival
Despite their small size, the seeds can lie dormant and survive for many years on the muddy margins oflakes, ponds and ditches, until low water levels expose the bare mud and disturbance brings them up tothe light, triggering germination. Five of the 13 records quoted in the survey of NW European soil seedbanks reckoned that R. sceleratus seed is long-persistent: one estimate reckoned survival ispossible for over 50 years (Thompson et al. 1997).
British and Irish occurrence
In suitable muddy habitats in other parts of Northern Ireland, Celery-leaved Buttercup is much morecommon than it is in Fermanagh, eg in Co Down (H38) and in all the vice-counties around Lough Neagh inparticular, ie Cos Tyrone, Armagh, Down, Antrim and Londonderry (H36-H40) (Harron 1986). It is also morefrequently found in coastal areas both around the province and in the wider British Isles. R.sceleratus is tolerant of brackish alluvial mud conditions and therefore it is also a frequentpioneer coloniser of mud on grazed or otherwise disturbed estuarine saltmarshes (NI Flora Website 2002;R.A. Fitzgerald, in: Preston et al. 2002). Overall, however, R. sceleratus occupies arather restricted niche habitat and it is not surprising that although it is widespread and frequent inC & SE England, the species is uncommon in many inland parts of Ireland, Wales and Scotland (Gray1970; Stace 1997).
European and world occurrence
In Europe, R. scleratus is widespread, especially in C & W areas, becoming scattered to rareand increasingly coastal to both the north and south of its range (Jalas & Suominen 1989, Map 1828).The species extends through the Near East, Siberia, C Asia, Japan to eastern N America (where it is infact also represented by an additional subspecies, subsp. multifidus) (Hultén 1971, Map 291). Afurther subspecies, subsp. reptabundus (Rupr.) Hult. is later mapped in N Europe and NW Siberia(Hultén & Fries 1986, Map 857). Hultén remarks that as the plant is so often apophytic (ie occupiesman-made or strongly man-influenced habitats) it is difficult to decide exactly where R.sceleratus is native. He is particularly suspicious about its native credentials in eastern NAmerica, and I would go further and say that it is always introduced in N America. In the southernhemisphere, Celery-leaved Buttercup is a certain introduction, for instance, in C & E Africa, NewZealand, Queensland and Tasmania (Hultén 1971; Jonsell et al. 2001).
Fossil record
Living as it does in nutrient- and nitrogen-rich muddy habitats and requiring disturbance to initiategermination, R. sceleratus is very often associated with human settlements. This naturally raisesthe possibility that in some places it might be an ancient introduction, ie an archeophyte. However,these muddy habitats also favour preservation of fossil pollen and achenes and this evidence clearlyindicates that R. sceleratus has been continuously present in Britain and Ireland from thePastonian stage onwards. Thus Celery-leaved Buttercup is very definitely a native species (Godwin 1975).
Toxicity
Like other buttercups R. sceleratus contains the glycoside ranunculin, which on hydrolysis breaksdown to yield an irritant oily substance protoanemonin, plus glucose (Saber et al. 1968).Protoanemonin is responsible for the toxicity of all Ranunculus species, and R. sceleratusis reputed to be the most poisonous of all. It is possible, however, that because of the rich,luxuriant, somewhat succulent growth of the plant, it may be eaten in larger quantities than otherbuttercup species. Protoanemonin poisoning is reported most frequently in cattle, the acrid juicecausing blistering of the mouth. In an experimental trial, one goat fed with R. sceleratus diedand two others became severely ill. Celery-leaved Buttercup does not normally invade pastures, but ahorse that grazed an area where it had access to R. acris (Meadow Buttercup) and R.sceleratus temporarily developed paralysis, convulsions and a loss of sight and hearing (Cooper& Johnson 1998).
Names
The plant's toxicity undoubtedly explains why it was given its Latin specific epithet 'sceleratus', whichmeans 'wicked' or 'vicious' (Gilbert-Carter 1964).
Apart from the aptly descriptive English common names, 'Celery-leaved Buttercup' and 'Celery-leavedCrowfoot', the plant is also known in N America as 'Cursed Crowfoot'. Presumably farmers whose animalsattempt to eat it, refer to it in this way. An alternative common name listed by Britten & Holland(1886) is 'Ache', apparently through a connection with the old French name for Parsley, and thus aconnection via 'Apium' to the vegetable, Celery. A further name 'Thiretelle' originates in twodictionaries of obsolete English which refer to, "The herb apium risus", which is identifiedby Britten & Holland (1886) as R. sceleratus. Another name these authors mention is'Blisterwort', which originated with Lyte (1578), and is a useful reminder that buttercup sap veryreadily causes burn-like blisters on skin.
Uses
In past times, beggars were said to use buttercup species commonly, and especially R. sceleratus,in order to induce sores on themselves to excite compassion and gain alms from the public (Lightfoot1777, p. 291; Vickery 1995, p. 63). According to Mrs Grieve who reports this nefarious use of the plant,after 'working' their 'con', the beggars afterwards would cure their blisters by applying fresh Mulleinleaves to the wounds (Grieve 1931, pp. 182 & 235). Funnily enough, Grieve does not mention thishealing property under her entry for Mullein (Verbascum species). Poor beggars! Grieve warns thatR. sceleratus, "is one of the most virulent of native plants: bruised and applied to theskin, it raises a blister and creates a sore by no means easy to heal". She goes on to indicatethat if the plant is boiled and the water discarded, it can be eaten as a vegetable, and was peasantfood in Wallachia (an old name for Romania). Grieve mentions a tincture, used in small doses, asan herbal cure for "a stitch in the side and neuralgic pains between the ribs".
R. sceleratus is widely used to this day in homeopathy and numerous internet websites deal withthis topic. YOU ARE STRONGLY ADVISED NOT TO MAKE ANY ATTEMPT TO MAKE USE OF THIS VERY DANGEROUS CAUSTICPLANT.
Threats
Due to the general nutrient enrichment of aquatic habitats, we may see this species increasing, providedthat climatic change does not reduce the water level fluctuation that provides bare mud forcolonisation.
Native, occasional, but easily overlooked and perhaps under-recorded. Eurosiberian temperate.
1836; Mackay, J.T.; Lough Erne.
May to September.
Growth form and habitat preferences
Greater Spearwort is an erect, robust, semi-aquatic, emergent perennial up to 120 cm tall with a creepingrhizome or stolon up to 50 cm long, bearing shallow fibrous roots (Jonsell et al. 2001). As such,it is a bigger plant that shares some of the characteristics of both R. flammula (LesserSpearwort) and R. repens (Creeping Buttercup), ie like R. flammula it is a helophyte,growing in soil frequently saturated with water or it stands in shallow water with its base submerged.Again, like R. repens, the underground stoloniferous stem branches and produces offsets, daughterplantlets or ramets, which can play an important role in vegetative reproduction, clonal local diffusionand longer-distance dispersal of the species within a lowland, wetland system (Johansson & Nilsson1993).
R. lingua is a plant of mesotrophic to eutrophic, rather nutrient-rich, sheltered, shallow water,rich-fen and swamp vegetation. It seems to prefer places where there is a fairly gentle inflow of streamwater, presumably adding nutrients. It is often rooted in organic, peaty, lime- or base-rich mineralmud, usually with a pH between 5 and 6.5 (Spence 1964; Garrard & Streeter 1983; R.A. Fitzgerald, in:Preston et al. 2002). As is the case with R. auricomus (Goldilocks Buttercup) and someother species, which in the British Isles frequent calcareous or base-rich soils, in Nordic countriesR. lingua appears indifferent to lime (Jonsell et al. 2001).
Fermanagh occurrence
Prior to 1975, there were a total of just 27 records for R. lingua in Fermanagh. However, thanksto the extensive recording in the VC from 1977 onwards, which has particularly focused on lowlandwetlands, there are an additional 200 post-1975 records for Greater Spearwort in the FermanaghFlora Database. This stoloniferous perennial has now been recorded in a total of 64 tetrads,12.1% of those in the VC.
In Fermanagh, R. lingua grows among tall, often dense, fen and marsh reed vegetation, especiallyaround the small inter-drumlin lakes that form the fretted margins of Upper Lough Erne. It is also foundaround a variety of other lakelets and ponds and on the muddy banks of rivers, canals and ditches, eg inmarl ponds along the River Finn, on the banks of the Swanlinbar River, the Old Ulster Canal and ingravel-pits at Gortaree. It typically associates with Phragmites australis (Common Reed),Typha latifolia (Bulrush), Carex elata (Tufted-sedge), Equisetum fluviatile (WaterHorsetail), Cicuta virosa (Cowbane) and Sium latifolium (Greater Water-parsnip). Itis present, but not a constant or even a very frequent species in one aquatic (A4) and six swampplant communities in the NVC listing (S1, S4, S17, S22, S24 and S27) Rodwell et al.1995).
Although R. lingua is stoloniferous and potentially clonal patch-forming, in Fermanagh it isalways a rather sparse and local component of the type of tall waterside vegetation it frequents. R.lingua is a decidedly inconspicuous plant until about mid-June when it comes into flower. Itcould thus be very easily overlooked during early season field recording. Seven Fermanagh tetradscontain only pre-1976 records, possibly indicating a loss of suitable habitat, or the need for moretimely recording.
Greater Spearwort is said to be intolerant of trampling and grazing (Sinker et al. 1985), andsince both these pressures must occur on the grazed water meadows along most of the Fermanagh lakeshores where the species occurs, these two factors might well be limiting its local occurrence.
Flowering
R. lingua begins flowering in late June and continues until September, reaching a peak in July.The rather large, creamy yellow, very glossy- petalled flowers are usually borne in a few-flowered cyme,but sometimes flowers are solitary. The flowers are protogynous (ie their female stigmas ripen beforetheir pollen) and they attract insect pollinators (mainly flies) by producing copious nectar (Claphamet al. 1962; Hutchinson 1972). Broad-leaved forms of R. flammula can easily be mistakenfor R. lingua, but they have much smaller flowers, about half the diameter of those of GreaterSpearwort.
Fruiting, overwintering and vegetative reproduction
R. lingua is described as 'thermophilous' by the Dutch botanists van der Voo & Westhoff(1961), meaning that it prefers relatively warm temperatures, or it requires such conditions to reallythrive (Holmes 1979). Further north in Sweden, fruit-set is often poor and recruitment from seed isregarded as very infrequent. It is possible that from time-to-time in a poor summer fruiting may also bepoor in parts of Britain and Ireland. I have not located any measurements or estimates of thereproductive seed capacity for this species in either Britain or Ireland and, clearly, there is need forfurther study in our latitudes. A Swedish study at around 60oN reported that individualplants (ie ramets) of R. lingua live for just one year, and that propagation occurs by means ofoverwintering rhizomes. These overwintering organs are up to 10 mm thick and 25-75 cm long and consistof 5-10 nodes. They are produced in late summer from basal stem nodes lying just below thesediment surface (Johansson & Nilsson 1993). This system of growth and perennation results in clonesof physiologically independent ramets or daughter plantlets. It was observed that each established rametwent on to produce one or two daughter ramets each year of the study (Johansson & Nilsson 1993).
Dispersal
The overwintering rhizomes or stolons are very efficient water-borne dispersal units, being perfectlybuoyant for long periods due to their having hollow internodes. In comparison, seed only floats for oneto two hours, which must severely limit their efficiency as water-borne dispersal units (Romell 1938,cited in Johansson & Nilsson 1993). There is, however, a high mortality of dispersed ramets duringthe first year after dispersal, so that successful establishment, even from organs as large as rhizomes,must be rare events. In their study the Swedes concluded that (at least in their area) R. linguais a "pseudo-annual clonal plant", and that annual clonal disintegration (of the individualplant) can be viewed as a form of (ecological and biological) risk-spreading (current author's insertedbrackets) (Johansson & Nilsson 1993). The probability of extinction decreases because some rametsare always devoted to dispersal to new sites. "In reality, however, only a few of all rhizomes aredispersed, and this can also be interpreted as a safeguard against local population extinction."(Johansson & Nilsson 1993).
Although seeds in many wetland species are ineffective for long-distance dispersal within water systems,their smaller size and longer life must still allow them to be the most effective means of dispersalbetween water systems by transporting agents such as birds and other animals (Smits et al. 1989).Only one reference is given in the comprehensive survey of soil seed banks of NW Europe (Thompson etal. 1997), and it suggests that seed of R. lingua is transient, surviving less than oneyear.
Changes in distribution
As mentioned above, R. lingua has been in decline for perhaps as long as 200 years (Harron 1986;Hackney et al. 1992), a situation demonstrated for the British Isles in the 1962 and 1976Atlases (Perring & Walters 1962, 1976). This decrease in range is also described for theNordic countries by Jonsell et al. (2001), where the plant has retreated south of the ArcticCircle (see their map, p. 277). Although changing temperatures may have had some effect on the sexualreproductive capacity of the species further north, it is much more probable that the main factorcausing losses in Britain and Ireland was land drainage, since R. lingua is not very sensitive toeither water pollution or eutrophication (nutrient enrichment) (Jonsell et al. 2001). However, atleast in Britain, if not elsewhere, previous losses have been reversed, as is clearly illustrated by themaps published in the New Atlas (Preston et al. 2002).
Introduction to gardens
During the last 40 years or so, Greater Spearwort has gained appreciation from horticulturalists and isnow considered a sufficiently decorative, appropriate and easily enough cultivated subject for plantingaround the fashionable, indeed almost obligatory garden 'water feature'. Thanks to this trend, therehave been so many 'escapes' and deliberate introductions of the plant to the wild in Britain, thatFitzgerald commented, "the distinction between native and alien populations is now hopelesslyblurred" (R.A. Fitzgerald, in: Preston et al. 2002).
Irish occurrence
The New Atlas shows that R. lingua is widely scattered and locally frequent in Ireland, butthe main areas of concentration are undoubtedly in the C & NE of the island. However, while it mayremain scattered in NE Ireland, the area around Upper Lough Erne now appears to constitute the mainstronghold of this rather scarce and local emergent aquatic species in the north of Ireland. Harron(1986) described Greater Spearwort as being widespread but very sparingly distributed around Lough Neaghand he considered it was probably decreasing there. Hackney likewise regarded it as rather rare in thethree counties in the FNEI 3.
British occurrence
Due to the increased use of the plant in gardens in recent decades, it is almost impossible todistinguish many native populations from introduced populations of R. lingua in S & C Englandin the New Atlas map (Preston et al. 2002). However, if we mentally subtract the obviouslyalien concentrations of R. lingua around the major conurbations, the encouraging impressionremains discernible, that the species has at least survived in those native areas where it appeared inthe earlier BSBI Atlas (Perring & Walters 1976). The better recording coverage in some areasalso helps offset some definite losses.
European occurrence
In Europe, the species is widely represented in W & C areas, declining to both N & S, and onlyoccasional and very scattered throughout the Mediterranean region. There have also been significantextinctions in Belgium, N and W France, S Germany and Hungary (Jalas & Suominen 1989, Map 1861).
World occurrence
Beyond Europe, Greater Spearwort ranges from SW Asian Russia (to the Altai), the Caucasus and Kashmir inW Asia. It has been introduced in New Zealand (Hultén & Fries 1986, Map 868). There is very littlevariation within the species and none of taxonomic importance (Jonsell et al. 2001).
Toxicity
Like other buttercup species R. lingua contains bitter-tasting toxins which undoubtedly help detercattle and other stock, although it is not specifically reported as being responsible for poisoning anysuch animals (Cooper & Johnson 1998).
Names
The Latin specific epithet 'lingua' means 'tongue' and refers, quite aptly in this instance, to the shapeof the stem-leaves (Gilbert-Carter 1964). The English common names 'Spear-wort' and 'Spear Crowfoot'were applied from the 14th century, again on the basis of the leaf shape, to both the more common R.flammula (Lesser Spearwort) and to R. lingua (Grigson 1974). Thus to distinguish them inmodern times, they have been called 'Lesser' for the smaller, narrower-leaved R. flammula and'Greater' for the larger of the two. The only additional name is 'Sparrow-weed', which Britten &Holland (1886) list only from Co Londonderry (H40).
Threats
Hyper-eutrophication of its habitat, or destruction of the vegetation surrounding the lakes where theplant grows.
Native, common, very widespread and locally abundant. European temperate, introduced in a few stations inW Asia and N America.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This is a very variable, abundant and widespread perennial of soft, wet, marshy ground found mainlyaround lakes and ponds and in seasonally flooded water-meadows. R. flammula is the type of plantdescribed in Raunkiaer's Life Form Classification as a 'helophyte', a term derived from Greek literallymeaning 'a marsh plant'. The term refers to a perennial with an underground storage organ which grows insoil saturated with water and which, therefore, has submerged winter buds and is thus typically shallowrooted like R. flammula (Raunkiaer 1934; Holmes 1979; Grime et al. 1988).
R. flammula occurs in situations where the dense shading effect of tall fen vegetation is excludedor kept minimal by a range of ecological factors, perhaps including nutrient limitation, but moretypically involving a combination of seasonal flooding and occasional spates, plus grazing, trampling,cutting or other forms of disturbance, including that generated by popular water-based human leisureactivities. It is also very frequent or almost constant in hollows in damp grassland, by flowing waterin streams, ditches, springs and in flushes on moors, bogs and upland woods, eg in Fermanagh's woodedCorrel Glen NR.
In Britain, R. flammula is mainly associated with moderately oligotrophic to mesotrophic waters(Preston & Croft 1997), but in Fermanagh it also features around or near some of our more decidedlyeutrophic waterbodies and it is particularly frequent around Upper Lough Erne. In coastal regions ofBritain and Ireland, R. flammula additionally occurs in dune slacks and on damp sea cliffs (Grimeet al. 1988).
Under terrestrial habitat conditions R. flammula typically grows erect, but it can also sprawlhorizontally to some extent (ie it can be decumbent). However, when the base of the plant becomessubmerged in water the habit often becomes creeping. Under these circ*mstances, it then roots at thenodes, ie it becomes stoloniferous, and given relatively open substrate conditions to colonise, such asbare mud or disturbed soil it may branch and spread to form a clonal patch (Cook & Johnson 1968;Grime et al. 1988). As with other waterside species, this feature undoubtedly plays an importantpart in the reproduction of R. flammula, since detached portions disperse readily in flowingwater.
Preferred soil types
In terms of substrate, Lesser Spearwort generally occurs on wet, moderately acidic, peaty mud, stonygravel or mineral soils, but in Fermanagh it is also very common in limestone and marl situations, egaround Lower Lough Erne and along the pools in the River Finn. This range departs from the British Islesnorm to some extent, since Hill et al. (1999) gave it an indicator value for soil reaction of'5', meaning, "of moderately acid soils, only occasionally found on very acid or on neutral tobasic soils". The original soil reaction value associated with the species by Ellenberg (1988),based on his experience in continental Europe, was '3', which is significantly (or strongly) inclinedtowards the acid end of the nine-point scale used in his soil classification. In the Sheffield region ofEngland, Grime et al. (1988) found R. flammula occurred, "mainly on mildly acidicsoils between pH 4.5 and 6.5, but extending locally to soils of pH up to 7.5". The latterrepresents a similar range to that noticed in Fermanagh. In contrast to our local experience, however,they went on to comment, "Rare on calcareous soils." Perhaps the interesting thing is that thespecies DID also occur on highly calcareous soils, as it very definitely does to a greater extent inFermanagh. Further north in Scandinavia, R. flammula is regarded as indifferent to lime (Jonsellet al. 2001).
Fermanagh occurrence
Lesser Spearwort is very widespread and abundant in Fermanagh and has been recorded in 438 tetrads, 83%of those in the VC, a situation very much expected in an area of the country with so many lakes and alarge expanse of marshy ground, fens, bogs and ditches. Although obviously very widespread throughoutthe county, it is especially frequent in the area lying south of Lough Erne.
British and Irish occurrence
R. flammula is also frequent and widespread throughout the whole of the British Isles, but hasdeclined to some extent in SE England, presumably due to the usual factors of pressure for land,development, drainage and intensive agriculture (R.A. Fitzgerald, in: Preston et al. 2002).
Is R. flammula really an aquatic plant?
The definition of what constitutes an aquatic plant is not an easy matter to decide, since the boundarybetween land and water fluctuates over several time scales. Lesser Spearwort is not really a true'aquatic species' in the ecological sense that fits all the related Water Crowfoots, including theRanunculus subgenus Batrachium species with the exception of R. hederaceus(Ivy-leaved Crowfoot) and R. omiophyllus (Round-leaved Crowfoot), ie which in Professor Cook'sview is limited to "species with a submerged phase during their generative history" (Cook1966). This very strong delimitation of an aquatic plant makes the submerged phase absolutelyobligatory. However, like the two previously mentioned water-crowfoot species, R. flammula isprimarily a terrestrial wetland species that very commonly is found standing emergent in water. It maybe temporarily submerged, either regularly, or only occasionally whenever local water table levels arehigher than normal. It is NOT quite patently a species that, "characteristically grows in waterwhich persists throughout the year", which is the definition of an 'aquatic plant' provided byPreston & Croft (1997). Despite the failure of R. flammula to comply with their definition,the latter authors included a species account of it in their excellent book, Aquatic plants inBritain and Ireland. At the same time Preston & Croft (1997) omitted other native speciescharacteristic of tall-herb fen, which in Fermanagh, and other parts of Ireland, are capable of formingpartially emergent floating mats in lakes and rivers. Many of these ignored or neglected species appearequally deserving of treatment as emergent aquatics, eg Cicuta virosa (Cowbane) and Siumlatifolium (Great Water-parsnip) (Cook 1998). As with all published selections of plant species,considerations of time, space and cost undoubtedly intrude and they can determine the resulting quorum.
Variation
Living in a fluctuating environment on or near the boundary between land and water, R. flammulashows a high degree of phenotypic variation in form with respect to changes in its environments (iemorphological plasticity). This affects a wide range of characters including size, habit, leaf shape,flower size and even achene shape (Jonsell et al. 2001). Leaf development is particularlyvariable and plastic in form, to the extent that when the leaf develops under aerial conditions it islanceolate, whereas when it develops under water, it is linear. This is also a case where heterophyllyis of a reversible kind, for although change in shape of an individual leaf is impossible, an extendingshoot can produce first one leaf form and then the other, in response to changing environmental growingconditions. It is important to distinguish this kind of heterophylly from that more often met, which isassociated with maturation of the plant or entering a flowering phase, where the change in leaf formfollows an irreversible one way sequence from youth to maturity (Cook & Johnson 1968).
Three subspecies of R. flammula are recognised by Stace (New Flora of the British Isles,1997), of which subsp. flammula is the common and widespread form, while the other two, subsp.minimus (A. Benn.) Padmore and subsp. scoticus (E.S. Marshall) A.R. Clapham, are very muchmore rare or under-recorded. We have not attempted to distinguish the subspecies in Fermanagh, althougha few old records of subsp. scoticus exist in the Fermanagh Flora Database (see separate accountbelow).
Possible identification problems
Large specimens of R. flammula, sometimes distinguished as var. ovatus Pers. (= var.major Schult.), can easily enough be mistaken for R. lingua (Greater Spearwort) (Padmore1957; Preston & Croft 1997). However, in Fermanagh the latter species is almost invariably foundgrowing around Upper Lough Erne and it generally occurs within the taller lakeshore vegetation thatR. flammula eschews. Despite the above mentioned phenotypic variation that occurs in R.flammula, after considerable field experience, we find that the small lanceolate leaves ofnon-flowering specimens are quite distinctive and recognisable.
The more procumbent phenotypes of R. flammula are also sometimes confused with its very rarerelative R. reptans (Creeping Buttercup), which is confined to two sites in Scotland and a few,perhaps transient sites in Cumbria (Padmore 1957; Gibbs & Gornall 1976). An equally rare hybrid alsooccurs between these two species (Gornall 1987; R.A. Fitzgerald & C.D. Preston, in: Preston etal. 2002). The hybrid, R. × levenensis (R. flammula × R. reptans), has been twice recorded in Northern Ireland, from LoughFea in Co Londonderry (VC H40), near Lough Neagh, which happens to be a major arrival site forwaterfowl, which are presumed to transport R. reptans (Gornall 1987).
Flowering
R. flammula flowers abundantly from June to August. It attracts insect visitors and being largely(but not absolutely) self-incompatible, it mainly carries out cross-pollination (Cook & Johnson1968; Gibbs & Gornall 1976). The flowers are described by Jonsell et al. (2001) as,"weakly protandrous" (ie the pollen matures first) and these Nordic authors also report thatselfing is possible when pollination is carried out by raindrops falling into the flower bowl.
Seed production, germination and dormant survival
Published estimates of viable seed production by R. flammula are unknown to the current author,but plants are known to produce between 0-20 flowers, each containing up to around 20 achenes (iesingle-seeded dry fruits), thus potentially each plant may produce up to 400 seeds. In a Canadian study,freshly collected seed kept moist germinated sporadically over a period of six months (Cook &Johnson 1968). In a survey of seed bank data in NW Europe, Thompson et al. (1997) reported 31estimates for R. flammula, of which six regarded it as ephemeral, eleven short-term, and sevenreckoned it produced a long-term seed bank (ie capable of surviving longer than 5 years).
Seedlings produce a rosette of leaves and if submerged they will also produce spreading stolons. InCanadian populations, flowering only occurred after the plant had been exposed to terrestrial conditionsfor an unspecified period of time (Cook & Johnson 1968).
European occurrence
R. flammula subsp. flammula is widespread in Europe except inboth the southernmost and northernmost areas. It is absent from Iceland, and occurs at isolated stationsonly in Greece, Turkey, the Caucasus and W Siberia (Jalas & Suominen 1989, Map 1856; Jonsell etal. 2001). It extends south as far as N Africa and eastward to W Asia (Hultén 1958, Map 147).R. flammula is rare and probably accidentally introduced on the E and W coasts of N America(Jonsell et al. 2001) and it is a definite alien in New Zealand (Preston & Croft 1997).
Medicinal uses
R. flammula was another buttercup used in herbal medicine as a rubifacient for blistering, thatis, for raising blisters. An ancient belief was that by irritating the skin and raising a blister,disease would be drawn out of the body. It was extensively used for this purpose during the bubonicplague in the 16th century, and was also used to treat scabs and running sores (Ui Chonchubair &Mhic Daeid 1995).
Threats
None.
Native, occasional or rare, possibly over-looked and under-recorded. Eurosiberian temperate.
1904; Praeger, R.Ll.; Shean Lough.
There are a total of ten records for this variant in the Fermanagh Flora Database, but they are allpre-1950 and questionable. When editing the Revised Typescript Flora, R.D. Meikle commented thatthis subspecies intergrades with subsp. flammula and that he considered it scarcely worthrecognition even as a variety. Webb & Scannell (Flora of Connemara and the Burren) commentthat Praeger (following E.S. Marshall's example), took far too wide a view of this variant, and that ifit occurs at all in Ireland, it is only in Co Mayo (H26 and H27).
The species account in the New Atlas comments that this segregate is little known, has beenconfused in the past with subsp. flammula and is almost certainly under-recorded (R.A.Fitzgerald, in: Preston et al. 2002). Five of the ten Fermanagh records are Praeger's own finds,one of which is dated 1934 and the other four 1904. In view of Webb & Scannell's comment, we willdiscount them unless vouchers subsequently emerge. Two records were made by R. Mackechnie, a highlyrespected Scottish field botanist, many of whose Irish records are supported by vouchers in EdinburghBotanic Garden (E). The remaining three records were made by Meikle and his co-workers in 1946,1947 and 1948, but again there are no vouchers and, as mentioned above, they clearly did not regard theplant as of much significance.
In the circ*mstances, we feel we can reiterate Hackney's comment in FNEI 3, ie "More fieldinvestigations of these creeping forms are required."
Threats
None.
Native, common, widespread and locally abundant. European southern-temperate, introduced in eastern NAmerica.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and habitat preferences
While R. ficaria is very commonly recorded in over half the area of Fermanagh (see below), westill regard this familiar yellow-flowered, patch- or carpet-forming tuberous perennial asunder-recorded to some extent. The plant is a geophyte, having its resting or over-wintering budsprotected by shallow burial in the soil. The reason we consider R. ficaria may be under-recordedis because the plant is pre-vernal, completing its annual growth and reproductive cycle early in theyear and then quickly dying down and becoming inconspicuous, usually by the middle of June or earlier insome sites (Grime et al. 1988). Dead leaves, however, can still be found and the speciesrecognised in suitable habitats later in the season and there is no month in which we do not have atleast a few records of Lesser Celandine in the flora database. The short growing season of the shootscharacterises R. ficaria as an ephemeroid perennial, the shoots disappearing before the summer(Rogers 1982).
R. ficaria occurs in a wide range of habitats and light environments, from deciduous mixedwoodland where it is most frequent (except in those on the most acid soils), to hedges, roadside banks,verges, stream-sides and damp pastures. It is also found in gardens and can become a persistent weed ofold lawns and flower beds (Taylor & Markham 1978; Sinker et al. 1985). Rapid growth veryearly in the season (ie its prevernal habit) enables the species to avoid the shade and competition ofother plants, particularly in deciduous woodland but also to some extent in its other habitats.
Lesser Celandine tolerates a wide range of soils in these islands, both chemical and physical and,indeed, it appears to show a wider amplitude in this respect in Britain and Ireland when compared withits behaviour in Continental Europe (Hill et al. 1999). Measurements of soil reaction by Taylor& Markham (1978) showed the pH of the substrates occupied by the species ranged from 4.4-6.9 in theBritish Isles and that it frequently occurs on seasonally wet or flooded situations. It does not grow onvery acidic mull soil (pH 3.9 or lower) and is absent from both permanently waterlogged or regularlydroughted dry soils.
The small tubers or bulbils and their roots are buried only shallowly in the top 5-10 cm of soil andlitter so that R. ficaria is confined to soils that are moist in spring, but which may possiblydry out later in the summer months. This dehydration does not matter as far as the plant is concerned,since by the time moisture stress occurs in the substrate, it is likely to be in its resting phase andtherefore completely resistant to drought.
Competitive ability
Given adequate spring moisture, Lesser Celandine is most frequent where soil and vegetation is averagewith respect to fertility, productivity, level of disturbance and extent of bare uncolonised soilsurface. Although it is a poor competitor in well illuminated sites, R. ficaria plants grow andperform better in ecologically open conditions in terms of producing a larger than average shoot, biggerindividual tubers in the soil and more of the plants produce flowers and fruit (Taylor & Markham1978).
The flora associated with Lesser Celandine typically has a relatively low level of diversity and this isparticularly so in damp mixed woodland. Here, under the shade and protection of tree trunks, barebranches and the occasional evergreen or wintergreen leaf and frond, as the population of R.ficaria increases and begins to form extensive patches, species diversity declines further as itbegins to oust other herbs. Thus under damp, shaded growing conditions, Lesser Celandine can sometimesform a more or less dominant, single-species carpet of ground vegetation (Taylor & Markham 1978;Grime et al. 1988).
Tolerance of disturbance
Plants can survive grazing and trampling by farm stock including horses, and mowing on grassy paths,roadside verges, and to a lesser extent in lawns. Due to the early season nature of its active period,annual growth and reproduction of R. ficaria plants is often near completion before these sortsof ecological disturbance reach any great level of intensity (Grime et al. 1988).
Phenology
Growth of R. ficaria is initiated by low temperatures in the autumn, and buds on the buried tubersbegin to develop in December. The long-stalked, deeply notched, cordate foliage leaves of the basalrosette are very variable in shape. They are also shiny, hairless and often blotched with radiating paleor dark markings. The leaves begin to appear above ground in mid-January and they are well deployed byearly February. Seedlings of the diploid, subsp. ficaria, require at least two year’s growthbefore they are capable of flowering (Marsden-Jones 1935).
Flowering
On established plants the flowering stem develops in February: it varies from 3-20 cm long and isspreading and weakly decumbent, branched and typically it bears one or two opposite leaves, similar tothe basal ones but generally more distinctly lobed. Solitary flowers are borne at the end of each branchand flowering reaches a peak in late March and continues into April. The flowers are 20-30 mm indiameter and appear in a range of yellow shades (Grime et al. 1988, 2007).
Compared with other buttercup species, individual flowers of R. ficaria vary much more widely inthe number of their floral parts, including the sexual organs. Early in the season many more stamens andcarpels are produced per flower than in the ones opening later (Lee 1902). Thus the flower is atypicalfor the genus Ranunculus, usually having just three sepals and anything between seven and 13petals instead of the more normal five of each organ characteristic of the genus (Sell 1994). Togetherwith the apparently opposite and unequal stem leaves, these floral part numbers have led sometaxonomists to separate the species off into its own genus and name it Ficaria verna Huds.(Hutchinson 1972).
There is not any concrete evidence suggesting that more open habitats favour flower and seed productionby individual plants, which is the particular strategy of subsp. ficaria, the diploid (but seebelow). Higher light levels do allow more diploid plants to flower, however, and as would be expected,very heavy shade prevents any flowers being initiated. The apparent preference of R. ficaria s.l.for damp habitats may well be related to the conditions required to maximise the formation and survivalof the perennating root tubers which are common to all ploidy levels (Nicholson 1983).
Pollination
However we decide to name this species, the flower structure is completely open and available to allinsect visitors and the presence of nectar attracts short-tongued bees, honey bees, small beetles andflies whose visits achieve pollination. In wet cold weather the flowers remain closed and no insectsvisit. The amount of sun per day is a significant factor affecting early season insect-pollinatedflowers, since whenever the sun becomes obscured there is always a sudden drop in temperature and aconsequent great falling off in the number of insect flower visits (Marsden-Jones 1935). In the absenceof visitors, self-pollination can occur, but the number of seed set is low in such circ*mstances, someflowers proving completely self-sterile and others producing very much reduced numbers of viable seed(Marsden-Jones 1935).
Fruit
The fruit of Lesser Celandine is a cluster of single-seeded achenes that are shed from the receptacle ofthe flower by early June. The number of achenes produced per flower varies with the fertility andsuitability of habitat growing conditions (see below). Between 15 and 35 achenes may form on thereceptacle of the flower, although often many of these potential fruit fail to develop or simply abort(Taylor & Markham 1978; Jonsell et al. 2001). Shortly after the achene clusters on the stemtips ripen and break up, the aerial parts of the plant wilt and quickly decay.
Genetic variation − two subspecies
It has been realised since the mid-1930s that Ranunculus ficaria occurs in the British Isles astwo races with different chromosome numbers, a diploid, subsp. ficaria L. (2n=2x=16) and analmost sexually sterile tetraploid, subsp. bulbilifer Lambinon (2n=4x=32) (Marsden-Jones1935). However, there is a problem distinguishing the diploid and tetraploid forms in the field, sincewhenever the species is flowering and is at its most conspicuous, the main character (the only fieldone), which distinguishes the two subspecies − the aerial bulbils of subsp. bulbilifer − have notyet begun to develop (Gill et al. 1972). In the 2nd edition of Stace's New Flora of theBritish Isles, additional vegetative distinguishing characters of a critical nature areprovided, but this involves microscopic examination to count the chloroplasts in leaf pore guard cells(stomatal guard cells).
Vegetative reproduction − two forms of bulbil
The diploid form of R. ficaria is fully fertile and generally sets seed, but unlike othersituations involving polyploidy where complete fertility attaches to even numbered chromosome sets, forsome reason in this species the tetraploid form has a very low level of sexual fertility. By way ofreproductive compensation, as it were, and undoubtedly encouraging or enabling their survival, thetetraploid plants possess aerial bulbils in addition to the subterranean vegetative bulbils that aretypical of the species. Aerial bulbils are not produced at all on the diploid plant (Nicholson 1983).The small white aerial tubers are produced in the angle between a stem leaf and the shoot bearing it andshould not be confused with the white or buff, fig- or spindle-shaped tubers which are produced justbelow ground level around the basal leaf rosette and which sometimes become exposed if the soil aroundthe plant becomes disturbed. Basal tubers occur in all forms of Lesser Celandine plant irrespective ofchromosome number and are both the perennating organ and a primary means of population increase anddispersal in many situations. The aerial bulbils are derived from swollen axillary adventitious rootsexactly as the basal tubers are, so the structures are distinguishable really only by their position onthe plant (Gill et al. 1972). It is clear from where they are produced on the plant that bothtypes of bulbil originate as modified buds.
Reproductive strategy
The reproductive strategy demonstrated by R. ficaria s.l. is very common in, or evencharacteristic of, Arctic environments where the frequency of polyploid species is much higher than isfound in the floras of temperate regions. It is estimated that 51% of the British and Irish flora is ofpolyploid origin, whereas the comparable figure for Iceland is 72% (Löve & Löve 1974). In Arcticregions, the season of growth in some years is sometimes too short for plants to complete their sexualprocesses and form viable seed. In such circ*mstances, natural selection favours the survival of specieswith a fall-back option of well-developed vegetative reproduction. The growing season of vernal herbs isalso brief in woodland and other deciduous seasonally shaded habitats, growth in pre-vernal and vernalherbs being confined to the light phase of the year, from autumn leaf fall to the expansion of the newleaf canopy the following spring. During at least part of the long dormant or overwinter period, lowtemperatures obviously restricts or prevents active plant processes (Taylor & Markham 1978). Whentemperatures again increase, the active growth phase of the pre-vernal plant is so early in the year,changeable weather conditions make activities like pollination uncertain and some processes, includingseed set, may actually fail. Therefore species such as R. ficaria, Hyacinthoidesnon-scripta (Bluebell) and, in Great Britain (but not in Ireland), Mercurialis perennis(Dog's Mercury), rely very heavily on vegetative reproduction for their increase and survival.
R. ficaria tetraploids have a greatly reduced seed output, but as discussed above, they have twomethods of vegetative propagation − aerial bulbils and basal tubers. In seasonally shaded, relativelyundisturbed habitats where opportunity for seed dispersal is poor, this may well prove the betterarrangement in terms of both plant survival and spread into new ground.
Fermanagh occurrence
In Fermanagh, R. ficaria s.l. is commonly recorded in 297 tetrads, 56.3% of those in the VC. Aspreviously mentioned, it is probably under-recorded to an unknown extent since its growth is earlyseason, pre-vernal and the plant dies down completely by June. The habit of distinguishing these twosubspecies only gained ground slowly in Britain and Ireland and it is regretted that they have not beendistinguished in any of the Fermanagh fieldwork to date.
British and Irish occurrence of the two subspecies
The distribution of subsp. ficaria is described by Stace as occurring, "throughout theBritish Isles", and for subsp. bulbilifer, "almost throughout, but apparently absentfrom Shetland and the Channel Isles" (Stace 1997). The British and Irish occurrence contrastsstrongly with that in Nordic regions, where the tetraploid is the common and widespread subspecies andthe diploid is very rare and confined to Denmark and the southern tip of Norway (Jonsell et al.2001).
In the 6th edition of An Irish Flora (the book relied on for most of the current Fermanagh surveyby Irish recorders), Webb (1977) indicated that a variant (ie subsp. bulbilifer) existed and was,"occasionally found, mainly in the E, and usually as a garden weed". This impression ispartially confirmed by the Flora of County Dublin, where subsp. ficaria was described asoccurring, "very common in woods and hedges, and very rare in gardens". In contrast, the samework described subsp. bulbilifer as being, "very common in gardens, disturbed roadsides, andalso in woods and hedges" (Doogue et al. 1998).
Triploid forms and aerial bulbils
Where the diploid and tetraploid overlap,
triploids result from their crossing (2n=3x=24) (Gill et al. 1972). Triploid plants may flower butthey are totally sterile. They sometimes produce aerial bulbils, but these are smaller and fewer thanthose of the subsp. bulbilifer tetraploid plants.
Comparison of subspecies with respect to light and shade
While R. ficaria s.l. is most commonly found in summer shaded habitats such as woods, scrub andhedgerows, as with the Wood Anemone (Anemone nemorosa) and the Bluebell (Hyacinthoidesnon-scripta), in the west of Britain and Ireland it also grows in open situations in meadows andrough pastures. Clapham et al. (1962) suggested that the diploid, subsp. ficaria, is themore commonly found form in the British Isles, especially in sunny sites and that the tetraploid, subsp.bulbilifer, is more restricted to shady places. In Yorkshire, Nicholson (1983) found that whilethe diploid occurred under the full range of light levels, the tetraploid was associated with moderateto heavy shade and was absent from high ground, including the Wolds and the cliffs at Flamborough Head,and it was also absent from low sandhills in the coastal region. This is a rather surprising claim sincethese areas appear to represent more physically demanding environments, and by analogy with the patternof arctic plants where higher degrees of ploidy and associated vegetative apomixis occur in the moresevere growing conditions. One would therefore expect the tetraploid form with its additional means ofvegetative reproduction to be favoured, rather than excluded from the more demanding environments.
Field observations and experimental measurements both suggest that subsp. ficaria is the morelight tolerant form of the two, rather than suggesting subsp. bulbilifer is more shadetolerant (Nicholson 1983). The observation that R. ficaria is able to persist in more open meadowconditions in the N & W of the British Isles is thus more probably due to the prevalent moist, coolconditions in these regions, rather than suggesting any requirement the subspecies has for shade.
There is not any concrete evidence suggesting that more open habitats favour seed production byindividual plants, which is the particular reproductive strategy of the diploid subsp. ficaria(but see below). However, higher light levels do allow more diploid plants to flower and very heavyshade will prevent flowers from being initiated. The apparent habitat preference of R. ficarias.l. for damp ground may well be related to the conditions required to maximise the formation andsurvival of the perennating root tubers, which are common to all ploidy levels of the plant (Nicholson1983).
Comparative seed production
In measurements of sexual reproductive performance, the mean seed output computed for sun and shadediploid plants was very similar (75 and 71 per plant), giving an overall mean of 73 ± 5.8.(Marsden-Jones 1935). Flowers on tetraploid plants in open conditions tended to be produced earlier thanthose on the diploids, but even if they become pollinated, only a low proportion of their carpels setviable seed. Marsden-Jones (1935) collected a total of just 23 apparently viable achenes from tentetraploid plants, a figure representing around 2% of the ovules produced by these plants. A largeproportion of the pollen of tetraploids is also non-viable, unlike that of the diploid form (Taylor& Markham 1978).
Seed dispersal
After fertilization in late May and early June, the head of achenes ripens and dries somewhat so that itbreaks up at the slightest touch. By this time, or soon after, the other aerial parts of the plant havealready begun to wilt and decay. There does not appear to be any special dispersal mechanism for theseed and presumably it does not travel more than a few centimetres. The fruiting stems do elongate,however, and become more horizontal as they age which helps dispersal, and some seed might be carriedfurther by rain wash, especially over bare compacted soil or along tracks and paths.
Seed dormancy and germination
Seeds of R. ficaria require a period of after-ripening as the embryo is only partly differentiatedwhen they are shed. A delay of four to six months and a period of low temperature is necessary to breakdormancy. Germination begins in the following spring and continues until early summer. Germination ratesare poor for both subspecies, ranging from 15-46% for the diploid and from 7-29% for the much rarerseeds of the tetraploid (Taylor & Markham 1978).
Seedlings have only one cotyledon
Seedlings of R. ficaria, like that of at least some populations of Conopodium majus(Pignut) are highly unusual for a dicotyledonous plant group, in having only one embryo leaf orcotyledon and not two (Taylor & Markham 1978). This fact has been known since the 1850s and was thesubject of much speculation among botanists for many years. The seedling's typically bilobed blade andthe venation in the single cotyledon suggested to some botanists that it might have arisen by the fusionof two cotyledons. However, in a beautiful piece of anatomical and embryological investigation carriedout at Kew Gardens, Metcalfe (1936) clearly and carefully argued and conclusively proved that thecotyledon is in fact a single foliar organ brought about by the suppression of one of the cotyledonsduring embryo development, and not the product of two embryo leaves becoming fused together. Part of theevidence for this is the rare occurrence of R. ficaria seedlings with two bilobed cotyledons, andtrilobed single cotyledons have also been observed. Metcalfe's discovery involved recognising theexistence of a rudimentary second cotyledon which normally fails to develop and thus seedlings onlyproduce a solitary embryo leaf or cotyledon.
A comparison might be made here with the genus Cyclamen (Primulaceae) members of which also haveonly one cotyledon, the other aborting during early development. Another case in point is the genusPeperomia (Piperaceae), where some species are heterocotyledous: one cotyledon being a green,aerial, assimilating organ, while the other is retained inside the seed when it germinates and servesprimarily for the absorption of food reserves from the seed (Metcalfe 1936). Other examples of this typeof odd phenomenon include some species in the genus Claytonia including C. virginica(Stebbins 1974; Mabberley 1997). [N.B. This very interesting paper by Metcalfe has been incorrectlyattributed to Marsden-Jones and given the wrong date by two recent authors, one of whom also got thevolume number wrong! It is correctly quoted, attributed and cited here.]
Tuber and bulbil dispersal
All clones of R. ficaria irrespective of chromosome number possess root tubers (around twelve percluster) and the tetraploid subsp. bulbilifer also has its aerial axillary bulbils. Fragmentationof the basal tuber cluster by soil disturbance is a very efficient means of vegetative propagation. Inaddition, the tetraploid produces up to 24 bulbils per plant that separate off and drop to the ground asthe shoot bearing them dies down. A heavy shower of rain will sometimes wash these aerial bulbils awayfrom the parent plant, heaping them together on the sides of run-off channels when the rain ceases. Insuch places, the quantity of bulbils aggregated is often so noticeable that the idea arose that they hadfallen from heaven with the rain, and the myth developed of 'potato rain', or a 'rain of wheat' (Grieve1931; Taylor & Markham 1978).
Tubers and bulbils provide the most rapid means of increase and spread for the species (and subspecies),as they are readily detached and may be spread by disturbance of any kind, but especially by digging,ploughing or mowing. Bulbils being considerably larger than seed, can regenerate the plant quicker andsome may even flower in their first season of growth (Marsden-Jones 1935).
Origin of polyploidy in R. ficaria: In view of the quite different ecologicalbehaviour and reproductive strategies of the diploid and tetraploid forms, it appears very likely thatsubsp. bulbilifer is an ancient autotetraploid, ie it is the result of mutant chromosomedoubling, followed by a long period of divergence of the two ploidy levels (Nicholson 1983). Geneticisolation barriers have not yet been established, however, since in Britain and Ireland (thoughapparently not in Nordic areas (Jonsell et al. 2001)), triploids are formed by interbreedingwhere the two subspecies overlap (Gill et al. 1972). Thus Lesser Celandine is a very good exampleof species evolution in action, and indeed in other parts of Europe several additional subspecies arerecognised (Sell 1994; Jonsell et al. 2001).
Toxicity
R. ficaria plants contain low levels of the toxin irritant protoanemonin, an unstable compoundderived from the glycoside ranunculin. The concentration of this toxin increases during growth and it isat its highest during the flowering process. Unlike many other buttercups it has not been known topoison grazing stock animals (Cooper & Johnson 1998). Despite the presence of the toxin, youngleaves of R. ficaria often have holes eaten in them in early February, and we have seen entireblades removed at this time of year, presumably eaten by slugs before the protoanemonin levels become afunctional deterrent.
European occurrence
External to the British Isles, the diploid R. ficaria subsp.ficaria is restricted to W Europe, north to SW Norway and Denmark, where it is extremely rare,and south to The Iberian peninsula and the W Mediterranean region (Jalas & Suominen 1989, Map 1836).The bulbil-bearing tetraploid, R. ficaria subsp. bulbilifer isalso confined to Europe, but it is very much more widespread in C and S parts of the continent than thediploid form (Jalas & Suominen 1989, Map 1835; Taylor & Markham 1978).
Fossil record
There is no fossil record of R. ficaria s.l. as the pollen and possible macrofossils areindistinguishable from other Ranunculus species (Taylor & Markham 1978).
Names
The specific epithet 'ficaria' is derived from the Latin name 'Ficus', the fig, and it means 'small fig'.This is a reference to the supposed fig-like shape of the root tubers (Gledhill 1985).
Two of the numerous English common names are 'Figwort' and 'Pilewort', 'fig' and 'pile' being alternativenames for haemorrhoids, which the root tubers resembled. The similarity in appearance led herbalists bythe 'Doctrine of Signs' to believe that a plant organ that looked like a haemorrhoid could be used tocure the complaint (Grieve 1931; Grigson 1987). Certainly the herb contains an astringent, but whileVickery (1995) includes a folklore recipe for a skin cleanser, it should be left well alone since thesap can cause blisters on or in the body (Grieve 1931).
Other English common names such as 'Brighteye', 'Butter and Cheese', 'Golden Stars' and 'Goldy Knog',refer to the shiny, yellow petal colour. Grigson (1987) also suggests the root tubers were regarded asreminiscent of a cow's udders, and hence the milk and butter references in some common names. An earlybotanical name for the species was Chelidonium minus, which translates as 'Lesser Celandine'.This erroneously linked the plant to the unrelated Chelidonium majus (Greater Celandine). Thename 'Chelidonium' was derived from the Greek 'chelidon', a swallow (the bird), supposedly because ofcoincidence between the time of the plant flowering and the arrival of the migrant bird.
Threats
None.
Native, occasional or locally frequent. Suboceanic southern-temperate, also rarely present on the easternseaboard of N America, but possibly introduced.
1884; Barrington, R.M.; eastern shore of Lower Lough Erne, near Enniskillen.
Throughout the year.
Fermanagh occurrence
In Fermanagh, Ivy-leaved Crowfoot has been recorded in 89 tetrads, 16.9% of those in the VC although, asthe distribution map indicates, it is widely scattered throughout the county; ten of the tetrads containpre-1976 records only, which could be argued as indicating some degree of local decline during the pasthalf-century.
Growth form and habitat preferences
R. hederaceus regularly occurs in Fermanagh as something of an invading, ephemeral, companionspecies of a number of quite different submerged floating or emergent vegetation communities. Thehabitats where this happens include the muddy banks of slow-flowing rivers, streams (together withsprings and upland flushes) and drainage ditches (especially in late autumn after ditch cleaning) andbeside relatively still waters in smaller lakes and ponds and on the sheltered shores of backwater baysof the larger loughs in the VC. It does not occur, or is only occasional and sparsely developed, wherewater stagnates. Typically, R. hederaceus requires sufficient seepage of ground water to createat least a slight fluctuation in levels and a consequence of this will be an additional in-flow ofoxygen and mineral nutrients.
R. hederaceus was also once found in muddy ground by temporary pools in an old quarry besideKeenaghan Lough and it also appeared on an urban waste tip in Enniskillen, which mirrors Segal'sexperience in Holland (Segal 1967). However, the most predictable habitat in Fermanagh of this small,prostrate annual or short-lived perennial appears to be by field gateways and along wet, muddy trackswhere water lies in shallow quagmire pools and puddles in mud that has been trampled, poached anddung-enriched by cattle. We find it both where the parent rock of the soil is acidic or of a calcareousnature, and it clearly tolerates a rather wide range of pH and appears indifferent to lime (Cook 1966b).
The authors of the critical Flora Nordica regard R. hederaceus as a calcifuge throughouttheir northern European region (Jonsell et al. 2001). In Great Britain, Hill et al.(1999), summarising the preferred environmental growing conditions of species, gave R. hederaceusan indicator value of '5' on a nine point scale, indicating that while it mainly occupiesmoderately acid soils, occasionally it ranges wider and it can occur on both very acid and moreneutral to basic (alkaline) substrates. In the latter situation, due to our very wet climate, a leachedcalcareous soil usually develops. This generally becomes overlain with a thin, acid, peat horizon, whichforms the substrate over which Ivy-leaved Crowfoot spreads and shallowly roots at its nodes.
Cook (1966b) made a significant point when he speculated that if R. hederaceus and theclosely related species R. omiophyllus Ten. (Round-leaved Crowfoot) are not directly competing(the latter does not occur anywhere in N Ireland), they may exhibit wider ecological amplitudes thanwhen they overlap with one another. The habitat description of R. hederaceus is rather similar tothat of R. sceleratus (Celery-leaved Buttercup), but the latter is more frequently found bylakes, or in somewhat less disturbed, more nitrogen-enriched, perhaps slightly better drained mud whichis more liable to drying out temporarily than that of R. hederaceus. Also, R. sceleratusnever takes on a truly aquatic 'submerged' – or more realistically – 'floating' existence, in the waythat Ivy-leaved Crowfoot is capable of doing in water even as shallow as 5 cm!
Apart from its freshwater habitats, R. hederaceus can also be found in sheltered coastal parts ofboth Britain and Ireland, usually on the upper edges of salt-marshes (Preston & Croft 1997).
Variability and identification
R. hederaceus is classified within the Subgenus Batrachium of the genus Ranunculus.Most species of this subgenus are notoriously variable, frequently hybridize and backcross, and thus areoften very difficult to identify (Cook 1966a; Holmes 1979; S.D. Webster In: Rich and Jermy 1998). R.hederaceus and its close relative R. omiophyllus (Round-leaved Crowfoot) (the latterapparently absent from Northern Ireland), differ from all the other members of the subgenus by beingessentially semi-terrestrial and by invariably having un-dissected, laminar, floating leaves. Unlike allother water-crowfoots in the subgenus, these two species do not possess thread-like submerged leaves(Cook 1963, 1966b). Thus, in our Fermanagh survey area, Ivy-leaved Crowfoot is very easily recognised,despite the species being remarkably variable in form in response to environmental changes, ie it has avery plastic phenotype with respect to many characters. This high level of variability is quitecharacteristic of emergent aquatic species (Segal 1967; Cook 1966a, b).
The creeping or surface floating habit of the plant, its small white flowers, and ivy-shaped leaves withdark markings following the veins are all very distinctive identification features, and the typicalplant species associates of Ivy-leaved Crowfoot include Montia fontana (Blinks),Callitriche spp. (Water-starworts) and Stellaria uliginosa (Bog Stitchwort).
Summary of environmental requirements
From the above, it is clear that R. hederaceus has the wide ecological tolerances that one wouldexpect of a species inhabiting situations where slight but significant fluctuating water levels are thenorm, and its requirements can be satisfied in a diverse range of habitats encompassing a mosaic ofvegetation communities, moist, wet, semi-aquatic and shallow aquatic, lowland and upland, coastal andinland. The most constant requirements of the species are for high levels of illumination, wet,moderately acid, waterlogged soil, or very shallow water only a few centimetres deep, of medium(mesotrophic) fertility, or richer more productive eutrophic levels, plus shelter from strong watercurrents.
Temperature limits and distribution
An equable, fairly low water temperature throughout the year is recognised by ecologists and plantgeographers as another very important environmental factor controlling the growth and occurrence ofR. hederaceus. Temperature is sufficiently significant to actively govern the wider distributionof the species, confining it to the truly oceanic or Atlantic region of W Europe. A summer maximum ofaround 16C, and mild winter temperatures with little in the way of severe frost, characterises therequired regime (Segal 1967).
In his study in Holland, Segal noticed that R. hederaceus tends to occur in specific landscapesituations where small scale water bodies lie on the junction between hilly, acidic, infertile(oligotrophic), non-calcareous soils and much more fertile, mineral-rich (mesotrophic to eutrophic)conditions, on lower ground at the base of slopes. In the habitat gradients that occur when two verydifferent ecological environments of this nature meet, and especially where the zone of contact is keptopen by some form of disruption, either physical (eg trampling and grazing), or chemical (eg a moderatelevel of pollution, including manuring and other forms of farm effluent run-off), or both of these, thenR. hederaceus and other interesting and quite scarce species such as Catabrosa aquatica(Whorl-grass) and Veronica catenata (Pink Water-speedwell) appear to find growing conditions totheir liking. On the other hand, in excessively enriched sites containing high levels of nitrogen andphosphorus, ground that tends to be rapidly overgrown by algae and by Lemna species (often L.gibba (Fat Duckweed)), R. hederaceus cannot compete for very long under such conditionsand becomes ousted.
Flowering and pollination
Ivy-leaved Crowfoot flowers early and for a prolonged period, often stretching from April to August.However, flowering is frequently curtailed by the habitat becoming overgrown later in the summer, orsometimes by it drying out. The small white flowers produce nectar, are sweet scented and areprotogynous (ie the female parts develop first, followed by the anthers). This difference in sexualtiming in the individual flower is generally considered an adaptation favouring or enablingcross-fertilisation between flowers, but in reality the blooms are highly self-compatible and theyappear to habitually inbreed. Indeed, self-fertilisation often takes place at the unopened bud stage,making it obligatory. Despite this sexual behaviour, the timing of flower opening depends on theprevailing weather and some level of opportunity for cross-pollination does exist in other flowers.However, observation also indicates that the flowers attract few insect visitors (Cook 1966b).
Occasionally R. hederaceus flowers are produced underwater. When this occurs, a gas bubble isformed within the bud, allowing pollination to proceed as normal (Cook 1966b). Since R.hederaceus is seldom submerged and, if so, then usually only for short periods, submergedpollination is not likely to occur very often.
Fruiting and dispersal
As the fruit develops the flower stalk bends away from the light, forcing the developing achenes into themud (Cook 1966b). This is another example of a negatively phototrophic movement by a fruit stalk (ie agrowth movement away from the direction of light), similar to that of the fruit stalk of Cymbalariamuralis (Ivy-leaved Toadflax). It appears odd that both species which commonly show this unusualphysiological feature, should possess ivy-shaped leaves!
Dispersal of the seeds (achenes – single-seeded dry fruits) is most probably achieved in mud byattachment to animals or vehicles. There might possibly be some degree of water dispersal too, if a fastcurrent were to develop in the locality and dislodge seed shed onto the soil surface around the plant.
Germination and growth form
Seed germination is reported by Cook (1966b) to be very irregular if the achenes are kept wet – which onemight imagine would be the normal condition of the habitat. On the other hand, if the mud dries outafter the seeds ripen and the seeds themselves become dried, then Cook found that when they arere-wetted, the seeds gave nearly 100% germination. He also showed that, depending on the local regime ofwater levels and competing species, R. hederaceus (and also R. omiophyllus (Round-leavedCrowfoot)), can behave either as winter or spring annuals, or individual plants may persist andreproduce on multiple occasions for periods up to six years, thus achieving perennial status.
Overwintering and factors affecting survival
During the winter months, the resting plant survives as a small, tight, rosette of leaves. In this state,it is very resistant to freezing, desiccation and shade. However, in summer when individual plantsdevelop their normal spreading habit, they becomes very susceptible both to the three mentioned physicalfactors and to pressure from taller, more aggressive competing plant species (Cook 1966b).
Ignorance regarding seed output, reproductive strategy and longevity in the soil seed bank
In common with many other species in the flora of Britain and Ireland, after careful searching, we do notappear to have any figures for typical plant seed output. Nor do we know anything regarding the relativesignificance of seed versus vegetative reproduction for perennial populations of R. hederaceus,nor even if a persistent soil seed bank exists (Thompson et al. 1997).
British and Irish occurrence
In Britain, R. hederaceus has quite a distinct northern and western distribution, partiallycreated by the destruction of suitable habitats in much of the SE due to development, drainage and ashift from livestock to arable farming. The change to arable farming means that existing wetlands are nolonger subject to the trampling and disturbance of grazing animals that previously opened the ground tocolonisation by Ivy-leaved Crowfoot (Preston & Croft 1997; Preston et al. 2002). In Ireland,R. hederaceus, being essentially a lowland wetland species, has a somewhat scattered, almostdisjunct distribution, featuring a heavy concentration of records in northern counties, but it is morecoastal in the Republic and appears quite scarce in the Midlands.
European and world occurrence
R. hederaceus occurs throughout the Atlantic region of W Europe from Portugal to the S tip ofSweden (Jalas & Suominen 1989, Map 1884). It was previously present further N up the coast of Norwayat Tronheim and considered native there by some, but it became extinct there in 1950 (Jonsell etal. 2001). Cook (1983) regarded the species as endemic to Europe (and indeed one of only a fewancient 'palaeoendemics'), and he felt that it has declined throughout Europe, probably due to changesin agricultural practices over the past 50 or so years which involved the destruction of wetlandhabitats. Preston reminds us that the species was losing sites in SE England before 1900 for the samereason and also because of urban expansion (Preston et al. 2002).
The species is also present in disjunct locations in eastern N America where it has been known since 1821– in Newfoundland and the Chesapeake Bay region (Hultén 1958, Map 137). The general shape of thedistribution of R. hederaceus in N America and its history discussed in Cook (1983, 1985),suggests that it was introduced from Europe. Against this view there is the existence of two fernspecies, Schizaea pusilla Pursh and Woodwardia areolata (L.) Moore, which have verysimilar N America distributions to R. hederaceus and they are certainly not Europeanintroductions.
Names
The Latin specific epithet 'hederaceus' simply translates as 'ivy-leaved' (Gilbert-Carter 1964) and theplant, having no herbal or folk-lore usages, does not appear to have any English common names other thanthe one chosen by the committee of the Botanical Society of Britain and Ireland (Dony et al.1974).
Threats
None. The species may be benefitting, like R. sceleratus, from the current general eutrophicationof water bodies, and considerable physical disturbance helps it colonise by keeping the habitat open.
Native, rare, but easily over-looked and therefore possibly under-recorded. Circumpolar wide-boreal.
July 1946; MCM & D; Lough Eyes, 3 km NE of Lisbellaw.
May to September.
Growth form and habitat preferences
This aquatic species, which only produces dissected capillary leaves, is a spreading to erect perennialwhen growing in permanent water, but it becomes greatly dwarfed and behaves as a small annual if itsmuddy substrate dries out during the summer. It is most frequently found in still or almost stillshallow water less than 50 cm deep in small lakes or ponds and water-filled holes in disused quarries.
Thread-leaved Water-crowfoot roots mainly in bare clayey bottoms and it can tolerate a wide range ofwater chemistry, from oligotrophic to eutrophic, lime-rich or otherwise. It chiefly frequentsmesotrophic to eutrophic conditions in lowland water bodies (C.D. Preston, in: Preston et al.2002). R. trichophyllus also occurs in brackish waters around the coastline of Great Britain andIreland (particularly in Scotland and Wales), eg in dune-slacks and around sheltered estuarine bays(Preston & Croft 1997).
Like its close relatives, R. aquatilis (Common Water-crowfoot) and R. peltatus (PondWater-crowfoot), this species tolerates disturbance such as trampling by stock animals, human activitiesand/or a substrate which seasonally dries out. It can therefore behave as a pioneer colonist of bare mudaround temporary water bodies, including recently dug or cleaned pools and drains (Cook 1966a, p. 134).Later in the season, as the vegetation in such muddy habitats matures, R. trichophyllus and itspioneer associates tend to be ousted by increased shade and other forms of competition produced byrhizomatous aquatic plants such as Potamogeton species (Pondweeds), Hippuris vulgaris(Mare's-tail), or by any of the numerous vigorous emergent plants of swampy ground.
Fermanagh occurrence
In Fermanagh, there were only two records for R. trichophyllus prior to 1980 and there have beenjust eight additional finds of the species since then. There are records from nine Fermanagh tetrads,seven with post-1975 dates. The species is undoubtedly under-recorded here, as is also the caseelsewhere in Britain and Ireland. In the New Atlas, Preston suggests that part of the reason forthe paucity of records is the early flowering season of the species, and this, together with thewell-known discouraging taxonomic difficulties of the Batrachian group of Ranunculus, is veryprobably responsible for the dearth of observations (Preston & Croft 1997; New Atlas).
Most of the Fermanagh records for this species will have been determined on the basis of the flower sizebeing less than 12 mm in diameter, with the petals being non-contiguous. Non-flowering plants in thisgroup are almost impossible to identify, contributing to their all being under-recorded. Regrettably,none of the ten Fermanagh records of R. trichophyllus is supported by a voucher.
As the tetrad map indicates, most of the records are confined to the Lough Erne basin, with just fouroutlying stations, one at Green Lough turlough and three scattered further east. Although morefrequently found by smaller water bodies, it does also occur in well-sheltered areas around largerlakes, especially where shallow backwater bays are protected by islands close to shore. Local examplesof the latter habitat are the West Point islands and Lowery Island, both at the W end of Lower LoughErne.
Flowering and pollination
Thread-leaved Water-crowfoot is the first Batrachian Ranunculus species to flower in Britain andIreland, which it does from April through to July. The flowers are habitually self-pollinated, thisoften (but not always) taking place in the bud (ie the flowers are cleistogamous). As a result of this,seed set is perfect (100%) and completely assured (Cook 1966a, p. 183; Hong 1991, p. 53).
Seed output
Statistics of mean seed production per plant do not appear to be available, or have not been located.
Germination
The seeds can germinate in wet conditions at any time of year, but in winter seedlings require theprotection of submerged conditions. If frozen under terrestrial conditions they die, but they cansurvive being frozen in ice for several months (Cook 1966a).
Survival in the soil seed bank
Seed longevity is unknown, but it is probably similar to that of R. peltatus which is consideredto be only short-term persistent, ie the seeds survive in soil for periods ranging between one and fiveyears (Thompson et al. 1997).
Vegetative reproduction
Like most other aquatic macrophytes, R. trichophyllus can reproduce vegetatively from plantfragments after disturbance or uprooting of an original established individual. An experimental study inFrance examined the fate of five different types of plant fragments of R. trichophyllus and fiveother wetland species. The plant parts used ranged from roots, to stems with or without nodes, to apicalbuds. After ten weeks greenhouse incubation floating in pans of shallow water (7-8 cm deep) andsediment, the R. trichophyllus results showed that within two or three weeks, between 30% and 50%of the aerial parts rapidly rooted and established into the sediment, while the remainder of thefragments of the species died off without showing any root development. None of the fragments of R.trichophyllus developed new buds before they rooted (Barrat-Segretain et al. 1998). TheFrench workers recognised that the six aquatic macrophyte species they studied reacted to disturbance byexhibiting one of two survival tactics: 1. the fragments either develop roots and establish rapidly intothe sediment (eg R. trichophyllus and Sparganium emersum (Unbranched Bur-reed), or 2. thefragments develop many new propagules that may be dispersed, but they fail to establish within the 10weeks of the experiment (eg Hippuris vulgaris (Mare's-tail) and Elodea canadensis(Canadian Waterweed)) (Barrat-Segretain et al. 1998).
The species examined in the French study appeared to present a trade-off between the plant's regenerationand colonisation abilities: each species practiced both tactics, but high levels of regeneration impliedlow colonisation ability and vice versa (Barrat-Segretain et al. 1998). One of the mostinteresting and unexpected conclusions of this work was that the different survival tactics involvingvegetative reproduction were not related to the morphological types of the species and thus they wereunpredictable. At the same time, it appears that species possessing narrow ecological amplitude promotedispersion of propagules, thus increasing opportunities of finding suitable habitats, whereas otherspecies (including R. trichophyllus), that are more tolerant of varying ecological conditions,try to anchor themselves as soon as possible at the nearest sites to the original population(Barrat-Segretain et al. 1998).
A subsequent similar experimental study showed that some aquatic species behaved quite differently withrespect to these two vegetative strategies in the spring and in the autumn, but this did not apply toeither R. trichophyllus or Sparganium emersum (Unbranched Bur-reed) (Barrat-Segretain& Bornette 2000).
British and Irish occurrence
In Ireland, R. trichophyllus is occasional in the C & SE, and rare and widely scatteredelsewhere including NI (Webb et al. 1996). In Britain, it is much more frequent in the SE ofEngland, below the line from Hull to Bristol, but it is widely scattered in the lowlands elsewhere,becoming distinctly coastal in the W and throughout Scotland (Preston & Croft 1997; Preston etal. 2002). Although the New Atlas contains a high proportion of pre-1970 records, theauthors consider that this does not indicate any real decline in what is a difficult enough species toidentify.
European occurrence
Beyond the British Isles, R. trichophyllus is probably the most widespread Batrachian species inthe world. The only reason for qualifying this with the word 'probably' is, of course, the great amountof variation and plasticity within the group, plus the infamous taxonomic difficulty of the subgenuswhich creates problems in terms of delimiting the species and their field recognition. For instance,when examining material of R. aquatilis (Common Water-crowfoot) and R. trichophyllus in SSweden, Hong (1991) found that all the distinguishing characters for these two species, includingnectary shape, were variable and readily modified. Also, sexual crosses between typical forms of the twospecies were found to be fully fertile, a feature undoubtedly allied to polyploidy within the complex.In S Sweden, both of these species are regarded as hexaploid taxa, whereas in his taxonomic review Cook(1966a) dealt entirely with tetraploid R. trichophyllus and hexaploid R. aquatilismaterial from C and W Europe. Reflecting this degree of variation, in Flora Nordica 2(Jonsell et al. 2001), R. trichophyllus is downgraded from a species to a variety of R.aquatilis, referred to as var. diffusus With.
Apart from these problems of polyploidy and status change, R. trichophyllus is regarded aswidespread throughout Europe, especially frequent in western regions and thinning eastwards. Unlike manyother members of the subgenus, it is even frequent in the Mediterranean basin and is present on many ofthe smaller islands, for example, Majorca, Malta, Cephalonia, Zante and Crete (Jalas & Suominen1989, Map 1895).
World occurrence
Beyond Europe, it is a very widespread species being both Arctic and circumboreal stretching across NAsia, the Himalaya, N and W China, Japan, SE Australia, Tasmania and central areas of N America (Cook1966a; Hultén 1974, Map 80; Hultén & Fries 1986, Map 874). R. aquatilis is the only otherwater-crowfoot that approaches this extent of world distribution, but it is not as well represented asR. trichophyllus in both northern regions and in the Mediterranean region.
Dispersal of aquatic species
We have to ask ourselves why should R. trichophyllus be so much more widely distributed than allthe other aquatic Ranunculus species? It is listed by Cook (1985, Table 1), along withPotamogeton pectinatus (Fennel Pondweed) and 24 other aquatic species widely distributed in theworld, where their native ranges are impossible to distinguish from their areas of introduction. Prof.Cook points out that it is surprisingly difficult to get recent, accurate information on thedistribution of aquatics and it is even more difficult to discover or decide where a particular speciesis native rather than introduced. In connection with dispersal ability, Cook also points out thataquatic plants in general lack very light wind-carried diaspores or propagules and they also lack largeseawater-resistant ones; they are thus ill-equipped for long-distance dispersal. The widespread plants,listed in his Table 1 of unknown native status, have probably spread more or less naturally, or at leastthey dispersed before botanists were around keeping records!
The majority of aquatic plant species have become mobile as man has increased his own globalmobility (Cook 1985). Reviewing the distribution of 172 aquatics, Cook found that marked imbalancesexisted between Old and New World regions, in terms of donor and recipient areas with respect tointroduced species. He concluded that the problems of successful establishment in new lands are quiteobviously very great and that it is extremely naive to regard the aquatic environment as globallyuniform, thus allowing or expecting aquatic macrophytes to have a wide geographical range. It is knownthat some aquatic macrophyte genera (eg Brasenia and Dulichium), became extinct in Europebefore or during the last Ice Age, but they persisted in N America. These plants have not re-establishedin Europe despite being in garden cultivation there, an observation, which lends strength to theargument that ecological niches in the aquatic environment are more complex than we commonly assume.Cook argues that we should not combine the terms 'introduced aquatic' and 'weed', pointing outthat weeds (ie undesirable plants) essentially tend to flourish in sites that are disturbed by man."Usually it is the nature of the disturbance that is more important for the establishment of theweed than its [powers of] mobility." (Cook 1985).
It is very clear that disturbance by man and his animals, and pioneer colonisation of bare muddy groundare two very important ecological features influencing the occurrence of many BatrachianRanunculus forms. In many cases, these species could not successfully reproduce sexually andestablish seedlings without the habitats they occupy temporarily drying out. Following Cook's advice,and maintaining a healthy scepticism regarding the real distribution of many difficult-to-identifyBatrachian Ranunculi, it appears to follows that his remarks regarding introduced aquatics must applyquite closely to R. trichophyllus, since it does not appear any better equipped for long-distancejump dispersal than R. peltatus (Pond Water-crowfoot), R. penicillatus (StreamWater-crowfoot) or, indeed, any of the other seed- and vegetative propagule-producing members of thiscomplex subgenus.
Names
The Latin specific epithet 'trichophyllus' is derived from the two Greek words for 'hair' and 'leaves',and thus translates as, "having leaves divided into hair-like segments" (Gilbert-Carter 1964).The English common name, 'Thread-leaved Water-crowfoot', is a straightforward invented 'book name',following the same line of thought.
Threats
None.
Native, rare or occasional, but difficult to identify and therefore over-looked and under-recorded.
1988-91; NI Lakes Survey; Lakeview Lough, 1 km N of Drummully.
June and August.
Since we have relatively few records of Batrachian Ranunculi for Fermanagh, these 20 undeterminedwater-crowfoot records from 17 tetrads, which need to be followed up, are simply included for the sakeof completeness.
The first seven records were made by the NI Lakes Survey between 1988 and 1991. In addition to the firstrecord listed above, the site details of their other finds are: Lough Ora, 1 km SSW of Ora More,Ballintempo Forest; Green Lough turlough, near Fardrum; and Fardrum Lough turlough. The remaining threerecords made by the Lake Survey were all made on unimproved neutral to calcareous grasslands on theshores of Lower Lough Erne shore as follows: E of Castle Hume, 11 June 1991; Inner shore, DulrushPeninsula, 2 August 1991; mainland shore opposite Dulrush Peninsula, 4 September 1991.
All the subsequent 13 records were made by RHN either alone or accompanied by HJN. The sites and datesare: Lough Skale, Clogtogle Td, 5 June 2006; Lackboy, S shore of Boa Island, 1 July 2010; Mill Loughnear Ballinaleck, 13 October 2010; Castle Archdale, 31 October 2010; all the remainder were around theshores of Lower Lough Erne in November and December 2010: mainland shore NW of Portinode Bridge; jettynear car park, Castle Archdale; Hill's Island; Rossigh Bay; riverfoot, just N of Rossclare jetty; nearRosscor Viaduct; mouth of Garvary River; Carrickreagh Bay; N shore of Tully Point, Sand Bay.
Native, rare. European temperate.
1884; Barrington, R.M.; Devenish Island, Lower Lough Erne.
May to August.
Growth form and preferred habitats
An annual or short-lived perennial that grows in shallow water in marshes, ponds and ditches and at theedges of slow-moving streams and margins of sheltered lakes. It prefers lowland sites with water that iseutrophic, somewhat base-rich and subject to a moderate degree of disturbance that helps keep thehabitat open. These conditions can occur where cattle graze lakeshore margins and water meadows, orpastures around muddy ponds.
Fermanagh occurrence
As emphasised in much of the literature, the heterophyllous and large-flowered water-crowfoot species,R. aquatilis, R. peltatus (Pond Water-crowfoot) and R. penicillatussubsp. penicillatus (Stream Water-crowfoot) are not easily distinguished. This fact should beborne in mind when studying distribution maps of the three species, either ours or anyone else's forthat matter matter (Cook 1966a; Preston & Croft 1997; Preston et al. 2002).
Undoubtedly identification difficulty leads to under-recording, and we have only eleven records for R.aquatilis. Since a degree of reasonable doubt hangs over them, we have combined them as s.l. Wedo not mean to suggest by this that we are considering R. aquatilis s.l. as the old pre-Cook(1966a) species aggregate. However, it is probable that most of the Fermanagh identifications weredetermined using An Irish Flora (1977) as a field guide. This text used only two features todiscriminate this species: flowers between 12 and 17 mm in diameter and floating leaves present insummer. As Cook (1966a) and Webster (1991) have shown, flower size in this species group variesconsiderably with environmental conditions, and we now appreciate that it is not a reliabledistinguishing species character.
Keeping in mind the mentioned reservations, as the tetrad map indicates, the Fermanagh records of R.aquatilis are thinly and widely scattered across ten squares in lowland Fermanagh. Only onerecord for R. aquatilis pre-dates 1975, that of Barrington (1884), from a wet ditch on DevenishIsland. Soon after this record was reported (as R. heterophyllus Fries), Tetley wrote into hiscopy of Irish Topographical Botany in pencil, "I have no doubt this should be R.peltatus". While only two of the records (Barrington's and one by the NI Lakes Survey) aresupported by vouchers in BEL, the eleven records involve at least seven determiners, increasingthe probability that some of them are correctly identified. The habitats listed are typical for thisspecies, ie shallow, eutrophic, somewhat base-rich water in lakes, turloughs, rivers, streams andditches. Further investigation of Batrachian Ranunculi in Fermanagh is very obviously required.
British and Irish occurrence
The distribution map in Preston & Croft (1997) took a more conservative approach to R.aquatilis records than did the editors of the New Atlas (Preston et al. 2002).Recognising that uncertainty still clings to the identification and status of the taxon, the earlierapproach is undoubtedly the wiser one. This shows modern records of R. aquatilis extremely thinlyscattered throughout Ireland, while the Great Britain distribution displays the plant as more frequentand there are four or more discrete areas of concentration in England and Wales. At the same time, R.aquatilis becomes much rarer and almost entirely coastal in Scotland (Preston & Croft 1997).
European occurrence
R. aquatilis is widespread in Europe from S Scandinavia southwards to the Mediterranean, butwhile it reaches Sicily, it does not penetrate far into the Iberian Peninsula, or into Greece (Jalas& Suominen 1989, Map 1894).
World occurrence
R. aquatilis s.l. is also found in N Africa, and in the boreal zone it extends across Asia exceptin the south, plus W and C North America where it also occurs in the alpine zone of the Middle Rockiesand southwards into California and New Mexico (Scott 1995, p. 657). It also extends into the southernhemisphere, being present in both western South America (the Andes between 2,700-4,000 m) and SEAustralia (Cook 1966a; Jonsell et al. 2001). The world distribution of R. aquatilis isprobably second only to that of the closely related species R. trichophyllus (Thread-leavedWater-crowfoot). However, one has to remember that here we are dealing with species which are difficultto identify, and since they are plants which live in disturbed muddy habitats, they are undoubtedly andunwittingly spread by man, so that we do not have much of an idea as to their native ranges (Cook 1985).
Name
The Latin specific epithet 'aquatilis', means 'growing in water' (Gilbert-Carter 1964).
Threats
Shading from competitors and overgrowth by algae and/or other macrophytes due to excessive culturaleutrophication.
Native, occasional or rare. European wide-temperate.
1860; Smith, T.O.; Lough Eyes, 3 km NE of Lisbellaw.
May to October.
Growth form and preferred habitats
Like several other species and forms within the Ranunculus subgenus Batrachium, inpermanently wet habitats R. peltatus is entirely perennial provided it is not overgrown or shadedby larger or more robust species. When growing in water, the plant normally produces both floating lobedlaminar leaves and finely-dissected submerged capillary ones. However, if the site R. peltatusoccupies dries out in summer, the species behaves as an annual and becomes both dwarf and prostrate, oreven somewhat tufted in appearance (Cook 1966a; Webster 1988). As far as we know the annual form of theplant has never been reported as such in Fermanagh, but at least in grazed ground in and aroundturloughs (ie vanishing lakes rarely occurring in limestone districts), it could very well be the normalmode of the species.
In Ireland, R. peltatus is most typically found in more or less unshaded stretches of slow tomoderately-flowing streams and rivers in calcareous districts. However, it can also occur in similarflow rates in much more acidic situations over peat or clay substrata and it appears to occupy a ratherbroad range of habitats in terms of nutrient status.
It should be noted that despite the species epithet 'peltatus', none of the organs of the plant,including the floating leaves, are ever peltate in form. The botanical name is misleading and a definitemisnomer, arising as an 18th century error made by Prof. Carl Linnaeus himself! Further remarks on thistopic are made below in the section 'Names'.
Occurrence in Fermanagh
In Fermanagh, this perennial or sometimes annual aquatic has been recorded in a total of just 14 lowlandtetrads, 2.7% of those in the VC. It grows in shallow water in calcareous rivers, streams, lakes andponds. The main site in Fermanagh for R. peltatus appears to be the Colebrooke River, where bothR. peltatus and R. penicillatus subsp. penicillatus(Stream Water-crowfoot) occur. R. peltatus is also found at the Green Lough turloughs (ielimestone lakes or ponds that drain vertically – often referred to as 'vanishing lakes', since they dryout quite regularly), and it is found in an additional five or six rather scattered locations in the VC.
Although locally R. circinatus (Fan-leaved Water-crowfoot) appears in four more tetrads than R.peltatus, the latter is probably the commonest aquatic species of Ranunculus subgenusBatrachium in the VC, provided that is, R. hederaceus (Ivy-leaved Crowfoot) is discounted,since at least in our survey the latter is more often a terrestrial rather than an aquatic species.
Variation and identification
According to An Irish Flora (1977), specimens of R. peltatus with semi-circular to circularfloating leaves produced in spring and summer (ie in addition to the constantly present finely divided,submerged, capillary leaves) and with large flowers (at least 18 mm across), should be reasonably easyto recognise as this species. In reality, however, there is considerable environmentally and seasonallyinduced variation in R. peltatus and it overlaps with R. penicillatus subsp. penicillatus (Stream Water-crowfoot), both in terms ofmorphological characteristics and ecology. Thus these two forms, in particular, are very readilyconfused (Cook 1966a; Holmes 1979; Webster 1988, 1991; Rich & Jermy 1998, p. 60).
Prior to Cook's 1966 monograph on Batrachian Ranunculi and for many years afterwards, due in part to theinadequacies of the Irish field Flora in widespread use here (An Irish Flora 1977), the fewrecords we have of this closely related group of species were regularly lumped as R. aquatilisagg. or R. aquatilis s.l. (Common Water-crowfoot). Three truly aquatic members of the subgenusBatrachium, namely R. peltatus, R. aquatilis and R. trichophyllus(Thread-leaved Water-crowfoot), all exhibit very similar patterns of variation, both genetic and inresponse to the environment (ie genotypic and phenotypic variation), which Cook (1966a) believesindicates their close parental affinity. When these three species are in a submerged or terrestrial,vegetative, wholly divided-leaf state, they are in fact morphologically indistinguishable. More work isdefinitely required to sort out the differences and clarify the distribution of the water-crowfootspecies in Fermanagh and, since the problem is universal, the same is true everywhere in the BritishIsles and, indeed, throughout Europe (Preston & Croft 1997; New Atlas).
Nutrient ecology and tolerance of disturbance
In the Sheffield area of England, Grime et al. (1988) believed R. peltatus was biasedtowards mildly acidic (often peaty) habitats, and that it was more or less absent from calcareouswaters, where it was replaced by R. trichophyllus. This is definitely NOT the case in Fermanagh.Like R. hederaceus and R. lingua (Greater Spearwort), R. peltatus also appears tofavour lowland aquatic sites which are subject to a degree of disturbance, appearing for instance indrainage ditches, quarry pools, and streams and ponds in areas that are regularly grazed and trampled bycattle or other stock animals, or in waters polluted from time-to-time with manure and agriculturalchemical run-off (Cook 1966a; Hong 1991). One is here reminded of Grigson's aside in his essay on alienplants entitled 'The Wandering Flower', where he writes, "You may not be able to say so to abotanist, but nature interfered with by man is rather more fascinating than nature left to itself."(Grigson 1952, p. 114).
Cook (1966a) regarded R. peltatus and its two very closely relatives R. aquatilis and R.trichophyllus, as being characteristic pioneer plants of freshly dug or recently cleaned poolsand ditches. All three rarely occur in water deeper than 100 cm and they do not tolerate deep shade orvery swiftly flowing water. Cook also regarded R. peltatus and R. aquatilis as beingconfined to eutrophic waters although in Fermanagh we regard R. peltatus as a plant of more orless mesotrophic conditions. Cook never found these two species sharing a habitat, while both of themmay frequently be intermingled with R. trichophyllus (Cook 1966a, p. 134).
In addition to the above habitats, in low-lying coastal areas of Britain and Ireland R. peltatusalso occurs in dune-slacks and in lagoons. In S England, it frequents the uppermost reaches ofcalcareous rivers, which like the Irish turloughs, tend to dry out in summer (C.D. Preston, in: Prestonet al. 2002).
The effect of drought on wetland plants
While numerous studies exist on the tolerance of wetland plants to flooding, in comparison very littleresearch has looked at the effects of drought periods on such species. A worthwhile exception is Volderet al. (1997), who carried out a pot experiment on R. peltatus from wetlands in the FrenchCamargue. This study found that the stage of development at which drought onset occurred had a majorimpact on the growth form and seed output of the species. R. peltatus plants subjected to themost severe experimental drought treatment (ie air drying of the soil with no added water), changedtheir vegetative growth form and survived for up to two weeks. Despite the very severe droughtconditions, their flower, fruit and seed production, while very low, was not zero. The ability of R.peltatus to survive and produce viable seed under a wide range of hydrological conditionsensures a regular input to the seed bank. This fact helps explain the success of the species intemporary marshes and other wetland habitats, where exposure to drought stress is a regular or even anirregular occurrence (Volder et al. 1997).
Flowering and pollination
How the breeding system of R. peltatus operates is a matter of minor dispute. Cook (1966a)describes pretty well all of the Batrachian Ranunculi as being protogynous (ie female organsdevelop first) and self-pollinated, with a tendency to cleistogamy (a term literally meaning 'closedmarriage', ie flowers self-pollinate in the bud). As he describes it, there is a variable length of timeduring which un-pollinated stigmas are exposed and outbreeding might occur. In R. peltatus, Cookreckons this duration can be up to 48 hours, but it is dependent upon the prevailing weather conditionsbeing warm and bright. The flowers produce nectar and do have a scent similar to Cratageusmonogyna (Hawthorn) blossom (Cook 1966a, p. 183).
In a study of Batrachian Ranunculi flowering behaviour in S Sweden, Hong (1991) agreed with Cook'sfindings for R. aquatilis and R. trichophyllus, but found no open anthers in the flowerbuds of R. peltatus. When the flowers opened, the stamens spread horizontally and gave goodseparation (c 5 mm) from the stigmas. In other Batrachian species Hong examined, the stigmas witheredwithin a few days, but in isolated flowers of R. peltatus they remained fresh, swollen and lookedreceptive for periods up to 15 days. Beetles were observed visiting the flowers and Hong considered thatthey were the real agents of pollination in this species (Hong (1991) cited in Proctor et al.1996). Other observations led Hong to suggest there might be a measure of self-incompatibility, such asdefinitely exists in R. acris (Meadow Buttercup). Jonsell et al. (2001) go further in thismatter, stating that in Scandinavia R. peltatus is an obligate outbreeder, but somehow it retainsa low degree of self-compatibility.
Overall reproductive strategy
As is often the case, nothing is known regarding the relative significance or frequency of successfulestablishment of sexual versus vegetative reproduction in R. peltatus, but as with R.penicillatus, a successful vegetative process is certainly easier to imagine happening.Certainly the species forms large stands and these probably consist of one or just a few clones,propagating simply by fragmentation (Jonsell et al. 2001).
British and Irish occurrence
The New Atlas map shows R. peltatus having a widely scattered, decidedly patchy, althoughquite frequent British and Irish distribution (Preston et al. 2002). However, the data must bequalified with an appreciation of the fact that all the Batrachian Ranunculi species are difficult torecognise and distinguish, and errors occur at an unknown frequency. We must therefore approach all mapsof their distribution with caution and do not regard the presented pattern as totally reliable.
Having said this, the New Atlas hexad map for Ireland shows more frequent recording of R.peltatus in NE Ireland than elsewhere, a pattern that probably reflects recorder effort. InGreat Britain, the species presence appears to thin out and become increasingly confined to coastalsituations in both SW England and to the north of the Glasgow-Edinburgh conurbations in Scotland. InScotland, at least, this probably reflects increasing altitude, but it will also mirror the number ofavailable expert field recorders capable of identifying the species of this difficult plant group.
European occurrence
The maps in Cook (1966a, Fig. 15), Jalas & Suominen (1989, Map 1888) and Jonsell et al. (2001,p. 261), show R. peltatus widespread throughout Europe from the Mediterranean to the far N ofScandinavia, but with an overall distinct western concentration of records.
World occurrence
Beyond Europe, R. peltatus occurs only in the coastal area of N Africa and in parts of Asia Minoradjacent to SE Europe, but a closely related form called R. sphaerospermus, occurs in SW Asia(Cook 1966a; Hultén & Fries 1986, Map 872; Preston & Croft 1997; Jonsell et al. 2001).
Names
The Latin specific epithet 'peltatus' is derived from the Greek 'pelta' meaning 'a shield'. In plants,the technical term 'peltate' most often refers to the attachment of the stalk or petiole directly to theunder-surface of the leaf blade, rather than to the blade margin in the normal manner (Gilbert-Carter1964; Gledhill 1985). As mentioned above, the name 'peltatus' was given to this species in error by CarlLinnaeus in 1751, since no member of the Batrachian Ranunculi ever develops peltate leaves orother organs (Cook 1966a, p. 123). However, since the species name complies with the international rulesof botanical nomenclature and is correct in every other respect, it has priority and must be retained.In this instance, the name is best considered merely a label, rather than as is more usually the case,descriptive, reflecting a specific character of the plant.
Threats
Shading and overgrowth by more vigorous plants encouraged by eutrophication.
Native, very rare. European temperate.
1892; Praeger, R.Ll.; Ardy More, Ballycassidy River.
May to July.
Identification difficulties and under-recording of subgenus Batrachium
Although there has been a huge amount of survey work on the aquatic flora of Fermanagh since 1986, almostall of it has concentrated on the lakes and comparatively little focus and attention has been given torivers and streams in the county. The resultant plant records could give the impression that fieldworkers appear to have avoided Ranunculus subgenus Batrachium. If this really is the caseand the group is under-recorded, it will largely be on account of the difficult taxonomic nature of thesubgenus, the range of variation and the confusion engendered by frequently changing names within thecomplex. Another contributory factor is the fact that the Irish field Flora in use for much of theperiod 1975-97 (ie An Irish Flora 1977) did not handle this plant group at all well. For example,it failed to mention R. penicillatus and completely ignored the detailed monographic treatment ofthe subgenus carried out by Cook (1966a).
Growth form and habitat preferences
R. penicillatus s.l. is by far the most common and one of the most robust perennial species ofwater-crowfoot in the flora of Britain and Ireland. It frequently forms large, conspicuous rafts ofdense branching stems rooted in the bed of moderately flowing rivers and streams, but it can also occurin waterways that are liable to occasional, torrential spates (Haslam 1978, p. 31; Holmes 1980; Webster1988). The waters that subsp. penicillatus frequents range from oligotrophic to mesotrophicnu*trient status (ie they are capable of supporting poor to medium levels of plant and animal growth).
Under favourable water conditions with stable bottom substrates of well-cemented rounded gravel and silt,R. penicillatus subsp. penicillatus can become the dominantaquatic vascular plant, its dense floating mats representing a significant structural feature of thewater channels it occupies. It has been observed covering around 70% of the stream bed and modifyingwater flow, promoting silt deposition, and providing both shelter for other plants and animals, and foodfor invertebrate animals and fish (Cook 1966a; Haslam 1978, p. 40; Ham et al. 1982; Preston etal. 2001).
In an Irish study of water quality indicator species, Caffrey (1985) placed all of the BatrachianRanunculi in his group of macrophytes that are most sensitive to increased nutrient forms of pollution,ie especially high levels of nitrogen and phosphates. We believe that all or most of the previouslyluxuriant rafts of R. penicillatus in the Colebrooke River have disappeared in recent times,almost certainly due to increased nutrient levels in the water originating from run-off of agriculturalchemical fertilisers, slurry and manure applied to adjacent field swards.
Heterophylly and identification features
Throughout the year, plants of R. pencillatus bear dissected, thread-like submerged leaves and,during the summer months, in response to longer day-length, they develop additional floating leaves thatare 3-5 lobed and either distinctly or bluntly toothed. Thus the plants become heterophyllous, iebearing leaves of two quite different types. The teeth on the laminar floating leaves may end in longslender filaments, which actually are the leaf veins extending beyond the margin. The submerged leavesof R. penicillatus subsp. penicillatus are always limp orflaccid, and when removed from the water they collapse to form a very slender tassel. When fullydeveloped these leaves are generally longer than the adjacent stem internode (Webster 1991; Webb etal. 1996). This feature, used with care, helps separate the species from R. peltatus(Webster 1991).
Fermanagh occurrence
Whatever is or was the situation regarding the difficulty of identifying Batrachian Ranunculi, relativelyfew records have been made of the water-crowfoot group as a whole in the Fermanagh survey (Cook 1966a;Webster 1991). There are a total of just eleven records for R. penicillatus in the FermanaghFlora Database, the species being present in eight post-1975 tetrads in the VC. Ten of the records needchecking since only one voucher specimen has been made by a field worker (R.M. Barrington). A few findswere originally determined as R. peltatus subsp. pseudofluitansusing An Irish Flora (1977), but we can reassign them to R. penicillatus since theycarried with them information that floating leaves were absent (which could be a seasonal effect), andthat the submerged leaves were longer than the internodes.
Webster (1991) looked at a wide range of subgenus Batrachium material throughout Ireland and, withthe exception of one site on the River Roe in Co Londonderry (H40), she recognised all the R.penicillatus found in Ireland as belonging to subsp. penicillatus. Our Fermanagh recordsundoubtedly are of this subspecies and therefore we have transferred all our R. penicillatusrecords to this taxon. The Fermanagh tetrad distribution map shows the subspecies well represented alongthe Colebrooke River and scattered in several other rivers feeding into Lough Erne, most notably theSwanlinbar River, also referred to as the Claddagh River discharging into Upper Lough Erne, and theBallycassidy River entering Lower Lough Erne.
Flowering
The white or very pale pink flowers are frequently produced in large numbers and R. penicillatusis arguably the most conspicuously beautiful of all the Batrachian Ranunculi. Although the flowers ofsubsp. penicillatus are usually slightly larger than those of R. peltatus, like many othercharacters in this subgenus, the flower size varies considerably with the environmental conditions, andin this case they do not allow reliable distinction between the two species (Cook 1966a).
The flowers are self-compatible and there is a tendency towards pollination in the bud (ie cleistogamy),so that seed is regularly set and can be independent of external pollinating agents (Cook 1966a).Flowering material of subsp. penicillatus from Ireland and Wales is reported to be very fertile,but it is usually far less so in the rivers of the English Lake District and SW England (Holmes 1980).
Anchorage in swift-flowing waters
Like other Batrachian Ranunculi species living in more or less rapidly flowing water, R.penicillatus anchors itself firmly in stream bed shingle or lightly silted gravel by means ofprofuse clusters of fine branching roots produced at the lower stem nodes (Sculthorpe 1967, p. 155).Unlike R. fluitans (River Water-crowfoot), which also frequents moderate- to fast-flowing waters(although this latter species is only very rarely recorded in Ireland), R. penicillatus doesnot form a prostrate compact overwintering stem firmly rooted to the bottom substrate. Despitelacking this specialised structure, R. penicillatus is capable of developing fresh roots fromnumerous lower stem internodes throughout the year, and thus anchors and maintains itself verysatisfactorily, even on somewhat unstable calcareous substrata (Cook 1966a).
Finding a 'safe site' and surviving long enough to become established in a suitable habitat is generallythe most hazardous phase in the life history of any plant species, terrestrial or otherwise. Itis almost impossible to imagine just how minute the odds of achieving successful anchorage must be for aplant propagule in the bed of a moderate or swift-flowing stream. It is similar to the stretch inimagination required to comprehend the improbability of long-distance jump dispersal to remote oceanicislands (Carlquist 1974). Nevertheless, in both instances, somehow the extremely unlikely event doeshappen! In the case of aquatic macrophytes, downstream dispersal is never a problem since the water flowsupplies the transporting vector. Rather the difficulty occurs in the plant propagule settling out fromrapid or even moderately swift flowing water, making the initial anchorage, and rooting securely withoutbeing dislodged again by the current.
Reproductive strategy
In contrast to the fruits and seeds of emergent waterside plants, the propagules of submerged andfloating-leaved flowering plants generally have no great powers of flotation. They either sinkimmediately after release from the parent plant, or after a few hours, or a day (Sculthorpe 1967).Clearly there must be some method of establishment, since aquatic Ranunculus species do occur andsurvive in relatively fast-flowing streams. Do propagules find sheltered sites in eddies behindboulders, or do they temporarily lodge in the slower flow amongst established plants or other debrislodged in the riverbed? Or do they grow out from the relative shelter of the riverbank?
Perhaps for aquatic species found in faster flowing waters, the major or maybe the only role for seed hasto do with long-range or jump-dispersal from one water system to another. Most probably this involvesanimal vectors and, in particular, water birds (Sculthorpe 1967, pp. 331-2; Cook 1988). In swift or evenmoderately flowing waters, colonisation and establishment within the same (ie original) riversystem or water body by vegetative means involving fragmentation of an existing clone, appears very muchmore likely, straightforward and achievable, than from seed. The stems of R. penicillatus breakat quite low forces (Haslam 1978, p. 49) and it is easy to imagine detached rafts snagging downstream onrocks, firmly 'planted' sticks or detached branches of trees, thus re-establishing the macrophyte in afresh site. According to Haslam (1978, p. 163), such trailing fragments must be held sufficiently closeto the stream bed for a month or two in order for them to root and become anchored.
In comparison with this vegetative process, it is impossible to conceive of a suitable site for seedgermination occurring in a moderate or swift flow of water. Indeed, except in almost perfectly stillwater, it is difficult to imagine how a Ranunculus or any other seedling might produce sufficientroots to achieve anchorage.
Clonal longevity
Well established aquatic clones probably survive for many years, but they undergo vegetative cycling, theclump increasing and decreasing on an annual basis with the seasons, a process referred to by ecologistsas 'wash out' (Haslam 1978; Ham et al. 1982).
Really basic research is still required on the reproductive biology and ecology of aquaticRanunculus species in order to definitively answer the sort of questions posed here. Let us hopethat airing the topics here, will help stimulate the necessary work.
British and Irish occurrence
Subspecies penicillatus is rather local and has a decidedly western distribution in Britain andIreland. It generally occurs in Britain in rivers over base-poor, acidic igneous rocks, but in Irelandit tolerates a much wider range of alkalinity, both base-poor, pH 6.1, and base-rich, up to pH 8.5(Webster 1988). In Britain, on the other hand, in base-rich waters subsp. penicillatus isreplaced by subsp. pseudofluitans and the latter is much more widespread and abundant than theformer (Webster 1991; Preston et al. 2002).
European occurrence
The distribution of R. penicillatus in Europe is even less well known than is the case in Britainand Ireland for exactly the same taxonomic and identification reasons. However, it appears to bequite widely scattered in middle European latitudes, the distribution thinning out considerably to bothnorth and south. For instance, apart from two provinces in Denmark, it is entirely absent from theNordic countries (Cook 1966a, Fig. 29; Jalas & Suominen 1989, Map 1892; Jonsell et al. 2001).
Names
The Latin specific epithet 'penicillatus' means, 'furnished with a tuft of hairs, like a paintbrush'(Gilbert-Carter 1964), presumably a reference to the tassel-like submerged leaves.
Population control measures
Where aquatic macrophytes including Batrachian Ranunculi become too productive and begin to formextensive mats obstructing river and stream drainage, the best method of achieving control is not tophysically cut the vegetation, but to shade the banks by planting trees. If cutting does becomeessential, then where aquatic Ranunculus species are dominant, the most successful modificationof subsequent growth is achieved when the mats are cut in full flower. This usually means cutting anddredging in June, but the timing of flowering is dependent on both site altitude and latitude (Dawson1980; Ham et al. 1982).
Threats
Pollution of rivers and streams and over-zealous drain clearance operations.
Native, very occasional or rare. Eurasian temperate.
4 July 1977; R.H. & Mrs H.J. Rossole; lough, south of Enniskillen.
June to August.
Growth form and habitat preferences
Unlike most other aquatic Ranunculus species, R. circinatus is an easy species to recogniseand ought to be well recorded. R. circinatus is a long-lived perennial of clear, base-rich,meso-eutrophic to eutrophic, still or slow-moving waters. It has no floating leaves, producing onlysubmerged, finely dissected ones, the short, rigid segments of which are arranged in a very distinctiveflat fan shape 1-2.5 cm across (Parnell & Curtis 2012). Typically, it lives rooted in mud or claysubstrates, submerged in fairly deep (around 1-3 m), sheltered lake water, or in ditches in grazingmarshes or fens. It is only found in shallower situations provided that the water is permanently presentand never dries out. This is because unlike many other British and Irish species of BatrachianRanunculi, it has no annual, terrestrial life-form, or only very rarely forms one on regularly floodedshorelines. Thus normally it cannot tolerate even a brief period of desiccation and therefore cannotcolonise and exploit exposed bare mud (Cook 1966a; Preston & Croft 1997; Jonsell et al.2001).
In other parts of Britain and Ireland, R. circinatus has colonised man-made still-water habitatssuch as reservoirs, quarry pools, gravel pits and even garden ponds (Preston & Croft 1997), but itdoes not appear to have done so in Fermanagh. In Baltic Scandinavia, R. circinatus is quitefrequently found in brackish conditions, where it can occur down to depths of 5 m (Cook 1966a; Jonsellet al. 2001).
Fermanagh occurrence
Preston & Croft (1997) note that R. circinatus often fails to flower, or does so onlysparingly, which may at least in part account for the paucity of records prior to the very thorough1988-91 NI Lakes Survey. There were no records at all for this species in Fermanagh prior to 1977, and22 of the 34 existing records were made by the NI Lakes Survey. This water-crowfoot has been found in atotal of 22 tetrads, 4.2% of those in the VC. As the tetrad map indicates, most of the records are inLower Lough Erne with outliers on the shores of Upper Lough Erne and at Rossgole Lough on the outskirtsof Enniskillen.
Reproduction and dispersal
R. circinatus is known to flower only occasionally in Britain and Ireland, typically doing sobetween May and August. When flowering does occur, it appears to produce very little or no seed. Withthis background we may safely assume that plants or clonal colonies of R. circinatus reproducealmost exclusively by vegetative means, involving fragmentation and re-rooting of stems (Hong 1991).When autumn arrives and the main shoot is in the process of dying off at the end of the growing season,shoot tips consisting of a short length of the stem apex plus a few green capillary leaves are formedand released from the parent plant. These vegetative fragments float away and have the potential todisperse and propagate the plant (Jonsell et al. 2001). Similar stem fragments may beaccidentally transported, for instance by attachment to boats or to the gear of anglers, and of coursethey may also disperse by attachment to water-living animals including birds (Cook 1966a; Preston &Croft 1997).
Colonising behaviour
In an interesting experiment, recolonisation by macrophytes of a stretch of river in France previouslycompletely cleared of such plants, showed that early colonisation by R. circinatus combined twodistinct strategies of invasion. Firstly there was the 'border effect', involving vegetative outgrowthfrom adjacent intact vegetation, while the second mechanism (without implying secondary importance toit), comprised random recruitment from detached propagules (ie viable vegetative fragments or seeddiaspores). The survey technique used could not distinguish between these two forms of propagule, makingit impossible to distinguish sexual from asexual reproduction (Barrat-Segretain & Amoros 1996).However, we know that in Britain and Ireland, reproduction of R. circinatus is almost exclusivelyvegetative.
The two strategies of aquatic colonisation that the French workers detected, mirror the terrestrial'phalanx' and 'guerrilla' approaches described by Lovett-Doust (1981) for clonal terrestrialRanunculus species such as R. repens (Creeping Buttercup).
Overwintering
Plants of Fan-leaved Water-crowfoot overwinter submerged as a prostrate length of branching stem lyingalong the bottom substrate attached by roots formed at numerous stem nodes. In the summer, the prostratestems develop erect, unbranched stems that grow up towards the surface of the water. (Cook 1966a).
Irish occurrence
In the Republic of Ireland, R. circinatus is very thinly and widely scattered in the N and C ofthe country (New Atlas). North of the border in Northern Ireland, it is concentrated in threediscrete areas: the limestones of Fermanagh and S Tyrone, around Lough Neagh and the adjacent Upperreaches of the River Bann, and finally on the River Quoile in E Down (NI Vascular Plant Database 2015).
British occurrence
In Great Britain, R. circinatus is quite frequent S of a line between Hull and Bristol, but itrapidly declines to rarity towards the N and W of the island.
European occurrence
R. circinatus occurs chiefly in W, C and E areas of Europe, but is very rare in the Mediterraneanbasin and reaches its northern extremity in both Sweden and Finland at a latitude of 63oN inthe Gulf of Bothnia (Jalas & Suominen 1989, Map 1898).
World occurrence
Elsewhere, R. circinatus is found in C and E Asia, and it, or more likely a closely related form,also occurs at similar latitudes in N America (Cook 1966a; Preston & Croft 1997; Jonsell etal. 2001).
Names
The Latin specific epithet 'circinatus' means 'circular' or 'rounded', presumably an inaccurate referenceto the fan-like shape of the dissected submerged leaves (Gilbert-Carter 1964). The English common name'Fan-leaved Water-crowfoot' gives a rather better description of the most characteristic feature of theplant, but is a pure 'book name' of no folklore merit.
Threats
R. circinatus is intolerant of both drainage and excessive cultural eutrophication and has been indecline in both Britain and Ireland, probably for 80 or more years (Grime et al. 1988; Preston& Croft 1997; C.D. Preston, in: Preston et al. 2002). In Fermanagh, further investigationwill be required to discover whether the species is actually spreading at present, particularly in LowerLough Erne, in the way that Potamogeton pectinatus (Fennel Pondweed) did, perhaps due to slow butprogressive effects of cultural eutrophication. It is also possible that shortly after the populationwas recorded it began to decline, following the pattern that appears to be the case elsewhere.
Aquilegia vulgaris L., Columbine
Possibly both native and an introduced, neophyte, garden escape; occasional. European temperate, butintroduced in N America and New Zealand and part of a circumpolar species complex.
1899; Tetley, W.N.; Dunbar House, Fintonagh Td.
January to September.
Growth form
Aquilegia vulgaris is a tall, erect rosette-forming perennial with a short, stout, erect,rather woody, perennating rootstock. According to Clapham et al. (1962), the rootstock is oftenbranched, yet the plant appears to produce only one rosette of basal leaves and a solitary floweringstem which can reach 100 cm, but more usually measures about 60 cm tall. The plant does not possess anypowers of vegetative reproduction, relying entirely on seed for population increase and dispersal.
Native and garden forms and their preferred habitats
Throughout Britain and Ireland, native populations typically grow in woodland glades and fairly openscrub, by woodland rides and streamsides, in damp stony grassland and fen and on scree slopes. Gardenescapes tend to be naturalised in more disturbed or man-made sites, eg in quarries, on roadsides andrailway banks and on or near old walls. Sometimes, the colour, size or hybrid form of the plant makes itobviously a garden escape, but on other occasions in Fermanagh, as for instance on Knockninny or atCarrickreagh, the typical blue or deep purple form of the species occurs and it appears to be a wellestablished member of a stable, semi-natural plant community. It is then probably best to consider itnative.
Columbine is generally associated with limestone rock and calcium- or base-rich soils, though at Lisblakein Fermanagh it grows on acidic peaty roadside banks. It is frost tolerant down to around -25C and canbe thought of as a plant of cool conditions, generally preferring deciduous woodland or hedgerow shadeand a moisture retentive, though not wet, soil. In gardens, it does not tolerate heavy clay soils(Huxley et al. 1992).
As with Hyacinthoides non-scripta (Bluebell) and other species, it is generally assumed totolerate and prefer shade conditions; under the frequent grey, cloudy skies that are the norm in westernIreland during much of the growing season, A. vulgaris also grows both in the open on rockypastures and in disturbed habitats, eg on roadsides and around buildings, as well as occupying partialshade in more natural relatively undisturbed habitats, such as Hazel scrub and lakeshore woodland.
Fermanagh occurrence
A. vulgaris is rare or occasional in Fermanagh, typically occurring either in very sparsepopulations or as isolated individuals. It has been recorded in a total of 26 tetrads, 4.9% of those inthe VC. As the distribution map indicates, it is widely and thinly scattered in the lowlands. Localhabitats include woods, shaded banks and lakeshores, open limestone rocks including quarries, roadsides,a churchyard and a graveyard.
Longevity and persistence of garden escapes
Observation shows that in Britain and Ireland some plants are ephemeral while others definitely persistfor many years. Garden forms are generally considered short-lived, perhaps surviving two or three yearsin cultivation (Grey-Wilson 1989). The longest persistence known to us locally in Fermanagh is the standat Carrickreagh woods on the shore of Lower Lough Erne, which was first recorded by Carruthers over 55years ago and is still there (Forbes & Northridge 2012).
Increasing occurrence in Britain and Ireland
The New Atlas survey completed in 2000 found that A. vulgaris had increased in GreatBritain and Ireland in the 40 years since the original Botanical Society of the British IslesAtlas (Walters & Perring 1962), with a calculated Change Index of +1.70. The change inspecies presence may be attributed to a combination of better recording and a presumed increase in thefrequency of the plant escaping from garden cultivation (R.A. Fitzgerald, in: Preston et al.2002). The editors of the New Atlas recognise that the native distribution of the species is nowtotally obscured, and they therefore mapped all A. vulgaris records as if they were native.
Flowering
Flowers are produced in May and June and copious nectar, secreted in five long hooked spurs, attractsbumblebees with sufficiently long tongues as pollinators (Garrard & Streeter 1983). In the absenceof pollinators, the plant carries out self-pollination. The wild form of the plant (often blue or purplein colour) has no fragrance, but white forms and other garden varieties are often sweetly perfumed, thescent being reminiscent of cloves. This is probably due to the presence of the genes of A.pyrenaica and other perfumed species, being incorporated through the efforts of plant breeders.
Although the flowers are pendulous, after fertilisation the follicular fruits are held stiffly erect.When ripe, the four or five follicles split open at the top of their inner sides so that the many black,shiny, smooth seeds are shaken out when the wind is strong enough to sway and jerk the tall fruitingbranch like a censer (Melderis & Bangerter 1955).
Seed and germination
As is often the case, reproductive performance statistics do not appear to be available in the literaturefor the species, but the individual seeds are quite large (2-2.5 mm), their average weight varying from0.78 mg (Grime et al. 1981) to 1.95 mg (Salisbury 1942). In germination tests, only 4% of freshlycollected seed germinated, indicating that such seed is dormant, perhaps immature, and requiresstratification. After dry storage for six months chilled at 5C, mimicking the overwinteringstratification process, only 30% of seed germinated (Grime et al. 1981).
Dispersal
Given the limited height of the fruit above ground and the passive release of the wind dispersed seedswhich have no appendage to assist flight, nor any form of edible secondary dispersal lure to entice antsor other animal vectors, one would not expect the seeds to travel from the parent plant more than a veryfew metres at most. However, A. vulgaris regularly manages to escape into the wild from gardens,indicating that it does possess considerable powers of mobility.
Further studies required
An obvious need exists for population and reproductive ecology studies of this familiar species, since adeep well of ignorance persists regarding the behaviour of the plant. Even the question of survival inthe soil seed bank does not appear to have been investigated as the species does not feature in themajor survey of this topic in NW Europe (Thompson et al. 1997).
Irish and British occurrence
The distribution of A. vulgaris in the New Atlas shows the species (wild and introducedrecords combined) widely but fairly thinly scattered in Ireland and nowhere common (Webb et al.1996). In Great Britain, the map indicates a much greater presence in the S and W, although again it isvery widely scattered throughout England and Wales, but thinning markedly towards the north andespecially so in Scotland (Preston et al. 2002).
European and World occurrence
Beyond the British Isles, opinions on the taxonomic circ*mscription of A. vulgaris sens lat. arefar from settled, with some authorities including A. atrata and A. nigricans, and othersnot (Jalas & Suominen 1989). Taking it in the broad sense, A. vulgaris s.l. is considerednative in much of W and C Europe, but is regarded as a naturalised introduction in Holland, N Germanyand the Nordic countries (Jalas & Suominen 1989, Map 1907). A. vulgaris s.l. is also regardedas native in N Africa, and it is naturalised widely in N America and in temperate areas of the SHemisphere including New Zealand (Jonsell et al. 2001, p. 312). Together with closely relatedspecies, sometimes considered subspecies or varieties, it forms a circumpolar polymorphic complex(Hultén & Fries 1986, Map 879).
Toxins
Aquilegia species are reported to contain toxic alkaloids similar to those in the genusAconitum (Monk's-hood), ie the cyanogenic glucoside triglochinin (Tjon Sie Fat 1979). However,there is no evidence of recent poisoning by any form of Columbine in Britain and Ireland (Cooper &Johnson 1998). The toxins contained in A. vulgaris are destroyed by heating or drying.
Medicinal and folk uses
The species was formerly employed in herbal medicine, mainly for its antiscorbutic effect. It has fallenout of favour and is little used nowadays, although the root and leaf sap are still sometimes usedexternally in poultices to treat ulcers, swellings and the commoner skin diseases (Grieve 1931; Vickery1995). However, a homeopathic remedy is made from the plant and is used in the treatment of nervousafflictions including hysteria. In the past, dried and crushed Columbine seed was used as a parasiticideto rid the hair of lice and to kill external body parasites (Plants for a Future Database Websitehttp://www.pfaf.org/ accessed 2016).
The flowers are so well supplied with nectar they were used to make a tea and were also eaten in salads.However, since the plant contains poisons it is not recommended for any such purpose (Mabey 1972).
Folklore reports that in the Middle Ages, Columbine was believed to be the food of lions, and those whorubbed the sap of the plant on their hands became gifted with the courage of a lion (Grigson 1987).
Names
The origin of the genus name 'Aquilegia' is doubtful but is sometimes said to be derived from theLatin 'aquila' meaning 'an eagle', the flower spur supposedly resembling an eagle's claw (Johnson &Smith 1946). The Latin specific epithet 'vulgaris' means 'common', which the species is not, at least inthe wild in Ireland. The English common name 'Columbine' is derived from the Latin 'columba' meaning'doves'. It alludes to the appearance (especially of the short-spurred form of the plant) which lookslike five doves drinking (Stearn 1992).
A large number of alternative common names exist in folklore. Grigson (1987) lists 35 of these names,many of which compare the flower shape to that of a style of bonnet worn by elderly ladies, for instance'Granny's bonnet' and 'Old Woman's nightcap'. Other folk names refer to various items of ladies' wear,including shoes, slippers and petticoats, and a few of male apparel, eg 'Batchelor's buttons' and 'Ragsand Tatters' (Grigson 1987).
Threats
None.
Native, very rare. European boreo-temperate.
1884; Barrington, R.M.; Killygowan Island, Upper Lough Erne.
June to August.
Growth form and preferred habitats
T. flavum is a rhizomatous perennial of low lying, moist to wet, near-neutral, moderately fertilemeso- to eutrophic, base-rich, generally calcareous grasslands. In Fermanagh, some of these particulargrasslands were previously used as hay-meadows although they really are rather damp for this purpose.However, the species is also capable of colonising fen sedge peat soils, ie peat formed from the partialdecomposition of sedge and herb litter. This type of organic soil has a significantly higher nutrientand mineral content than the much more acid moss peat found in bogland.
The plant possesses a long, creeping, branching yellowish rhizome from which stout, annual, unbranched,furrowed stems arise, and clonal patches can develop if good growing conditions are met (Melderis &Bangerter 1955; Clapham et al. 1962; Hill et al. 1999). Common Meadow-rue generally growsin full sun, but occasionally it is found in light shade.
The erect habit and the fact that its ultimate leaflets are much longer than wide readily distinguishT. flavum from T. minus. In Fermanagh, T. flavum is often on the small side of itssize range, only around 50-60 cm tall unless it is forced to compete with taller, ranker vegetation,when it can become extremely lax and straggly. The plant can become so enmeshed with the surrounding,competing, grassy herbage, that it becomes difficult to spot, even when one knows it is present.
Flowering and pollination
The cream to yellowish flowers, which are produced between June and August, are largely composed of theslender filaments of the stamens. In Fermanagh, the stamens are sometimes much more sparse than normal,which again makes the plant easily overlooked even when in flower. The bunches of erect yellowishstamens attract small flies and bees (both bumble-bees and honey-bees). They manage to attract insectswithout the advantages of nectar, perfume or petals to advertise and reward the visitors; they simplycollect and feed on the plentiful supply of nutritious pollen (Proctor & Yeo 1973, p. 60; Garrard& Streeter 1983). Some authorities regard the flowers as fragrant, others not, so there might wellbe a measure of geographical variation in this particular respect. But then every nose is different!(Proctor &Yeo 1973; Jonsell et al. 2001).
In addition to pollination induced by movements of the insect visitors, very probably some degree of windpollination of the reduced flowers also takes place (Clapham et al. 1962).
Fruit
Each small flower of the more or less dense branched panicle inflorescence contains 3–12 free carpels(some of which are often abortive). The fruit is a small cluster of single-seeded, almost spherical,ribbed achenes (ie single seeded dry fruits) (Butcher 1961; Hutchinson 1972).
Seed survival
There is a solitary estimate of the soil seed bank longevity of the species in the NW Europe survey(Thompson et al. 1997). This suggests the seed is long-persistent (ie buried seed may persist forat least five years).
Conservation field studies
Information is lacking or difficult to locate on such matters as the scale of seed production, populationdynamics, the balance of vegetative versus sexual reproduction, or the competitive ability of thespecies, studies of which would assist any serious attempts to conserve this declining plant species.The various bodies involved in species conservation in both Ireland and Britain give the impression(deserved or not) that attending meetings, drawing up reports, endlessly revising species lists, drawingboundary lines on maps and arguing for resources are all that they are required to do in order toeffectively manage threatened wild populations of plants and animals – work they regard as a businesslike any other (Marren 2002, p. 307). One wonders when the basic natural history studies, let alone thedetailed ecological field studies and localised site observations and analysis that should inform anddirect active conservation management are going to be carried out. The conservation industry, if we cancall it that, appears much more concerned with words than deeds. Hands and boots need to get dirty toreally make a change in species fortunes on the ground.
Fermanagh occurrence
T. flavum occurs very rarely in fens, ditches in wet meadows and in marshy grasslands on lakeshores. It has been recorded in a total of just nine Fermanagh tetrads, seven of which have post-1975records. As the tetrad distribution map indicates, it currently occupies six stations on both shores ofthe southern half of Upper Lough Erne, but has not been refound at either of the two earliest sites inthe VC which lie closer to Enniskillen. In 1996, an interesting new station, quite remote fromthe seven previous ones, was discovered by a survey team from the EHS on the shore of another of thelarger lakes in the county, at the NW end of Lough Melvin. In July 2010, another new site was discoveredby RHN and HJN at Derrymacrow Lough, where a solitary plant grew alongside Thelypteris palustris(Marsh Fern).
The other record details are: in damp meadow, W of Lough Digh, 1950, MCM & D; Lough Corby, 3 km W ofNewtownbutler, 1983, RHN, also 6 June 2002, RHN, RSF, J.S. Faulkner & I. McNeill, and 11 July 2010,RHN; Drummully Td and Croostan Td shores, 2 km NE of Derrylin, 23 July 1986, P. Corbett & P.J.T.Brain; Derryad Td shore, 13 August 1986, S.A. Wolfe-Murphy & L.W. Austin; S shore of Trasna Island,19 August 1986, T. Waterman & P.J.T. Brain; SE corner of Corraharra Lough, Mullynacoagh Td, 30 June1991, RHN, also 15 August 1994, RHN & RSF, 25 July 2004, RHN, and 8 July 2010, RHN & HJN; Sshore of Derrymacrow Lough, 11 July 2010, RHN & HJN; shore of Lough Melvin, NW of Garvos, oppositethe end of Bilberry Island, 26 June 1996, EHS Habitat Survey Team.
Irish occurrence
The Fermanagh records represent the only nine stations (seven extant) for this rare perennial plant inNorthern Ireland outside the Lough Neagh and Lough Beg basin and the connected River Bann drainagesystem. Even in the latter areas, where once it was frequent, T. flavum is now regarded as beingon the decline due to drainage and widespread changes in and intensification of agricultural practices(Harron 1986; Hackney et al. 1992).
In the Republic of Ireland, T. flavum is widely but very thinly scattered, (mainly along thecourse of the River Shannon) and it is locally frequent in a few central areas of the island (Webb etal. 1996; Preston et al. 2002).
British occurrence
In Britain, the distribution is greatly concentrated to the SE of a line between Swansea and Whitby witha few rare outliers, chiefly coastal, stretching as far north as Edinburgh in the east and Kintyre onthe west coast (Preston et al. 2002). With the decline of the species on agricultural land, inmany areas of Great Britain, Common Meadow-rue is no longer common. Rather, it has become restricted torelict land, often occurring along linear landscape features such as river banks and roadside ditches(R.A. Fitzgerald, in: Preston et al. 2002).
Significant local factors affecting the widespread decline of Common Meadow-rue in Britain and Irelandinclude the almost universal move away from hay to silage production, the widespread use of herbicidesand the ploughing and reseeding of pastures, very often subsequently fertilised by slurry sprayingand/or agrochemical application. These measures have fundamentally altered grassland ecology, increasingproduction and heightening interspecific competition in ways that force some plants towards localextinction.
Fossil record
The fossil record of T. flavum is almost exclusively based on identification of theircharacteristic and abundant achenes. Pollen grains can really only be distinguished to the genus level,but the fruits do prove that T. flavum has persisted in the British Isles throughout the lastfour interglacial periods and the last two glacial periods − at least in the southern part of Englandand Wales (Godwin 1975).
European occurrence
In Europe, T. flavum is widespread throughout N, W and C regions, stretching N to within theScandinavian Arctic Circle and S to the toe of Italy and to the Spanish pre-Pyrenees, but thedistribution thins markedly further E and it is absent from Greece and all the Mediterranean islands(Jalas & Suominen 1989; Jonsell et al. 2001).
Uses
T. flavum is quite a popular garden border perennial, especially a very fine, large, robustvariety called 'Illuminator' (Griffiths 1994). Despite this, there is very little evidence of it as agarden escape in the British Isles, certainly in comparison with several other members of the family(Preston et al. 2002).
The leaves of T. flavum in the past were used to allay fevers, and the root was sometimes used indyeing (Melderis & Bangerter 1955); supporting evidence for these uses of the plant has not beenlocated. Rather surprisingly, Grieve (1931) makes no mention of the species whatsoever in her verycomprehensive herbal.
Toxins
In addition to containing the toxin protoanemonin, common to all members of the Ranunculaceae (Cooper& Johnson 1998), T. flavum contains at least six alkaloids, the principal one being berberine(Velcheva et al. 1992). There is a tremendous amount of current pharmaceutical research going oninvolving European species of Thalictrum. RSF found that 80 out of 81 scientific papers that werelocated by a major Internet database searching back as far as 1981 were on this topic alone.
Names and folklore
The genus name 'Thalictrum' is an ancient Classical Greek name first given by Dioscorides to amember of the genus, possibly in the T. minus species aggregate (Gilbert-Carter 1964). The namemay be linked to the Greek 'thallo', meaning 'to flourish' or 'to abound in', perhaps a reference to thenumerous bundles of stamens in the inflorescence (Johnson & Smith 1946). The Latin specific epithet'flavum' is straightforward by comparison, translating as 'yellow'.
The plant has four English common names in Britten & Holland (1886), of which 'Meadow-rue' is themost well known, being given on account of the finely divided rue-like leaves. The suggestion made byGerard (1633, p. 1252) is that some old herbalists confused this species with Ruta graveolens(Rue) which had medicinal uses. Indeed, T. flavum was previously called 'Ruta palustris',or 'Fen Rue', from the appearance of its leaves and its typical place of growth (Prior 1879, p. 77).
The two remaining names in Britten & Holland (1886), refer to T. flavum as 'False Rhubarb' and'Meadow Rhubarb'. According to Lyte (1578), this was because like Rhubarb the plant has laxativeproperties and also because it has roots that are yellow like those of Rhubarb! It was also called'Great Bastard Rhubarb', or 'English Rhubarb' (Gerard 1597, 1633), and it was supposed to cure oldulcers, be good for the belly and act as an astringent 'without biting' (Gerard 1633, p. 1252).
Threats
The Corraharra site could well become overgrown by conifers and the Lough Corby site has already beendamaged by drainage.
Native, rare. Eurasian boreo-temperate.
1850-80; Smith, T.O.; Ardunshin Bridge, Colebrooke River.
May to August.
Growth form and habitat preferences
T. minus is a extremely variable, clump-forming stoloniferous perennial species or speciesaggregate, the differing growth forms being produced by the orientation and length of its rhizome, whichcan be either ascending and short, or horizontal and long (Clapham et al. 1987). The plantproduces erect stems, variable in form and colour and up to 120 cm tall, but in Fermanagh often muchless, most typically around 50-60 cm in height. Variation in the species also extends to the widevariety of habitats it occupies. Again, in a local context, these range from fairly dry to quite moist,lime- or base-rich conditions. The three or four times pinnately divided leaves are also very variable,but the ultimate leaflets are generally about as long as broad, a fact which clearly distinguishes thespecies from T. flavum, which is much less finely divided and generally grows under quitedifferent ecological conditions. In comparison, T. flavum is a plant of very much wetter and moreopen sun-lit, marshy grasslands and fen-like habitats.
In Great Britain, T. minus is also represented in at least three rather different types of habitatthat are identified by Clapham et al. (1962), as: i. limestone rocks and grassland; ii. coastaldunes; and iii. streamside or lakeshore gravel and shingle. To the first of these we should also addcliff ledges, screes and the gryke (or grike) crevices of limestone pavement (Halliday 1997).
Fermanagh occurrence
In Fermanagh, T. minus has been recorded in a total of eleven tetrads (2.1%), eight of which havepost-1975 records. It is chiefly distributed around Lough Erne and Lough Melvin and along stretches ofthe Colebrooke River. In Fermanagh, T. minus typically occurs as very small populations of just ahandful of scattered individuals, or as isolated clumps growing in fairly dry, rocky or sandy lakeshoresituations, or in similar soils on river banks under the shelter and shade of scrub. Unless activeconservation measures are taken to reverse the present decline, these small, fragmented populationsrepresent the end of the line for T. minus in Fermanagh. Small, isolated populations inevitablysuffer from poor cross-pollination, consequent inbreeding depression and a loss of fertility.Undoubtedly all such rare and scattered species populations are on the downhill slope towards localextinction at some future date. In the case of T. minus, the existence of some degree of apomixismay delay matters for a while, but eventually fragmented species like it are fated to fade out anddisappear.
Variation
The New Flora of the British Isles (Stace 1997) accurately describes the species or speciesaggregate as, "very variable and little understood". Stace also points out that while up toeight subspecies or other rank of taxa have been described on the basis of fruit, habit and hairs, thevariation has not been properly studied. Until it is, his opinion chimes with my own, "thesevariants are not worth recognising". Matters are complicated by the fact that some races within thespecies or species aggregate appear to be apomictic and can thus set asexual seed, which is a rathersophisticated form of vegetative reproduction.
By whatever means the plant manages to reproduce, once it becomes established in a site T. minusbecomes very persistent. It still survives, for instance, on the banks of the Colebrooke River whereSmith found it and at Gubbaroe Point in crevices in the limestone pavement shore of Lower Lough Erne,two sites where it was originally found in the second half of the 19th century.
Irish occurrence
T. minus used to have other inland stations in Northern Ireland, but with the exception of theFermanagh records and a few stations in the Mourne Mountains, Co Down (H38), practically all theremaining Northern Irish stations for the species are on coastal sand dunes, where they are sometimesdistinguished as subsp. arenarium (Clapham et al. 1987; FNEI 3).
In the Republic of Ireland, T. minus is confined either to the coast or to the shores of largerlakes in the west and in the Central Plain, with a few additional stations, mainly in the mountains ofCos Sligo (H28), Leitrim (H29) and the Connemara region in the mid-west of the country. In thesemountain stations, Lesser Meadow-rue occupies rocky ground and mountain ledges where it can avoidintensive plant competition; at the same time it is less accessible to browsing sheep (Walters &Perring 1962; Preston et al. 2002).
Toxicity and medicinal uses
Browsing this herb must be a mixed pleasure for stock animals, since like other members of theRanunculaceae T. minus contains the bitter tasting and highly irritant toxin, protoanemonin(Cooper & Johnson 1998). In common with T. flavum, T. minus is the subject of a greatdeal of current pharmaceutical research since both species contain several additional poisonousalkaloids, including berberine, although thalactamine is the main ingredient in the toxin co*cktail inT. minus. Berberine is used in modern medicine to treat stomach and gall bladder ailments(Urmantseva et al. 2000).
Garden cultivation and occurrence in Great Britain
T. minus has also had a long career in Britain and Ireland as a garden border perennial. It iscertainly not grown for its flowers, but rather for its attractive finely dissected foliage which isgreatly appreciated by florists and flower arrangers (Robinson 1909, p. 888). Due to these widespreadgarden populations, T. minus also regularly jumps the garden wall and occurs as an establishedintroduction. In Great Britain, these garden escapes sometimes become intermixed with supposed nativeplants, mainly in the Midlands, S & SE England and in Wales, although it can perhaps more rarelyoccur in this way as far north as Glasgow (Preston et al. 2002).
European occurrence
T. minus is widely but unevenly spread throughout continental Europe, being rather less frequenton the Atlantic coastline than the extent of the species or aggregate distribution in the British Isleswould suggest. It occurs up to 70N in Scandinavia but is absent from much of the rest of Norway, Swedenand Finland, and, indeed, it appears to have declined in the latter two countries. On the IberianPeninsula, T. minus penetrates to the southern Spanish Sierras and it also occurs rarely onCorsica, Sardinia and Sicily, yet is absent from the Peloponnese, Crete and the smaller Mediterraneanislands (Jalas & Suominen 1989).
World occurrence
Beyond Europe, the distribution of T. minus stretches to N, E and S Asia, Ethiopia, S Africa andAlaska (Jonsell et al. 2001).
Names
The Latin specific epithet 'minus' is a comparative of 'parvus', and means 'smaller than', presumablysmaller than T. flavum, which generally is the case (Gilbert-Carter 1964). The plant appears tohave been insufficiently known or was ignored as being of little or no interest or use, so it does notappear to have accumulated much in the way of English common or local names, apart from 'LesserMeadow-rue' (see T. flavum species account). Gerard (1633, pp. 1251-2), however, distinguishes itfrom T. flavum as, 'Small Bastard Rhubarb', and for an explanation please again see the T.flavum account.
Threats
The Colebrooke River sites could easily be destroyed by river bank improvements and the Muckross site bytourist developments.
Introduction, neophyte, deliberately planted, very rare. Eurosiberian temperate, widely naturalisedincluding in N America and New Zealand.
1947; MCM & D; Gubbaroe Point, Lower Lough Erne.
March and August.
Growth form and preferred habitats
The deciduous, sharp spiny shrub B. vulgaris typically grows in hedgerows, scrub banks,coppice woodland and on waste ground. Being a very spiny shrub, at quite a late stage in history,Barberry was widely planted for hedging.
Secondary host of plant pathogen
Barberry happens to be the secondary host plant of Puccinia graminis, the pathogenic fungus whichcauses black stem rust of wheat and other grasses. When this was realised in 1865, Barberry waseradicated in some countries (eg Denmark and in parts of N America), thus providing some degree ofcereal disease control. According to Rackham (1986, pp. 42-3), Barberry is not necessary to thelife-cycle of the fungus in England (and presumably elsewhere in Britain and Ireland). Although farmershave from time-to-time removed the shrub from field hedgerows, Rackham (1986), who appears to thinkchiefly or entirely in terms of S England, considers it unlikely that B. vulgaris was evercommon, nor was it much diminished by these intermittent eradication measures.
Flowering
The drooping bunches of small lemon-yellow flowers which appear in May and June are sweetly anddelicately scented, and their abundant nectar readily attracts bees and flies as pollinators (Genders1971; Lang 1987). Interestingly, the six anther filaments in each flower display an instantaneousreaction, moving inwards when touched by an insect visitor and thus dabbing pollen on to it, ie they arenot just elastic and under tension, but rather they are tactile or 'touchy', a feature we associate verymuch more with animals than with plants! In his book The action plant, Simons (1992) deals withthe phenomenon of flower 'irritability' in great detail. He describes the mechanism of the reversibletouch-sensitive stamens of Berberis involving biochemistry and transferred electrical impulsesand motor cells, exactly comparable to these processes in animal nerve tissue but, of course, minus theactual nerve cells (Simons 1992, pp. 35-39 & 240).
Fruit dispersal
The red berries attract birds and at least nine species have been recorded eating them (Lang 1987),chiefly members of the thrush and crow families (Ridley 1930). They are, however, very acid, andapparently are not popular with them for this reason, but cattle, sheep and goats are known to eat them,and in N America after the shrub was introduced by settlers, stock animals are reputed to have dispersedthe seed with their dung (Ridley 1930, p. 365).
Culinary, medicinal and other uses
Barberry was a much cultivated shrub in medieval times, the ripe red, egg-shaped berries borne inclusters of about ten, making a delicious, if somewhat tart jam or dessert jelly. The berries were alsocandied and eaten as sweets and they were used to flavour punch. The bark has an unforgettable brightyellow 'blaze' on its interior when cut with a knife. It was used to tan leather and to colour ityellow. Above all else, in terms of uses for the plant, the yellow inner bark of both stem and root wasmedicinally prized as a cure for jaundice following the well known 'Doctrine of Signs' or 'Signatures'(Grigson 1987; Vickery 1995). The Berberis jaundice cure was also extended in unspecified areasof Ireland from humans to cattle (Allen & Hatfield 2004).
Toxicity
While the ripe berries are edible and contain high levels of vitamin C, unripe they are distinctlypoisonous, containing the alkaloid berberine and at least three other toxins. The highest content ofberberine is actually present in the bark, but the seed (especially that of related cultivated speciesand varieties), should also be avoided. Berberine (which is also present in species ofThalictrum) is an antibacterial gastric and mucosal irritant, causing vomiting and diarrhoea iflarge quantities are ingested. Berberine can depress respiration by acting on the central nervoussystem, and it can also cause uterine contractions. THUS IT IS DANGEROUS TO EAT THE BERRIES, ANDESPECIALLY SO FOR PREGNANT WOMEN (Lang 1987). Having said this, no authenticated cases of either animalor human poisoning by any part of the plant were uncovered by Cooper & Johnson (1998).
Barberry it is still very much used in herbal medicine and homeopathy for a wide range of complaints (seebelow). In its favour, Cooper & Johnson (1998) report that the pharmacological activity of berberineto treat some cancer tumours is currently being investigated.
Fermanagh occurrence
There have only ever been four stations recorded for the plant in Fermanagh. The one found by Meikle andco-workers at Gubbaroe Point on the shore of Lower Lough Erne very probably fell victim to forestryplantation operations in the vicinity soon after its discovery. The details of other three more recentstations are: hedge E of Clonagore Td, near the old Ulster Canal, 23 August 1996, RHN; hedge atClonelly, NW of Kesh, 17 March 1999, RHN & HJN; and several large bushes in front of old castle atCastle Caldwell, 11 November 2006, RHN.
British and Irish occurrence and status
For all of the above reasons, in Ireland, B. vulgaris is regarded (at least probably) as adeliberately planted introduction, whereas in Britain it might either be a native or an archaeophyte(T.D. Dines, in: Preston et al. 2002). In Britain, the 1962 BSBI Atlas and its 1976revised edition, recorded B. vulgaris as being quite a frequent and widespread species. Indeed,the New Atlas displays the plant as even more widespread in Britain than the earlier BSBIAtlas map, thinning out now only N of Glasgow and Edinburgh (BSBI Atlas; NewAtlas).
Formerly in Ireland, Barberry was much more frequent, but it is now very rare and widely scattered,having been most successfully extirpated for disease prevention during the 19th century (Cat Alien PlIr).
European and World occurrence
Beyond the British Isles, B. vulgaris is considered native in C Europe by Jonsell et al.(2001), but it also extends over most other temperate parts of the continent, although absent in the farnorth and around the Mediterranean (Lang 1987). Other botanists consider it a native of Asia's middleand western mountains (Royer & Dickenson 1999). I believe it is best considered an archaeophyte inmost of W Europe (ie it is an ancient or very long-standing introduction). Grieve (1931) regarded therange of the species included N Africa, but the N Hemisphere distribution published by Hultén &Fries (1986) does not support this idea (Hultén & Fries 1986, Map 886). In all these regions, as inBritain and Ireland, it is difficult to distinguish where the shrub is naturalised, from areas where itis supposed or imagined to be native. Barberry is introduced in N America, and it has spread there fromcoast to coast. It has also been recorded in New Zealand.
Names
The genus name 'Berberis' is thought by some to be the Latinised form of the Arabic word for the fruit ofthe plant, 'berberys' (Chicheley Plowden 1972; Stearn 1992), in some way signifying a shell (Grieve1931). Other writers consider it simply a geographical reference to the Barbery coast of N Africa (egGledhill 1985). Gilbert-Carter (1964) plays it safe, regarding the name as, "a medieval Latin wordof doubtful origin". The Latin specific epithet, 'vulgaris', meaning 'common' is certainlyinappropriate in Ireland, and perhaps this is becoming the case elsewhere since the species is regardedas declining in Great Britain (Change Index calculated as -0.61) (T.D. Dines, in: Preston et al.2002).
There are at least nine English common names for the plant, several referring to its supposed jaundicecure mentioned above, eg 'Jaundice Tree', 'Jaunders Tree' and 'Jaunders Berry' (Grigson 1987). There arerecords of this jaundice cure from all over Britain and Ireland (Allen & Hatfield 2004).
The name 'Barberry', which we have already mentioned in connection with its Latinised genus name, hasseveral local variants around the British Isles, for example, 'Berberry', Barbaryn', 'Barboranne' and'Berber'. The remarkable yellow inner bark and wood is undoubtedly the origin of the name 'Guild' or'Guild Tree', while 'Woodsour', 'Woodsore' and 'Woodsower' are applied to both B. vulgaris andOxalis acetosella (Wood-sorrel) on account of their woodland habitat and sour taste (Britten& Holland 1886).
The remaining suite of about eight variant names centres on 'Piperidge' and includes 'Pipricks','Piprage' and 'Pepperidge'. Prior (1879) reckons that this is derived from the French 'pepin', a pip and'rouge', red, a reference to the fruit which he considers rather small and lacking in juice, and hence apip rather than a berry!
Threats
Deliberate destruction in the mistaken belief that it is necessary to do so to limit the spread of cerealrust.
Introduction, neophyte, a very rare garden escape.
18 August 1987; EHS Habitat Survey Team; hedgerow on Staff Island, Upper Lough Erne.
The solitary Fermanagh record of this spiny, evergreen shrub is very probably bird-sown on this small,well-wooded island. The species, which reproduces entirely by seed, is capable of becoming naturalised,but we do not have any further evidence of its behaviour at this particular station. Elsewhere inIreland, B. darwinii has been rarely recorded in just six other widely scattered VCs, includingthe northern Cos Tyrone, Down and Antrim (H36, H38 & H39) (Cat Alien Pl Ir).
Introduced, neophyte, a very rare casual garden escape.
6 April 1996; Northridge, R.H.; roadside hedge at Toura Crossroads, near Drummully.
This spiny evergreen shrub native of Western N America is very familiar as a garden plant. It produces anedible, grape-like black berry in July, a time when fruit demand is high from birds such as juvenileBlackbirds and Mistle Thrushes. Thus seed is very readily carried over the garden wall into 'wild'situations and deposited with bird faeces (Snow & Snow 1988, p. 108).
In Ireland, the New Atlas records M. aquifolium in just four hectads in NI, but none at allelsewhere on the island. The solitary Fermanagh record listed above is very close to the internationalboundary with Co Cavan (H30). Reynolds (Cat Alien Pl Ir) lists the plant as a rare garden escapewith details of just three records (omitting the Fermanagh occurrence): one from Glendalough, Co Wicklow(H20), despite this being missing from the New Atlas, and one each from Cos Tyrone (H36) andAntrim (H39). The latter record was listed in the Flora of Urban Belfast, somewhere in southBelfast near Dunmurry.
The New Atlas shows that the shrub has become very widely naturalised in Britain, especially in C& E England, but it has also escaped from cultivation as far north as Easterness, near Inverness (VC96).
Introduction, archaeophyte or neophyte, an occasional escape from cultivation.
1902; Abraham, J.T. & McCullagh, F.R.; Pettigo.
May to October.
Growth form and preferred habitat
This is the only poppy, other than the Welsh Poppy (Meconopsis cambrica), which is in any wayfrequent in Fermanagh. It is a distinctive, tall, blue-green, fleshy summer annual of lowland, ruderalhabits, most typical of roadsides, waste ground and rubbish tips, but occasional also in disturbed soilin gardens. In the latter case, it may have been accidently imported with container plants purchasedfrom garden centres. It flowers in July and August, but even when not bearing its large white, lilac orred terminal flowers, often with a basal purple blotch on each petal, this poppy is very easilyidentifiable. The only plant it might possibly be confused with is Glaucium flavum (Yellow HornedPoppy) but since it is a strictly maritime plant (certainly with no inland stations in Ireland (Prestonet al. 2002)), and Fermanagh has no coastline, we do not meet with any such problem.
Like the other 'cornfield' red poppies, P. somniferum is principally found in light, dry to moistbut typically well-drained soils, always in disturbed habitats. In Britain and Ireland, it isalways a weed of cultivation, a garden escape or discard (sometimes double-petalled), or a bird-seedintroduction (Clement & Foster 1994; Cat Alien Pl Ir).
There are two forms of opium poppy, often but not always given subspecies rank as subsp.somniferum and subsp. setigerum (DC.) Arcang. The former is the common, usual form, whilesubsp. setigerum, which is more conspicuously bristly on its stems, leaves and sepals, is a rarecasual only in Britain and Ireland (Stace 1997).
Fermanagh occurrence
In Fermanagh, P. somniferum has been recorded in a total of 28 tetrads, 5.3% of those in the VC.Twenty-three Fermanagh tetrads contain post-1975 records. It is confined to the lowlands of the countyand is found especially in and around the Enniskillen and Maguiresbridge 'conurbations', although bothare hardly large enough to be called such. Previously, when Fermanagh still had railway transportconnections, it grew along lines and in stations, but now Opium Poppy is only found as a weedy escapealong roadsides, in or near gardens and on disturbed or waste ground.
Irish occurrence
While P. somniferum has been recorded at least once in 30 of the 40 Irish VCs (Cen Cat Fl Ir2; Cat Alien Pl Ir), the New Atlas hectad map shows it is much more frequentlyrecorded in N Ireland than further south. Apart from a concentration around Co Dublin (H21), it appearsvery thinly scattered across the Republic of Ireland (New Atlas). I regard this Irishdistribution pattern purely as a reflection of recorder hours in the field. The restricted distributionin northern Britain and to lower ground everywhere is undoubtedly due to the susceptibility of youngplants to late season frost.
British occurrence
The New Altas distribution shows P. somniferum as a frequent and widespread casual inlowland areas of Great Britain, mainly occurring S of a line between Anglesey and Hull, and becomingmore confined
to coastal habitats further north. The restricted distribution in the north of these isles, and to lowerground everywhere, is undoubtedly due to young plants being susceptible to late season frost.
Flowering
The flowers are self-compatible and pollination probably takes place in the bud. Again like otherwidespread poppies, P. somniferum is phenotypically very plastic and it has a high seed output (amean of between 6,000 and 7,000 seeds/capsule). The seed is long-persistent in the soil seed bank(Salisbury 1964; Jonsell et al. 2001).
History of cultivation and uses
P. somniferum has been in cultivation for medicinal, culinary and decorative purposes sinceancient prehistory, and images have been preserved from the civilisation of ancient Sumeria (4000 BC)depicting the poppy along side other images indicating euphoria. Sumerian knowledge of the plant waspassed via the Assyrians and Babylonians to the Egyptians, who around 1300 BC began cultivating P.somniferum var. album, referred to as the 'Opium Thebaicum', 'The White Lotus'. The LateBronze Age Mediterranean trade routes of the Phoenicians and Minoans moved this profitable item fromEgypt to Greece, Carthage and Europe. Figurines of Late Minoan and Mycenaean goddesses dating from about1350 BC, show them wearing poppy decorated crowns (Hood 1978, p. 109; Baumann 1993, p. 69).
Writing sometime between c 460-377 BC, Hippocrates, the Greek father of medicine, dismissed the magicalattributes of opium. However, he acknowledged its usefulness as a narcotic and a styptic in treatinginternal diseases, the ailments of women and epidemics. By 300 BC, however, opium was being widely usedby Arabs, Greeks and Romans, both as a sedative and as a soporific.
For many years, the Christian church and the associated state authorities banned opium, probably in partbecause of its 'devilish' eastern origins and, rather surprisingly, it fell into disuse. In 1527 AD,during the height of the European Reformation, opium was reintroduced into medical literature by theSwiss alchemist and physician Paracelsus (1493-1541). He very cleverly renamed the drug 'Laudanum'(Tincture of Opium in alcohol), and thus circumvented church and state disapproval. The black pills or'Stones of Immortality' Paracelsus prescribed were a compound of opium, citrus juice and essence ofgold, and they were used as painkillers (Webpage 'Opium throughout history' at:http://www.pbs.org/wgbh/pages/frontline/shows/heroin/etc/history.html, accessed April 2016).
The Romans are reputed to have sprinkled the top of their bread with the nutty flavoured poppy seeds, theonly part of the plant which contains no, or very little alkaloid (Le Strange 1977).
Grieve (1931) provides much information on both cultivating the plant and the history of its herbalmedicinal uses. Rather surprisingly there does not appear to be any great fund of plant lore associatedwith this interesting, long cultivated and heavily used plant (Vickery 1995).
Nowadays, P. somniferum is a frequent escape from cultivation throughout its range, where it isusually grown either for oil-seed, or as a pharmaceutical crop plant for the latex from its unripecapsules. In past years, most recently in the period between 1880 and 1930 when agriculture wassuffering one of its periodic cycles of decline in the British Isles, a variety with pale lilac flowersand whitish seeds was grown for opium in England, chiefly in Lincolnshire (Grieve 1931; Thirsk 1997).
However, and in spite of this, in most of the British Isles P. somniferum contains very littlealkaloid due to our less than ideal climate (too little sunshine) for the species. It is mainly grown,therefore, as a garden ornamental of some minor culinary and herbal medicinal use. Nine garden varietiesare listed, and these and others illustrated in Phillips & Rix (1999) book, Annuals andbiennials. The garden or cultivated form is subsp. setigerum, occasionally referred to assubsp. hortense (Hussenot) Corb., but not so by Griffiths (1994) in his RHS Index.
Geographical origin of the two subspecies
Having been in cultivation so long, the native territory of P. somniferum naturally has becomeobscured, but it is generally agreed to have originated somewhere in SW Asia, probably in Anatolia(Kadereit 1986a). It has been suggested that subsp. setigerum may be a native of W & CMediterranean, Cyprus and probably or at least possibly the Atlantic Islands (Kadereit 1986 a, b).
Fossil history
The earliest fossil remains of apparently cultivated P. somniferum in Europe were found in SHolland and in adjacent regions of W Germany, but according to Kadereit (1990) there does not appear tobe any reliable information about where cultivation first began on the continent. I suggest that themost obvious place to look would be around the coastal area of the Tyrrhenian Sea, ie S Italy and ESicily, and especially in Campania, since this was the most important early trading area in southernEurope of the ancient Greeks and Phoenicians. These traders certainly brought the product from Egypt andvery probably introduced the plant along with it (Boardman 1980; Grant 1987).
In Britain and Ireland, P. somniferum first appeared as seeds in the Bronze Age archaeologicaldigs and it is common from the Iron Age onwards (Godwin 1975; P.J. Wilson, in: Preston et al.2002).
Toxins
A total of about 25 alkaloids are obtained by the pharmacist from the opium latex, but six of theseaccount for about 98% of the total. The six are: morphine (named after Morpheus, one of the sons of thegod of sleep, Hypnos (Radice 1973)), codeine, papaverine, narcotine, narceine and thebaine (Stodolaet al. 1992). Morphine is the most potent and the most present of these alkaloids. Althoughnowadays opium is chemically purified and separated, none of the drugs obtained have been superseded byany synthetic product.
Stock animals are very unlikely to graze the plant, but Opium Poppy IS EXTREMELY AND FATALLY POISONOUS,AND IT SHOULD NEVER BE USED FOR SELF-MEDICATION (Cooper & Johnson 1998).
Names
The origin of the genus name 'Papaver' appears to be obscure but it is a Latin name for a poppy, probablythe medicinal P. somniferum, Opium or White Poppy (Gilbert-Carter 1964). It may have been derivedfrom the Celtic 'papa', meaning 'thick milk', or Latin 'pappa' meaning 'milk', either way an obviousreference to the milky latex members of the genus contain (Melderis & Bangerter 1955; Hyam &Pankhurst 1995).
The Latin specific epithet 'somniferum' is a combination meaning 'sleep-bringing' or 'soporific' from'somnos', sleep and 'fero', to bear (Gilbert-Carter 1964).
Apart from the English common name 'Opium Poppy' which refers to the origin of the drug, Britten &Holland (1886) list eight additional common names including 'Balewort', said to be derived from 'bealo'or Old English 'bealu', meaning 'bale', as in evil, mischief, woe, injury or pain (co*ckayne 1864-6). Thenames 'Cheesebowl', 'Chasbol', 'Cheesebouls' and 'Chasse' all derive from the shape of the ripe capsulebeing considered as resembling round cheeses (Prior 1879). The name 'Mawseed' has some connection withthe fact that the seed is provided for garden birds in winter. In German, it is called 'Magsamen', andin Polish, 'mak' (Prior 1979).
Threats
None.
Introduction, archaeophyte, a rare casual. European southern-temperate, but very widely naturalised inboth hemispheres.
1902; Abraham, J.T. & McCullagh, F.R.; Greenhill, north of Ardunshin.
July to October.
Growth form and preferred habitats
Common Poppy is a summer annual which in Britain and Ireland has always been tightly restricted to open,disturbed, artificial habitats. Before the advent of selective herbicides, which over the last 30 yearsor so have almost exterminated the species from farmland, the distinctive red poppy flower was closelyassociated with fallow field margins around arable land, especially where the ground was under cerealcultivation. Nowadays, P. rhoeas is confined to other unsprayed forms of open or disturbed,artificial habitats, such as waysides, quarries, building sites, soil heaps, waste land and on walls(McNaughton & Harper 1964; Grime et al. 1988). The soil which suits Common Poppy best is afairly light, warm, sandy and free draining one with a pH between 6 and 8. The ideal soil is of mediumfertility and moderate to rich in calcium or other bases (Sinker et al. 1985).
Status as an ancient introduction
This ruderal annual species has an interesting and much studied biology, ecology and history. Along withnumerous other arable weeds, populations of which were once considered stable and resilient, P.rhoeas populations have rapidly become impoverished and in Co Fermanagh the species has declinedto rarity. Thus Papaver rhoeas has become of greater interest and concern to someconservation-minded people, while other botanists consider it a mere accidental contaminant of cropseed, albeit that of migrant Neolithic or Bronze Age farmers, 5500-2500 B.P. (Grieg 1988; Kadereit 1990;Wilson 1992). Plants of this origin in the British Isles have recently and belatedly (see eg Stace 1997)been recognised as 'archaeophytes', ie ancient, pre-1500 AD 'synanthropic' introductions (ie associatedwith man). A total of 149 such species are now acknowledged in the flora of Britain & Ireland,including P. rhoeas, by the editors of the New Atlas (Preston et al. 2002), andthis figure is likely to grow.
Unlike other archaeophyte arable weeds, for example, Scandix pecten-veneris (Shepherd's-needle)and Centaurea cyanus (Cornflower), P. rhoeas is not yet a rare plant on an entire BritishIsles basis, but it has already become a rare casual in Fermanagh, there being a total of only twelverecords in the Flora Database, three of which are derived from sown seed mixtures (see details below).On account of its local rarity, the 'Common Poppy' is given more consideration here than might otherwisebe considered appropriate. After all, as Wilson (1992) reminds us, "weeds have accompanied thesecultivated annual grasses [he is here referring to cereals] since their wild days, and in a very realsense, the history of weeds is the history of mankind".
Geographical and cultural origin
Along with three other poppies which are also archaeophytes (ie P. dubium (Long-headed Poppy),P. lecoquii (Yellow-juiced Poppy) and P. argemone (Prickly Poppy)), Common Poppy probablyoriginated in SE Europe. The fossil history of P. rhoeas stretches back to middle and lateNeolithic times in C Europe, and to the late Bronze Age in the British Isles (Godwin 1975). Kadereit(1990) examined the distributions of these poppies and compared them with their nearest 'wild' (ienon-synanthropic) relatives. His study suggests that P. rhoeas originated in the easterncoastline of the Mediterranean, where it appears to have differentiated from one of three closerelatives, or from a hybrid between them (although without involving any change in chromosome number).It probably did so only after disturbed and cultivated ground in sufficient extent was provided by manas a habitat. Thus it seems possible that not only was P. rhoeas widely distributed by man withhis cereals, but that as a taxonomic entity, the species may also be of synanthropic origin (Kadereit1990).
P. rhoeas is chiefly a summer annual, a lifestyle which in itself might suggest acircum-Mediterranean origin, although in mild areas of the British Isles or in exceptional winters withlittle severe frost, autumn germinated seedlings may sometimes survive (Greig 1988). Summer annuals likeP. rhoeas are capable of maturing and flowering after a short spurt of rapid vegetative growth inthe spring, or failing this, flowering even when the plant is very small. Having this short, orrelatively short, growing season, poppy plants are certainly favoured by our mild, damp oceanic climate,rather than a hot, dry continental one, which curtails their growth and development during the summermonths (Greig 1988).
Fermanagh occurrence
For this weedy species, there have only ever been a total of twelve records spread over seven Fermanaghtetrads, four of which have post-1975 records. Until RHN discovered an individual plant flowering onroadside waste ground near Gublusk Bay, Lower Lough Erne in September 2003, Common Poppy had not beenseen 'wild' anywhere in the county since 1950. However, there were two instances of it beingdeliberately sown in so-called 'wild-flower' seed mixtures in 1996 and 2003 in garden and in roadsidesettings. Since this very colourful and conspicuous species is well known to be capable of persistingburied and dormant in the soil seed bank for at least 80 years, and probably for more than a century(Salisbury 1964), it is always worth looking out for it on any freshly cultivated or recently disturbedpatch of ground.
Before RHN, the current author and our contemporary botanical associates began recording in the early- tomid-1970s, agricultural weeds and plant introductions of any type or origin tended to be completelyignored by previous field workers in Fermanagh. Apparently earlier recorders considered themunimportant, or at least a waste of time in comparison with the need to record more obviouslysignificant native plants, ie definite or perceived native species.
The details of the other eleven existing Fermanagh records are: football club car park, Newtownbutler,1930-40, R.C. Faris & J.M. Cole; old railway station, Florencecourt, 1950, Meikle et al.;planted wild flower seed mix in garden near Levally House, 1 km SE of Roosky, October 1996 & October1997, RHN; roadside waste ground, Gublusk Bay, Lower Lough Erne, September 2003, RHN; sown wild flowermix, at road junction of Sligo and Ballinaleck Roads, Enniskillen, September 2003, RHN; one plant onQueen Elizabeth Road, Enniskillen, 17 July 2006, RHN; one plant in a garden at Goblusk Bay, Lower LoughErne, 14 August 2009, RHN; Clabby village, 26 June 2011, RHN; and Riversdale Forest, 29 June 2011, RHN.
Factors limiting distribution
Unsuitable soils
Apart from coastal situations, warm, sandy soils with the particular micro-environment that suits thispoppy are scarce in the north and west of Ireland, in Scotland and on higher ground anywhere throughoutthe British Isles. Most of these geographical areas combine a geological structure that producesstrongly acidic, high-silica soils that are frequently covered with a layer of nearly impervious glacialtill of similar rock mineral origin and chemistry. In addition, lying close to the Atlantic seaboard,many soils in the more westerly parts of Britain and Ireland have suffered high levels of precipitationthroughout the entire 11,000 years of the post-glacial period. The subsequent percolation of rainwaterhas resulted in soil mineral leaching, acidification, gleying and podsolisation, sometimes leading tothe formation of huge swathes of organic blanket bog peat. The resultant lack of suitable soils makes itunsurprising that P. rhoeas is a scarce or casual species in the most northerly and western partsof Britain and Ireland.
Decline of tillage
The scarcity of this previously common weedy species in the north and west of Britain and Ireland wasproduced and perpetuated when the low intensity arable agriculture that previously allowed the CommonPoppy's annual seed population renewal, gradually became increasingly uneconomical. Arable cultivation,especially of cereals, continued to decline further after the Second World War, eventually reaching avery low level, or near extinction.
A measure of disturbance
All poppy species, including P. rhoeas, require open, sunny habitat conditions, with bare patchesof soil for seedling establishment. Thus poppies are rarely found in closed, grazed turf. If they are,then disturbance such as grazing and trampling must be sufficient to minimise competition fromneighbouring plants, and provide sufficient openings in the soil surface for this ruderal species togerminate and establish. At the same time, the foraging disturbance must be limited to allow the speciessufficient time to grow, flower and set seed successfully.
Toxicity
In order to limit grazing pressure, P.rhoeas contains a number of toxic or narcotic alkaloids, egrhoeadine and rhoeagenine, which give off an unpleasant odour and make the plant unpalatable orsickening to browsing animals. There are no recent cases of animals being poisoned by P. rhoeasin the British Isles (Grime et al. 1988; Cooper & Johnson 1998).
Phenotypic variation and reproductive capacity
All Papaver species in Britain and Ireland are very plastic in their phenotypic response togrowing conditions. They are particularly sensitive to shading, nutrient deficiencies, lack of water, oran excess of it, and interference from non-related species. All of or any of these variables candrastically reduce the poppies growth and reproductive capacity. P. rhoeas is exceptionallyplastic in these respects, and McNaughton & Harper (1964) described finding depauperate plants insoils of very low fertility that were only 5 cm tall, bearing a single capsule containing four or fiveseeds. At the other extreme, under favourable conditions, an isolated individual might bear 300 to 400large capsules, each containing more than 1,000 seeds (McNaughton & Harper 1964; Harper 1966).
An unusual feature of P. rhoeas variation is that flower size is also among the characters whichare very plastic. Nutrient or water deficiency results in reduction of petal size, number of stamens,ovary locules and seeds. When species are as variable as poppies are, mean figures for properties suchas seeds/capsule, or per plant unfortunately become rather meaningless, so I will refrain from quotingany such figures (Salisbury 1942; McNaughton & Harper 1964).
Flowering and pollination
The flowering season of P. rhoeas is longer than for other poppies that occur in Britain andIreland. Although flowering begins rather late in the season in mid-June, and it peaks in early July,flower production still straggles on intermittently until October. The scarlet blooms attract a widerange of insect visitors, offering no nectar but easily accessible pollen in vast quantity. Honey beesare probably the most significant cross-pollinators. The flowers are outbreeders and are almost entirelyself-sterile. Self-sterility is unusual in poppies and, indeed, this is true for successful annual weedsin general (Baker 1974). For further comparisons and discussion of this matter, please see the followingP. dubium species account. Self-incompatibility in P. rhoeas is controlled by a complexmulti-allelic gametophytic system; those wishing to know more about this topic are recommended toconsult Richards (1997), Chapter 6, for a technical explanation of this rather involved geneticmechanism.
Seed dispersal
Seed ripens rapidly and is shed within four weeks of flowering, being quickly shaken out of the largepores at the top of the capsule which is borne on a long, swaying elastic stalk. This is often referredto in textbooks as the 'censer mechanism' of dispersal (McNaughton & Harper 1964). Dispersaldistance was measured in a German experiment which purported to represent "near-naturalconditions". However, the experimental setup could be regarded as rather artificial, since itinvolved dispersal over carefully maintained constantly bare ground. In any event, measurement showedthat the great majority of seed travelled less than 100 cm, and the maximum distance travelled was 3.5 m(Blattner & Kadereit 1991).
Competitive ability
A range of competition experiments within and between different species populations has clearly shownthat P. rhoeas reacts differently and in a complex manner to different sorts of neighbours −sometimes by mortality, sometimes by plasticity (Harper 1966). Generally, P. rhoeas is a weakaggressor when it comes to plant competition, but it is a "magnificent opportunist" (Harper1966, p. 34). Common Poppy reacts rapidly to fill any vacancy in the environmental 'space', should somefactor, eg disease, or selective browsing by an herbivore, happen to weaken neighbouring plant species.It is especially exploitive of conditions providing such openings when high nutrient levels areavailable (Jonsell et al. 2001). The competitive situation should be visualised, however, as aform of balance between coexisting species, rather than as one species replacing another, sincemortality and competitive exclusion are not always involved. From this standpoint, plant species areoften more sophisticated than we tend to allow. Biological scientists of most disciplines nearly alwayswant to simplify and reduce the subtle patterns we find in nature. Plant (and animal) species do notbelong in any fixed order of merit; "common species have their peculiar biology satisfied infrequent places; rare species may be superbly adapted to rarer sites" (Harper 1966, p. 35).
Irish occurrence
The New Atlas map shows that in Ireland, P. rhoeas is much more locally frequent toabundant and continuously distributed SE of a curve linking Newry in the north with Cavan, Longford andAthlone in the Midlands, and running south to Cork City. Elsewhere in the Republic of Ireland (RoI),P. rhoeas is now a very rare and scattered casual, while in Northern Ireland (NI) it is currentlyvery thinly and widely distributed. The number of pre-1970 symbols on the hectad map indicates that ithas declined from being a rather local plant and become a rare casual species (NI Flora Website,accessed 2002; Preston et al. 2002). It might well be argued that P. rhoeas is well downthe road towards local extinction in NI, as Hackney et al. (1992) have suggested for the threecounties of NE Ireland.
The pattern of occurrence in SE RoI does not appear to correlate all that well with geology, soils orclimate (except perhaps a lesser number of rain days than elsewhere). However, it does reflect ratherwell the distribution map of 1970 tillage, and even better the distribution of wheat fields at the samedate. Both of these features are mapped in the Royal Irish Academy's Atlas of Ireland (Haughtonet al. 1979). Unfortunately, the same connection cannot be made in NI, where the speciesdistribution pattern is really very fragmentary and does not appear to correlate or compare with anyrelevant factor, even if we include figures of rural land valuation.
Opposing the observed marked decline in the species occurrence across Ireland, however, two recent P.rhoeas Fermanagh sites arose from a sowing of 'wild flower seed'. This mode of origin is aphenomenon that in recent years has become rather common throughout the British Isles, and it isdistorting the previous 'natural' distribution achieved by the species, derived from agriculturaldisturbance and previous crop-associated sowings. As both forms of the plant's occurrence arise entirelyas a result of human activity, perhaps we should not get too agitated about this new pattern ofintroduction, of what is, after all, a beautiful, easily eradicated, weedy species.
British occurrence
In Britain, P. rhoeas is shown in the New Atlas to be much more common and widespread inthe Midlands and SE England, extending more or less continuously up the E coast to around Dundee.Elsewhere in Britain, it is more scattered and occasional, becoming very rare and local in N Scotland.Being largely associated with arable agriculture, Common Poppy is essentially a lowland species,generally occurring below 200 m. Above this altitude, it is probably more casual than established (Grimeet al. 1988; Preston et al. 2002).
European occurrence
In Europe, P. rhoeas is a common and widespread archaeophyte in W and C regions from theMediterranean north to around 55N. However, it rapidly declines to casual status beyond this in manyparts of Scandinavia. The species just reaches 60N on the Baltic coast of Sweden (Jonsell et al.2001). P. rhoeas is largely absent from E Europe (Jalas & Suominen 1991). As elsewhere,populations rapidly declined in Europe from the 1950s onward due to use of selective herbicides, butsince the 1980s it has recovered somewhat due to fallowing or set-aside (Jonsell et al. 2001).
World occurrence
Worldwide the distribution continues into SE Asia, N Africa, N & S America, Australia and New Zealand(Jonsell et al. 2001).
Uses
Corn Poppy carries with it numerous historical, folklore and medicinal traditions and uses. The seedshave a pleasant nutty flavour, are processed for Poppy seed oil, and are commonly sprinkled on cakes andbread for decoration and flavouring. Parts of the plant have also been used as a source of black dye (Vickery 1995; Grieve 1931, p.651; Le Strange 1977, p.199-200; Mabey 1996).
Names
The origin of the genus name 'Papaver' appears to be obscure but it is a Latin name for a poppy, probablythe medicinal P. somniferum, Opium or White Poppy (Gilbert-Carter 1964). It may have been derivedfrom the Celtic 'papa', meaning 'thick milk', or Latin 'pappa' meaning 'milk', either way an obviousreference to the milky latex members of the genus contain (Melderis & Bangerter 1955; Hyam &Pankhurst 1995). The Latin specific epithet 'rhoeas' used to be the genus name. It is possibly derivedfrom the Greek 'rhoia', meaning or referring to the 'pomegranate', Punica granatum, which both inthe colour of the flower and shape of the fruit, P. rhoeas does indeed resemble (Johnson &Smith 1946; Gilbert-Carter 1964).
Grigson (1987) lists an overwhelming 50 English common names, many of which refer to the red flowercolour or warn of the supposed bad luck or ill health (for example, thunder and headache, or blindness),consequent upon touching the plant. These myths were possibly used to keep children away from thepoisonous plant, or out of the field to prevent trampling the corn (Vickery 1995). The gardeners''Shirley Poppy', which has a wide range of colour shades from scarlet to white, grey and lavender and adouble form, was bred and selected from a sport of P. rhoeas (McNaughton & Harper 1964; Mabey1996; Phillips & Rix 1999).
Threats
None.
Introduction, archaeophyte, a very rare casual. Eurosiberian southern-temperate, but very widelynaturalised in both hemispheres.
1902; Abraham, J.T. & McCullagh, F.R.; around Newtownbutler.
August to September.
Growth form and preferred habitats
As with P. rhoeas (Common Poppy) dealt with above, nowadays the weedy annual P. dubium andits two subspecies only merit the status of rare casuals in the flora of Fermanagh. The same is true inmany other areas of Britain and Ireland where this once common plant has markedly declined, being almosteliminated by the combination of modern agricultural seed cleaning and the use of herbicides. Thestations of P. dubium now tend to be rare or scarce, and generally the plant is transitory andconfined to recently disturbed soils on roadsides, urban or residential areas, disused farmland, loughshores, calcareous screes and gravel pits.
The most recent edition (8th) of the Irish field botanist's An Irish Flora (2012), listssubsp. lecoqii (also regularly spelt lecoquii) on the basis that it differs from P.dubium in containing yellow rather than white latex, and bearing brownish to bluish anthersrather than yellow anthers. However, Parnell & Curtis (2012) remark that "other supposedlydistinguishing characteristics appear to be unreliable". Since the BSBI's New Atlas mapsP. dubium species and its two subspecies separately, we will follow it and list the recorddetails of the eleven Fermanagh records in that manner. There are sufficient similarities between thesubspecies, however, to make it sensible to discuss them together in this one species account.
Soil requirements
The substrate requirements of P. dubium and its subspecies are fairly dry, or at leastfree-draining, light, sandy or clayey soils, which again, as for P. rhoeas, absolutelymust be subject to some degree of disturbance. The brief species accounts in the New Atlassuggest that subsp. dubium favours light calcareous soils and that subsp. lecoquii differsin colonising heavier ones (P.J. Wilson, in: Preston et al. 2002). It is uncertain just howreliable a feature this is, but it might possibly assist identification decisions based on themorphological characters of latex and anther colour mentioned above.
On account of the essential pioneer colonising behaviour of the species, and the necessity of soildisturbance and open ecological conditions for its growth, P. dubium, like Common Poppy, used tooccur frequently in arable land before the advent of herbicides, and previously these two Papaverspecies regularly coexisted.
Variation and identification difficulties
Whatever the ranking of the taxa, the morphological similarity between P. dubium and P.lecoquii is such that most recorders experience great difficulty in distinguishing them, despitethe fact that they do have different chromosome numbers – ie subsp. dubium is hexaploidwhile subsp. lecoquii is tetraploid. Kadereit (1990) advocates their separation mainly by thecolour of the latex, "This is white or cream when fresh, and brown to black when dry in subsp.dubium, but mostly yellow or turning yellow when fresh and red when dry in subsp.lecoquii." However, another study undermines this identification character and echoesStewart's (1888) doubts on the matter in N Ireland. Some forms or races of otherwise P. dubiummorphology have been encountered in Europe that contain yellow latex (Koopmans 1970). The separation ofP. dubium and related taxa is certainly difficult and, from the above, clearly reliance on thecolour of the contained latex colour alone is unwise.
Kadereit (1990) also points out that despite the difference in ploidy levels, hybrids between the twoforms can easily be produced. Furthermore, the hybrid form carries out regular but unequal meiosis,indicating that chromosome hom*ology and pairing does occur. It therefore seems likely that subsp.dubium originated as a hexaploid directly from tetraploid subsp. lecoquii without theparticipation of any other taxonomic form.
Other distributional differences between the forms in C Europe suggested to Kadereit (1990) that subsp.dubium might be a native of SE Central Europe, and subsp. lecoquii of W. Anatolia and SEEurope. This idea is very much built upon his assumption, however, that Papaver species in thistaxonomic section of the genus predominantly have vicarious distribution patterns (ie they occupyseparate areas that do not overlap to any great extent). On the other hand, it is very clear from thestudies of McNaughton & Harper (1964) and of Harper (1966), that the five archaeophyte (ancient,introduced) forms of 'cornfield' poppy which are found in the British Isles, "represent aprogression of more and more northerly distribution, but that there is nothing in the distribution ofthese species which suggests any geographical or a regional displacement of one species byanother." (Harper 1966, p. 27).
Wild and crop weed forms
Is it the case that the wild relatives of our cornfield poppies have separate, distinct species ranges,while the several forms closely associated with man and with cultivation typically co-exist and overlapin distribution with one another? As a measure of the distributional overlap and association betweenpoppy species in S and SE England, McNaughton & Harper (1964) tabulated the number of stations inthe total sample of 75 in their study, where the species co-existed; P. rhoeas and P.dubium did so in no less than 33 of these stations, while in an additional four stations theywere joined by P. lecoquii (McNaughton & Harper 1964, p. 771, Table 3).
Fermanagh occurrence
As the coincidence tetrad map indicates, both poppy species, P. dubium and P. rhoeas, arewidely scattered across Fermanagh, but they display a slight concentration in the SE of the county.There are just six records of P. dubium s.l. in the Fermanagh Flora Database, and a total ofseven for the two subspecies. The details of the other four records of P. dubium s.l. are: pileof topsoil beside a house, Lisnagole, 2 km S of Lisnaskea, 3 July 1995, RHN & RSF; disused farm toSE of Clonmackan Bridge, 25 August 1995, RHN; roadside at Whitepark, 2 km NW of Lisnaskea, 8 September1995, RHN & HJN; roadside N of Killymackan, near Teemore, 25 July 2000 and 11 July 2004, RHN.
The details of the three records of Papaver dubium subsp. dubiumL. are:
- scree slope above Doagh Lough, 1986, S.J. Leach & S.A. Wolfe-Murphy;
- NW shore of Derryvore Td, Upper Lough Erne, 11 September 1986, T. Waterman;
- Gortaree gravel pit, Slieve Rushen, 11 September 1988, RHN, voucher in DBN.
The details of the four records of Papaver dubium subsp. lecoquii(Lamotte) Syme (Yellow-juiced Poppy) are:
- Belleek village, 1950, MCM & D;
- about twelve plants in a field of rye, halfway between Newtownbutler and Galloon Bridge, 1950, MCM& D;
- Enniskillen town, 16 May 1988, RHN;
- roadside E of Killymacken Lough, September 2004, N.F. Stewart.
Northern Ireland occurrence
In the pre-1940 period, in the 2nd edition of the Flora of the North-east of Ireland which coversCos Down (H38), Antrim (H39) and Londonderry (H40) – an area of Northern Ireland at that time very muchbetter recorded than Fermanagh – P. dubium and P. lecoquii were regarded as being aboutequally frequent (Praeger & Megaw 1938). Fifty years previous to this, in 1888, dealing with thesame NE region of Ireland, S.A. Stewart wrote of P. dubium occurring, "on sandy cultivatedfields, and sandy or gravelly waste ground; abundant in Down, less common in [Cos] Antrim andDerry." Sagely, he also commented, "Our plant seems to be var. lecoquii of Lamotte, butscarcely distinguishable. The colour of the sap does not appear to be a reliable guide." (Stewart& Corry 1888). In the latest, 3rd edition of the same Flora, Hackney et al. (1992)wrote of P. dubium agg., noting that it was, "less frequent than before", and was,"recorded from only eight 10 Km squares [ie hectads] in Co Antrim."
Interestingly, in view of the slight tendency for Fermanagh sites of P. dubium to occur in townsand villages, in their survey of the Urban Flora of Belfast, Beesley & Wilde (1997) foundP. dubium was widespread on disturbed and waste ground in the city.
The effect of competition
In the late 1950s, when McNaughton and Harper's pioneering work on plant populations was carried out, itrequired very detailed experimental study and careful analysis to discover that density dependentself-thinning of poppy populations was more intense within a single species than betweenspecies (Harper 1966). This selection pressure, added to the range and degrees of plastic response whichpoppy species produce in many individual characters (including some normally quite conservative floralcharacters) when they are reacting to various forms of environmental stress, taken together often enableor allow plant survival in the population, rather than necessarily always leading to thinning mortality.This in turn then helps us to account for the frequency with which mixed stands of poppy species arefound in the field.
The broad similarity in range of habitats and the weedy characteristics shared by all poppy species,contrasted with the subtle differences in their requirements, their ways of using the environment and intheir breeding systems, is also very striking. This is very clearly reflected in the distributions inBritain and Ireland of P. rhoeas (a strongly self-incompatible outbreeder, extremely plastic inphenotype) and P. dubium subsp. dubium – which is facultativelyself-compatible and almost exclusively inbred, but not quite so phenotypically variable as P.rhoeas (Rogers 1971).
These two poppy taxa are remarkably alike in their ecology, both being opportunist colonisers ofrelatively dry, bare ground, and prior to the use of selective herbicides, both plants were widespreadmajor weeds of annual arable crops in Britain and Ireland. For these reasons, they extensively overlapin their distribution, but P. dubium is the better represented of the two in Wales and furthernorth, and especially so in Scotland. It is also better able to transcend the altitude limit of arableagriculture (Preston et al. 2002).
Flowering
P. dubium flowers from May to July, and the flowers are fully self-compatible, although in anexperimental natural selfing, seed set was measured at only between 20 and 30%. The anthers are bornewell below the level of the stigmatic disk in the flower, an arrangement which clearly assists thepossibility of outbreeding. Even so, Rogers (1969) found that the seed set possible by assistedself-pollination was 41% of the average set in normal 'open pollination' involving insect visitorswhich, in view of the relative positions of anthers and stigma already mentioned, is a surprisingly highfigure. There was, however, great variation in the numbers of seed set as a result of selfing betweentrials, and this may indicate population differences in the degree of inbreeding success that are likelyto be encountered within this species.
In P. lecoquii, the flower's sexual organs are at the same level and self-pollination, commencingin the dangling unopened flower bud, is more likely to be the norm. In this case, Rogers (1969) foundthe average number of seed set by self-pollination in P. lecoquii was 61% of the open pollinatedaverage (Rogers 1969, p. 56, Fig. 1 and p. 59, Table 2b).
Reproductive strategies and genetics of colonising weed species
Success as an opportunist coloniser of bare, disturbed or cultivated ground requires a high degree ofadaptation, and the gene combinations which confer this on plant species are generally maintained bythree inter-related mechanisms: self-compatibility, rigorous inbreeding and low levels of geneticrecombination (Baker 1959). Certainly P. rhoeas does not conform to this pattern of breedingbehaviour, and to a much lesser extent P. dubium and P. lecoquii do not fit it welleither. Rogers (1971, p. 274) argues that while the premium for the opportunist coloniser is onadaptations such as rapid germination and establishment, a short vegetative phase and a highreproductive capacity (ie large seed output), the most fundamental characteristic enabling success inthis particular form of weedy lifestyle is flexibility. This especially means the ability tomature successfully and reproduce, irrespective of seasonal, climatic or soil variations, or indeed theeffects of interspecific competition. Baker (1974) describes this combination of characteristics as,"releasing the weedy species from restrictions on its range of tolerance of abiotic environmentalvariation", and he refers to the ideal weed characteristics as the development of, "a generalpurpose genotype (or genotypes)".
Some degree of genetic heterozygosity permitting the flexibility found in these poppy species (and whichis particularly marked in the strongly outbreeding and ecologically diverse P. rhoeas) would alsobe of great advantage to opportunistic colonising species. Research indicates that the balance betweeninbreeding and outbreeding and how it is achieved is different in each species. For instance, in thecase of P. dubium, the degree of inbreeding has been shown to vary from one population to another(Rogers 1971). It is likely that in some cases, only an occasional outcrossing event is required inorder to maintain the required level of heterozygosity that enables adequate phenotypic flexibility.This is particularly so if the species has both a very high seed output and a long-lived dormant soilseed bank as a fallback genetic resource (Snaydon 1980). All poppy species appear to meet these twocriteria. P. rhoeas is rigorously outbreeding and so far there is no evidence of any breakdown inits incompatibility mechanism. The fact that it has a high degree of genetic heterozygocity is obviousfrom the wide range of variation shown in many different morphological characters. The reallysignificant question we must ask is, How does P. rhoeas maintain a genotype, or more likely, arange of genotypes which confer adaptive advantage in open, disturbed habitats typical of cropsituations, without having to resort to inbreeding? (Rogers 1971; Baker 1974).
Firstly, it should be clear that the characteristics of the ideal weedy species, as delineated by Baker(1974), do not require that the weed population should contain only a single genotype, and individualscertainly do not need to be completely hom*ozygous, even when selfing and inbreeding prevails andselection pressure is at its most severe (Baker 1974). If this is the case, then the same thing mayapply in reverse, ie outbreeding, and the associated chromosomal heterozygosity, need not implyabsolute gene heterozygosity. We can then postulate that over time, selection pressure wouldgradually remove less well adapted recessive hom*ozygotes, reducing the frequency of recessive alleles inthe populations of the particular weed species. In turn, this would also favour a low mutation rate inthe dominant habitat-adapted alleles. Most phenotypes would then as a consequence approximate to theoptimum for strongly selected characters, while still maintaining a higher level of heterozygosity inother characters that are subjected to less rigorous selection pressure (Rogers 1971).
Survival of buried seed
In an experiment studying seedling emergence and seed survival of four poppy species (P. rhoeas,P.dubium, P. lecoquii and P. argemone), seedling numbers of all four were greatestin the first year following their production and burial in the soil. Thereafter, germination decreasedexponentially from year-to-year and while all species had viable seed after five years, P. dubiumshowed the slowest rate of decline in seedling emergence. The numbers of seed which remained dormant andviable tended to be greatest for P. lecoquii and P. dubium (Roberts & Boddrell 1984).It is probably safe to conclude from this, that seeds of P. dubium and P. lecoquii arecapable of remaining viable in the soil seed bank for a long period, certainly for more than five years,and possibly as long as has been shown for P. rhoeas, ie 80 years or more.
Genetics, numbers and survival
The seed output of poppies such as P. rhoeas and P. dubium is so vast, and their seedlongevity in the soil is sufficiently long, that these poppy species generally have an enormous surplusof genotypes available to provide the next generation. The proportion of the seed which produces thenext generation is correspondingly small, and a small proportion of ill-adapted phenotypes, even if theywere somehow to survive and breed, would be unlikely to have any significant effect on the ultimatepopulation size, or on the success of the species as a weed (Rogers 1971).
Pollination and seed dispersal
The flowers of P. dubium and P. lecoquii attract mainly bees and hover-flies which feed onthe plentiful pollen mostly in the early morning. After pollination, the capsule requires five to sixweeks to ripen before the pores open to shake out the seed. The pores are quite large in relation toseed size, but even so dispersal distances are likely to be only a few metres even under optimum windconditions for release and transport. The seed show marked dormancy, with few or none germinating withinthe first months after shedding (McNaughton & Harper 1964).
Toxicity
As with other poppy species, plants of P. dubium and P. lecoquii are undoubtedly able todeter some browsing herbivores by means of their toxic or distasteful alkaloid content.
European occurrence
In Europe, P. dubium is widespread over much the same area as P. rhoeas, from theMediterranean, through W & C regions and with more of a presence in the north, extending on bothshores of the Baltic to 60N and slightly higher than this in on the Swedish shore (Jalas & Suominen1991, Map 1989). According to McNaughton & Harper (1964), the northerly spread of P. dubiumin Scandinavia is due to the discharge of ships' ballast containing the seed.
World occurrence
Beyond Europe, again like P. rhoeas, P. dubium s.l. stretches from N Africa, through SEAsia and eastwards to reach Nepal. As a previously very common crop seed weed contaminant, it has alsobeen widely introduced by agriculture to both N and S America, Fennoscandia, Australia, New Zealand andTasmania (Hultén & Fries 1986, Map 888; Jonsell et al. 2001).
Names
The Latin specific epithet 'dubium' means 'doubtful' or 'uncertain' which in view of the identificationproblems associated with it and P. lecoquii, is all too apt (Gledhill 1985). The name 'lecoquii'is clearly called after someone, and the most likely candidate appears to be Henri Lecoq (1802-1871), aFrench botanist or plant geographer. There is a genus of the Apiaceae (= Umbelliferae) called 'Lecoqia','Lecoquia', or 'Leco*kia' found in the eastern Mediterranean, which perhaps is another such memorial(Willis 1973).
Threats
While both P. rhoeas and P. dubium have clearly declined over their entire synanthropicrange following the advent of modern seed cleaning procedures and the use of selective herbicides, itstill remains possible that these poppy species may produce herbicide resistant forms, as has alreadyhappened in over one hundred other crop weeds (Warwick 1991; Briggs & Walters 1997).
Native, but with the potential for additional garden escapes to occur, occasional. Oceanicboreo-temperate, also widely naturalised.
1904; Praeger, R.Ll.; limestone scree slope above Doagh Lough.
April to November.
Growth form and preferred habitats
M. cambrica is a slender, hairy, long-lived montane perennial with a deeply penetrating tap-rootor tapering root-stock (Butcher 1961). Sometimes, as at the Hanging Rock NR, Welsh Poppy occurs in greatabundance, forming a yellow swathe on rocky slopes under trees, but more usually it occurs as isolatedclumps in slightly damp areas. In Fermanagh and elsewhere in these islands, native occurrences aretypically found on moist, shady, rocky slopes, cliffs or screes, sometimes under trees in woodland orscrub and usually on limestone or base-rich soils (Stewart et al. 1994). The species also occursquite frequently as a garden escape in somewhat disturbed ground sites, and since it is very capable ofnaturalising itself in the wild, distinguishing the 'escapees' from native plants is not always an easymatter.
Flowering
The solitary, fragrant, lemon-yellow flowers are produced from June to August. Like other poppies, theblossom is nectar-less but the flowers offer visiting insects a plentiful supply of protein-rich pollenas an alternative food source. Large numbers of small, black, finely pitted seeds are shaken out of thecapsule and are the only means of reproduction open to the species.
The facts that M. cambrica can become a weed in the garden and that it regularly escapes into thewild, both attest to its great fecundity and manifestly efficient powers of dispersal. It is the easiestMeconopsis species to grow in the garden: all the other members of the genus are Asiatic inorigin and, in the experience of the current author (Ralph Forbes), demand a degree of skill and anelement of luck to cultivate successfully in our climate.
Fossil record
There are no fossil records for Meconopsis cambrica since its pollen is not distinguishable fromthat of Papaver (Godwin 1975).
Fermanagh occurrence
M. cambrica has been recorded from a total of 16 Fermanagh tetrads (3.0%), 15 of which havepost-1975 dates. There are fairly frequent records from localities on the upland limestones, but justfour questionable lowland stations in the VC that might be non-native occurrences. These records fromsuspect sites (eg by roadsides, on waste ground or tips, or at Boho Church) are omitted from theFermanagh tetrad map. However, three of the four sites are in the near vicinity of upland limestoneswhere the species occurs naturally, and it is certainly possible that they may have spread to thesesites naturally. This leaves only the record on waste ground near a bridge on the Ballycassidy River asa very probable garden escape.
N Ireland and the Republic of Ireland occurrence
Elsewhere in N Ireland, M. cambrica is generally considered a rare, native plant. Apart from itsFermanagh occurrence, it has only about nine listed indigenous stations in N Ireland − one in CoLondonderry (H40), two in Co Down (H38) and six in Co Antrim (H39) (FNEI 3). In the Republic ofIreland, M. cambrica has isolated stations on the hills in ten scattered VCs at altitudes from15-500 m, but it is strangely absent from likely sites in counties such as Kerry (H1 & H2), Mayo(H26 & H27) and Donegal (H34 & H35) (The Botanist in Ireland; BSBI Atlas; CenCat Fl Ir 2). It is also surprisingly rare in the limestones of the Burren, Co Clare (H9), whereone would readily imagine the terrain to be ideally suited to it. In this exceptionally well-exploredkarst limestone region on the shores of Galway Bay, M. cambrica was first discovered as recentlyas the mid-1970s. It appears perfectly indigenous in the two known sites there (Webb & Scannell1983; Nelson & Walsh 1991). Elsewhere in the Republic, the New Atlas map indicated thespecies is thinly scattered mainly across the Midlands and the east in non-native stations thatpresumably are of garden origin (Preston et al. 2002).
British occurrence
In Britain, this species is regarded as a scarce native in Wales, N Devon (VC 4) and N & S Somerset(VCs 5 & 6) (Stewart et al. 1994). Everywhere else in Great Britain (from the south coast toShetland), it is considered a very common and widespread garden derived introduction.
European occurrence
M. cambrica is endemic to W Europe and is distributed from W Ireland southwards to the French andSpanish Pyrenees (Jalas & Suominen 1991, Map 2023). It is an established garden escape in otherparts of the British Isles, France and southern Scandinavia, but not truly established in Germany,Netherlands and Switzerland (Jalas & Suominen 1991; Jonsell et al. 2001).
Names
The genus name 'Meconopsis' is derived from the Greek 'mekon', meaning 'poppy' and 'opsis', meaning'like' or 'appearance' (Johnson & Smith 1946; Gilbert-Carter 1964). The Latin specific epithet'cambrica' means 'of Wales', since the first site reference for the plant in the British Isles was,"in many places of Wales", in Parkinson (1640), a fact also commemorated in the English commonname.
Threats
Clearance of the woodland under which it occurs.
Introduction, archaeophyte, a rare or occasional escape from cultivation. Eurasian temperate, widelynaturalised including in eastern N America and New Zealand.
1884; Barrington, R.M.; Old Crom Castle.
April to November.
Growth form and preferred habitats
This conspicuous bright-yellow flowered, short-lived perennial or biennial species has a branching,short, woody rootstock and brittle stems that contain acrid and very poisonous orange latex which burnsand stains the skin when handled (Cooper & Johnson 1998). Possession of latex is a feature sharedwith other members of the family Papaveraceae, but the four-petalled yellow flower, the elongatedpod-like fruit capsule and the almost-pinnately lobed leaves all rather strongly resemble members of theclosely related Cabbage Family, the Cruciferae (or the Brassicaceae as we must now learn to call itunder new taxonomic rules!). Nevertheless, because of other details such as the one-celled capsule,C. majus really is a poppy (Webb et al. 1996).
The widely used English common name 'Greater Celandine' is another source of possible confusion, but intruth the plant is completely distinct from 'Lesser Celandine', Ranunculus ficaria of the alreadymuch too large and varied Ranunculaceae! Adoption here of the common name widely used in N America,'Celandine Poppy', would be an easy way to avoid any confusion (Kang & Primack 1991).
C. majus is typically found in sunny, sheltered, lowland sites, which are kept open by some levelof human-related disturbance. It is always found close to habitation and the gardens from where itoriginates, which suggests that it has only rather limited powers of dispersal. The type of sitesinhabited by C. majus includes roadside verges, hedgebanks, waste ground and rubbish tips. Italso occurs on the tops of old walls, or in crevices in them and even more frequently in the sparseblown soil and litter accumulated along their base. C. majus is essentially a species of moderatesoils with respect to such factors as moisture, fertility, acidic-alkaline reaction (ie pH), but itdefinitely prefers base-medium to base-rich or calcareous conditions (Sinker et al. 1985).
Distribution and variation
The native distribution of Greater Celandine has been obscured by innumerable escapes from cultivationand it is now found in gardens mostly as a weed. C. majus does not show great variationthroughout its total range, but a small number of subdivisions have been made, most notably separatesubspecies in Europe (subsp. majus), C and E Asia (subsp. grandiflorum) and in coastalChina and Japan (subsp. asiaticum) (Jonsell et al. 2001).
Fermanagh occurrence
Greater Celandine is only an occasional or rare species in Fermanagh, there being reports from a total of20 tetrads, 14 of them with post-1975 records. As the distribution map indicates, it is rather thinlyscattered around the county, but chiefly occurs to the east of Lough Erne. While its populations areusually not very large, they may seed themselves freely and sometimes (perhaps rarely) they become wellestablished and long persistent. For instance, the population at Old Crom Castle appears to havesurvived for well over a century.
N Ireland occurrence
Considering the position of the plant in the three VCs of the NE of Ireland (Cos Down, Antrim andLondonderry), Hackney et al. (1992) more definitely stated that Greater Celandine, "rarelypersisted to become truly naturalised".
Fossil record
The few fossil finds of the species in the current interglacial warm period are confined to sites ofhuman settlement and their dates suggest that it was first present in the British Isles in Roman times.We should therefore regard it as an ancient introduction by man (ie a pre-1500 AD 'archaeophyte') (P.J.Wilson, in: Preston et al. 2002). The fact that it has been in use in herbal medicine (and morerecently in homeopathy) for a long time and has been cultivated for this purpose, strongly reinforcesthis argument. However, the fossil record also indicates that Greater Celandine was present in Britainin the previous Ipswichian interglacial, when it must have arrived by entirely natural means ofdispersal without the assistance of man. In turn, this suggests that it might have been able to disperseto Britain, if not to Ireland, under its own power during the current Flandrian interglacial period (inIreland known as the Littletonian) (Godwin 1975, p. 129; Mitchell 1986).
Flowering reproduction
Flowers are produced all summer from May to August. In addition to being visited and pollinated by beesand flies, both selfing and cleistogamy occur (the latter process involves selfing within the flowerbud) (Jonsell et al. 2001). An interesting American study of C. majus flowering behaviourfound that most of the variation in the sizes of reproductive characters occurred within individualplants, instead of among plants, or between populations. Flower and fruit sizes as well as seed numberper fruit declined significantly during the flowering season, while mean seed size per fruit remainedmore stable (although even this normally extremely conservative characteristic dropped significantlyduring the season in one population studied). This suggests that maternal plants may have a strategy forconserving resources by a gradual reduction in the size of some reproductive characters, possibly inorder to prolong the period of seed production (Kang & Primack 1991).
This study also showed that (as expected) larger plants not only produced more flowers and fruits thansmaller plants, but that these organs were also consistently larger. Thus an important generalconclusion was that, in ecological and evolutionary studies, characters such as seed size and numberthat are commonly taken to reflect plant fitness, should not be viewed in isolation from vegetativecharacters. Flower and fruit sizes matter, if the goal is to understand the mechanism of naturalselection in wild populations (Kang & Primank 1991). Those contemplating a study of floweringperformance would do well to read this important paper carefully.
Seed form and dispersal
The elongated capsule contains white-tipped black seeds. The white outgrowth on the seed coat is anutritive elaiosome oil-body which attracts ants (and possibly also snails and birds), which helpdisperse the seed to some unmeasured extent (Ridley 1930, p. 522).
Toxicity and medicinal uses
Greater Celandine contains several toxic alkaloids including chelidonine, hom*ochelidonine, chelerythrine,sanguinarine and protopine. Cooper & Johnson (1998) remark that the plant rarely causes poisoningbecause it is so unpalatable, having an acrid taste, a pungent foetid smell and caustic sap. Despitethis, cattle and horses have been known to browse the plant, and it is particularly dangerous, or evenlethal, when it is in seed (Cooper & Johnson 1998, p. 169).
NONE OF THE FOLLOWING HERBAL USES SHOULD EVER BE PRACTICED WITHOUT QUALIFIED MEDICAL SUPERVISION. Adilution of C. majus sap was formerly widely used for the treatment of sore or cloudy eyes.External application of the sap was also used to treat warts, corns and ringworm, although beingcaustic, it is reputed to damage any skin that it touches. However, Cooper & Johnson (1998) reportedthat repeated external application of the sap failed to cause any skin damage, and they concluded thatthe reputation of Greater Celandine as a severely toxic plant is dubious. A tincture is used inhomeopathy and an herbal ointment is used to treat chronic eczemas. The alkaloid chelidonine affectscell division (like colchicine does) and in Russia it has been investigated for possible use as ananti-cancer drug.
European occurrence
C. majus subsp. majus is widespread in temperate areas of W, Cand E Europe, thinning to both N & S, although it does occur on all the islands of the WMediterranean, and it just reaches 65N (Jalas & Suominen 1991, Map 2028). As is the case in Britainand Ireland, this subspecies is not always considered native on the continent. In Scandinaviancountries, it is regarded by most botanists as an archaeophyte.
World occurrence
It is also present in N Africa (status undetermined), while it definitely is an introduction in both NAmerica (where it stretches across the temperate zone) and in New Zealand (Jonsell et al. 2001).
Names
The somewhat unusually prolonged flowering period typical of the species is said to be reflected in thegenus name, 'Chelidonium', which is derived from the Greek 'khelidon', meaning 'a swallow' (ie thesummer visiting bird), supposedly because the arrival and departure of swallows is thought to coincidewith the flowering period of the plant (Johnson & Smith 1946; Gilbert-Carter 1964).
C. majus has quite a number of English common names: Grigson (1987) lists 14, and Britten &Holland (1886) no less than 19. Several names as usual are merely different dialect spellings, forexample, 'Celandine' and 'Celidony', plus 'Saladine', 'Seladine' and 'Sollendine' (Britten & Holland1886). The name is derived similarly to the Latinised genus name, either from the original Greek or
from Old French, 'celidoine', referring again to the swallow. One legend had it that the mother bird usedthe orange sap of the plant to restore sight to her blinded young (Grigson 1987).
The use of C. majus for cleansing the eyes is certainly older than the explanation or thecivilization of the Greeks, since it was equally a part of ancient Chinese medicine (Grigson 1974). Thejuice of Celandine took away cloudy spots on the eyeball which were called 'kennings' and the plant wasintroduced to NE America under the name 'Kenning-wort' (Grigson 1987). The Americans also call the plant'Swallow Wort', a name known to both Gerard and Lyte (Grigson 1987). The name 'Yellow spit' obviouslyrefers to the sap, and names involving 'wart' remind us of another major use of it, for example,'Wartflower', 'Wart plant', 'Wartweed', 'Wretweed' and 'Kill Wart' (Grigson 1987). Another name in theold herbals is 'Tetter-wort', tetters being some form of skin disease involving a running sore or wound,which the sap was reputed to heal (Prior 1879).
Threats
None.
Introduced, neophyte, a rare garden escape. European temperate, native range restricted to foothills ofSW and C European Alps.
1946; MCM & D; Newtownbutler.
May to September.
Growth form and preferred habitat
A rhizomatous perennial, endemic to the lower slopes of calcareous parts of the Swiss and ItalianSouthern Alps, in its native ground P. lutea is a plant of shaded rocks and screes (Landolt &Urbanska 1989; Grey-Wilson & Blamey 1995). In Britain and Ireland, it typically grows in the mortarof old walls and prefers half-shade. Dainty and undemanding, Yellow Corydalis has been a popularhorticultural choice for walls and beds in shady corners of gardens in the British Isles since the endof the 16th century (Grey-Wilson 1989). The only problem with the plant is its ability once establishedto seed itself prolifically or indeed excessively and become a weedy nuisance (Hansen & Stahl 1993).
Fermanagh occurrence
This conspicuous garden escape has been recorded just six times in Fermanagh and only twice in the past50 years – the last observation having been made by RHN in 1989. The details of the other five Fermanaghrecords are as follows – the first four being made by Meikle and co-workers, the last two by RHN: onwest bridge, Enniskillen town, 1946; ruined house near Sand Lough, 1949; bridge between Belcoo andBlacklion, 1952; waste ground beside Ballycassidy River, 100 m N of road bridge, 9 May 1988, RHN; wallof old Railway Station, Maguiresbridge, 3 September 1989, RHN.
Flowering
P. lutea has an extremely prolonged flowering period (March to November). The flower isself-compatible and is pollinated by bees (Jonsell et al. 2001). Initially the flower is closed,and it opens with a once-only 'explosive' mechanism, similar to that of Gorse, Broom and other membersof the Pea family. All of these plants are pollinated by vigorous bees of sufficient weight to triggerthe flower opening mechanism (Proctor & Yeo 1973, Fig. 58, p. 205).
Seed and dispersal
The seed of P. lutea is rather too large and heavy to be readily wind-borne, but it has anattached white elaiosome oil food body which attracts ants and possibly birds. This probably explainshow it manages to colonise walls (and cliffs in its native area), at heights well above ground level(Salisbury 1964). Ridley (1930) noted it growing approximately 5 m up a wall, and he suggested that antswere responsible for transporting the seed!
Fossil record
I am not aware of any fossil record for this species. I believe that there are very few fossil recordsfor the related Fumaria species, some only at the generic level, and most or all of the recordsare associated with sites of human settlement or cultivation (Godwin 1975).
British occurrence
In England and Wales, P. lutea is fairly widespread and commonly naturalised S of a line betweenCarlisle and Berwick-upon-Tweed. However, N of this region Yellow Corydalis becomes somewhat morecoastal and diffuse, except around the Edinburgh conurbation where it is again more frequent. Since the1962 BSBI Atlas, P. lutea has increased in some areas in Great Britain (calculated ChangeIndex = + 0.59), eg in Wales and SW England (P.J. Wilson, in: Preston et al. 2002).
Irish occurrence
The New Atlas shows the species in Ireland is very much more rare and widely dispersed than inBritain, but as with Papaver somniferum (Opium Poppy), in N Ireland it appears somewhat morefrequent, although still decidedly sparse (Preston et al. 2002). Between them the Irish FloraCensus Catalogue 2nd edition and Reynolds's list of alien plants Cat Alien PlIr indicate that P. lutea has been recorded at least once from a total of 18 IrishVCs (excluding Fermanagh, which has not had records published until now). The great majority of theIrish P. lutea records emanate from the north. I believe that this scattered pattern ofoccurrence simply reflects the greater number of recorder hours spent per hexad in the northern part ofthe island, since this level of recording enables casual species and infrequent garden escapes to bediscovered along with the more common plant species.
European occurrence
In W Europe, P. lutea is widely naturalised beyond its narrow endemic range, but apart from nearNarbonne in S France, it is completely absent from the Mediterranean, as well as from the entire IberianPeninsula. Yellow Corydalis is better represented in Scandinavia than shown by Jalas & Suominen(1991, Map 2059) and, indeed, Jonsell et al. (2001) records that it is spreading northwards incoastal areas of both Norway and Sweden where it was first recorded as late as 1922.
Names
The genus name 'Pseudofumaria' combines the Greek 'pseudo' meaning 'false' and the genus name 'Fumaria',which is ultimately derived from the Mediaeval Latin 'fumus terrae', meaning 'smoke of the earth', iearising from the ground like smoke. This is considered a reference to the diffuse foliage of somespecies of the genus (Gilbert-Carter 1964; Hyam & Pankhurst 1995). The Latin specific epithet'lutea' means 'yellow', being derived from 'lutum', the ancient name of Reseda luteola (Weld),which produces a yellow dye (Gilbert-Carter 1964).
Threats
None.
Introduced, neophyte, a very rare casual, probably accidentally imported with other garden subjects. Veryprobably locally extinct. Oceanic temperate, native range restricted to W Europe.
8 May 1993; Northridge, R.H; derelict garden at Clifton Lodge, near Lisnaskea.
Growth form and preferred habitat
C. claviculata is a delicately stemmed, much branching annual, whose branched leaf-tendrils giveit a decidedly limited climbing, scrambling and trailing ability since the plant usually only extends upto around 80 cm or so in height. In terms of soil preference, it is markedly calcifuge in character.Most unusually for an annual species, its most natural habitat in the British Isles is in shady orsemi-shade sites in rocky hillside glens, or sometimes along drier areas of scrub-lined river or streambanks. In such 'wild' situations, acidic mineral or peaty soils are well-drained by being on fairlysteep slopes. Here, the plant typically scrambles amongst rocks and over other plants in half-shade orin the better light of clearings in deciduous woods, scrub or conifer plantations, especially when theseare protected from grazing and disturbance. Sometimes it may also occur in gorse or other scrubby heathvegetation (again, particularly when they are protected from browsing), and it may occasionally be foundon heathy hillsides or roadside banks that have become infested with bracken colonies or bramble patches(McMullan 1972; Chater 2010). C. claviculata becomes especially conspicuous after the associatedwoody species have been cut or burnt in order to harvest, control or regenerate heath or woodlandvegetation.
Flowering
The plant flowers from June to September; the small flowers, borne in racemes of five to ten, are paleyellow or cream to almost white in colour and they produce nectar which attracts bees. They are,however, self-compatible and probably mainly self-fertilising (ie they are autogamous) (Clapham etal. 1962; Jonsell et al. 2001).
Seed dispersal
The fruit capsule produces between two and four seeds which (depending on the authority cited) may (Stace1997), or may not (Jonsell et al. 2001) possess a fleshy aril or elaiosome food body. The latterattracts ants and possibly other animals and appears adapted to effect some degree of seed dispersal. Aninteresting project would be to clarify the true position regarding the existence of a seed oil bodyand, if one does exist, to arrive at an estimate of the distance ants (or possibly snails, or birds)transport the seed (Ridley 1930).
Fossil record
Again, as with Pseudofumaria lutea, I am not aware of any fossil record for this species. Ibelieve that there are very few fossil records for the related Fumaria species, some only at thegeneric level only, and most or all of these are associated with sites of human settlement orcultivation (Godwin 1975).
Fermanagh occurrence
There is a solitary recent record of this rare casual in the Fermanagh Flora Database as listed above.The overgrown derelict garden of an abandoned dwelling of architectural interest contained aninteresting collection of around 30 weeds, some of which, like this species, are rare, unusual or seldomfound and recorded in Fermanagh, or indeed elsewhere in most of Ireland. Since the Fermanagh record wasmade, the house has been restored and reoccupied and therefore it is very likely that the garden hasalso been brought into cultivation and the species ousted.
Irish occurrence
Mackay (1836) described the species in his book Flora Hibernica as occurring, "on rocks,walls, and on tops of old thatched houses". He mentions stations occurring for instance, "onthatched cabins between Ballinteer and the little Dargle (river)", and by "the wayside"(ie under hedges or on hedgerow banks), "between Dundrum and the Dublin mountains".
At the end of the 19th century, Colgan & Scully (1898) described C. claviculata as occurring,"chiefly in the counties of Dublin and Wicklow, with two out-stations in Waterford andDonegal". Three years later, Praeger (1901), who had meantime found an additional station for theplant upstream from Waterford on the River Barrow in Co Kilkenny (H11), was writing of C.claviculata as being, "a plant of the SE, with one outlying station in Donegal". Withthe addition of two further localities in the south-centre of the island in N Tipperary (H10) (McMullan1972), this still remains very much the position taken in the Census Catalogue of the Flora ofIreland by Scannell & Synnott (1987). The latter reference lists the plant as Corydalisclaviculata, which the authors regard as indigenous and recorded at least once in seven IrishVCs, not at that time including Fermanagh.
The couple of records of C. claviculata that exist in the north of Ireland, including the oldCulmore, Co Donegal record of Hart (1874), and another northern 1988 record in a cleared plantation woodin Co Monaghan (Alan Hill, pers. comm., January, 1990), on examination appear decidedlysynanthropic. They were found growing either close to or upon dwellings, or occurring as gardenweeds or in estate plantation woods. Since the earliest mentioned Cos Dublin (H21) and Wicklow (H20)stations also fit this pattern closely, I believe there definitely is a case for describing C.claviculata as a naturalised, neophyte alien throughout Ireland. It was probably introduced withsoil on the roots of garden plants and plantation tree stocks, most likely during the late 18th andearly 19th centuries.
British occurrence
In Great Britain, C. claviculata is widely dispersed throughout the island and is locally commonin the west. Although it is chiefly a lowland plant, it does reach 430 m in S Northumberland (VC 67).Despite the amount of forestry plantation that has gone on across Britain since 1970, the distributionhas remained stable when compared to that in the 1962 BSBI Atlas. It is, however, better recordednow than previously was the case (Walters & Perring 1962; P.J. Wilson, in: Preston et al.2002).
European occurrence
Beyond Britain and Ireland, the distribution of C. claviculata is restricted to W Europe, ie it isendemic to that region (Hultén & Fries 1986, Map 894). The species extends in a slightly disjunct,discontinuous manner from N Portugal along the Atlantic coastline to W Norway, reaching its northernlimit just beyond 60N. The European distribution closely overlaps that of heathland in W Europe(Gimingham 1972) and the perimeter of both these distributions fits well with Koppen's (1918) 'Cfb', iea constantly moist and mild 'Oceanic' type of climate.
C. claviculata is regarded as indigenous in Denmark and populations are expanding, encouraged bythe plantation of conifers. The species is also considered indigenous in coastal parts of S Norway.Elsewhere in Norway, however, established alien populations are known to occur. Furthermore, in SSweden, Climbing Corydalis arrived in the late 1950s and is now recognised as an established alienvariously transported along with imported timber, with Rubus shrubs, or as a weed amongst otherhorticultural stock. It already occurs in S Sweden as a garden escape (Jonsell et al. 2001).
Names
The genus name 'Ceratocapnos' is a difficult combination to decipher, and it does not really appear tomake much sense. 'Keras' is Greek for 'a horn', and 'capnos' (or in Greek, 'kapnos'), means 'smoke'. Thelatter is possibly a reference to its related genus (and family name), 'Fumaria', which is reputed tocomes from the Mediaeval Latin 'fumus terrae', meaning 'smoke of the earth', a supposed reference to thediffuse (perhaps, smoke-like), foliage of some members of the family. Another possibility is referenceto a supposedly smoky, irritating smell the plant is said to give off (Stodola et al. 1992). TheLatin specific epithet 'claviculata' means 'having tendrils', the reference here being to the tendrilsof the Vine and also to the branched tendril resembling a small key (Gilbert-Carter 1964).
Threats
None.
Probably introduced and an archaeophyte, a very rare casual, but possibly under-recorded.Submediterranean-subatlantic.
3 September 1988; Northridge, R.H.; roadside at Glenross Td/Rossgweer Td, N of Killadeas.
Growth form and preferred habitats
This fairly tall, diffuse or scrambling summer or occasionally winter annual has large purple-tippedcreamy white flowers. The New Atlas map and the species account by David Pearman and ChrisPreston in Scarce plants in Britain both indicate that in Britain and Ireland, F.capreolata tends to be much more prevalent at the coast, on open scrub, hedge banks, old wallsand cliffs. It is only occasionally found inland in lowland situations, where the habitats then includearable farmland, gardens and open waste ground. It is, however, less frequently found as a weed ofdisturbed ground than other species of fumitory (Stewart et al. 1994).
Variation
F. capreolata has been split into two subspecies, the one commonly found throughout Britain andIreland is the endemic form, F. capreolata subsp. babingtonii(Pugsley) Sell. The more southerly, continental form of the species is subsp. capreolata, whichis found on the Channel Isles and possibly also on the south coast of England, as well as on theEuropean mainland.
Flowering
The flowers of F. capreolata are self-compatible and they habitually and automaticallyself-pollinate in the bud before they open (ie they are cleistogamous). This floral mechanism ensures anabundant seed set (D.A. Pearman & C.D. Preston, in: Stewart et al. 1994).
Seed survival in the soil
There is very little evidence available on the seed bank potential of the species, but Thompson etal. (1997) list one German reference which suggests that seed survival is merely ephemeral.
Fermanagh occurrence
F. capreolata has been recorded on three occasions in recent years. It was first recorded alongwith F. muralis on disturbed waste ground by the roadside as listed above. The details of theother two records are: in the grounds of the Carlton Hotel, Erne Bridge, Belleek, 12 July 2006, RHN; andS shore of Rossigh Bay near the Point, 2007, I. McNeill.
Status in Britain and Ireland
In Britain, this weedy species is regarded as indigenous, but in Ireland it has long been regarded as aprobable accidental introduction. The beautifully illustrated Fumitories of Britain and Ireland(BSBI Handbook, no. 12) did not appear until the great bulk of the current Fermanagh survey wasfinished, but in it Murphy (2009) refers to this plant as F. capreolata subsp. babingtonii (Pugsley) P.D. Sell.
Changes in distribution
Comparison of the hectad maps of F. capreolata in the BSBI Atlas and the New Atlasindicates that this fumitory's distribution is stable at its maritime sites. However, the same maps andthe prevalence of older record date classes on them suggest that the species is declining at its inlandsites, to the extent indeed that it is probably only an ephemeral casual in the latter (BSBIAtlas; P.J. Wilson, in: Preston et al. 2002). The frequency of occurrence of F.capreolata in Northern Ireland compared to the Republic of Ireland is quite marked in the NewAtlas map, but this probably is an artefact simply reflecting recorder effort.
Fumitories of Britain and Ireland (Murphy 2009) features hectad distribution maps with increasingdot size for more recent records. This visual representation makes it even more obvious that fumitoriesare much more actively sought in some areas of Britain and Ireland than elsewhere.
Names
The genus name 'Fumaria' is derived from the apothecaries' Mediaeval Latin 'fumus terrae', meaning 'smokefrom the earth', a poetic allegory of the way F. officinalis spreads its pale blue-green, diffusefoliage across the soil surface supposedly like smoke when seen from a distance (Grigson 1974). TheLatin specific epithet 'capreolata' means 'with tendrils', although in fact the plant has no tendrils,scrambling and climbing instead using its twisting or coiling leaf stalks or petioles (Murphy 2009;Parnell & Curtis 2012). There are no specific English common names for this species in Britten &Holland (1886), but see entry under Fumaria officinalis. The name 'White Ramping-fumatory' is abook name of recent origin.
Threats
None.
Probable introduction, very rare. Mediterranean-Atlantic, also widely naturalised.
1902; Abraham, J.T. & McCullagh, F.R.; Enniskillen Town.
Growth form and preferred habitats
This very variable, diffuse to robust scrambling annual is closely associated with disturbed arable orhorticultural sites, most frequently appearing alongside other Fumaria species among spring-sowncrops, generally on acidic, free-draining soils. Much more rarely, it may be found on hedge banks alsooffering a well-drained habitat, or on waste ground or roadside verges providing suitable, recentlydisturbed substrate conditions (P.J. Wilson In: Preston et al. 2002). It can sometimes also occurin dry soils at the base of walls. Everywhere it is confined to the lowlands.
Fermanagh occurrence
Until Ian McNeill recorded it in 2000 and 2002, this very variable weedy annual had been distinguishedjust six times in Fermanagh and it had not been listed by anyone since 1949 (Revised TypescriptFlora). This could possibly be due to the marked local decline of arable farming almost toextinction in our survey area or, perhaps more likely, to difficulties faced by field recorders in itsproper identification. This problem is commented on here under F. muralis (CommonRamping-fumitory), the plant with which F. bastardii is most often confused by everyone includingmyself. The publication in 1989 of Peter Sell's account of the Fumaria bastardii/F.muralis complex in BSBI News 51, and his subsequent key to the British and Irishmembers of the genus in the BSBI Plant Crib (Rich & Jermy 1998), has made identificationsmuch more reliable. The publication of the BSBI Handbook Fumitories of Britain and Ireland withits excellent illustrations will doubtless prove invaluable for future identification (Murphy 2009). Thelatter recognises three distinguishable varieties, one of which is endemic in western Britain andIreland, var. hibernica (Pugsley ex Praeger) Pugsley (Murphy 2009). This variety has not yet beenrecorded in Fermanagh.
There are a total of eight Fermanagh records scattered across the lowlands of the VC. The record detailsadditional to the first finding listed above (which incidentally was originally identified as F.confusa Jord.) are as follows – in all cases the sites are only loosely characterised: nearLisnarrick, 1942, R. Mackechnie; Belleek, 1948, MCM & D; Lisbellaw, 1948, MCM & D; DonaghCrossroads near Lisnaskea, 1948, MCM & D; sandy fields below Gortaree, Slieve Rushen, 1949, MCM& D; Killee Lough, Ballinamallard, 2000, I. McNeill; Necarne estate, near Irvinestown, 2002, I.McNeill.
On account of taxonomic confusion and identification difficulties described below in the F. muralissubsp. boraei account, I have decided to amalgamate the recordsfor these two taxa and simply map them as F. muralis agg.
Fumaria muralis subsp. boraei (Jord.) Pugsley, CommonRamping-fumitory
Probable introduction, rare. Oceanic southern-temperate, also widely naturalised.
1942; Mackechnie, R.; near Lisnarrick.
April to August.
Fermanagh occurrence
In the 1962 BSBI Atlas, Fermanagh was shown as being close to the NW limit in Ireland of F.muralis and the New Atlas confirms this impression. It is significant and should be noted thatthis species (or subspecies when listed as subsp. boraei) was recorded only once before 1960 inFermanagh, yet 13 times subsequently, whereas F. bastardii (Tall Ramping-fumitory) was recordedsix times before 1960, and just twice since (Revised Typescript Flora).
In the most recent (7th) edition of An Irish Flora, the editors remarked that in their view it isdifficult, if not impossible, to separate Irish material of F. muralis from F. bastardii.Irish material of this species aggregate (ie the F. muralis agg.) is extremely variable and itdoes not readily conform to earlier descriptions based entirely on material from Britain. This isparticularly so with the several subspecies of F. muralis and in the light of this experttaxonomic opinion and of my own difficulties experienced when attempting to key out the two forms(especially when using the earlier 6th edition of An Irish Flora 1977 as field guide). Iam certain there has been confusion in the identification of the Fermanagh records for these two commonlarge-flowered (corolla length over 9 mm) fumitories (ie F. muralis subsp. boraei and F. bastardii).
Nevertheless, for what it is worth, in these circ*mstances, RHN and the current author (Ralph Forbes) arelisting in this work the relatively few records we currently have in the Fermanagh Flora Database foreach of these taxa separately. With the above qualification the 13 records for F. muralis subsp. boraei, additional to the first one listed above, are:garden weed, Ballinamallard, July 1984, RSF & RHN; Glenross Td/Rossgweer Td, 3 September 1988, RHN;waste ground at Doon Td, SW of Derrylin, 18 July 1991, RHN; Lackaghboy, near Killyvilly, 10 May 1994,RHN; garden at Magheranageeragh, 1 October 1994, RHN; potato field 300 m W of quarry, Aghakillymand Td,17 August 1995, RHN & HJN; Drumbrughas North, NW of Lisnaskea, 26 April 1996, RHN & HJN;roadside N of Killymackan, 25 July 2000, RHN; waste ground at Tullycleagh, 7 October 2000, RHN; graveltrack at Corraslough Point, 17 August 2004, RHN & HJN; roadside waste ground at Tamlaght village, 3July 2005, RHN; Enniskillen town, 18 August 2008, RHN; and disturbed roadside at Tattynuckle Td, 21 June2010, RHN.
Obviously the situation regarding these fumitories is very unsatisfactory, and the whole genus requiresdetailed local re-investigation. The revision of the genus by Peter Sell in the BSBI Plant Crib(1998) will, I hope, provide a better basis upon which to work, and Murphy's Fumitories of Britainand Ireland (2009) handbook will certainly prove essential in future. The publication of Volume1 of the critical Flora of Great Britain and Ireland by Sell & Murrell (2018) should alsogreatly enable better discrimination of the members of this genus.
Because of the taxonomic confusion and identification difficulties described above I have decided toamalgamate the records for this and the previous species and to map them as F. muralis agg.
Introduction, archaeophyte? A rare casual, but probably over-looked and under-recorded.
1947; MCM & D; a potato field at Donagh Crossroads, near Lisnaskea.
Growth form and preferred habitats
F. purpurea has a loose inflorescence of pinkish-purple flowers, the petals and wings tipped witha darker purple, and it differs from the other large-flowered species, F. capreolata, in the factthat the raceme of flowers is as long as the stalk that carries them, not shorter. Like all the otherspecies of Fumaria, it frequents disturbed, acidic, free-draining soils, and in this case it hasa tendency to appear sporadically at some sites.
Fermanagh occurrence
The above is the only Fermanagh record of this plant (Revised Typescript Flora) and despite itsdate, it does not appear in the 1951 Typescript Flora.
Irish occurrence
As is the case with the slightly more common F. capreolata (White Ramping-fumitory), most recordsof F. purpurea in Ireland tend to be located at or near the coast. It is perhaps best thought ofas merely a casual introduction in all or most of Ireland, occurring chiefly at E coast ports. In manyinstances, it is synanthrophic (ie closely associated with man and with artificial habitats) (NewAtlas).
In Northern Ireland, F. purpurea is the rarest fumitory species by quite a long margin, havingpost-1970 records from just seven hectads in Cos Antrim and Londonderry (H39 & H40) (NI VascularPlant Database).
British occurrence
F. purpurea is confined to Great Britain, Ireland and the Channel Isles (Guernsey), ie it is ascarce endemic species, but at a majority of sites it is merely a casual introduction (Jalas &Suominen 1991, Map 2089; Stewart et al. 1994; Murphy 2009). It is thinly distributed acrossBritain in the whole range of latitude from the Channel Isles to Orkney, but it does not reach Shetland.Again, as in Ireland, it is chiefly but not entirely coastal in its British sites. However, all commentson distribution have to be made guardedly, since very probably this species is overlooked or mistaken bymany recorders and is therefore under-recorded (Preston et al. 2002).
Conservation status
Being endemic, F. purpurea is given a high priority in terms of conservation despite its weedyhabit and its association with agricultural or industrially disturbed ground (Stewart et al.1994).
Threats
None.
Introduction, archaeophyte, rare. European southern-temperate, widely naturalised in both hemispheres.
1947; MCM & D; disturbed waste ground at Derrygonnelly village.
April to October.
Growth form and preferred habitats
An erect to diffuse, much branched annual, germination of F. officinalis occurs in the spring andvegetative development and flowering is rapid in reasonably fertile, preferably light mesic soils, givenopen, sunny, warm growing conditions (Salisbury 1964; Sinker et al. 1985). Having said this, oneof the English common names F. officinalis is 'Beggary', a name similar to 'Beggar Weed', that isoften applied to such plants as Polygonum aviculare (Knotgrass), Heracleum spondylium(Hogweed), Spergula arvensis (Corn Spurrey) and Galium aparine (Cleavers), either becausetheir presence denotes poor soil, or because they are such noxious weeds they manage to beggar thefarmer (Britten & Holland 1886; Grigson 1987).
Fermanagh occurrence
Although this is indeed the commonest fumitory in Fermanagh, it is still a decidedly rare plant in thispart of the world. It has only ever been recorded in 13 widely scattered Fermanagh tetrads, 2.5% ofthose in the VC. Just eleven tetrads contain post-1975 records. Apart from the occasional potato orturnip field, there really is no arable farming carried on now in Fermanagh. Annual ruderal weeds likethis one have become confined to other forms of disturbed soil, eg in gardens, on piles of earth dumpedbeside roadside workings, on waste ground or building sites where they temporarily appear as casuals. AsSell (1991) pointed out in connection with both Fumaria and Papaver (Poppy) species, inthe past agricultural methods allowed weeds to flourish in almost any crop. Nowadays, however, since theadvent of modern arable management techniques, weeds appear only in crops that do not require heavyapplication of herbicides, eg onions and potatoes.
Variation
F. officinalis is variable and plastic with respect to the environment in which it grows, and itis also genetically variable – two subspecies comprising four varieties having been described by Sell(P. Sell, in: Rich & Rich 1988; Sell & Murrell 2018). Stace in New Flora of the BI (1997& 2010) recognises just the two subspecies, as does Murphy (2009) in her Fumaria handbook.Our local Fermanagh form of the plant is probably subsp. officinalis, the most widespread form inboth Britain and Ireland, but subsp. wirtgenii (W.D. Koch) Arcang. may also be present here. Aplant collected at Lisnaskea was keyed down to this by RHN in 1988 but, unfortunately, no voucher waskept. This form has also been found at least once in Co Londonderry (H40) (FNEI 3).
Irish occurrence
The Northern Ireland Vascular Plant Database (2002) shows F. officinalis as having been found in58 hectads, making it the second most frequently recorded fumitory species in the six northern counties.It is led only by F. muralis (Common Ramping-fumitory), although it must be said that we havereservations about the accuracy of the representation of the latter species due to the identificationproblems between it and F. bastardii (Tall Ramping-fumitory).
In the Republic of Ireland, F. officinalis is most frequently found in the Midlands and in a wideplain around Dublin − what was once referred to as, 'the English Pale'. This region represented thebetter farmland of the island that was earliest colonised by the ruling English settlers from the 14thcentury onwards.
Flowering, pollination and fertilisation
Common Fumitory plants flower throughout the summer from June to September. The raceme inflorescenceconsists of between 10-40 (or occasionally more) flowers. It begins life fairly dense and compact, butit elongates as it ages. The flowers are large for a fumitory, up to 8 or 9 mm long, the petals pinkishpurple having blackish purple tips (Jonsell et al. 2001). Like other Fumaria species,F. officinalis normally self-pollinates and self-fertilises. However, the flowers still doproduce nectar and thus they attract flies and bees, which must also produce some degree of crosspollination − if not necessarily cross-fertilisation (Fitter 1987; Richards 1997).
In flower buds and young flowers, the stamens and the solitary style are positioned close together. Asthe flower develops and the style grows and elongates it pushes between the anthers, collecting pollenon the stigma as it does so, thus effecting self-pollination even before the flowers fully mature(Murphy 2009).
The flower has four petals, of which the uppermost one is largest and has a long prominent spur at itsbase. Nectar is secreted into the spur by long backward-pointing processes on the filaments of the upperstamens. The two lateral petals are curved inwards at their margins and are fused together at theirtips, forming a sheath or hood that encloses the rigid style and stigma. The stigma is large and lobedand when mature it is already covered with pollen released from the anthers which wither before theflower opens. Bumble-bees and flies occasionally visit Fumaria flowers, probing for nectar in thespurred upper petal. In doing so, they dislodge the hood: in younger flowers, the anthers have justopened and the proboscis of the insect becomes dusted with pollen. In older flowers, the stigma is fullymature and is receptive to pollen transferred between flowers by the insect visitor.
Having just described insect pollination in Fumaria species, it must be said that the vastmajority of flowers in the genus are self-fertilised. Do you mean self-pollinated? This is undoubtedlyresponsible for the observed myriad of locally distinct forms that occur in Fumaria species, someof which have received taxonomic recognition (Murphy 2009).
Fruit and seed
The fruit is a slightly heart-shaped nutlet, brown and rough in texture, and each contains only a singleoval seed (Salisbury 1964). Although the seed production of this annual is not normally large, it formsa persistent soil seed bank which sometimes allows the species to become a troublesome and abundantweed.
Seed longevity
Seed survival of over 60 years has been reported in soil buried beneath pastures (Chippindale &Milton 1934) and, indeed, 'ancient seed', buried for over 660 years was also found to be viable (Odum1965; Thompson et al. 1997).
Native or alien status
F. officinalis was listed by Webb (1985) along with 40 other species previously assumed to benative in Britain and Ireland which he considered "probably introduced" and whose status hesuggested required further investigation. The Cen Cat Fl Ir 2 has also long regarded F.officinalis as being a probable introduction (Scannell & Synnott 1987). The editors of theNew Atlas having studied the botanical findings of archaeologists have recently accepted Webb'sview of all but seven of his species (and even these are accepted as being a mixture of both native andalien). Furthermore, they have added a further 108 species previously assumed native species to Webb'sshortlist of probable or definite introductions (Preston et al. 2002). F. officinalis isone of the 149 species now understood to be ancient, pre-1500 AD accidental human introductions in bothBritain and Ireland (Preston et al. 2002).
The Northern Ireland Flora Website (2002), shows F. officinalis as being present in 58 gridhexads, making it the second most frequently recorded fumitory species in the six northern counties. Itis led only by F. muralis, although it must be said that we have reservations about the accuracyof the representation of the latter species due to the identification problems between it and F.bastardii −(see the F. muralis species account).
British occurrence
F. officinalis is widespread throughout lowland areas of England and Wales, but in Scotland itbecomes very much more eastern and coastal, although not exclusively so (Preston et al. 2002).
European occurrence
The European range of the species is very wide, indeed it is almost cosmopolitan, stretching from theMediterranean (although much less well represented in Portugal than might be expected), to well withinthe Arctic Circle in Norway (Jalas & Suominen 1991, Map 2096).
Centre of origin and world occurrence
This mainly Eurasiatic species probably originated in and is native of central and southern parts ofEurope, N Africa and W Asia. As a weed it is distributed worldwide, to areas including Fennoscandia,Ethiopia, S Africa, N and S America, Java, Tasmania and New Zealand (Hultén & Fries 1986, Map 900).
Toxicity and medicinal use
All parts of Fumaria species contain poisonous alkaloids such as fumarine, plus fumeric acid,tannins and mucilage, and they have a long history of use for medicinal and cosmetic purposes. F.officinalis is still widely used by herbalists for eczema and other skin diseases, livercomplaints, colic and constipation (Launert 1981; Stodola & Volak 1992; Bartram 1995). Large dosesof the alkaloids can cause severe diarrhoea and even respiratory failure. It should only be takeninternally under the supervision of a qualified medical practitioner or herbalist.
Names
The genus name 'Fumaria' is derived from the apothecaries' Mediaeval Latin 'fumus terrae', meaning 'smokefrom the earth', a poetic allegory of the way F. officinalis spreads its pale blue-green, diffusefoliage across the soil surface supposedly like smoke when seen from a distance (Grigson 1974). TheLatin specific epithet 'officinalis' reminds us of the medicinal use of the plant, it being kept in the'officina', the druggist's or apothecary's shop (Gilbert-Carter 1964). In reference to the Englishcommon name 'Fumitory', Britten & Holland (1886) relate how the old authors (ie of the English andcontinental herbals), believed the plant was produced without seed from vapors or smoke rising from theearth. They also remark that, "it is rather curious that the root when fresh pulled up gives off astrong gaseous smell, remarkably like the fumes of nitric acid, hence probably the belief in its gaseousorigin."
Grigson (1987) lists an additional twelve English common names, several of which refer to the Virgin (eg'Lady's Lockets' and 'Lady's Shoe'). Another very curious name is 'Wax Dolls', which appears to derivefrom the fact that the foliage is rather waxy and difficult to wet, or possibly also from the somewhatwaxy appearance of the flowers, and thus the plant is being compared with the then familiar texture of aVictorian wax doll (Grigson 1987).
Threats
None.
Native, but also planted, occasional, but during this survey, the majority of older trees were killed orvery seriously affected by disease. European temperate.
1900; Praeger, R.Ll.; Co Fermanagh.
May to January.
Growth form and preferred habitats
A tall tree, more or less orbicular in outline, of lowland woods and hedges, most frequent in limestoneareas where in the past it sometimes became dominant. It often occurs around lakes and along rivers andstreams, but also grows below cliffs and occasionally in quarries.
Identification
Wych Elm has leaves which on their upper surface are very harsh in texture because of the presence ofminute, dense, bristly hairs. The specific epithet 'glabra' does not refer to the leaves but tothe bark on the twigs, which remains smooth for many years once it loses its initial downy pubescence(Hadfield 1957). The leaf base is one of the most distinctive vegetative identification features, itbeing very strongly asymmetric so that the ear-like lobe on the longer side actually conceals the short(2-5 mm) hairy leaf stalk or petiole. Before the devastation of Dutch elm disease killed mature Wych Elmtrees, they alone of all elms seeded themselves abundantly every two or three years, the papery circularwinged fruits being efficiently and widely dispersed by wind. U. glabra also differs from otherelm species in failing to sucker, and these characters taken together enable it to be easily enoughdistinguished from the other forms of elm which are very much more critical and difficult todistinguish. The woodland historian Rackham (1980) reckoned that elms are the most difficult criticalgenus in the British flora.
Variation
Stace (2010) mentions that two 'ill-marked' regional subspecies are sometimes recognised: subsp.glabra, with broadly obovate leaves, more southern in distribution, and subsp. montanaHyl., with narrowly obovate leaves, a form that is more typical of the N & W of Britain and Ireland.
Native or introduced status
From the 18th century onwards, U. glabra was frequently planted along with oak and ash in woods onlimestone in Fermanagh, as was common elsewhere in Ireland (McCracken 1971). However, while it isundoubtedly native in some of its more remote and inaccessible upland sites in the county, other treesmust be self-sown from introduced planted stock, with the result that it is no longer feasible todistinguish between the two modes of origin.
Fermanagh occurrence
Whatever its origin, the tree was undoubtedly very frequent and widespread in Fermanagh until about 1990.It was recorded in both woods and hedges, most frequently in limestone areas of the county, often aroundlakes and along rivers and streams, but also in woods below cliffs and in quarries. It has been recordedin 127 tetrads, representing 24.1% of those in the VC, and 121 of them contained post-1975 records.There are just twelve post-2000 records in the Fermanagh Flora Database. The tree was very widespread inlowland Fermanagh, especially about Upper and Lower Lough Erne, but mature trees became increasinglyrare from about 1995 onwards. The fate of these trees can only be reckoned a most dreadful pity, sinceU. glabra is the only undisputed native elm in Britain and Ireland (Mitchell 1996). While itnever had the same dominance in our landscape when compared with the situation in S England, Wych Elmwas still a significant and often very beautiful tree. In the Irish 8th century 'Laws of Neighbourhood'coding, it was given a strong level of protection in the second rank of the woody plant usefulnesshierarchy, being regarded as, "a commoner of the wood" (Nelson & Walsh 1993).
Apart from the very rare isolated trees on remote rocky hillsides or lake shores, all the older specimensof Wych Elm previously recorded in our survey area have now died off from the virulent form of Dutch Elmdisease which hit Fermanagh from the early- to mid-1980s onwards. While many younger Wych elms continueto die, others seem to be surviving and avoiding the attentions of the bark-boring beetle that is thevector carrying the fungal pathogen from tree-to-tree. Younger trees do not possess a sufficiently thicklayer of bark to interest and attract the beetle, and thus for a time they avoid infection.
Dutch Elm disease
At the very end of the 19th century, elm trees in Europe were first noticed to be inexplicablydying. The timing of the tree deaths around The Great War, and the way apparently healthy trees wouldsuddenly up and die in midsummer, led many to assume that the deaths were related to nerve gas used bymilitary combatants. Research led to the discovery of a fungal pathogen, and the ailment was named DutchElm disease because the species involved was first isolated and described by a young researcher in theNetherlands in 1921. When an elm becomes infected with the fungus, mycelial threads invade the waterconducting tissues and eventually block them with fungal by-products (ie gums, plus irregular shapedgrowths of callus tissue (called 'tyloses'), and gas bubbles which break the water columns). When itsxylem conducting tissues are completely blocked, the tree wilts and eventually it dies a slow death.Essentially the tree dies of drought, although fungal toxins, including a type called 'cerato-ulmin',also play a limited but significant part in its demise (Hadfield 1957; Ingram & Robertson 1999, p.136).
An earlier attack of took place in Ireland during the 1930s (in Britain from 1927 onwards). In thatoutbreak, the affected trees often survived the attack, although the fungal infection did spread aroundthe world producing a 'pandemic' disease epidemic affecting elms.
Fossil record – the elm fall
In vegetation history, the famous (or infamous) so-called 'elm fall' or 'elm decline' described how thefossil representation of Ulmus pollen grains preserved in peat and lake sediments in Britain,Ireland and indeed over the whole of NW Europe, suddenly dropped – not quite like a stone – around about5200 years BP. The elm pollen decline was characterised by its abrupt and almost simultaneous occurrenceacross a wide area of western Europe (Smith & Pilcher 1973). The cause of the massive decline in theelm tree population, which these pollen records indicated, presented a major mystery to science 50 yearsago when I was an undergraduate student.
While all research into prehistory inevitably views such distant vegetation changes through a glassdarkly, modern dating methods have tightened up the focus, revising and sharpening the date of the elmpopulation decline and giving us some idea of the duration of the process. A study by Sylvia Peglar inNorfolk places the classic major 'elm decline' at about 5000 BP. The fossil pollen record at Peglar'ssite demonstrates that the impact on the tree population did not produce a sudden, isolated populationfall, but rather it happened over a period of six to seven years (Peglar & Birks 1993). Danish andSwedish workers have put the date of the same phase of elm decline in their areas of NW Europe occurringbetween 5790 and 5745 BP, and they suggest that it took place over a period of about 40 years.Additional, more detailed work carried out in Denmark, identified four periods of elm decline at datesof 6530, 6130, 5870 and 5410 BP, with 5870 BP identified as the 'classic' phase of elm decline. In NEIreland, a study at Sluggan bog has also shown four phases of elm decline occurring after 5050 BP, theclassic decline occurring at 4900 BP (Smith & Goddard 1991).
It now appears clear (although more work will need to be done before the following hypothesis iscompletely proven), that the most likely cause for the sudden and widespread loss of theelm pollen (indicating the death of mature trees) was not so much the arrival of Neolithic farmersclearing the land of forest to make fields (although this was definitely happening and obviouslyit did not help matters), nor a change of climate (ie it was towards the end of the 'Atlantic' period)but, rather, the trigger was a bout of fungal infection, similar in effect to the latest phase of 'DutchElm disease' that infected and killed elms throughout Britain and Ireland in the 1970s, specificallyattacking the genus Ulmus (Molloy & O'Connell 1987; Peglar & Birks 1993; Mitchell 1995;Ingram & Robertson 1999; Pilcher & Hall 2001, pp. 33-35; Hall 2011).
Peglar & Birks (1993) make the perfectly feasible suggestion that perhaps human disturbance prior tothe elm fall weakened and damaged the elm trees, so that they became more susceptible to attack by thedisease-causing organism. They have provided circ*mstantial evidence in the form of pollen and charcoaldata to support this idea. The discovery in Europe of fossil elm timber of mid-Holocene age containingcharacteristic elm bark beetle galleries, and more importantly, the discovery at a site in London (onHampstead Heath of all places!), of wing cases of two individual beetles of the species (Scolytusscolytus (F.)), has proved for the first time that the insect carrier of Dutch elm diseasewas present in England prior to the time of the elm fall (Girling & Greig 1985; Parker etal. 2002).
Apart from the similar pattern of sudden and nearly synchronous elm disappearance across Britain andIreland observed recently and in the fossil pollen record, circ*mstantial support for the hypothesisthat disease was the main cause of the elm decline in the mid-Holocene comes from a fossil studyby Peglar and Birks (1993) based in SE England, which found that Corylus avellana (Hazel), and toa lesser extent Taxus baccata (Yew), expanded after the pre-historic elm fall, just as hazel hasdone in the 1970s and 1980s in British woodlands where Wych Elm declined as a result of the modern Dutchelm disease epidemic (Rackham 1980).
Irish occurrence
Prior to the most recent fungal endemic in the 1960s and early 1970s, U. glabra was very common inN & E Ireland, although as already mentioned, it was not always indigenous. However, at the sametime Wych Elm was scarce to absent in the wetter, more peaty soils of the west and far south of Ireland.
British occurrence
In Great Britain, likewise, U. glabra was ubiquitous throughout England, Wales and most of lowlandScotland, but now most mature trees in British and Ireland have been killed, except perhaps in Scotland(C.A. Stace, in: Preston et al. 2002).
European occurrence
The native European distribution shown by Jalas & Suominen (1976, Map 311), showed U. glabrawidespread over W & C Europe from the British Isles eastwards to the Urals, while in the north itreaches 60N in Russia, and 67N up the coast of Norway. In SE Europe, the distribution also reachesTurkey, and stretches eastwards to the Caucasus Mountains and N Iran (Jonsell et al. 2000). DutchElm disease devastated the elm population of parts of Denmark and S Sweden in the late 20th century, andit eventually reached the Oslo area of Norway in the 1990s.
Names
The genus name 'Ulmus' is the old Classical Latin name for the tree, very possibly derived from, orcognate with, the Celtic name 'ulm', and the Old Norse 'almr' (Johnson & Smith 1946; Grigson 1974).Grigson (1955, 1987) suggested that 'elm' was an Old English borrowing from the Latin 'ulmus', but in avery detailed linguistic thesis, Friedrich (1980, pp. 80-5), showed that the name 'elm' has acomplicated ancestry in three stocks of Western Indo-European language: Italic, Celtic and Germanic. TheLatin specific epithet 'glabra' means 'without hairs', or 'smooth' in this case (Gilbert-Carter 1964).
The English common name 'Wych Elm' bears no connotation of witches, but rather goes back to the OldEnglish 'wice' and 'wic', meaning a tree with pliant or switchy bark, branches and timber, that could beused for weaving. Etymologists claim that elm names like 'wych' in English and 'vjaz' in Russian areclosely linked to words meaning bending, binding and weaving (Heybroek 1993, p. 4-5; Friedrich 1980, p.82-83). Elm tissues were thus used to weave ropes, baskets and other structures. The flexibility to bewoven is also the case for some species of Sorbus, and most notably, numerous Salix treesand shrubs. Friedrich (1980) points out that Baltic and Slavic peasants and earlier tribal peoples ofthe same geographical area, used elm bast for making mats and footwear, customs that he speculates mightgo back to Proto-Indo-European prehistoric times. Nordhagen (1954) mentions German references toelm-bast being used for weaving second-rate (ie rough) textiles. Grigson (1987) lists eight associatedor variant names that refer to these uses, including 'Quicken', 'Switch Elm' and 'Witan Elm'.
Uses
In the distant past, elm was used in many ways other than for timber, knowledge of which has almostcompletely died out in this part of the world. Across Europe for instance, Neolithic farmers cut elmbranches and used the dried foliage and twigs as winter fodder (Nordhagen 1954; Heybroek 1963 &1993). The latter Dutch author also lists other functional uses of the elm, eg as fodder for pigs, or aschopped and soaked bark fed to calves. Elm bark was even used as famine food by humans. Elm, closelyfollowed by ash, was widely regarded as providing the best fodder of all for stock in winter, thebranches being lopped, dried and stored, often by being stuck into the forks of the tree. A farmers'proverb from W Norway sums up the situation as, "Rowan nourishes, Elm fattens" (Troels-Smith1960; Heybroek 1963). Scientific analysis confirms that dried elm leaves have a "starchequivalent" nutritive index of 50 to 64, while the comparable figures for good to very good meadowhay are only 48 to 57 (Heybroek 1963, p. 6).
Apart from the ancient and important historic uses already mentioned, most of which have long since beendiscontinued, mature Wych Elm gave (sadly past tense), excellent, beautiful, durable timber, suitablefor the manufacture of furniture (especially chairs), and even for external windowsills of buildings. Itwas also the preferred timber for threshing floors, for the shafts of carts. Since it was suitablyflexible, elm was also used for bow making - although in England it was secondary to imported yew forthis particular arms manufacture (Grigson 1987).
The Danish archaeologist, Rolf Nordhagen has described how ancient people in Scandinavian countries andin N Russia, when cereals were in short supply made a flour substitute, scraped, pounded, sieved andcarefully prepared from the inner bark of several tree species, of which Wych elm was the very muchpreferred "bread-tree". This practice was regularly revived right up into the twentiethcentury whenever disaster struck the cereal harvest. Nordhagen reports talking to old Norwegians whor*membered bread wholly or partially baked with elm flour, or porridge made from it. The bark wasstripped from young stems, not more than two or three years old, for this purpose (Nordhagen 1954, p.301-303).
It has been found that harvesting branches and young shoots of elm at intervals of less than six yearscompletely prevents the tree from flowering, so that prehistoric human activity could depress the pollenoutput, and hence modify the regional fossil pollen record. It is obvious that extensive reliance on elmbranches and bark would immediately affect the reproductive and ecological success of the local treepopulation, and if the trees were regularly used, or excessively harvested from even once, this wouldseriously affect their survival, and in the longer term it could disrupt the species distribution.Nordhagen instances exactly this happening to U. glabra in some districts of E, W and N Norway.Fear of this outcome resulted in a cultural taboo developing against the felling or over-exploitation ofWych Elm in some parts of Norway, and to compensate for such pressure, elms were often planted nearhouses and they became venerated as an important bread reserve (Nordhagen 1954, p. 303).
At the same time there is strong support amongst palynologists for the belief that mid-Holocene humanpopulations were too small to alone give rise to the nearly synchronous, sudden elm declineacross Britain, which the fossil record demonstrates.
The medicinal use of elm was found by Allen and Hatfield (2004) to be an almost exclusively Irishphenomenon. The commonest use was of the slimy inner bark as a salve for burns and scalds. Thismuscilage was also used for skin problems in general, and more specifically, to treat swellings andsprains. The leaves were sometimes employed instead of the bark for swellings and inflammation. Otherrecorded uses in Ireland included to staunch bleeding, to cure jaundice, and to counteract ulcers,cancer and other "evils". In England, by comparison there were only two folk medicinal uses ofelm; a tea brewed from the wood taken for eczema in Hampshire, and the inner bark crewed raw or madeinto a jelly and used for colds and sore throats by villagers in Wiltshire. The only veterinary usem*ntioned in folklore was for the expulsion of afterbirth in cows, and came from the NE of England andScottish borders (Johnston 1853; Allen & Hatfield 2004, p. 358).
Threats
Dutch Elm disease has destroyed almost all the older trees of this species in Ireland, as it has doneeverywhere in England and Wales. However, there is a degree of resistance within the species, and youngtrees with bark unsuitable for the beetle do escape infection, at least for a time.
Introduced, neophyte, a deliberately planted cultivar, very rare.
1912; Druce, Dr G.C.; planted by roadsides near Enniskillen.
Fermanagh occurrence
This hybrid has only been recorded once in Co Fermanagh (as U. major Smith (= U. ×hollandica Miller = U. coritana Melville × U. glabra Huds.)), by the famous Englishbotanist Clarence Druce. The solitary Fermanagh site is simply recorded by him as, "Enniskillen,County Fermanagh" (Druce 1912), but it is worth looking out for along roads around Enniskillen,where specimens of it might well survive. Mature trees of this cultivar display somewhat greaterresistance to Dutch elm disease than does Ulmus glabra (Wych Elm), our only indigenous Irish elmspecies.
Identification problems
Hybrid elms are very variable and difficult to unravel, and there has been very considerable, yetperfectly understandable confusion between this hybrid and U. ×vegeta (Loudon) Ley, the Huntingdon elm (a single clone, and therefore a cultivar), both ofwhose parent species, U. glabra and U. minor (Small-leaved Elm), are shared byU. × hollandica. Although all U. x hollandica hybridsare called 'Dutch Elm' by some authorities, the cultivar 'Major' is a convenient reference to theparticular type of this natural hybrid that was usually planted in England, and it also provides ahorticultural distinction between Dutch Elm and the Huntingdon Elm, or indeed the Belgian Elm 'Belgica'(More & White 2003, p. 403).
In his account of this hybrid, Melville in Stace (1975) concluded that U. × hollandica (or U. × hollandica'Major') also includes U. plotii in its ancestry, a belief indicated by the smaller leaveswith blunter serrations (R. Melville, in: Hybridization; Mitchell 1996). However, in theirnew book on the subject Hybrid Flora of the British Isles, Stace et al. (2015) regard thisopinion as rather questionable, on the grounds that U. x hollandica is known to beabundant on the Continent and U. plotii is considered endemic to England.
There has also been confusion in nomenclature, as is already indicated by the list of names given inbrackets above. In his posthumously published book Trees of Britain, the English treeidentification expert Alan Mitchell considered that, "The group name for the many hybrids of thisorigin has, of course, been a botanical plaything." And again, "…there were some half a dozennames in use before Miller’s 'hollandica' was adopted as the group name for all the Dutch elmsand 'Vegeta' was placed among them." (Mitchell 1996, p. 351). The problem of elm identification isencapsulated in several statements made by Oliver Rackham. "There are arguably more kinds of elm inEngland than all other native trees together. Wych-elm (Ulmus glabra) is a 'normal' species; itis not clonal, grows from seed, and coppices. Clonal elms [on the other hand] generate a host of'microspecies', rather as brambles and dandelions do. There are many intermediates and possiblehybrids." (Rackham 2006, p. 29)
Irish occurrence
No thorough investigation of unusual, often small-leaved forms of Ulmus has ever been attempted inN Ireland, and very probably this is the case throughout the whole of Ireland. However, in Cat AlienPl Ir, Reynolds mentions that on the basis of his own observations Dr D. Kelly of TrinityCollege Dublin shares the opinion of the English elm specialist Dr R.H. Richens, who examined the Irishsituation in the 1970s. Both these authorities believe that U. ×hollandica and U. minor s.s. are widely planted elms in Ireland, and that U.procera (English Elm), on the other hand, is a scarce and very local tree here. Nevertheless,the Cat Alien Pl Ir published in 2002 lists a total of only five records for U. × hollandica from three Irish VCs, with dates ranging from 1973 to1986.
On account of the critical identification difficulties mentioned, it is not really surprising that theNew Atlas shows only a couple of Irish hectads with records of U. × hollandica, and there are no records at all of U. × vegeta plotted for Ireland in the atlas. This is the case despitethe fact that the Huntingdon cultivar grows very vigorously without suckering, and it displays evengreater resistance to Dutch elm disease than its already fairly resistant U. glabra parent does(New Atlas).
Anyone wishing to re-establish beautiful and useful mature elms anywhere in Ireland might, in RSF's view,try planting the Huntingdon cultivar, U. × hollandica'Vegeta', or one or more of the new disease resistant varieties being bred at present.
British occurrence
In marked contrast with the Irish situation, elms with this parentage are frequent and widespread inEngland and Wales and especially so in SE England. They occur in ancient woodland, roadside andstreamside copses, in hedgerows and in amenity plantations in both rural and urban settings (Stace etal. 2015). A survey by Jeffers & Richens (1970) concluded that this hybrid was, along withU. glabra, U. minor and U. procera, one of the four principal constituents of theEnglish elm flora. The hybrid is regarded as certainly commoner than U. glabra in some areas of EEngland where this parent is an uncommon tree, and the same applies in the Channel Isles (Stace etal. 2015).
Introduced, neophyte, deliberately planted, occasional, but sometimes mis-identified and very probablyover-recorded. European temperate.
2 July 1986; Brain, P.J.T.; north shore of Lusty More Td, Upper Lough Erne.
January to November.
Fermanagh occurrence
It seems rather strange that there were no records for this tree species before 1986 and that there havebeen quite so many records since. The Fermanagh Flora Database currently contains records from 73tetrads (13.8% of those in the VC). On this basis, English Elm appears to be widely spread across thelowlands in woods along lakeshores and riverbanks, in field hedges and those along roadsides.Nevertheless, for reasons already mentioned in the U. glabra account above, the true identity ofFermanagh's planted elms undoubtedly requires further, more critical investigation. This is particularlythe case for those forms of elm with small leaves, ie U. procera, U. minor (Small-leavedElm), plus the rare or almost unrecorded U. plotii (Plot's Elm) and the hybrid U. × hollandica (Dutch Elm) – all of which have at least a fewFermanagh records (New Flora of the BI; Cat Alien Pl Ir). However, it must be rememberedthat the majority of these trees, if not quite all of them, have subsequently fallen victim to Dutch Elmdisease and died, rendering their identity a purely academic question.
Dutch Elm disease
Even those elms which sucker and manage to re-grow after an attack by the fungus has killed off themature stem, tend to succumb to secondary assaults of the disease when they get to a stem size of around5 cm diameter. As a result they do not get the chance to flower and fruit and, furthermore, the normalleaf dimensions and other required identification characters are generally absent (Flora of CoDublin; Pilcher & Hall 2001). It is also the case that, as with all other suckering species,leaves on suckers, or on epicormic side shoots are extremely variable and they provide totallyunreliable features for identification purposes. Thus while some elm suckers do survive in hedgerows,they really have become impossible to properly identify (Stace 1997, 2010; Parnell & Curtis 2012).
Threats
Dutch elm disease has or will destroy most, if not all of these trees.
Introduced, neophyte, deliberately planted, very rare, although possibly under-recorded, now locallyextinct.
1912; Druce, Dr G.C.; roadside hedges between Enniskillen and Dromore Td.
There is only one field record for this species in the Fermanagh Flora Database, but since it wasrecorded by G. Claridge Druce on a 'motor journey' through the county (Druce 1912), on which he probablywas accompanied by Praeger, we feel it is worthy of mention. The rather vague site is absolutely typicalof the time, being given only as, "Between Enniskillen and Dromore, Co. Fermanagh."
U. plotii is an endemic species of hedgerows in the English southern midlands, although it wasdeclining even before the current Dutch Elm disease pandemic developed (C.A. Stace, in: Preston etal. 2002). Like U. minor (Small-leaved Elm) it is particularly susceptible to the morevirulent form of Dutch Elm disease that arrived in England in 1967 with a cargo of timber from NAmerica.
We believe Plot's elm is extinct in Fermanagh, but we freely admit that all the small-leaved elms wouldhave repaid more time spent on them before the disease more or less wiped them out. The New Atlasmap shows no records at all of U. plotii in Ireland, so this interesting old published recordmust have been overlooked.
Native, common, very widespread and locally abundant. Eurosiberian boreo-temperate, but very widelyintroduced in both hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Synopsis
The Stinging or Common Nettle is probably one of the first plants we learn to recognise as children,unfortunately all too often a lesson we learn the hard and tearful way. As Pigott (1964) rather drylyand somewhat heartlessly points out, "If our early contacts with nettles lead us to dislike them,we also acquire a respect for one of the few plants which makes its presence felt." Mechanicaldeterrents like prickles and spines we can see and avoid, but Urtica dioica (and its closerelative U. urens (Small Nettle)) are the only hairy plants in Britain & Ireland that punishus for touching them. When touched they administer a painful, hot stinging sensation that can last foran hour or so, sometimes accompanied by a blistering weal. The attack is reminiscent of that of an angrybee or wasp.
Growth form and preferred habitats
This dioecious, rhizomatous and stoloniferous perennial is represented in a wide range of damp, disturbedhabitats from woodland to fens, ditches, along the banks of rivers and streams, in hedgerows, roughgrassland, including on roadside verges, plus numerous moderately disturbed sites associated with man,his habitation and grazing stock. This includes any ill-managed or unmanaged waste ground, especiallywhere materials of any sort are stacked or heaped, or where rubbish is discarded (Bates 1933;Greig-Smith 1948; Godwin 1975).
On deep, damp, nutrient-rich soils in unshaded sites, Common Nettle forms large, conspicuous, dominant,clonal patches, often around 150 cm tall, but occasionally forming a canopy up to 255 cm in height underoptimal conditions (Oliver 1993a). Under drier, more shaded, or less favourable nutrient conditions,nettle plants may only achieve a height of around 30-50 cm and they display very much lower reproductiveand competitive vigour than their taller counterparts. Established tall-growing nettle patches oftendominate sites that are naturally well placed to receive nutrient inputs, eg at the base of slopes oralong waterways. These clones are sometimes quite old (possibly 50 or more years in age). Indeed somestands, in old, long-established woodland, could even prove to be of ancient origin.
Common Nettle occurs on almost any soil except waterlogged ones, but it is only very rarely found on acidpeat, or on heavily disturbed, regularly cultivated or frequently mown ground (Bates 1933; Greig-Smith1948). Excessive repeated disturbance is the best method of extirpation of this terrible weed.
Fermanagh occurrence
Common Nettle is the ninth most widespread vascular plant in Fermanagh being present in 491 tetrads, 93%of those in the VC. In tetrad frequency it ranks ninth, just behind Angelica sylvestris (WildAngelica) and immediately ahead of Ranunculus acris (Meadow Buttercup). In terms of recordnumbers, it is ranked much lower however, being the 29th most frequently recorded species in the VC.
British and Irish occurrence and status
The BSBI Atlas and the New Atlas both demonstrate that U. dioica is abundantthroughout Britain and Ireland, being absent from only a few hectads on high and boggy ground in the N& W of Scotland. A Common Plants Survey organised by the conservation charity 'Plantlife' in GB andcarried out for the first time in 2000 by volunteers (albeit on a small scale and totally unscientificin terms of coverage), nevertheless found that U. dioica was the most frequently recordedvascular plant in the country, being present in 93% of the sampled locations (Harper 2001).
U. dioica is probably definitely native only in fen carr alder-willow scrub and in associated'tall herb' wetland communities. It might possibly also be indigenous in other forms of woodland,particularly under ash-alder, blackthorn scrub, or oak in some regions, although even here there maystill sometimes remain an association with past human habitation (Tansley 1939; Rodwell et al.1991a). Very occasionally, in decidedly wet or constantly humid environments, Urtica seedlingsmanage to colonise soils, and in similar conditions they may also grow in crevices on sheltered oldwalls or even as an epiphyte on trees. Otherwise we may safely consider Common Nettle to be a pronouncedfollower of man, occupying damp, fertile, moderately disturbed sites.
Common Nettle nutrition
U. dioica often grows luxuriantly around farms and houses, particularly beside gateways whereanimals wait to be brought in, along roadside verges where they are driven, and anywhere manure andexcrement has been heaped. The association of Common Nettle with litter, manure, disturbed ground,habitation and rubble led to debate as to which factor most encouraged the growth of the species.Studies carried out by Pigott and Taylor showed that nettles only really thrive on luxury: when comparedwith many other wild plants they require, like most crop plants, large supplies of all plantnutrients. Although they are especially associated with high nitrogen and phosphate levels, CommonNettles are more 'greedy' than 'gourmet' when it comes to mineral nutrition (Pigott 1964; Pigott &Taylor 1964).
The association of Common Nettle with litter, manure, disturbed ground, habitation and rubble led todebate as to which factor most encouraged the growth of the species. Was the principal factor soil richin nitrogen (Olsen 1921; Tansley 1939, p. 283), or was soil of loose texture – easily penetrated by thecreeping, rhizome or stolon, most encouraging (Bates 1933; Greig-Smith 1948; Ivins 1952)? Pot and fieldexperiments both showed that U. dioica always demonstrates a high demand for nitrogen, but it isat least equally greedy with respect to its phosphate requirement (Pigott & Taylor 1964). The rarityor absence of nettles from very acidic soils is explained by the fact that at pH levels below 6.0,soluble phosphate becomes chemically limited, and therefore is less available to plant roots.Furthermore, below pH 5.5 phosphate becomes almost insoluble, forming chemical complexes with iron, orwith aluminium, which lock it away and render it completely unavailable to plants. In alkaline soilswith a pH above 7.3, phosphate also becomes very tightly locked into calcium compounds, again making itunavailable to plants (Abeyakoon & Pigott 1975).
Soil phosphate availability, plant nutrition and nettle growth rate
Very wet, dry or cool soils also limit the availability of nutrients to plant roots in general, since therelease of minerals from organic sources by the activities of micro-organisms slows down under suchcirc*mstances. Phosphate availability to plant roots is optimal between pH 6.0 and 7.0, and it isgenerally at a maximum at pH 6.5, but in reality other factors, eg the proportion and type of clayparticles and the organic content of the soil, complicate the availability of all nutrients, and indeedthese factors create a state of ever-changing flux in the soil-plant interface. This makes phosphatepositioning in the soil important, since it does not migrate readily, or travel more than a fewcentimetres in the soil solution towards plant roots. On the plus side, this means that phosphate is noteasily leached out of the soil, as happens with nitrogen, potassium and other more soluble, mobile plantnutrients. Finally it should be realised that the pH at the root surface can be significantly lower thanthat of the surrounding soil: organic acids are excreted by the roots themselves or by the bacteria andfungi in their rhizosphere (ie the zone around the root surface), and this also affects plant nutrientavailability and obscures our understanding of it (Abeyakoon & Pigott 1975).
Growth of U. dioica colonies, eg in typical woodland soils, often becomes limited by the speciesreaching a point where it is unable to absorb any additional phosphate for its needs from the verydilute soil solutions that often occur in such situations. Pigott (1964) found this behaviour contrastedvery strongly with that of other woodland species such as Mercurialis perennis (Dog's Mercury)and Deschampsia cespitosa (Tufted Hair-grass), which were much more able to take up phosphatefrom low soil concentrations. Healthy and vigorous plants of M. perennis may have as little as 40mg of phosphorus per 100 g of dry-leaf tissue, while experimental pot-grown plants of Common Nettle showdeficiency when the concentration on the same basis is around 200 mg. He found that wild grown nettleplants often contained as much as 700 mg phosphorus. This may to some extent also reflect a differencein phosphorus metabolism between the species (Pigott 1964). Most soils supporting more or less 'naturalvegetation' in Britain and Ireland, when judged by the performance of nettles, are phosphate deficient.However, the plants that naturally grow on these soils are obviously able to tolerate low phosphateconcentrations, and often they do not respond to experimental additional supplies of the compound.
The soils from habitats where U. dioica appears to be most likely native are usually very rich insoluble phosphate, eg moist soils in lowland alder woods, often beside streams or rivers. On the otherhand, disturbed sites, especially those associated with past or present human habitation, are alsophosphate enriched, due to the presence of the chemical compound in bones and other forms of discardedfood waste. Rorison (1967) showed that additional calcium reduced the permeability of Common Nettle rootcells to phosphorus, and that relative growth rates of the nettle and other species are correlated withsoil phosphate levels. At low levels of phosphate availability, U. dioica shows deficiencysymptoms and grows poorly and slowly (Nassery 1970).
Phosphate deficiency is most obvious in unfertilised upland rough pastures, particularly in the N and Wof Britain and Ireland where the majority of soil parent material is highly siliceous, and where strongleaching occurs due to high levels of precipitation. Together these factors produce nutrient-poor,strongly acidic substrates unsuitable for U. dioica growth. Young stock farm animals, whichrequire a large amount of phosphorus for their bone growth are often grazed on such ground. When theyare eventually marketed, their removal from the land represents an export of phosphorus from the soil.
Severe phosphate deficiency has an important indirect effect on the supply of nitrogen in the soil, andthis can also significantly affect the distribution of U. dioica. Around sites of humanhabitation soils are always phosphate enriched due to the incorporation of bones, crop-derived manureand household waste, including faeces. For this reason, Rackham (1990, p. 136) described man as a"phosphate gathering animal", and it explains why nettles are so commonly associated with farmbuildings. Free-living, soil borne nitrogen-fixing Azotobacter bacteria appear to have a highphosphate requirement, and clover or other legumes, which have captive nitrogen-fixing bacteria in theirroot nodules, are often sparse or even absent from phosphate-deficient pastures (Pigott 1964).
European ecological studies of U. dioica, reviewed by Srutek & Teckelmann (1998), have shownthat increasing levels of nitrogen fertiliser up to 240 kg/ha always increased both leaf and shootbiomass of the species. At high levels of nutrient supply there is a shift in assimilate partitioning inCommon Nettle towards additional leaf production (Weiss 1993). On the other hand, self-shading givesrise to permanent leaf abscission, a phenomenon observed in U. dioica stands in the wild. Theyoung nettle shoot builds a dense canopy as early as the seven expanded leaf stage, but it continues toproduce new leaves and discard older ones as it grows, so that the total leaf canopy of the shoot isreplaced three times during each growing season. Despite translocation of around 60% of the nitrogencontent of the plant, the leaf pool of this nutrient has to be replaced from the soil twiceduring each growing season (Teckelmann 1987). Measurements of this kind underline the exceptionally highnitrogen demand U. dioica makes on the soil for this very soluble and mobile chemical element.
Are nettle colonies an ecological invasive problem or not?
The striking question as far as the widespread distribution and abundance of U. dioica inFermanagh is concerned, is how to relate this and the species' high levels of nutrient demand to thevery widespread infertile, gleyed or shallow calcareous ranker soils that are so characteristic of largeareas the county (Cruickshank 1997, 2012). Clearly there must be very many local soil pockets ofsufficient fertility to support Common Nettle, or it simply would not be present to the extent that itis. Have changes in pastoral agriculture practices over the last 50 years or so, involving impressiveadditional local drainage measures plus the massively increased use of fertilizers, both artificialchemical and even more commonly, the widespread use of organic manure and slurry, resulted in recentgreatly augmented soil nutrient status throughout the county? Upland afforestation and the use offertilizers and lime on the more acidic peaty soils where this activity typically occurs, would alsopoint in the same direction. Would the increase in soil fertility be localised, or could run-offnutrients become sufficiently widespread and available to benefit weeds like U. dioica to theextent that it spreads into more than 90% of tetrads in the county? In other parts of Britain andIreland, downwind of large-scale industrial regions, airborne nitrogen-rich pollutants might also assistnettle growth and perhaps enable the species to spread (eg in the Ochil Hills, Clackmannanshire (VC 87),Welch 2000). Although this factor (while present worldwide) is well recognised, it is most unlikely tobe significant in Fermanagh or in other areas of W Ireland where the prevailing wind and weather isdominated by the proximity to the Atlantic ocean.
I believe that, while undoubtedly there have been increases in soil fertility, and particularly innitrogen and phosphate levels in Fermanagh and elsewhere in Ireland, in general the soil fertilityeffects derived from the atmosphere are confined to soils of less extreme pH, to lower slope and valleyground levels, and they would be much too temporary in their duration to drastically alter plant growingconditions throughout our regional area. We might concede that U. dioica should have benefittedfrom such fertility factors to some unmeasured extent, and probably it has become taller, andpossibly more abundant and dominant in its existing lowland sites. However, the county is notoverrun with a plague of Stinging Nettles, and in numerous tetrads where infertile soils are very muchthe norm, the species remains scarce, sterile and depauperate, and it has to be actively searched forwhen plant recording!
Winter vegetative growth and horizontal spread
Irrespective of the height to which nettle stems grow locally, they die down every December after thefirst really hard frost of the winter, sometimes leaving slender, bare, grey ghost shoots standingthroughout the remainder of the season. While frosty midwinter conditions halts the vertical growth ofthe nettle, it does not prevent continuing horizontal growth of its rhizomatous underground stems. Thesespread out under the loose but bulky sheltering soil litter layer, formed by the fallen leaves andwithered shoots of the previous year. The buried, creeping rhizomes send up cream, pink or redsurface-running stolons, and in milder spells of weather during the four coldest winter months,individual stolons can achieve a total horizontal spread stretching to somewhere between 10 and 120 cm.The stolons branch frequently and root freely at their numerous nodes (Oliver 1993b, 1994). Fieldmeasurements show that an overwintering individual nettle plant can spread up to 2.5 m in diameter fromits original starting point entirely by this vegetative means (Oliver 2001).
In contrast to the appearance of the slender stolons, older portions of rhizome and deeper, establishedroots are covered with an extremely distinctive bright yellow, furrowed corky layer. With increasingage, these tissues become rather woody and mechanically very tough (Olsen 1921). Given time, the rhizomematerial in soil can become very extensive, eg a riverbank study in England which sampled one squaremetre of soil unearthed a total rhizome length of 63.41 m (K.G.R. Wheeler, pers. com., 1995, quoted byOliver (1997)). Similar rates of midwinter horizontal stem growth and creeping dispersal are possibleunder any form of sheltering and insulating debris, organic or mineral. All winter extensiongrowth of this type consumes stored energy present in the rhizome and root tissues (Bates 1933; Oliver1993b, 2001).
Spring growth
Fresh green vertical shoots, generally unbranched, arise from the rhizome and stolon system, usuallyappearing in early March. In 2004, which was an exceptionally mild winter, these young shoots firstappeared in mid-February. The leaves on the early shoots are extremely variable in size, shape anddegree of hairiness, yet because of their unforgettable stinging ability, we quickly learn to recognisenettle leaves in all their guises
The roots usually lack mycorrhizas, and Abeyakoon & Pigott (1975) found none in over 20 root systemsthey sampled from natural habitats. The aerial stems are generally unbranched, although often later inthe season some lateral branches may be produced towards the top of the stems. This typically occurs asa response to frost injury of the bud at the stem apex, or as a result of other physical damage,including trampling or browsing when the stems are young and still palatable to animals (Greig-Smith1948).
Nettles spread both vegetatively, by means of its underground rhizome and overground creeping stem, andalso by seed production. The annual spread of nettle rhizomes is between 35 and 45 cm but, in addition,the rhizome also branches frequently, enabling quite rapid and effective colonisation and almostsimultaneous dominance by the tall aerial stems to take place (Salisbury 1942, p. 216-7).
Flowering
The age of the plant at first flowering is not known, but it does not flower in the first year(Greig-Smith 1948). Typically, flowering occurs from late May or early June through to September. Sincethe species is normally dioecious (ie having separate male and female plants), some colonies may beunisexual. Some of these unisexual colonies might perhaps have formed from a single individual, so thatseed production is sometimes localised. However, U. dioica has been widely introduced to manycountries, and as a result, gene exchange has taken place with other closely related Urticaspecies. The increased genetic variation that this gives rise to means the traditional taxonomicdistinguishing characters between species have become unreliable. Sometimes the variation is so greatthat only quantitative differences can be made between forms, and the species complex becomes unclear,with transitional forms and numerous named varieties being proposed (Hultén 1974, p. 294). This applieseven to normally very conservative characters, including those governing the reproductive strategy ofthe species. Consequently, in some areas of the world U. dioica has become so variable thatmonoecious forms occur.
In any event, huge numbers of male flowers are produced, up to about 1,200 per stem node, and they eachrelease their pollen explosively (Hickey & King 1981). Efficient wind dispersal of the vast numberof pollen grains normally ensures adequate fertilisation of the female flowers, each of which produces asolitary oval seed (ie an achene), after obligatory cross-pollination (Greig-Smith 1948).
Seed production, dispersal and longevity
Seed is shed from late June onwards, although often some viable seed still remains on dead stems throughuntil the following January (Greig-Smith 1948). Each achene is enclosed by four roughly hairy (iehispid) persistent perianth segments of the female flower. These structures allow the achene to adherelike a burr to fur, feathers and cloth, and this same structure probably also assists with winddispersal (Ridley 1930). The occurrence of nettles sometimes growing high above ground level on wallsstrongly supports the premise of wind dispersal, but it is possible that seed ingested by birds couldalso be transported to such elevated sites. The fruits are probably quite often dispersed in multiples,since the inflorescence may remain intact and be dispersed as a unit. Dispersal may also occur afteringestion by animals, achenes having been recorded in the faeces of cattle, fallow deer and magpies(Ridley 1930).
U. dioica produces large quantities of seeds (some large clones probably generating billions ofseeds (Bassett et al. 1977)). The seed or achene develops a persistent soil seed bank (Odum 1978,quoted in Grime et al. 1988). This being the case, dispersal of the species by seed and rhizomefragments in transported soil is also highly probable.
Germination
Germination is stimulated by direct sunlight, or in a shaded site by fluctuating temperatures.Germination rates are rather variable, even under favourable conditions, typically falling between 26%and 96% (Greig-Smith 1948). Seedlings are found mainly in the spring on open, disturbed ground,particularly on soils that have previously been very wet (eg areas where puddles have formed)(Greig-Smith 1948; Grime et al. 1988).
Variation and taxonomy
The opposite and stalked, coarsely toothed, rough textured leaves are extremely variable in size, shapeand degree of hairiness. Almost hairless (ie subglabrous) forms also exist (Stace 2010). The slender,tapering stinging hairs are made of silica and are mingled with non-stinging ones on the leaves andstems of the plant, their density being very variable. A rare almost completely non-stinging form ofnettle occurs in Britain. It was at first almost exclusively associated with Wicken Fen and ChippenhamFen both in Cambridgeshire (VC29), but later reported in Berkshire (VC22), Norfolk (VCs27 &28), SEYorkshire (VC61) and Angus (VC90) in E Scotland. In the past, the non-stinging nettle has been given thenames, var. subinermis Uechtr. and var. angustifolia (Butcher 1961; Pollard & Briggs1984a; Cook 1997; Beckett & Bull 1999). Most recently the non-stinging form has been elevated tospecies rank as U. galeopsifolia Wierzb. ex Opiz., by the Russian botanist D.V. Geltman. Heregards the non-stinging English nettle as part of a mainly E & C European segregate which heconsiders "presumably diploid" (2n=26). Geltman regards the much more widespread stingingCommon Nettle, U. dioica, as being a tetraploid with 2n=52 or 48 chromosomes (Geltman 1992). Theexistence of transitional intermediates, however, obliges Geltman to admit the possibility that U.galeopsifolia may not be a "completely good" species in terms of the species conceptof Flora Europaea (Tutin et al. 1, 2nd ed., 1993). Therefore Geltman recognisesthat many botanists may prefer to accord the non-stinging nettle subspecific rank.
Geltman (1992) also suggests that U. dioica is probably of hybrid origin, the likely parents beingU. galeopsifolia (or a species closely related to it) and U. sondenii (Simm.) Avrorin exGeltman, a form which occurs in W & C Siberia and in N Scandinavia (see Jalas & Suominen 1976,Map 323 – where it appears as U. dioica subsp. sondenii).
The stinging hair and its chemistry
The stinging hair or trichome (or "stinging emergence", as E.L. Thurston (1974) prefers to callit after his careful study of its fine structure), consists of a fine capillary tube calcified at itslower end and silicified at its upper end. It is closed at the tip by a tiny bulbous swelling. Thesilica-rich upper portion of the hair is brittle like very thin glass (Salisbury 1964), and the bulb atthe hair tip readily breaks off along a pre-determined line when it comes in contact with skin. Thebreak produces a fine, needle-like point formed by an oblique fracture at a line of weakness in theupper tapering region of the hair. It only requires very slight pressure for this extremely sharp,slender needle to penetrate the skin, and the attendant compression of the unsilicified bladder-likehair base injects the contained fluid into the minute wound (Emmelin & Feldberg 1947).
The burning pain of a nettle sting is so strong that Germans call the plant 'Brennessel','brenn' meaning 'burning, branding or stinging', and 'essel', the equivalent of our,'nettle' (Betteridge 1957; Simons 1992). The proverbial advice to 'grasp the nettle' is good advice,since whenever the nettle plant is handled roughly the hairs tend to be broken lower down rather than attheir tip, and thus they are not so sharply pointed and do not penetrate the skin (Salisbury 1964).
The nature of the sting has been a topic of investigation ever since Robert Hooke examined the hairs withhis microscope in 1665, yet despite a great amount of biochemical and pharmacological research over thepast 120 years, the precise nature of the nettle sting toxin still remains something of a mystery(Thurston & Lersten 1969; Pollard & Briggs 1984b). It is still commonly thought by many membersof the general public that the active chemical producing the sting is formic acid. However, this is nowrecognised as being incorrect. Formic acid is almost certainly absent from the stinging fluid, or elseit is in much too low a concentration to produce the painful stinging effect (Thurston 1974).
The first investigators to use pharmacological techniques involving in vitro bioassays to test theeffects of stinging hair extracts on living systems were Emmelin & Feldberg (1947). Their studyfound a combination of histamine and acetylcholine present, which they concluded produced the stingingsensation, the former irritating the skin and the latter producing the burning sensation. They alsoshowed that acetylcholine on its own had little irritant action, but in combination with histamine itproduced an immediate stinging pain (Emmelin & Feldberg 1947). A few years later, a third substancein the sting fluid was identified as 5-hydroxy-tryptamine (serotonin), which like histamine andacetylcholine is also present in animal tissues and causes inflammation and a rash on the skin (Collier& Chester 1956). In animal tissues, these three chemicals are neuro-transmitters that can inducecontractions in smooth muscles, accompanied by a fall of arterial blood pressure and inhibition of theheart muscles. In plants, they appear to exist purely to sting and deter herbivore browsers(Starkenstein & Wasserstrom 1933; Emmelin & Feldberg 1949). However, it should be noted that anUrtica cell extract completely free of these three chemical compounds still elicits a painfulresponse on the skin, indicating that additional compounds are involved in producing the characteristicdeterring reaction (McFarlane 1963).
Chemical constituents have been found in Rumex obtusifolius (Broad-leafed Dock), which inhibit5-hydroxytryptamine, and this helps explains why rubbing dock leaves on a nettle sting is so soothing(Brittain & Collier 1956).
Further doubts and debate regarding the chemical nature of the sting consider the quantities of the threesupposed pain-inducing compounds are much too low to produce such a significant irritant effect (seePollard & Briggs 1984b, pp. 508-9). Similar investigation of the stinging irritants in the relatedgenus Laportea carried out by MacFarlane (1963) found the same three chemicals present, butanother unidentified substance that is not dialyzed through cellophane, appeared to be much more activein producing pain than acetylcholine, histamine and 5-hydroxytryptamine.
Leucotrienes
In 1979, biochemists discovered and named compounds called leucotrienes (also spelt 'leucotrines' in somepapers) in animal tissues. These substances are capable of inducing persistent cutaneous wheals afterinjection into human skin, even when present in only very minute quantities below the nanogram permillilitre level. Leucotrienes have also been found in insect venom and in the stings of sea-animals(Czarnetzki et al. 1990a). Using RP-HPLC (reverse phase high pressure liquid chromatography) andRIA (radioimmunoassay), Czarnetzki et al. (1990b) were able to show high levels of leukotriene B4and leukotriene C4 and histamine in the urticating (ie stinging) fluid of U. urens. Leucotrienesare a family of biologically active compounds described as eicosanoid inflammatory mediators. They werefirst discovered in mammalian leukocytes, being produced by the oxidation of arachidonic acid (AA) andthe essential fatty acid eicosapentaenoic acid (EPA) by the enzyme arachidonate 5-lipoxygenase. Theyhave since been found in other immune cells. In animals, they participate in host defence reactions andpathophysiological conditions, such as immediate hypersensitivity and inflammation. In mammals, thesecompounds have potent actions on many essential organs and systems, including the cardiovascular,pulmonary and central nervous system as well as the gastrointestinal tract and the immune system. Inaddition, leukotriene production is usually accompanied by the production of histamine andprostaglandins, all of which act as inflammatory mediators (https://en.wikipedia.org/wiki/Leukotriene).
Budavari (1996) describes the leukotrienes as potent broncho-constrictors with a role in immediatehypersensitive reactions and some as potent chemotactic agents. She suggests it is the chemotactic roleof the leukotrienes that gives a longer, stronger stinging effect to the nettles. Budavari (1996)characterizes histamine as a potent vasodilator involved in allergic reactions.
In common with the other compounds involved in generating the stinging effect, exactly how thesechemicals are produced from fatty acids in plant cells remains mysterious. It is certainly beyond thechemical understanding of the present writer. However, the fact that these several different biochemicalcompounds have been located in stinging hairs of both U. dioica and U. urens, and theirrole in animal cells is known to involve or include the induction of inflammation, makes it very likelythat they are actively involved in producing the nettle sting.
Further studies indicate that the chemical co*cktail in the Urtica trichome includes significantlevels of tartaric and oxalic acids, both of which induce a pain reaction and help extend the durationof the pain experienced when histamine, acetylcholine and serotonin are present (Han Yi Fu et al.2006).
Treatment for stings
When stung, to minimise the pain it is important to avoid touching the affected area for at least 10minutes. The best approach is to wash the stinging fluid off the skin without touching it, using liquidsoap and lukewarm water. Applying the juice from a leaf of an Aloe vera plant, or using amanufactured product with a high concentrations of aloe vera, can help to manage the red and inflamedskin area and reduce the painful burning sensation. Cold compresses or bathing in tepid water are alsorecommended as ways of relieving the burning skin reaction (Cooper & Johnson 1998).
The deterrent effect
While many invertebrates (particularly insect larvae, slugs and snails), can attack nettle leaves withimpunity, mammalian herbivores (eg rabbits, sheep and horses) are positively deterred by the numerousirritant hairs. The greater the density of stinging hairs, the more the plant is avoided, a learnedbehavioural reaction which occurs to the extent that clonal nettle patches in pastures become free toexpand, unless they are mechanically cut or otherwise managed (Pollard & Briggs 1984b). Protected inthis way, dense nettle colonies may smother out grass and reduce the grazing area available in pastures,since unlike the case of isolated thistles or many other poisonous or distasteful weeds, the stockanimals cannot browse vegetation between the plants, for fear of the burning sting (Bates 1933).
The protective function of the sting
The highly specialized structure and chemistry of the nettle stinging hair suggests that it is unlikelyto have any function other than defence against herbivores (Pullen & Gilbert 1989). While it mayappear obvious that possession of the sting affords such protection, it took considerable care andeffort to design and execute the experiments which proved that many invertebrates (particularly insectlarvae, slugs and snails) can attack nettle leaf tissues with impunity, yet mammalian herbivores (egrabbits, sheep and horses) are positively deterred by the irritant hairs.
Having said this, the situation is not one of complete mammal avoidance. In common with some animals'reaction to toxins in poisonous plants, mammals will browse stinging or mechanically protected plants ifthey are sufficiently hungry. It has been reported that some breeds of domestic cattle avoid nettles,while others eat them readily (Uphof 1962). Stinging hairs of U. dioica have also been found inthe faeces of a number of mammalian herbivores (see Seed Dispersal below). In general, however, it hasbeen shown that the greater the density of stinging hairs the more the plant is avoided by browsingmammals, a learned behavioural reaction that occurs to the extent that clonal nettle patches in pasturesexpand unless they are cut or otherwise managed (Pollard & Briggs 1984b).
Significant variability in stinging hair density exists within many examined nettle populations, and ithas been shown to have a genetic basis (Polland & Briggs 1982). Later experiments by these workersfound that the interaction of large animal herbivores with variation in stinging hair defences could bean important selective force in nettle populations displaying a typical range of variation. While theexperimental results do not suggest that stinging is unimportant to invertebrate herbivores, thestinging mechanism does seem particularly well suited to deter larger animals. Large herbivores cannoteat 'around' stinging hairs in the way that insect larvae or molluscs can, and the immediate deterrenceproduced by a sting's burning sensation will usually act before significant quantities of plant biomasshave been consumed by the larger grazing animals (Pollard & Briggs 1984b).
After browsing damage by vertebrate herbivores, or after mechanical clipping to manage or control nettlepatches, the density of stinging hairs on regrowth stems and leaves is significantly higher than on theinitial nettle growth. Since stinging hairs are presumably energetically expensive for the plant toproduce, it would be strategically advantageous for an individual to be able to produce only as many ofthem as existing herbivore pressure necessitates. The observed increase in stinging hairs after grazingis thus an example of an induced response to environmental pressure (Pullin & Gilbert 1989).
Subsequent experiments using degrees of leaf and stem apex clipping to mimic grazing showed that thereare differences in response even between the sexes of plants, eg with respect to regrowth, branching,reproduction and stinging hair density. In the case of the latter, density was higher on the new leavesof female plants than on males, which might be explained by the greater demand for defence in femalesdue to their higher and longer allocation of resources to reproduction (Mutikainen et al. 1994).Similar earlier experiments by Pullin (1987) found that there was an increase in nitrogen levels andwater content in fresh leaves re-grown by nettle plants after clipping, when compared with matureleaves. This increase in leaf quality allowed higher growth rates of the specialist herbivore larvae ofAglais urticae (Small Tortoiseshell butterfly).
The authors of the original Typescript Flora of Fermanagh noted that Common Nettle wasparticularly abundant (and viciously stinging!) on the screes below the limestone cliffs at Knockmore, asite long frequented by feral goats whose droppings over many generations must certainly have encouragedthe plant's growth (Meikle et al. 1957).
Environmental factors encouraging nettle population spread
Changes in pastoral agriculture practices, common in Fermanagh and elsewhere, over the last 60 years,have involved impressive additional land drainage measures, plus the massively increased use offertilisers, both artificial chemical and organic. The widespread, regular spraying of fields withliquid manure and slurry has resulted in greatly augmented soil nutrient levels throughout most of thecounty. In addition, upland afforestation and its use of fertilisers and lime on the more acidic, peatysoils, has also led to further soil nutrient enrichment downstream. The increase in soil fertility maysometimes be very localised, but due to high rainfall levels, nutrient run-off also occurs and issufficiently widespread to benefit common weeds like U. dioica. The question remains, has soilnutrient enrichment occurred to the extent that it encouraged U. dioica to spread into more than90% of Fermanagh tetrads?
I believe there have been widespread, significant increases in environmental nutrient enrichment offreshwater bodies and soil fertility, and especially in nitrogen and phosphate levels. However, thegreatest effects of this are confined to waters and soils with less extreme pHs, and to those situatedon lower slopes and in valleys. The effects would be much too temporary in their duration to drasticallyalter plant growing conditions throughout our whole area. U. dioica will certainly have benefitedfrom the increased fertility to some unmeasured extent, and probably the plant has become taller andperhaps more abundant and dominant in its existing lowland sites. However, the county is notoverrun with a plague of Common Nettle, and in numerous tetrads where infertile soils are the norm, thespecies remains scarce, sterile and depauperate, so that it has to be actively searched for during plantrecording.
Dominance and Plant Associations
Nettle patches in unshaded, relatively undisturbed sites are often either pure (ie single species)stands, or else they have very low species diversity, reflecting the very strong competitive ability ofthis often dominant, tall, herbaceous plant. Frequently, the only closely associated species in densenettle clumps is the clinging climber Galium aparine (Goosegrass), which is so vigorous it canscramble over and sometimes smother the tall supporting stems of Urtica dioica. The NationalVegetation Classification (NVC) recognises this vegetation as OV24, the Urtica dioica-Galiumaparine community (Rodwell 2000, 5, p. 406-9). The NVC also lists Common Nettle as acomponent in 18 other communities of open vegetation, and it is a constant species in another one ofthem, OV25, the Urtica dioica-Cirsium arvense community. Open vegetation nettle patches,such as these communities represent, might give way to a more closed canopy woody vegetation, but thedense habit of the dominant nettle plant – associated with its tangle of rope-like rhizome, a deep layerof litter in autumn, and rapid spring canopy regeneration – all tends to greatly hinder the invasion ofU. dioica's territory by any other plant species, including woody ones (Srutek & Teckelmann1998).
Nettle stems frequently provide mechanical support for Galium aparine, but in parts of Britainthey are also quite often entangled and parasitized by Cuscuta europaea (Greater Dodder), forwhich the Stinging Nettle appears to be the primary host plant (Holland 1981). This parasitic annual orrarely perennial species does not occur in Ireland; it is regarded as native in S England, but as anintroduction it may be spreading northwards in Britain, while at the same time apparently declining insome of its southern stations (F.J. Rumsey, in: Preston et al. 2002).
European occurrence
U. dioica is shown in the Atlas Flora Europaea as commonly and continuously presentthroughout Europe, the distribution thinning only towards the east – although, in reality, this mightmerely reflect recording effort in those parts of the continent (Jalas & Suominen 1976, Map 322).U. galeopsifolia and all its synonyms fail to feature in this particular European Atlas,being simply subsumed in U. dioica. Beyond Europe, the distribution of U. dioica s.l.,which includes up to eleven forms variously ranked as species, subspecies or varieties, is shown byHultén (1974, Map 285) as being almost completely circumpolar in the temperate regions of the NHemisphere.
World occurrence
In the stricter sense (often recognised as U. dioica subsp.dioica), the Stinging or Perennial nettle is mapped by Hultén (1974) and Hultén & Fries(1986, Map 635) as occurring as a native form throughout Europe and along the coast of N Africa,extending east to Lake Baikal where it meets the usual E Asia form, U. dioica var.angustifolia. U. dioica subsp. dioica also extends southwardsinto Asia Minor and eastwards through Iran to Pakistan. This form of the species has also been recordedas an introduction in South Africa, St Helena, Ethiopia, as well as in N & S America, includingMexico.
In N America, north of Mexico, in addition to three native annual Urtica forms recognisedat the species level, there is just one perennial species, U. dioica, and within it threesubspecies (Woodland et al. 1982). Of these, the American Stinging nettle is U. dioica subsp. gracilis (Ait.) Selander; it exists in both diploid andtetraploid forms (2n=26 & 52), the latter having a strictly western distribution – the RockyMountains forming an effective barrier between the two ploidy levels. Subspecies gracilis extendssouth to Louisiana, New Mexico, Arizona and California (for a map of these N American forms of U.dioica occurring in Canada, see Basset et al. 1977, Fig. 3).
In N America, the 'European Stinging nettle' (ie subsp. dioica) is considered a relatively recentintroduction into or invading the range of subsp. gracilis. Subspecies dioica was firstrecognised from Stone Mills, Bay of Quinte, Ontario in 1877, and since then has been collected onlyrarely in scattered locations in E Canada & USA, usually near seaports, including abandoned fishingports, ballast heaps and railway yards. Very likely it arrived with ships’ ballast or as a seedcontaminant in cargo, and has become accidentally spread inland by man. Many herbarium specimens appearto bear aborted or underdeveloped female flowers, perhaps due to a lack of pollen, a fact which suggeststhat subsp. dioica may have increased its range largely by asexual means, ie throughtransport of fragments of rhizome (Woodland 1982; see his Fig 2 for distribution map of subsp.dioica and subsp. gracilis in N America).
The third N American form of U. dioica is subsp. holosericea (Hoary Nettle), a nativediploid (2n=26), which is a polymorphic complex displaying considerable phenotypic variation. It isconfined to western states where it scarcely ever meets subsp. dioca (see map, Fig. 1 in Woodward1982). In experimental pairings, tetraploid U. dioica subsp.dioica was genetically compatible with other tetraploid taxa, but incompatible with diploids(Woodland et al. 1982).
U. dioica subsp. dioica and subsp. gracilis, plus U. urens, are allpresent in New Zealand as introduced plants colonising waste places and cultivated land. The Europeanform, subsp. dioica, which was first recorded there in 1870, is more widely scattered thanthe N American subsp. gracilis, which wasn’t recorded until 1944. New Zealand also has fivenative Urtica species, of which the woody shrub U. ferox Foster (Tree Nettle), has anextremely vicious sting – indeed it is said to have actually killed people (Webb et al. 1988; Royet al. 1998).
Hultén (1971, p. 294) remarked that where U. dioica s.s. (also known as U. dioica subsp. dioica) meets other Urtica taxa native of therespective country, gene exchange apparently often takes place, and very difficult taxonomic problemsare created. Subsequent biosystematic study of Urtica in N American by Woodland et al.(1982) and Woodland (1982) greatly clarified and simplified the situation there, as detailed above.
Uses
Young nettle shoots are often boiled and eaten like spinach, and nettle broth is a well-known, delicious,healthy food. In medical situations, when nettle stinging hairs come into contact with a painful area ofthe human body, they can actually decrease the original pain of the patient. Scientists think the nettlesting ingredients do this by reducing levels of inflammatory chemicals in the body, and by interferingwith the way the body transmits pain signals. Stinging nettle has been used for hundreds of years totreat painful muscles and joints, eczema, arthritis, gout, and anemia. Today, many people also use it totreat urinary problems during the early stages of an enlarged prostate (called benign prostatichyperplasia or BPH). It is also used for urinary tract infections, hay fever (allergic rhinitis), or incompresses or creams for treating joint pain, sprains and strains, tendonitis, and insect bites (Konradet al. 2000; Safarinejad 2005; Schneider & Rubben 2004 and many references available online).
In the past, the strong fibres present in stem tissue were used to make good quality cloth and paper, apractice remembered in fairy stories and revived during the First World War (see Grieve 1931, pp. 574-9,for a full account of many such uses, and see also Vickery (1995)).
Names
The genus name 'Urtica' is derived from the Latin 'uro', meaning 'to burn', the reference to the stingbeing all too obvious (Hyam & Pankhurst 1995). The Latin specific epithet 'dioica' is a Latinizedform of two Greek words, 'di' and 'oikos', meaning 'two households'. This refers to the nettle plantsbeing unisexual (Gilbert-Carter 1964).
The English common name 'Nettle' is derived from the Anglo-Saxon and Dutch word 'netel', which accordingto Prior (1879) is the "instrumental form" of 'net', itself the passive participle of 'ne', averb common to most Indo-European languages, meaning 'to twist', 'to spin' or 'to sew'. 'Nettle' is thusconnected with the plant's long, strong fibres providing good quality thread and cloth, which it didfrom prehistoric times up until the Industrial Age, when it was replaced by linen and cotton (Grigson1987).
Nine alternative common names are supplied by Grigson (1987), several of which refer to the sting, eg'Tenging-' or 'Tanging-nettle'. Names such as 'Heg-beg', 'Hidgy-pidgy' and possibly even 'Hoky-poky',originating in places as far apart as Scotland and Devon, may possibly be derived from the Anglo-Saxon'hege' or 'haga', meaning 'hedge', often the place where nettles are found growing. Names such as'Devil's Leaf', 'Devil's Plaything' and 'Naughty Man's Plaything' suggest connection with Danish nettlefolklore which held that nettle patches marked where elves lived, and that stings were a protection fromsorcery (Grigson 1987).
Threats
None.
Introduced, archaeophyte, rare. Eurosiberian temperate, but very widely naturalised and now circumpolarand almost worldwide.
1901; Praeger, R.Ll.; Co Fermanagh.
June to September.
Growth form and preferred habitats
Apart from the fact that through being an annual U. urens is easily uprooted, it is not easy todistinguish from the closely related, ubiquitous perennial Urtica dioica (Common Nettle), sinceboth species tend to occur mainly in disturbed sites and both are very distinct 'followers of man'.Small Nettle stings in exactly the same manner and with the same burning effect as U. dioica(Emmelin & Feldberg 1947). The Latin specific epithet 'urens' means 'burning' or 'stinging',something which all true nettles do, even when they are dried and very old on a herbarium sheet (Stearn1992)! Like U. dioica, U. urens is also considered a nitrophile (ie it has a high demandfor nitrogen). U. urens is regarded by farmers as an indicator of soil deficient in lime(Greig-Smith 1948).
Distinguishing characters
Even under optimum growing conditions, Small Nettle only reaches a maximum height of 75 cm. More normallyit stands around 60 cm tall and thus is generally a lot smaller in stature than U. dioica. SmallNettle often occurs as isolated plants or in small patches, again unlike the sometimes very extensivecolonies of Common Nettle (Greig-Smith 1948).
The shape of the leaf base of basal leaves and the relative length of the leaf stalk against the bladehave often been used to make the distinction between the two commonest Urtica species: U.urens leaves are not cordate (ie heart-shaped at the base) like U. dioica, and theblades of the lower leaves tend to be shorter than their stalks.
In his New Flora of the British Isles, Stace (1991, 1997, 2010) uses differences in the relativelength of the terminal leaf-tooth and its adjacent laterals to separate the two common nettles found inBritain and Ireland: in U. urens the terminal leaf-tooth is about as long as its adjacentlaterals, while in U. dioica the terminal leaf-tooth is longer than the adjacent teeth.
When in flower, U. urens is seen to be monoecious, each inflorescence consisting of many femaleand a few male flowers, whereas U. dioica, as its name indicates, normally has separate male andfemale plants (Garrard & Streeter 1983; Webb et al. 1996). However, since U. dioicahas been widely introduced to very many countries, gene exchange has taken place with otherUrtica species, so that due to increased variation the traditional distinguishing charactersmentioned have become unreliable, and they now allow only quantitative differences to be made (Hultén1971). This applies even to normally very conservative characters, including the reproductive strategyof the species, which has become variable in U. dioica, so that monoecious forms regularly occur.
Fermanagh occurrence
There are records of Small Nettle in just eight widely scattered tetrads in the Fermanagh Flora Database,only five of them with post-1975 records. It appears to frequent cultivated ground and light sandy soilson lakeshores. Apart from the first find listed above, the details of the other eight records are: Kesh,1947-53, MCM & D; shore of Lough Melvin near Garrison, 1947-53, MCM & D; Hanging Rock NR,1973-5, J.S. Faulkner, D.L. Kelly & W. McKee; E shore of Cargin Lough, Upper Lough Erne, 19 August1986, L.W. Austin & A.S. McMullin; shore of Derrydoon peninsula, Upper Lough Erne, 28 August 1986,A.S. McMullin; S shore Mill Lough, Upper Lough Erne, 19 September 1986, A.S. McMullin & A. Farr;lakeshore and garden Gublusk Bay, Lower Lough Erne, 6 June 1987 & 22 September 1997, RHN; burialground N of Tattynuckle at Tullynakerran, 26 April 2007, RHN.
The fact that Fermanagh records of U. urens are so few and so widely scattered in both time andspace, suggests it may well be regularly overlooked or taken for young, or poorly grown U.dioica.
Irish occurrence and status
Elsewhere in N Ireland, U. urens is described as "uncommon on waste ground and byhouses" (Hackney et al. 1992), and in Cavan, only two records have ever been noted, dated1987 and 1995 (Reilly 2001). Clearly the distribution and status of this plant needs furtherinvestigation, and not only so in Fermanagh. In the past, U. urens has been recorded in all theIrish vice-counties, and has generally been regarded as introduced in Ireland (Scannell & Synnott1987).
British occurrence and status
U. urens has also been recorded as frequent in all Great Britain vice-counties, but is more commonin the east of the country, where it typically occurs as a weed of broad-leaved crops such as sugar beetand potatoes (Grime et al. 1988; Stace 1997, 2010).
Webb (1985) included U. urens in his list of 41 plants previously considered native in the BritishIsles, but in his view, probably introduced. The New Atlas editors agree with Webb, and they nowrecognise it as an 'archaeophyte', ie an ancient (pre-1500 AD) introduction (Preston et al.2002). Stace (1997) considered U. urens as "probably native", but he has revised hisopinion and now recognises it is an archaeophyte (Stace 2010).
European and world occurrence
U. urens is widespread throughout western and central Europe, thinning eastwardsinto eastern regions. It extends northwards from the Mediterranean to well within the ArcticCircle in Norway, Iceland and Finland (Jalas & Suominen 1976, Map 326). Hultén & Fries (1986)suggest U. urens probably originated and is native in central Europe and the Mediterranean area.They conclude it is so widely introduced that its distribution is now almost worldwide, although theknown occurrences in the southern hemisphere are still scattered. It is believed that it was introducedto S Australia with hay from Tasmania as early as 1840 (Kloot 1983). The species is also introduced inNew Zealand and is regarded as locally common on both main islands (Webb et al. 1988). With itspresent distribution, Small Nettle is circumpolar (Hultén & Fries 1986, Map 637).
Common names
Four English common names for this species are listed by Britten & Holland (1886), but none areunique to the plant and they all refer to the burning sting.
Threats
None.
Very probably introduced and locally very rare. Submediterranean-subatlantic, but also present in C Asia.
1934; Praeger, R.Ll.; old wall in Enniskillen Town.
Growth form and preferred habitats
This easily recognised small to medium sized perennial is common elsewhere in Ireland, especially in thefar south and around Dublin and the Wicklow area. A lowland species, it grows mainly on old walls, butoccasionally on coastal rocks and shingle beaches. It generally prefers dry, sunny, relatively shelteredsituations. Under suitable growing conditions this stress-tolerant species can become both frequent andabundant and clearly demonstrates pronounced competitive ability. It has a short, woody rootstock and isreputed to have long, slender, but tough roots that worm their way deeply into mortar or down intoshingle or rock crevices looking for moisture, thus providing the plant with strong anchorage. Thereddish stems are annual. They are either erect or prostrate, and bear alternate, downy leaves thattaper at both ends. In urban areas of Britain and Ireland the species typically grows in tufts out ofcracks in walls or in the mortar between stones or bricks. The numerous silky, stingless hairs on theplant tend to become covered with air-borne dust, usually derived from nearby roadways.
Flowering
From June to September the small, simplified flowers are produced in tight, greenish cymose clusters inthe axils of leaves, especially towards the top of the red stem. The perianths of the flowers becomereddish in fruit. The flowers are mostly unisexual, but sometimes they are intermingled with a fewbisexual ones. The female flowers are terminal and the males lateral (Clapham et al. 1987). Ineither event, unisexual or not, the flowers are wind-pollinated (Melderis & Bangerter 1955). Thefruit is a smooth, shiny, black, ovoid achene (ie a single seeded dry fruit), sometimes referred to as anut. It is truncate (ie flattened at one end), measures 1.5 × 1.0 mm and is enclosed by the reddishbrown persistent calyx segments (Butcher 1961).
Fruit dispersal
As described, the achene or nut fruit has no obvious adaptation enabling or assisting dispersal, exceptit is small in size and light in weight. Undoubtedly it is to some extent wind dispersed, but as notedabove, the plant typically prefers relatively sheltered growth sites. Having said this, Ridley (1930, p.29) includes Parietaria (as P. officinalis), ranking it last (and therefore possessing theleast obvious means of transport) among a long list of species recorded by O.J. Richard in 1888,observed high up on the walls and towers of churches in Poitiers, France. The plants listed were allrecorded at heights on the walls of the studied buildings, in sites above any buttresses and porches.One can therefore assume the sites were exposed to wind. P. judaica is commonly recorded insimilar elevated sites throughout Britain and Ireland and, again, wind appears the most probable meansof seed transport.
The achenes of some Parietaria species appear to be attractive to ants and have been found intheir nests. They include P. lusitanica, the fruit of which is reported to have a small swellingat its base acting as an eliasome or food body reward for the ant vector, and P. diffusa (= P.judaica) which does not appear to possess a food body, yet has also appeared in ant nests(Ridley 1930, pp. 520-4). This is not considered a likely dispersal mechanism for P. judaica inmost wall or cliff environments.
Fermanagh and N Ireland occurrence and status
Strangely, until 2011 P. judaica was considered virtually absent and about to become declaredextinct in Fermanagh, having previously been seen only once, in Enniskillen in 1934 by Praeger (1934c).Then, in June 2011, Robert & Hannah Northridge rediscovered a well-established colony on the wallsof Monea Castle, a National Heritage site. Since then it has been found again by the Northridges in agarden at Killyreagh House near Tamlaght in 2014 and 2015.
Paul Hackney in FNEI 3 concluded that P. judaica is almost certainly not native in the NEof Ireland. He decided this on the basis that almost all of its occurrences were on the mortar of oldwalls, it was unknown on natural rock outcrops, and the single coastal shingle record in Co Down wasprobably a secondary habitat.
Beesley & Wilde's survey of the flora of urban Belfast located the species in seven 1-km squaresaround the city. They described the occurrence of the plant as being, "occasional on oldwalls", although they also recorded it on waste ground in the city, and they regarded its status asintroduced and naturalised (Flora of Urban Belfast). Thus Pellitory-of-the-wall is scarce or rarein NI, having recent records in only 15 hectads, most of which are either coastal, around Lough Neagh,or confined to old walls in the larger towns (NI Vascular Plant Database).
Republic of Ireland and British occurrence and status
The New Atlas map shows the species concentrated in the S and E in both Ireland and Great Britain,becoming rarer and more restricted to coastal sites in E Scotland as one travels northwards, indicatinga well-marked climatic limitation on its distribution. The species does, however, extend very muchfurther north in Britain than in Ireland in terms of latitude (Preston et al. 2002). It is notobvious why it is so scarce in the N of Ireland and in Fermanagh in particular, but clearly we have apuzzle here which could repay further investigation.
Praeger commented that the species appears "unquestionably native in the Central Plain and the South[of Ireland]" (text in square brackets is my addition to clarify Praeger's view) (Praeger &Megaw 1938). The New Atlas hectad map plots the vast majority of P. judaica records inBritain & Ireland as being native, including all of the Irish records. An Irish Flora (1977,1996, 2012) and the Cen Cat Fl Ir 2 also fail to discriminate between the status of northern andsouthern occurrences of P. judaica, treating them all as native.
In relation to the status question and also the species known distribution, it is significant thatPellitory-of-the-wall (previously known as P. officinalis L.), has had a long history of use inherbal medicine for the treatment of urinary complaints. It was certainly cultivated by medieval monksin their physic gardens, and very probably this happened throughout Britain & Ireland, and possiblymuch further afield, perhaps worldwide (Grieve 1931; Harvey 1981; Darwin 1996). It is thereforeunsurprising that it would be transported by man and introduced to regions well beyond its nativeoccurrence. For instance, both P. judaica (commonly) and P. officinalis (two sites only)have been reported as introductions in New Zealand (Webb et al. 1988), and also in N America.
European and world occurrence
P. judaica is native and widespread in most of W Europe the Mediterranean region. It is alsorecorded in Macaronesia (Madeira and associated islands), very possibly a western outlier where Isuggest it might have been introduced as a medicinal plant (Townsend 1968; Press & Short 1994). Asthe species name suggests, it is also known from SW Asia, including a quite remote eastern outlier atTian Shan in S China, together with Pakistan, Iran, Iraq, Syria, Palestine, Lebanon, Egypt and NW Africa(Townsend 1968). As noted above, P. judaica is certainly introduced very widely around the globe,including the southern hemisphere.
Medicinal uses
Pellitory-of-the-wall was regarded by herbalists as a most useful remedy for stones in the bladder,gravel, dropsy, stricture and other urinary complaints. It was given as an infusion of the plant (1ounce to 1 pint of boiling water), and taken in wineglassful doses (Grieve 1931, p. 624). Gerard'sHerball listed several other medicinal uses, including cleansing the skin from spots, freckles,wheals and sunburn (Gerard 1633).
Names
The genus name 'Parietaria', is derived from the Latin 'paries', meaning 'a wall' or 'a house-wall', intowhich this weedy plant often grows (Prior 1879). The specific epithet 'judaica' means 'of Judaea', andthe previous and perhaps more familiar epithet, 'diffusa', is Latin meaning 'spreading', or 'looselyspreading' (Gilbert-Carter 1964).
The English common name 'Pellitory-' or 'Paritory-of-the-wall' was so named to distinguish it from theoriginal 'Pellitory', 'Pellitory-of-Spain', the composite Anacyclus pyrethrum of SE Europe. Thename 'Pellitory' is derived from 14th century Old French 'peletre' or 'piretre', from the Latin'pyrethrum', from Greek 'purethron', which comes from 'puretos' meaning 'fever'. In the 14th centuryParietaria diffusa (= P. judaica) was known by the Old French name 'Paritarie', from theLatin already detailed above, and similarity of sound combined the two plant names into 'Pellitory',which then had to be distinguished from one another by habitat and geography (Grigson 1974).
Threats
None.
Introduction, neophyte, a rare garden escape. Submediterranean-subatlantic.
1952; MCM & D; damp walls and paths, Castle Coole, Enniskillen.
August to October.
Growth form and preferred habitats
This creeping, red-stemmed, mat-forming, evergreen perennial with minute rounded bright green leaves lessthan 6 mm in length, is an endemic species of Corsica, Sardinia and the Balearic islands. It wasintroduced to English gardens in 1905 as low-growing ground cover for damp, dark conditions where littleelse would survive. Recommended by Farrer (1930) as, "especially delightful for rambling about in acool and shady rock", gardeners subsequently realised S. soleirolii is an almostineradicable weed and it is particularly unsuitable for rock gardens (Ingwersen 1978). Farrer was wellaware that the plant grows with "devastating vigour", yet he considered that as ground coverit had, "high value and charm".
S. solierolii was first introduced to greenhouses where it rapidly over ran damp floors, walls andthe pots of choice plants. The species then probably both escaped and was ejected from gardens as beingtoo vigorous to control. While slightly frost sensitive, it quickly naturalised and established inmilder parts of Britain & Ireland in lawns, on damp walls, shady banks, roadsides, in gravel,churchyards and as a persistent garden weed, often growing at the base of walls (Ellis 1993; Clement& Foster 1994; S.J. Leach, in: Preston et al. 2002).
Despite its invasive tendencies, the plant popularly known as, 'Mind-your-own-business' and'Mother-of-thousands', is still cultivated and features in garden manuals including those published withthe imprimatur of the Royal Horticultural Society, often without any mention of its weedy growthpotential (Brickell et al. 1989). We may expect it to continue 'escaping' and spreading to newareas.
Flowering and vegetative spread
Lateral growth of the brittle prostrate shoots which root at their nodes spread the plant to formmat-like colonies and seed production assists this spread into suitable ground. The unisexual flowersare tiny, pinkish-white and very inconspicuous. They are produced in the leaf axils from May to Octoberin the main. The fruit is a small, hard, achene or nut (Ellis 1993).
Fermanagh occurrence
S. soleirolii is rare in Fermanagh, having been recorded from only eight tetrads, seven withpost-1975 records. There is a record from the roadside at Tamlaght, one at Goblusk Bay, Lower LoughErne, another from a caravan park at nearby Castle Archdale, while the other five records are fromgardens (one derelict) or on waste ground, mainly around Enniskillen.
Irish occurrence
In Ireland, it was formerly established largely in the W, but has spread to at least 21 vice-counties,including those in the N, and in the E, especially around Dublin (Scannell & Synnott 1987; Webbet al. 1996; Reynolds 2002). Around Belfast city, Beesley & Wilde (1997) found it in 17 1 kmurban squares, which was much more frequent than they expected from previous experience (Hackney etal. 1992).
The evidence appears to show that S. solierolii is still actively spreading, yet while present formany years, it is rarely recorded and occurs almost exclusively in gardens and near habitation. Thus itsinvasiveness appears limited.
British occurrence
S. soleirolii is widespread from the Channel Isles to Shetland, but is most frequent in the SW& S of England and Wales, becoming scattered and more coastal further north into Scotland (NewAtlas). Compared with the 1962 BSBI Atlas, it is much more frequent and is spreadingquite rapidly.
European occurrence
While regarded as endemic to islands of the W Mediterranean – most notably Corsica and Sardinia (statusunknown in Majorca), S. soleirolii as a neophyte introduction is mainly recorded in NW France,Britain and Ireland. Otherwise it appears to have just two very isolated introduced stations in theNetherlands and Portugal (Jalas & Suominen 1976, Map 338).
Names
In taxonomic terms, S. soleirolii has previously been included in the genera Helxine andParietaria (Tutin et al. 1993). Nowadays, both genus and species are renamed in memory ofJoseph Francois Soleirol (d. 1863), who botanised extensively in Corsica (Ingwersen 1978). The plant ispopularly known by the English common names, 'Mind-your-own-business', 'Baby's Tears' and'Mother-of-thousands', the latter an obvious reference to its vegetative vigour. This name is sharedwith at least seven other species, including Achillea millefolium (Yarrow), Cymbalariamuralis (Ivy-leaved Toadflax) and Pseudofumaria lutea (Yellow Corydalis).Mind-your-own-business might refer to the self-effacing nature of a plant that creeps around in darkcorners or sprawls over other plants in pots (Grigson 1974).
Threats
Although a vigorous ground-cover species, this garden escape has not yet invaded any semi-naturalvegetation, being apparently confined to damp, shady sites, typically at the base of walls.
Native, frequent and fairly widespread in upland, acid terrain. Suboceanic boreo-temperate, also in EAsia and N America.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Generally a small, much branched, deciduous shrub with oblong-obovate leaves, 2-6 cm long, under idealconditions Bog-myrtle can grow up to 2.5 m tall, although usually it reaches only 1.5 m in height. Theshrubs are grazed by sheep, goats and hares and on occasions may be reduced in height to 50 cm or lessby such pressure. Twigs and leaves are covered with yellowish, resin glands and crushed leaves give offa pleasant resinous fragrance. When crushed and rubbed on skin they are very effective in keeping midgesaway, a tip worth remembering when walking or working on bogs in summer. Midges are paralysed byMyrica oil, and ants are also repelled by it (Skene et al. 2000).
Bog-myrtle is a species of wet heaths and blanket bogs whenever the peat is shallow (50-80 cm), but italso occurs in swamps and fens and can spread into adjacent marshy ground, even when there is some shadefrom willows and birch. The species is very much associated with oceanic climates where rainfall is bothplentiful and regular, typically occurring around 200 days per year. It grows best in wet, butwell-aerated soil, conditions often associated with water movement on slopes. Roots bear annualnitrogen-fixing nodules and plants can tolerate acidic conditions as low as pH 3.8. Near the sea, shrubscan spread from acid peat to colonise adjacent soils, thus displaying some degree of salt tolerance(Skene et al. 2000).
Fermanagh occurrence
As the tetrad distribution map shows, in Fermanagh M. gale is scarce and widely scattered in thelowlands, but it is very frequent over the whole of the upland Western Plateau boglands and takentogether it occurs in 196 tetrads, 37.1% of those in the vice-county. It most typically grows inlakeshore swamps, fens, bogs and wet heathy moorland, but it does not survive on very exposed highground, such as for instance the summit ridge of Cuilcagh.
Vegetative reproduction
M. gale reproduces vegetatively by suckers to produce thickets which dominate the ground theyoccupy. The suckers are often referred to as rhizomes, since although they become woody with age, theyact as important over-wintering food stores maintaining the plant.
Flowering
The flowering strategy of M. gale is mainly dioecious (ie separate sexed bushes), but within apopulation monoecious bushes and bisexual flowers may also occur, so the species is really'subdioecious' (Lloyd 1981). In April, the small, ovoid red-brown buds and the distinctive and strikingred catkins and flowers appear on the bare branches before the leaves open (Skene et al. 2000).The flowers are wind-pollinated, but the amount of seed set is unknown. Establishment from seed isdescribed as rare (Skene et al. 2000). Further work is needed to measure seed production.
A remarkable feature is that individual plants which bear flowers of one sex during a particular seasonmay change to the other sex in the following year (Hutchinson 1972). In N Wales, Lloyd (1981) found thatpredominantly male stems were about 20 times as frequent as predominantly or strictly female stems. Thepattern of gender variation appears to be environmentally induced, but the cause or causes remains amystery, as does the advantage to the species.
When M. gale produces seed, it is water dispersed. The transverse bracts associated with the fruitact as swim bladders prolonging flotation (Skene et al. 2000). Biologically, the fruit is asmall, green, resinous, 2-winged drupe-like nut containing one smooth, ovoid seed. It is fleshy outside,stony inside and secretes a considerable quantity of wax (Melderis & Bangerter 1955; Butcher 1961).The greasy wax covering of the fruit provides an aromatic tallow from which 'Bayberry' (an alternativecommon name) candles are made, the fruiting catkins being boiled in water to produce a scum beeswax(Grieve 1931; Stearn 1972).
Grazing pressure: M. gale is grazed by sheep, goats and hares, and shrubs may bereduced in height to 50 cm or less by such pressure (Skene et al. 2000). Interestingly, theFermanagh farmer who owns the Carrickbrawn Erica vagans site near Belcoo, believes that sheep donot browse Bog-myrtle as hard as cattle do. The increased plant competition between M. gale andErica species following removal of cattle from that site may help explain the observedcontraction of the Cornish Heath population (see that species account).
British distribution: Bog-myrtle is very decidedly a plant of the N and W of Britain and Ireland,although it does also occur less frequently and intermittently in the S and E (Skene et al. 2001;New Atlas). Scotland is the principal area of distribution in Great Britain, and the BSBI LocalChange Survey, a repeat sampling of the Monitoring Scheme survey carried out during 2003-4 showed a 2%mapped decline of recorded M. gale. This gave a Relative Change index of 10% overall. The editorsof the report (Braithwaite et al. 2006), considered this a modest loss at the fringe of thedistribution, readily explained by changes in land use associated with drainage and afforestation. In afew scattered sites in England, M. gale is recorded as an introduction (New Atlas).
European occurrence
In Europe, M. gale has a decidedly Atlantic and north-western distribution, co-incident with thelowland heathland region of Gimingham (1972). However, the distribution extends further north than this,stretching up the Atlantic coast of Norway and all around the shores of the Baltic Sea (Skene etal. 2000).
World occurrence
Beyond Europe, M. gale s.s. is widespread in N America, making it an amphi-Atlantic species. It issaid to inhabit riverbanks and freshwater ponds in temperate parts of N America, a rather differenthabitat compared to that which it usually frequents in W Europe. A variety, M. gale var.tomentosa, is also widespread in E. Asia and NW America (Hultén & Fries 1986, Map 618).
Uses
In addition to the wax candles mentioned above, the use of M. gale as an insect repellent has beenpart of folklore for centuries (Grieve 1931). Midges are paralysed by Myrica oil, and ants arealso repelled (Skene et al. 2000). Dried Bog-myrtle leaves were used to perfume drawers wherelinen was stored and to repel moths. Most interestingly, branches were used to flavour beer before hopsbecame available in the 9th century (Simpson et al. 1996; Skene et al. 2000). The bark canalso be used to tan calfskin, or to dye wool yellow (Grieve 1931). While the leaves are fragrant, theytaste bitter and astringent, yet in China they are used to make tea, and used medicinally as a stomachicand cordial (Grieve 1931). A review of the uses of M. gale has recently been published by Simpsonet al. (1996), in which it is suggested as a treatment for Herpes zoster.
Names
The name 'Myrica' is derived either from the Greek 'myron', meaning 'perfume' (Skene et al. 2000),or from the Greek 'myrike', the Classical name for Tamarisk (Stearn 1992). The specific epithet 'gale'is probably derived from Anglo-Saxon, Old English or German, 'gagel' or 'gagol', a vernacular name forBog-myrtle, adopted as the specific name by Linnaeus (Grigson 1974; Stearn 1992). Additional commonnames include 'Sweet Gale', 'Flea-wood', 'Candle Berries', 'Bayberry', 'Wax Myrtle' and 'Golden Withy',the latter word referring to the flexibility of stems (Grigson 1987; Stearn 1992).
Threats
None.
Introduced archaeophyte, both deliberately planted and occasionally self-sown and naturalised, verycommon and widespread. European temperate, widely cultivated and naturalised beyond its native range.
1934; Praeger, R.Ll.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Beech are large trees up to 30 m or more in height and they can grow on a very wide range of soil pH,from 3.5 at the extreme acid end of the spectrum, to very much rarer occurrences at pH 7.5 in alkalinerendzina calcareous soils, or even pH 7.9 on softer, chalk and oolite substrates (Tansley 1949, pp.361-6 & 421-6; Rackham 1980, Chapter 19). Beech seedlings or saplings are occasionally found on acidbogland, and we have records of the tree in Fermanagh on or very close to raised-, blanket- and cut-overbogs. The tree also grows in calcareous areas, for instance in County Fermanagh (H33) at Hanging RockNNR, at Marble Arch and on the Florencecourt estate. However, beech has not been planted, and it has notcolonised in other limestone areas, including around Knockmore mountain, nor on Knockninny hill. Ingeneral, beech trees do not thrive on calcareous soils, and they certainly never approach anything likethe dimensions they achieve on deeper, more fertile, free-draining moderately acid, brown earth soils.
Beech litter and gradual soil deterioration
Beech trees normally cast a heavy leaf, twig and branch litter each year, each tree typically shedding upto 10% of its terminal branch material annually (Thomas 2000). The slow decay and incorporation of beechlitter gradually leads to the development of a strongly acidic, infertile, sour, mor type of humus insoils beneath beech trees. This degrades the soil through the leaching out of minerals, and particularlyin our moist oceanic Irish climate, it creates a less fertile podsol-type of soil profile. Podsolizationinvolves the formation of a rust coloured 'iron-pan' of re-precipitated iron at depth in the soilprofile, and this impedes the previous usual free drainage of the substrate (Ingrouille 1995). As thesoil gradually becomes podsolized, it turns wetter and becomes subject to waterlogging, so thatultimately, beech itself ends up at a disadvantage, since by this process it creates soil surfaceconditions that are unfavourable to the germination and establishment of its own seedlings (Tansley1949, p. 361).
Environmental factors limiting beech longevity
The twin enemies of the mature beech are late frost, which can defoliate the tree and kill both buds andseedlings, and prolonged summer drought, which may do the same. In Fermanagh and in most of Ireland,long droughts are extremely rare. Beech is not a long-lived tree, and being shallow-rooted, it issusceptible to wind-throw during severe storms. Thus specimens over 250 years old really areexceptional. Wind-throw of large, old trees is especially likely if they are growing in open parkland orin an avenue rather than in the shelter of other woodland. Old trees generally become hollow, whichmakes them more resistant to wind since the tubular trunk is actually more stable and stronger than whenit is solid (Milner 1992, p. 16). A moving account by Thomas Packenham in the introduction to hisbeautiful book, Meetings with remarkable trees, describes the effect on him of the toppling ofseveral giant beeches at his Tullynally estate in Co Westmeath (H23). Accounts like his should make usrealise the need for active replanting of beech for landscape reasons in many parts of these islands(Packenham 1996).
Native and alien (introduced) status in Britain and Ireland: Beech was one of the last woodlandtrees to expand in Britain after the last glacial period. Although present as a rare species from around4450 BP in the Sub-boreal pollen zone VIIb, F. silvatica does not appear abundant in the fossilpollen record until around 3000 BP. Thus having increased and spread at a time when competing vegetationwas well established, and agriculture had already been developing for around 2000 years, the pace ofbeech colonisation was fairly slow. Indeed, in many areas its natural dispersal may have absolutelyrequired the destruction of pre-existing forest by man, a process that was already occurring atthis time in Britain (Godwin 1975; Ingrouille 1995, p. 195) and also in other parts of Europe, eg SSweden (Bjorkman 2001).
F. sylvatica plantation has occurred so much in Britain within and beyond the likely indigenousrange in SE England and SE Wales, that native beechwoods and trees have become very local anderratically distributed. Planting has happened to the extent that, except in very general terms, it isimpossible to accurately delimit beech's native occurrence (Rackham 1980; T.D. Dines, in: Preston etal. 2002). It is therefore puzzling as to why the editors of the prestigious BSBI NewAtlas decided to publish a map of F. sylvatica that totally ignores the questionof status, and misleadingly and knowingly and erroneously plots and displays beech as a native speciesthroughout Britain and Ireland. In doing so, they include mainland Scotland, the western and northernisles and Ireland, all perfectly distinct geographical areas where nobody imagines the tree isindigenous. If for some production reason a single status map was obligatory, then surely anall introduced map would have been a more accurate picture than the one published in the NewAtlas.
The same apparently nonsensical, misleading approach has been taken with Carpinus betulus(Hornbeam) and Tilia cordata (Small-leaved Lime), but NOT with Tilia platyphyllos(Large-leaved Lime) or Acer campestre (Field Maple), both of which have perfectly sensible mapsthat discriminate native from introduced areas. None of these trees are indigenous in Ireland, andobviously it is important for botanists in both Britain and Ireland to know this fact, as knowledge ofit colours their impression of what types of woodland there will be on the smaller, more ancient island(Rackham 1986; McCracken 1971).
Woodland 'Climatic Climax' vegetation and dominant tree species
Despite its questionable status, beech is a tall, potentially dominant tree, and on the more fertilesoils at least, it is capable of outgrowing and replacing old, well-established oak, Quercusrobur (Pedunculate Oak) and Q. petraea (Sessile Oak). Oaks have been long regarded as thedominant native species of the 'Climatic Climax' woodland vegetation of these islands (Tansley 1949;Newbold and Goldsmith 1981). The notion of a single dominant form of vegetation covering the wholeclimatic region (ie the concept of a 'Climatic Climax' vegetation of a region or large geographicalarea) has tended to go out of fashion as we learn more about the requirements of plants and of thedynamic and sometimes cyclical nature of vegetation change (Watt 1947; Gimingham 1972). However, whilewe should approach the ecological concept of climax vegetation with caution, it is still a usefulbackground scheme to assist the interpretation of much that we observe in vegetation dynamics, and italso remains a tool for the prediction and direction of habitat management and active conservation.
Beech in competition with Oaks
The potential of beech to outgrow oak is partially due to the fact that the former is essentially bothshallow-rooting and mycorrhizal, and thus it makes better use of, and more fully occupies, the mostfertile upper horizons of the soil. This feature also allows beech to colonise and thrive on shallower,more rocky substrates than those preferred by oak, or required by the latter if it is to competesuccessfully with other tall tree species, including beech. The competitive benefit of shallow rootingallows beech trees to restrict the amount of water and mineral nutrients available in the soil to otherspecies. This is also the case with Tilia spp. (Limes), another tree genus that did notreach Ireland in the post-glacial 'plant steeplechase' (Mitchell 1986, p. 72).
F. sylvativa casts such a heavy shade and its leaf litter is so resistant and slow to decay, thatit restricts to a minimum the number of species that can maintain themselves in the field- andground-layers of the woodland structure beneath its canopy (Watt & Fraser 1933). Another aspect ofshallow rooting, however, is that it makes the beech tree vulnerable to summer drought, since it may notput its roots down deep enough to tap ground water sufficiently under conditions of prolonged dryweather. Whenever (or in Fermanagh, if ever!) drought occurs, large trees and their seedlings mustcompete for scarce water in the upper soil layers, which restricts the species ability to regenerate(Rackham 1980).
Invasion by F. sylvatica poses a threat to the purity of native or long-established plantation oakwoodland in places like Killarney, where it likewise threatens the even rarer Yew woodland (Reynolds2002).
Sexual reproduction, mast years, fruit predation and dispersal
Beech is not all that well equipped for seed dispersal. The tree reaches sexual maturity when it issomewhere between 30 and 80 years old, depending on the shade environment it inhabits. It then producesits tiny, green, wind-pollinated, monoecious flowers each April, at around the same time as the leavesare opening. Heavy seed crops of beech (and of Q. petraea (Sessile Oak) and Q. robur(Pedunculate Oak)) are sporadic, the so-called 'mast years' occurring roughly every five to 15 years.The timing of mast years, in beech at least, is dependent upon a good warm summer the year beforefruiting, involving high July temperatures, low rainfall and plenty of sunshine. This must then befollowed by a mild, frost-free spring. These special seasonal conditions together induce a heavy beechfruit crop. However, even within the best mast years, the quantities of fruit produced on individualbeech trees varies greatly (Matthews 1955; C.D. Pigott, in: Milner 1992, p. 93).
The one or two seeds (triangular nuts), in their woody pericarp husk covered with coarse bristles fallbeneath the tree, and birds such as rooks, jays, pigeons and pheasants, plus squirrels, mice and voles,collect them as food. The irregular, sporadic, heavy fruiting mast years, it is now realised, are thetrees' evolved response to limit its losses due to this heavy seed predation by birds and mammals. Inthe intervening years between 'masts', beech trees produce very few or no fruits. It follows that thepopulations of birds and rodents decline due to
food shortage in these lean years. Then, when a mast arrives, the quantity of fruit is suddenly somassive that the diminished animal population cannot eat it all, and a high proportion of the trees'seed crop escapes predation. Many trees in the world with large edible fruit use this strategy, but inbeech it is absolutely vital for survival, for only the seed released in mast years contribute tospecies regeneration. The few fruit and seedlings produced in the intervening years are all eaten (C.D.Pigott, in: Milner 1992, p. 92).
While a high proportion of the nuts collected by wild animals are eaten and digested, it is the few nutsthat are transported but which the animals fail to eat that reproduce the species. Apart from thepossibility of some degree of wind dispersal of the triangular nut, which might roll on a suitablesurface, this is the only means of dispersal open to the tree (Ridley 1930; Hadfield 1957).
Toxicity of the fruit
The beech tree does attempt to protect its fruit in another way, but obviously this is not effectiveenough. The nuts and the pericarp contain saponins plus a poisonous uncharacterised substance called'fa*gin', which in sufficiently high dosage is toxic enough to kill horses and cattle. Previously oil wasextracted from beech nuts and the residue made into animal feed cake. This cake fodder, or the beechnuts themselves, have been known to poison both horses and cattle, the former being much moresusceptible. Humans have also suffered after eating up to 50 or more nuts, a diet which producedsymptoms of headache, soreness of the mouth and throat, vomiting and other unpleasant and dangerouseffects. THE NUTS SHOULD THEREFORE BE REGARDED AS POISONOUS AND BE COMPLETELY AVOIDED (Cooper &Johnson 1998). Despite this, in his popular guide book to wild flowers and trees, Mabey (1972) givesinstructions for extracting beech oil from the nuts, and also mentions the difficult labour of shellingthem to eat.
Seed survival, germination and establishment
The seed may survive a year or so, but there is no evidence of prolonged viability or of a soil seedbank. Seed germinates in spring but seedlings are very susceptible to late frost, which may indeed bethe limiting factor determining the northern and eastern boundaries of the indigenous distribution ofthe species in Europe, if not in England. The seedlings are very shade tolerant and are often foundunder bushes or other plants which undoubtedly provide them shelter from frost. For this same reason,beech seedlings and saplings never establish in open conditions (Jonsell et al. 2000).Nevertheless, saplings do not long survive under the dense shade cast by the beech itself (Grime etal. 1988), and their success in replacing existing trees is heavily dependent upon the frequencyof light gaps appearing in the woodland canopy.
Variation
Although as a species F. sylvatica in Britain and Ireland does not vary greatly, and from around1955 onward much of the seed of broadleaved trees planted here has been imported from continentalsources (Gordon & Fraser 1982), a number of distinct forms of horticultural interest have arisenover the years. The most notable varieties probably are the purple 'Copper Beech', several pendulousforms, an erect fastigate form often used as a street tree ('fastigata'), and most interesting of all,the 'Fern-leaved Beech' (= 'Cut-leaved Beech'), named 'heterophylla' (= 'laciniata'). This is an exampleof a 'chimaera', an unusual type of graft having inner tissues of the ordinary beech overlain by tissueof the cut-leaved form (Mitchell 1974).
Fermanagh occurrence
It may seem very odd that the first record of this species in Fermanagh dates from 1934! However 19thcentury Irish botanists knew that Beech was alien and widely planted so they did not deem it worthrecording (eg Cybele Hibernica 1866, 1898). In Fermanagh, Beech has now been recorded in 276tetrads, 52.3% of those in the VC. F. sylvatica is very commonly found throughout the lowlands:it is more frequent in the east of the county and especially so on the larger demesnes. Although we donot have any pure beech woodland as such, merely small copses in mixed species plantations, the oftenlinear representation Beech displays on the tetrad map reflects the fact that it is commonly planted inroadside hedgerows and along other linear geographic features. These include slopes and river banks (egthe Tempo, Colebrooke and Swanlinbar Rivers), plus along estate avenues (eg Castle Archdale and CastleCoole). Frequently on such estate lands it is possible to come across fine individual specimen trees,occasionally up to 30 m in height and some possibly over 200 years in age.
British and Irish occurrence
The New Atlas hexad map shows F. sylvatica common and widespread throughout most of bothislands, but the distribution thinning to the north and west in both Scotland and Ireland. As discussedabove, the information available does not allow accurate discrimination of minority native occurrencesthat possibly occur in SE England and SE Wales, so the map shows all sites as native.
European occurrence
The distribution pattern of F. sylvatica in Britain and Ireland suggests that exposure to wind andto frost are significant factors limiting the species occurrence (Preston et al. 2002). However,from what we know of the requirements for successful regeneration, it is clear that we really need toconsider the map of the indigenous European distribution of the species (Jalas & Suominen 1976, Map284). This shows that beech is widespread throughout W and C Europe and extends northwards to60oN on coastal Norway. Southwards, the presence of F. sylvatica thins and fades awayin the Mediterranean basin, although the tree is recorded in S France and down the length of Italy toSicily, the only other Mediterranean island it reaches is Corsica. In the east, the species reachesMoldavia and the Crimea (Jonsell et al. 2000).
Uses
The usually white, close grained wood is hard, smooth and strong and is widely used for furniture, toolhandles, sports equipment and kitchen utensils. Milner (1992) contains interesting information on boththe uses of the timber and other parts of the tree, and the extremely limited folklore that isassociated with it.
Names
The genus name 'fa*gus' is the ancient Roman name for the tree (Gilbert-Carter 1964), but it may bederived from the Greek verb 'phagein' meaning 'to eat', since in famine times beech mast was eaten bystarving people, and must have made many of them ill (Milner 1992). Grigson (1974) has however pointedout that in Greek 'phegos' refers to oak not beech, and the local Greek oak, Q. macrolepis(Valonia Oak) does have very large acorns that were sometimes eaten by man. The Latin specific epithet'sylvatica' means 'growing in woods' from 'silva', 'woodland' (Gilbert-Carter 1964).
The English common name 'Beech' is derived from the Old English 'bece', 'boc', 'beoce', words that mean,with a difference in gender only, 'a book' and 'a beech tree'. The connection between the two is thatthe Runic tablets on which people once wrote were made of beechwood (Prior 1879). 'Bece' and itsassociated dialect forms are word elements in many place names, including some well beyond theindigenous area of the tree, eg Beckwith in W Yorkshire (previously 'Becwudu', meaning 'beechwood', around 972 AD), and Bitchfield in Northumberland (Hadfield 1957; Grigson 1987).
Alternative English common names include 'Buck', transferred directly from the Old English. Another nameis a reference to a rough-barked variety of the tree, 'Hay Beech'. The fruit of the tree is well knownby the name 'Mast', and other forms exist as 'Buck Mast' and 'Buck's Mast'. In Hampshire, when pigs areturned out into the beechwoods in autumn to feed on the beech nuts, they are said to be turned out tomast (Britten & Holland 1886).
Research literature
There is a huge wealth of published research on beechwoods in both the British Isles and in Europe.Rodwell (1991a) provides a comprehensive bibliography and is a suitable entry point to the literature onthis and other forms of British Isles woodland and scrub for anyone who requires further reading.
Threats
As mentioned above, F. sylvatica invades native Irish oak woodland, but it is under noconservation threat itself since it is an introduction. However, in Fermanagh and other areas of Irelandat least, the beech population is old and very definitely living on borrowed time due to frequent windthrow and a general lack of regeneration. The frequency of wind throw appears to be increasing as globalwarming makes our climate more extreme. Being a familiar and attractive tree, additional beech plantingis surely desirable. The invasive fungal pathogen Phytophthora ramosum that recently arrived in NIreland is known to have infected and severely attacked beech trees in England, but this has not yetoccurred here.
Castanea sativa Mill., Sweet Chestnut
Introduced, neophyte, deliberately planted and rare or very occasional. European temperate, alsocultivated and widely naturalised in Europe.
17 September 1986; McMullin, A.S. & Corbett, P.; Mullynacoagh Td shore, Upper Lough Erne.
January to November.
Growth form and preferred habitats
This is potentially a very large deciduous tree, up to 30 m with a wide, broad crown. Older trees havedeeply fissured bark on the trunk, the longitudinal fissures often spirally curved, allowing the speciesto be easily recognised at any time of year. Ecologically, C. sativa can tolerate mostsoils, although it thrives best on moist, acidic, sandy ones (T.D. Dines, in: Preston et al.2002). In season, the leaf canopy casts a heavy shade, while the deep, persistent leaf-litter creates amor humus, often with a pH of 4 or under (Rackham 1980, p. 332). In Fermanagh, conspicuous, distinctiveand often very large old trees of this species are almost always associated with parkland or treecollections (arboreta) in landed estates such as Castle Coole, Crom or Colebrooke. Unlike thequasi-native chestnut woods in Kent, E Essex and SE Suffolk described by Rackham (1980, pp. 332-9) whichare or were coppiced every 12-14 years for small timber, in Ireland C. sativa never formedwoodland stands. A possible reason for this is the more recent date of introduction and plantation ofthe tree in Ireland, probably sometime before the 17th century – although nobody really knows a definitedate (Nelson & Walsh 1993, p. 105).
Flowering reproduction
Trees flowering in June and July, the minute flowers being borne on long catkins held erect at the tipsof shoots. Individual catkins are 10-20 cm long, with female flowers at their base and more numerousmale flowers on the remainder of the catkin length. Various insects pollinate the flowers and thecharacteristic prickle-covered green chestnuts develop until the autumn. In most of Ireland, the vastmajority of ovules abort and it is doubtful if any of the chestnuts produced in northern counties arefertile and capable of germination as self-sown trees are very rare here. The survey of the Belfasturban area in the 1990s produced a solitary seedling at the edge of playing fields (Beesley & Wilde1997; Reynolds 2002).
European region of origin
C. sativa is a native species of the more humid areas of eastern Mediterranean countries.However, selected cultivated forms, some of them grafted with superior fruiting scions, have long sincebeen introduced into more northerly regions of Europe, and the tree and its edible nuts and otherproducts have been present in Britain since Roman times (Zohary & Hopf 2000, p. 189). Despite thefamiliar 18th century English common name 'Spanish Chestnut' and the present day frequency of the treein the Pyrenees, the species is unlikely to be native as far west as Spain (Rackham 1980, p. 329). Thenative range of C. sativa is controversial. The editors of Flora Europaea took aconservative view, regarding it as indigenous in the Balkans (Albania, Greece, Jugoslavia) and Turkey,and less probably in SC Europe (Tutin et al. 1993). It is common in Italy as well as in southernFrance and Calabria, where it is extensively coppiced (Rackham 1980, p. 332).
British and Irish occurrence
The New Atlas hectad map demonstrates that C. sativa is much more thinly scattered inIreland than in Britain, and especially so when compared to S England and Wales. The tree flowers andfruits readily, and in suitable, warmer conditions in S England, if not elsewhere at present, the nutsgerminate in the spring after their production. Unlike Beech and Oaks, losses due to seed predationappear rare in England. As it cannot regenerate under its own shade, transport into gaps, or intosuitable sites under other species is necessary, the likely vectors being rooks (Rackham 1980, pp.334-5).
Fermanagh occurrence
This is a rare or very occasional tree in Fermanagh, having only been recorded in 15 tetrads, 2.8% ofthose in the VC. It is confined to estate parks and woodland plantations, plus a few outliers nearby.Only one or two of the records occur on the margins of the demesnes mentioned above and, although Igreatly doubt it, these might just be self-sown trees. However, in all our years of field recording RHNand I have never found either seedlings or saplings of Sweet Chestnut anywhere in Fermanagh.
In truth, the reproductive biology and local behaviour of the exotic (ie non-native) trees and shrubs inN Ireland have had almost no attention paid to them. In many cases we do not know whether they cansuccessfully set seed, disperse and establish in our part of the world, as many of them undoubtedly doin S England (T.D. Dines, in: Preston et al. 2002).
A well-known English tree expert, Alan Mitchell, suggested that only a few exotic, introduced trees"plant themselves" in Britain and Ireland. He highlighted one very obvious exception"that proves the rule", as Acer pseudoplatanus (Sycamore). Mitchell went on to listSweet Chestnut along with several introduced conifers that he believed could produce natural seedlings,although he reckoned these were likely to be "very local" (A. Mitchell, in: Milner 1992, p.158). In Fermanagh, as elsewhere, in the autumn we can certainly find beneath the trees shiny chestnutsinside the leathery, green or brown, heavily-spined husks which split into four valves. However, it isvery doubtful if they contain viable seed in Ireland as far north as Fermanagh. This contrasts with thesituation in Essex and Kent where Rackham (1980) found C. sativa regenerating successfully, andprobably even capable of invading woodland of other species. A study of the ability of exotic,non-native trees to regenerate or not, would be another example of a feasible school project, where avery little study could fill a gap in our knowledge regarding the performance and the local behaviour oflong-introduced plants.
Uses
The tree is valued for its nut crop. For many generations it formed an important part of the traditionaldiet of farming communities, and it was also fed to stock animals.
Names
In S England, Sweet Chestnut survived throughout the Dark Ages, and acquired an Anglo-Saxon name, 'cyst'or 'cisten', the latter form (pronounced 'chisten'), being a derived survival from the Classical Latinname, 'Castanea'. The tree name then made the transition to a word element in at least a few Englishplace names, for instance, 'Cystewde' of 1272, which is today's 'Chest Wood' near Colchester (Rackham1980, p. 330).
Threats
None.
Introduced, neophyte, deliberately planted, very rare but possibly over-looked and somewhatunder-recorded. European temperate, cultivated and naturalised in Europe.
14 June 1987; Northridge, R.H.; roadside at Garvary, near Teemore.
A distinctive tree, native of S Europe and SW Asia and which is usually evergreen or semi-evergreen inFermanagh on account of our relatively mild oceanic climate. It is certainly under-recorded since inrecent years source material has become much more widely available in garden centres in some parts of NIreland, and it has again become quite a popular, fashionable tree to plant in gardens. Thelong-standing policy of ignoring garden escapes and deliberately planted exotic trees should come to anend, so that field botanists can begin to be aware of the local behaviour of such plants.
Fermanagh occurrence
There are only three records for Q. cerris in the Fermanagh Flora Database. The details of theother two are: old estate wood and derelict garden, Waterfoot, Lower Lough Erne, 17 August 1990, RHN& RSF; hedgerow, Cornamucklagh Td, NE of Brookeborough estate, 12 April 1996, RHN & RSF.
British and Irish occurrence
The New Atlas map shows that Turkey Oak is very rare and thinly represented in Ireland compared toBritain. There has been a dramatic increase in records in Britain since the 1962 BSBI Atlas, dueto both a genuine increase and better recording of aliens. The fact that Q. cerris, at least in SEngland and Wales, regenerates freely and naturalises itself in a range of open habitats, represents athreat to native or semi-native communities in some sites there (T.D. Dines, in: Preston et al.2002). In the far south of Ireland and northwards to around Dublin, there are a few scattered records ofself-sown seedlings and saplings found near parent trees. However, few if any established trees arisefrom these seedlings in a naturalised setting (Cat Alien Pl Ir; Green 2008). There do not appearto be any genuinely naturalised trees in N Ireland.
European and world occurrence
Q. cerris is native of S and SC Europe from the SE coastal France through Italy and Sicily toAustria, Hungry, Alabania, Greece, Romania and the Black Sea coast. Beyond Europe it extends eastwardsto Syria and throughout Asia Minor (Jalas & Suominen 1976, Map 294; Clapham et al. 1987;Tutin et al. 1993).
Threats
While RSF has seen parasitic Knopper wasp galls on Turkey Oaks in parks in the Belfast area, none haveyet been noticed on Fermanagh oaks.
Introduced, neophyte, deliberately planted, very rare. Mediterranean, planted well beyond its nativerange.
15 October 1987; Waterman, T.; Inish Rath Island, Upper Lough Erne.
The fact that there is just one record in the Fermanagh Flora Database for this species does notaccurately indicate the frequency of the tree in the county. Rather, it reflects the fact that localbotanists have always ignored this dark foliaged, evergreen Mediterranean oak, since it is only everpresent in Fermanagh as deliberately planted specimens in demesne parkland, large gardens and ondriveways. I have no doubt that the tree on Inish Rath is also a planted specimen.
The New Atlas map shows Q. ilex in Ireland occurring extremely thinly scattered, almostexclusively on or near the coast. In Britain, by contrast, it is heavily concentrated in the area southof the Severn-Wash line, steadily declining and becoming increasingly coastal northwards, and rare inScotland although it does stretch to an outlier on the extreme N coast in W Sutherland (VC108).
Q. ilex seeds and regenerates freely in S & E England where it can aggressively invade naturalhabitats and threaten semi-natural vegetation with its very dark evergreen shade (T.D. Dines, in:Preston et al. 2002). However, even in Norfolk where the tree is especially abundant, in a localcounty survey James et al. (1981) found 87% of their Evergreen or Holm Oak sites were largegardens, estate parkland and churchyards. While the species grows to produce very large specimen treesin lawns and appears perfectly hardy around larger houses in the north of Ireland, as far as I am aware,the only place where it reportedly naturalises itself in Ireland, is in two sites in Co Waterford (H6)in the far south of the island (Green 2008).
Native occurrence
Q. ilex is prevalent from Portugal to Italy along the northern Mediterranean coastal belt, andfrom Morocco to Tunisia along the southern Mediterranean coast. There are two subspecies recognised;Q. ilex subsp. ilex is native on the northern shore of theMediterranean from northern Iberia to Greece, and subsp. rotundifolia, is native in the SW of thespecies range from central and southern Iberia to NW Africa.
Holm Oak occurs in mixed species montane forests with Cedrus atlantica (Atlas Cedar) in the AtlasMountains in Tunisia, Morocco and Algeria. While it can obviously tolerate existing conditions on slopesin the latter mountains, Q. ilex is generally confined to more definitely maritime situations,where it avoids cold continental winters. Holm Oak is introduced and grown in California.
Native and frequent. European temperate.
1934; Praeger, R.Ll.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
A relatively long-lived deciduous tree, Q. petraea can form the dominant top canopy species inmixed woodland, especially in more upland woodland or in coppice. It performs best on well-drained,shallow, moderate to strongly acidic mineral soils.
Fermanagh occurrence
Although this species is nowadays regarded as the native Irish oak, in the 19th century, probablypartly for reasons of identification difficulty, it was considered a rarity (eg Cybele Hibernica1898). Currently the Fermanagh Flora Database indicates that it is less frequent in the VC than theclosely related Q. robur (Pedunculate Oak), representing only 35% of the total combined recordsof these two common oak species. Q. petraea has been frequently recorded in Fermanagh, but it iswidely scattered across just 82 tetrads, 15.5% of those in the VC. It is mainly distributed in woodlandson demesnes around the major lakes, with a major concentration of records around the Crom Castle estate.
In comparison, Q. robur is represented in 173 tetrads, 32.8% of the squares in the county.Clearly, Q. robur is much more generally scattered throughout our woodlands than Sessile Oak. Twopublished studies have involved oaks in Fermanagh: the first was a comparative study of oak leafcharacters across 35 woods in N Ireland, nine of them in the VC, carried out by Rushton (1983); andsecondly and more recently, a genetic study by Kelleher et al. (2003) of Irish oak material fromacross the island that included samples from the Crom Castle National Trust estate.
Regarding oaks in Fermanagh, one of the oddest things is that Q. petraea is so well representedaround the shores of Lough Erne, and particularly in the low lying area around the Lower Lough wherelimestone rock outcrops very frequently. It may be that these particular soils are too shallow and dryto support Q. robur, but clearly Q. petraea can tolerate the peculiar set of environmentalconditions they offer. Nevertheless the two common oak taxa overlap in Fermanagh as they do elsewhere,both in their ecology and distribution. A species coincidence map of the Fermanagh data displayed 25sites and subsites in the VC where the species had been recorded together.
Oak variation, hybrids and identification difficulties
Species overlap of Q. petraea and Q. robur is also the case throughout their geographicalrange in Europe (the distribution overlap being partially due to past phases of timber plantation inwhich Q. robur was the preferred timber tree). On account of the amount of variation expressed,many oak trees are very difficult to assign in the field to either Q. petraea or Q. robur.Indeed, many specimens may be fertile hybrids or extreme forms of one or other of these two geneticallyintrogressive species (Cousens 1963, 1965; B.S. Rushton, in: Rich & Jermy 1998). Q. petraeais generally regarded as a relatively good, hom*ogenous species at least in S Ireland and possibly alsoin S England. On the other hand, Q. robur is more variable and has been 'tainted' with gene flowfrom Q. petraea, and probably also from other European oak species earlier in history, probablyduring one or more of the 'Ice Ages' when all oak species were forced to migrate south into refuge areasin S Europe (Cousens 1965).
While there are areas in Britain and Ireland where one or other of these two native oak species is themore prevalent, they obviously overlap to a considerable extent in both their ecological andgeographical ranges, and undoubtedly their natural distribution is modified by past timber plantation.Oak hybrids between the two species are also very variable and they further complicate the question ofidentification. The hybrids are frequent, fertile and are known as Q. ×rosacea Bechst. As a result of introgression and backcrossing of the hybrid with both itsparents, the precise limits of the three taxa (ie the two parents and their hybrid) are controversial,and they have been the subject of investigation for many years (Stace et al. 2015).
Some taxonomists faced with the oak identification problem have even tried to redefine the whole speciesconcept in plants (Burger 1975). Van Valen (1976) has emphasised the notion of 'multispecies', a form ofecological species concept embracing the ecocline and gene transfer, rather than thereproductive species concept based on genetic isolation. This is a deep and provoking topic, butit is interesting and worth mentioning since it arose out of the difficulties faced when identifyingoaks. The arguments and the pattern of variation involved in oaks has led to a continuing academicdebate as whether Q. petraea and Q. robur should be considered two separate species or astwo ecotypes within a common species (Muir et al. 2000; Thomas et al. 2002).
A study of the two oaks by Bacilieri et al. (1996) for instance, showed that both species werealmost completely out-crossing (ie there was a high degree of self-incompatibility). It found that therewas a considerable rate of unidirectional gene flow from Q. petraea to Q. robur, varyingbetween 17% to 48%. These workers regard Q. robur ecologically as a pioneer species, which is, orwas in past forests, progressively replaced by Q. petraea. They believed that the measured geneflow reinforces the succession occurring between the two species (Bacilieri et al. 1996).
At the same time these authors and others query how it is, that despite the high level of interspecificgene flow, Q. petraea and Q. robur somehow remain differentiated to the degree that theydo throughout their natural geographical distribution (Zanetto et al. 1994; Bacilieri etal. 1996; Main et al. 2000). Herein lies the real mystery! As Stace (1975) and Staceet al. (2015) have shown in a British and Irish context, hybridization is a very frequent andnormal process in flowering plants, and it is especially expected and developed in long-lived and clonalspecies.
Timing of flowering (anthesis) and occurrence of hybrids: While as noted there are differences inthe distribution and ecology of the two oak species, there is also a small separation in the floweringtime between the species in the same locality. Flowering in Q. robur is earlier than in Q.petraea by between two and ten days, the difference diminishing at higher altitudes. Despitesome degree of timing difference, there is a considerable overlap in anthesis, and together with thesubstantial degree of self-incompatibility shown, this helps increase hybridisation frequency in mixedpopulations. Experimental hybridisation studies have been carried out by several workers in Britain,Germany and France, but the progeny typically appear small, weak and more prone to fungal disease, sothat few, if any hybrid acorns produce healthy seedlings, let alone saplings (Rushton 1977; Steinhoff1998; Bacilieri et al. 1996; Stace et al. 2015).
Selection of identification material
When attempting identification of these two oaks and their intermediate hybrids, it is vital to selectsuitable representative material from individuals for examination. Unfortunately the ideal material iswell-illuminated central crown leaves from mature individuals, which is by definition, generallyunobtainable from well-grown trees of the size to which these trees grow. When collecting samples, it isalso important to realise that oak trees produce two batches of leaves per season. The later, late-Julyand August opening 'Lammas leaves', are more variable and they differ in shape from the first leaves ofthe season that expand in May and early June. It is the early season leaves that are the 'correct' orbetter ones to choose for identification. It is also important to recognise that the leaves on seedlingsand on young trees are completely impossible to identify accurately (Jones 1959).
Based on his own work and that of several others, Brian Rushton has assembled a very useful table of thesignificant identification leaf characters used to separate the two oaks and their hybrid, together withhelpful illustrations published in Rich & Jermy (1998, pp. 74-6). Potter (1994) has also formulateda reasonably straightforward identification procedure for non-specialists using a leaf index based onseveral simple measurements to be carried out on ten leaves from three segments of the crown.
Studies of Irish oak variation
A study of Irish oaks by McEvoy (1944) concluded that Q. robur is native on the island, notingthat it is the principal oak of the central limestone area of the island in the rather limited areaswhere the rock has not become overlain by peat bog. McEvoy also found that the only extensiveremnants of semi-native oak woodland in Ireland are all peripheral to this central area of the country,growing mainly on soils derived from acidic, siliceous rocks. He also discovered that the trees in thesesemi-native woods are of undoubted Q. petraea affinity. This agrees with the work of Jones (1959)on the occurrence of oaks in Britain, where he found that in general, Q. robur predominates overQ. petraea on more base-rich, fertile, low-lying soils, and it is more tolerant of waterloggingthan the latter. On the other hand, Q. petraea is the principal species on both more acidic sandysoils, and on more upland terrain.
A more recent, detailed study of Irish oaks, which includes trees from the Crom estate in Fermanagh, isthat of Kelleher et al. (2003). These workers looked at the genetic diversity of 26 Irishpopulations using chloroplast DNA for an analysis of the 'genetic fingerprints' of the individual tree.Chloroplasts are inherited exclusively through the maternal line in oak acorns, and thus carry geneticmarkers identical to those of the mother tree. While European populations of Q. petraea and Q.robur have a total of 25 distinct chloroplast types (ie haplotypes) present in them, only twohaplotypes (haplotypes 10 and 12), were found in the Irish oaks sampled. Each of the two haplotypeoccurs in both of our oak species, with haplotype 12 present in 81% of Q. petraea individuals andin 62% of the Q. robur specimens examined. Using this type of data, indices of genetic diversitycan be calculated within and between oak populations, and in this respect the Irish oaks have provedless diverse than mainland European populations. Nevertheless, the Irish oaks are in agreement with theoverall pattern of haplotype diversity and distribution that is shown over the larger continentalgeographical range of the trees (Kelleher et al. 2003).
Studies of this type confirm that Q. robur and Q. petraea are closely related at amolecular level, and that they hybridize both experimentally and in nature, yet they do so withoutswamping the differences between them (except perhaps in parts of Scotland) (Cousens 1963, 1965). Thenearest we can come to a current explanation of this phenomenon is to point to a degree of ecologicalseparation between the species. Of the two oaks in question, throughout Europe Q. robur is thespecies of lowland, heavy soils, tolerant of waterlogging. In this larger geographical range, thedifference in terms of substrate base status appears less consistent than it does in Britain andIreland, but Q. robur is generally associated with the more fertile soils (D. Kelly, pers. comm.,June 2003).
Fossil history
Fossil pollen studies cannot tell us which oak species (ie Q. petraea or Q. robur, orboth), was or were initially present in the British Isles (Godwin 1975). However, after the last coldglacial period, oak first appeared in the SW tip of England around 9500 BP. Fossil material is firstrecorded in SE Ireland very shortly afterwards, around 9400 BP (Birks 1989). The two oak species andtheir hybrid appear to have migrated up along the western coast of France from a glacial refuge(refugium) in N Spain (Cantabria) or around the Bay of Biscay.
This was one of three such southern refugia shelters these Quercus species retreated to duringglacial conditions, the others being the Balkan peninsula to the Black Sea shore, and to a much lesserextent, the Italian peninsula south of the Alps (Palmer & Birks 1983, p. 354). Q. petraea hasclose taxonomic affinities in both the Iberia and Balkans, whereas Q. robur has only one closerelative in the Balkans, so the location of the refugia of the two oak species most common in Britainand Ireland is still a matter of discussion and macrofossil research.
Chloroplast DNA research has also shed light on the distribution and migration routes of oak species inthe early post-glacial. Kelleher et al. (2003) found two main chlorophyll DNA haplotypes inIreland (haplotypes 10 and 12). These correspond to the oak haplotypes that migrated northwards fromSpain after the end of the last ice age. Haplotype 12 was found in oaks in the north and on theperiphery of Ireland, while haplotype 10 was more central and southern in its occurrence. Five of thepopulations studied, including oaks on the Crom estate in Fermanagh, contained mixtures of these twohaplotypes. Only one population in Glencar, Co Kerry contained a single tree with a non-native haplotype(number 7). The pattern of chloroplast DNA genetic markers found in Ireland was found to be consistentwith that expected from a natural distribution of oak, and it was concluded that in the main Irish oaksare derived from indigenous (native) material, rather than from foreign, planted, introduced stock(Kelleher et al. 2003).
While the all Ireland study of Kelleher et al. (2003) only sampled a few Fermanagh trees at theCrom estate, in them it found both of the characteristic Irish genetic marker haplotypes 10 and 12. Thismade the Crom tree sample one of a minority of just five Irish oak populations out of the total 26populations sampled which contained more than one haplotype marker in its chloroplast DNA. This factindicates a higher genetic diversity in these five populations compared to the other Irish populations,yet the general levels of genetic diversity indicated by this study overall are low in comparison withBritain, where oaks have three main haplotypes and three subsidiary ones (Cottrell et al. 2002),and in France, where the oak populations of these two species contain a total of twelve haplotypes(Dumolin-Lapègue et al. 1999).
The fossil record shows oaks initially spreading into de-glaciated ground that was not bare, but alreadyoccupied by a mixture of birch, hazel, willow and juniper. Even though the ground was occupied, oaksinvaded at an extremely rapid rate, advancing 350 to 500 m per year until around 8000 BP, by which timethe species had reached S Scotland and N Ireland (Birks 1989). The rate of further spread then sloweddramatically to around 50 m per year, possibly because of the cooler summer temperatures typical ofthese more northerly regions.
Climatic limits to oak growth
To this day, climate remains a factor limiting oak growth in Britain and Ireland. For instance, a studyof tree-ring samples from oaks at 13 sites across both islands showed marked similarities in theirresponse to climate (Pilcher & Gray 1982). The study proved that high rainfall, particularly in thegrowing season, and high temperature in early summer, favour oak growth. High temperatures in theprevious winter, however, are detrimental to growth in the following season, a finding which Pilcher& Gray suggested could well be due to depletion of the tree's starch reserves.
Acorn dispersal
Since the acorns of our two oak species represent very large heavyweight seed, the rapid rate of treeadvance measured in fossil studies suggests that the species must have had considerable assistance withtheir dispersal. Most probably this involved a feeding relationship amounting to a mutualisticpartnership with birds. Bird species such as jays, rooks and wood pigeons are considered the most likelyoak seed vectors (Birks 1989).
Toxins
Oak trees have tannins in every tissue of the plant and there are relatively high levels of hydrolysabletannins in young leaves and in green acorns. This latter form of tannin breaks down in the gut ofherbivores to form gallic acid and pyrogalllol. Pyrogallol oxidises blood haemoglobin and it is said toalso attack the liver and the kidneys of animals. As toxicity levels are high in young tissues, animalpoisoning is seasonal, occurring mainly in the spring (Cooper & Johnson 1998).
Oak mast years
It is not clear to the current author whether or not the calculations of oak dispersal rates quoted abovefrom fossil studies allow for the occurrence of oak mast years which certainly occur nowadays. One wouldimagine that seed predation very probably did occur in the early post-glacial period as the large seedrepresents an important source of food for numerous woodland animals. In the case of beech and othertrees including oak species, masting behaviour is very important in setting limits to seed predation,and thus permits and enables tree regeneration (Matthews 1955) (see the fa*gus sylvatica speciesaccount).
Individual oak tree life-span
One of the myths associated with our oaks is their supposed longevity. The often met notion of the tree"being 300 years in the growing, 300 years in the being, and 300 years in the dying"' (Thomas2000, p. 266), stretches credibility very thin. In the real world the lifespan of the typical oak couldgenerally be truncated into more like 200 to 300 years in total, and the evidence suggests, or ratherproves, that only the very exceptional individual survives longer, unless we widen our definition toinclude the epicormic growth on pollarded trees and coppice stools. The biggest and oldest oaks are allhollow pollards, and the oldest apparent 'maiden tree' (ie not pollarded, - and we cannot even becertain of that status in this particular case), is the Q. robur 'Majesty' at Fredville Park,Kent, which is considered to be around 450 years old at most (Mitchell 1996, p. 312 & 313; Pakenham1996, p. 18 & 19).
Pilcher (1979) examined the ages of living oaks in nature reserve woodland slopes above Rostrevor, CoDown (VC 38). This woodland stands on steep rocky slopes and it contains trees growing under stress thatare similar in size and appearance to many found today in Fermanagh. Using tree-ring counts and allieddendrochronological techniques, Pilcher found that the oldest trees he sampled began life in the early1740s (ie maximum age 260 years at this sampling date). Historical evidence suggests that the price ofoak timber in Britain had risen to such an extent in the 1730s that felling became economically drivenand was happening extensively throughout Ireland (McCracken 1971). Thus Pilcher felt that the Rostrevortrees he studied probably represented regeneration that occurred after this particular spell of economictimber extraction. His results did not suggest that fresh planting occurred, but rather that naturalregeneration took place.
Tree-ring studies of modern oaks in Ireland associated with carbon dating and dendrochronology, indicatethat the majority of trees of both native oak species and their hybrid growing in semi-native habitatsare only 150 to 250 years old, and that they are very heavily constrained by human activity (Baillie& Brown 1995). The oldest oaks in N Ireland are (or were) at Shane's Castle, Antrim, where two largetrees felled in the 1980s, proved to be 307 and 340 years old. The well-grown oaks at Inisherk on theCrom estate in SE Fermanagh are almost as large as those surviving at Antrim, but they are quite a lotyounger. Tree ring boring samples of the trunks of the largest Crom oaks produced a planting date ofaround 1720, making them nearly 300 years old. Most of the older Crom trees are 19th century (Browne& Hartwell 2000).
As a result of their study, Baillie & Brown (1995) concluded, "Overall, with the exception of afew comparatively young oaks sampled in hedgerows, it was apparent that the nineteenth-century Irishlandscape outside enclosed estates, must have been almost devoid of oak trees." (myitalics). Furthermore, these authors also suggested that, "despite the wishful thinking of manypeople, there does not appear to be any evidence of existing relict ancient oak forest anywherein Ireland, including even those small patches on lake islands and inaccessible slopes." (again, myitalics). Other woodland history experts disagree with both these statements, especially with the latter(eg Rackham (1995), and D. Kelly, pers. comm., June 2003).
Irish Ordnance Survey maps of the 1830s-40s show where woodland survived through the period of greatestpressure on the land. However, for several reasons the current author (Ralph Forbes) will go further andstate that I reckon almost all woodland oaks in Fermanagh, as opposed to parkland trees, have a maximumage of around 200 years. When growing under stressed conditions, oaks do not develop massive boles.Observation shows them beginning to rot and break up while still of modest dimensions when compared withtrees in 18th or 19th century parkland, or when compared with estate woodland specimens in Glenarm, CoAntrim (VC H39) or Killarney, Co Kerry (VC H2). Trees in estate parkland generally benefit from open andalmost unrestricted growing conditions, so that uncut 'maiden' trees grow much faster and very muchlarger than upland trees growing on rocky slopes. Alternatively, estate trees may have been coppiced orpollarded at some stage and their trunk growth has thus been invigorated. Again, observation shows thatmost very old trees of any species, apart possibly from Yew (Taxus baccata), have been lopped inthis way at least once during their lifetime (Mitchell 1996; Pakenham 1996).
Undoubtedly part of the difficulty in understanding oak biology and ecology is the extent to which thetree is a relatively long-lived plant, and secondly - and probably the crux of the matter - ever sincethe arrival of Neolithic farmers in these islands, it has seldom been allowed to live out its biologicallife span of somewhere between 200 and 400 years (or maybe somewhat more in exceptional cases),depending upon growing conditions, in anything even remotely approaching a natural manner (Shaw 1974;Minihan & Rushton 1984; Ellenberg 1988, Table 9; Ingrouille 1995). Throughout history oak timber hasbeen far too valuable to allow the tree to die and fall naturally. Alternatively, the land on which thetree is growing becomes too valuable, or it becomes required for other purposes. Either way, managementsteps in and fells the tree or the entire woodland. Even if the individual tree survives into old age,during its long life some form of major disturbance of its growth almost inevitably occurs. Often thedisruption is perfectly natural, but increasingly it is due to human activities. In any event,disturbance curtails the natural performance of the tree, resulting in distortion of its growth or itspremature destruction.
The effect of past human pressure on woodland
It is easy for us today to forget that in the late 18th and early 19th century, Ireland's humanpopulation peaked at a level somewhere between 8.0 and 8.5 million. In comparison, the 2016 totalpopulation of Ireland (north and south combined) was 6.6 million. The great majority of Irish peoplenowadays live in urban conurbations, whereas in the 18th and 19th centuries, rural populations were verymuch larger and widespread. The Irish population doubled in the 60 years between 1780 and 1840,creating an enormous demand for all kinds of timber. Demand was particularly high for hard, durableconstruction timber, and throughout that period in rural communities shortages must have reached levelsof desperation that are inconceivable today (McCracken 1971; Mitchell 1986). Before the development ofthe railways in the mid-19th century (the Londonderry and Enniskillen Railway reached Enniskillen in1854), coal was very expensive and industrial and domestic fuel was almost entirely confined to wood andturf (dried spade-cut sods of moss peat). As a result of the demand for wood, even the mostnatural-looking and extensively visited and studied oak or mixed deciduous woodlands in Ireland today,are in reality totally secondary and planted. In some cases, as for instance in Killarney, secondary oakwoodland stands on ground that previously supported clear-felled woodland containing oaks (Kelly 1981;Rackham 1995; Pilcher & Hall 2001).
The best Fermanagh woodlands containing substantial oak populations are the nature reserves at CorrelGlen, Cladagh River Glen and Hanging Rock, plus the numerous wooded islands and shoreline woods of LoughErne and Lough Melvin. Most of these woods are associated with past or present landed demesnes thatbelong, or used to belong, to wealthy families. Even in the case of these secondary semi-natural mixedoakwoods, at present we cannot be certain regarding the native provenance of the seed sources used intheir plantation. However, modern allozyme coding genetic techniques may eventually allow us to discoverwhich tree samples represent native local Irish oak genome regeneration (eg Kelleher et al.2003), and which are derived from seed imported from Britain, or less likely, from continental suppliers(Gordon & Fraser 1982; Nelson & Walsh 1993, p. 117).
General oak ecology and biology
An enormous amount has been written about the ecology and biology of the two oaks native in Britain andIreland, Q. petraea and Q. robur, and for instance, a Web of Science Database search on 21Mar 2003 for 'Quercus petraea' turned up no fewer than 343 scientific papers dating from 1993onwards! On account of the huge volume of published research and the fact that there is reasonably easyaccess to the general information in summary form (eg in the Biological Flora account of Jones(1959); papers in the Botanical Society of Britih Isles conference volume edited by Morris and Perring(1974) 'The British Oak, its History and Natural History.'; Rackham (1980) 'Ancient Woodland,its history, vegetation and uses in England., Chapter 17.'; plus concise accounts in Grime etal. (1988) and Milner (1992), and the excellent summary and review of forest ecology byIngrouille (1995) in his book 'Historical Ecology of the British Flora.', pp. 170-203), I willnot rehearse here the basic details of oak ecology. Instead I wish to present a review of the recent(ie, post-1980) changes in perceptions regarding the biology, ecology and conservation situation ofthese two oak species, which will be for the most part applicable both to them and to the complex oftheir hybrids, ie, what might broadly be referred to as the British Isles oak species aggregate.
Lack of oak regeneration and oak population decline
The lack of regeneration in oaks in Britain and Ireland was quite widely appreciated from around 1910onwards (Watt 1919), and the near absence of young saplings and trees under 50 years of age is nowunderstood to be more general in its occurrence. This phenomenon is now recognised as a widespreadEuropean 'oak decline problem', akin to, but not identical with that at present being recorded in NorthAmerica (Thomas et al. 2002).
In C Europe at least, present day oak decline involves two syndromes characterised by either (i) cyclicepisodes of rapid mortality in local but widespread centres, followed by decreasing and slowermortality. Such episodes may last for up to 10 years and sometimes are preceded by a predisposing phaseof reduced growth; or (ii), general oak woodland decline which is characterised by increasing crownthinning of the trees in entire stands over large areas, but which involves only low levels of mortality(Thomas et al. 2002).
In Britain and Ireland we may be witnessing the latter form of oak decline, although it is not all thatapparent in some areas, including N Ireland. The sudden oak death scenario we have certainly notencountered here, although there are worries that a new virulent lethal form of the fungusPhytopthera introduced with ornamental plants such as Rhododendron and Viburnum isactively spreading in Britain and Ireland. While this fungal pathogen is capable of attackingQuercus species, it is not yet known to have done so.
Late onset of seed production
A factor adding to the reproductive limitation of the oak tree is that younger trees do not becomesexually mature until they are at least 40 years old, and acorn production is variable from year toyear, following the mast pattern which is also met in the more shade tolerant beech tree (Matthews 1955;Shaw 1974). Having said this, acorn production itself is not likely to be a significant limiting factorfor oaks, since, except in the very worst years, far more of them are produced than is required tomaintain existing oak populations in woodland. The perils undoubtedly lie with the fate of the acorn,the seedling and the young sapling, the most critically stage being the first year after germination,when seedling establishment is taking place (Rackham 1980).
Ecological advantage of large seeds
Having said this, the large seed, while susceptible to damage or destruction early on from drying out,frost, or hungry herbivores unless covered, hidden and protected either by the soil litter horizon or bythe surrounding vegetation, does provide the biological and ecological advantage of a relatively large"starting capital" for the tree embryo, making the seedling independent of soil nutrients forup to two years after germination (Ovington & MacRae 1960; Jarvis 1963). The large seed size alsopermits the pattern of seedling growth which provided the shade is not too dense places the taprootbefore the shoot and allows the former to thicken and accumulate storage materials. In turn this oftenenables the young seedling and the sapling to survive dieback and to resprout after meeting adversegrowing conditions, or even after the repeated attentions of browsing herbivores (Jones 1974). In deepershade on the other hand, the shoot etiolates and the root becomes too poorly developed to mechanicallysupport it, so that the tall, spindly seedling generally ends up lodging, i.e., falling oversideways and dying, being pushed aside by competing stems of such plants as e.g., bracken, or thegrass, Deschampsia flexuosa. If this latter fate doesn't materialise, root competition from oldertrees and from other species becomes of greater importance in limiting the growth of the young oakseedling than the low light intensities, but again, poor root development of the seedling does makes itvery susceptible to any occurrence of drought conditions (Jarvis 1964).
It has also been shown in cultivation experiments that the larger the acorn, the more rapidly it growsand therefore the more likely it will escape through the very vulnerable small seedling developmentstage. "In view of the large number of ways in which small seedlings may be killed or stultified,the ultimate advantage to the larger seedling may be out of all proportion to its initial sizeadvantage. For example, there is only a small margin between successful establishment and smothering bybracken or Deschampsia." (Jarvis 1963).
Saplings and regeneration
Saplings and young trees are occasionally present in more open woodland areas, for instance along rides,roads and wood margins. However, these saplings are usually present in numbers far too small to achieveeffective woodland regeneration (Rackham 1980, p. 296). This is especially the case when there ispressure on seed, seedlings and saplings from grazing by both farm-stock and wild or feral animals.Small scale herbivores such as wood-mice, voles and specialist feeding invertebrates are frequentlyinvolved, together with the more obvious seed predation and herbivory carried on by larger vertebrates,such as squirrels, rabbits and deer (Crawley & Long 1995; Kelly 2002).
The first year seedling growing in woodland shade needs all its leaf area intact in order to be able tomanufacture and store sufficient reserves for its overwinter survival, so that any damage is potentiallylethal during its early months of existence. Young leaves are particularly vulnerable since they aresucculent and initially they have low levels of the bitter tannins which make older, harder, moreleathery leaves highly unpalatable or indeed toxic to grazing animals of all descriptions (Cooper &Johnson 1998).
A 25-year experimental study monitoring Q. petraea sapling survival in Killarney, SW Ireland byKelly (2002) concluded that successful regeneration only occurred there in unshaded or lightly-shadedsites where grazing levels were kept low. Even so, oak mortality was high and the median proportion ofsaplings that survived in unshaded plots protected from grazing was extremely low, measured at just 0.2after 25 years study. The study found Oak saplings were soon over-topped by faster growing species,mainly Betula. At the end of the study the surviving oaks had a mean height of 3.1 m (a rangefrom 0.9 -7.0 m), and the birch which was suppressing them had developed a canopy at around 12 m (Kelly2002).
It has been suggested that the decline in oak regeneration noted in the early years of the twentiethcentury might stem from the arrival of oak mildew, Microsphaera alphitioides, which was unknownin Europe prior to 1907. Rackham (1980) hypothesised that this probably made oak seedlings and saplings"succumb to a degree of shade which they would formerly have survived". Some subsequentstudies however have failed to show this effect (e.g., Kelly 2002), and it is probably an oversimplification to expect a single factor rather than a specturm of interacting causes to be responsiblefor the syndrome of regeneration failure.
The problems that the two Quercus species face in regenerating under oak canopy (or the nearimpossibility of this happening!), suggests that perhaps oak woodland was not the climatic vegetation'Climax' in Britain and Ireland as suggested and believed by Tansley (1949), and was taken up by earlypalynologists interpreting their fossil pollen diagrams (Shaw 1974). Later fossil pollen studiessuggested that a more mixed deciduous woodland assemblage, including elm, and, at least in England wherethey are indigenous, other more shade tolerant tree species, such as lime, beech and hornbeam, mightco-exist with oak, or act as stages in some form of succession in which oak played just a part (Godwin1975, p. 277-281, and Mitchell 1986, p. 34).
On the other hand, historical studies of British woodland and its uses by Rackham (1980) have led him tosuggest that oak regenerated more freely in the historic past than it does now, and that perhaps somedetrimental change in the biology of our native oaks took place around 150 to 200 years ago whichreduced oak regeneration capacity. Rackham's researches found that medieval carpenters had an abundantsupply of "small oaks", with a rapid turnover in just that age-class, ie, trees of less thanfifty years growth. At present, this size of oak is very deficient in woodlands throughout theseislands. While most of this small bore timber would have come from oak coppice, Rackham still points outthat it wasn't until the eighteenth century that possible hints begin to appear in the written record,suggesting there might be a deficiency in the supply of oak timber from managed woods (Rackham 1980, p.295).
A possible more pioneer colonising role for oaks
Studies on the European continent suggest that in the past oaks might have undertaken a more pioneeringrole, similar to that of the light-demanding birches today, colonising woodland gaps, newly availabledisturbed ground, and drying out areas of fens and bogs (Björkman 2001). Here the more rapid growth ofaccompanying birch might allow it to nurse oak saplings which would have colonising the ground at thesame time as the birch with the assistance of avian seed predators. Oaks are very much slower to developand gain height than birch, but if they survive (even in a stunted form), eventually they shouldoutlive, outgrow and overtop the comparatively short-lived birch (Kelly 2002).
In the more cloudy oceanic and upland areas of Britain and Ireland, however, oaks are unable toregenerate under their own shade and they are constantly accompanied by a very rich variety of thephytophagous insect, lichen, fungal, bird and mammal communities they support. Oak trees under this sortof pressure eventually lose vigour and they would then very gradually be replaced in woodland byinfiltration of more shade-tolerant, high canopy species, ie elms, beech, hornbeam and limes (Rackham1980; Ingrouille 1995).
However the arrival of Neolithic man in Britain around 5000 BP and the onset of his woodland clearancesand the prolonged exploitation, management and especially the favouring over other species of oak fortimber usage, together with the more recent mismanagement and neglect of woodlands since other materialshave replaced timber, has modified the ecological picture of woodland out of all recognition. The morerecent changes in our use of timber and our appreciation of woodland have happened quickly, over just acouple of human generations, to the extent that we only now appear to be gaining some foggy notion at tohow woody species and the woodland community naturally behave during an interglacial stage. No doubt wehave a great deal more to discover before we can effectively manage upland and lowland mixed deciduouswoods in changed circ*mstances purely for the conservation of the trees and the communities theysupport. We just hope and pray that it is not already too late to achieve this goal without major lossof the associated biological diversity.
British and Irish occurrence
The Q. petraea map in the New Atlas clearly demonstrates the accuracy of these assessments.It shows that Sessile Oak predominantly occupies peripheral and upland siliceous terrain in Ireland,while in Britain it has a pronounced western distribution, correlated with the more ancient, acidicrocks and with a wetter oceanic climate. The New Atlas map of Q. robur indicates that atthe hectad level of discrimination this species is almost omnipresent on all soils in Britain, avoidingonly the most shallow, driest limestone soils, deep acid peat and high mountains. Since considerableareas of Co Fermanagh have an underlying Carboniferous limestone rock structure, even though this isfrequently buried beneath boulder clay or peat and the majority of the county is low-lying and with apreponderance of seasonally wet, clay soils, overall the existing data agree with the views of bothMcEvoy (1944) and Jones (1959) on the predominance of Q. robur under such conditions.
The New Atlas maps of the two common oak species clearly illustrate that the one area of Irelandwhere neither oak can survive is Co Mayo (H26 & H27) with its vast empty stretches of treelessbogland and quartzite mountain cones. In Britain, the same degree of absence is clearly apparent in theScottish Highlands and their vast north-western blanket bog areas (Preston et al. 2002).
Oak and Man
The Oak tree was regarded as a 'Noble of the wood' in the 8th century Irish Laws of Neighbourhood, onaccount of the trees' size and quality of timber. In addition, the acorn fruit was a useful andimportant seasonal food for pigs – ie autumn pannage (Edlin 1963). For these reasons, oak was protected,and rather severe penalties would have been due from anyone found cutting branches, boughs or felling atree (Nelson & Walsh 1993). Despite this, oak plays a relatively minor role in Irish traditions whencompared with ash or yew (see Nelson & Walsh 1993).
In a lengthy article on 'The Sacred Trees of Ireland', however, Lucas (1963) points out the connection ofsacred oak groves (in Irish, 'doire', from 'dair', meaning 'an oak tree'), with ecclesiastical sites. Itis also the case that 'doire' (anglicized as 'Derry') and its diminutive 'doiríin' (anglicized as'Derreen') are two of the commonest elements in Irish place-names, reflecting the fact that the forestsof ancient Ireland were chiefly of oak. Furthermore, McCracken (1971, pp. 24-5, Map 2), has mapped allof the Irish townland place-names containing the word 'doire', and then used this together with thewritten historical record to estimate the extent and distribution of major pre-1600 AD oakwoods acrossthe island.
Oak timber had very many uses in Ireland as elsewhere, including furniture, barrel staves, building andboat construction. In addition, less well grown pieces were converted into charcoal for iron smeltingand for gunpowder, or they were used for fencing or as planks for wetland and bog walkways. McCracken(1971) has provided a very full historical account of Irish timber use from Tudor times to the present,while Neeson (1991) has likewise provided a detailed history of Irish Forestry.
Names
Grigson (1987) provides derivation of the English name 'oak' from the Anglo-Saxon/Old English 'ãc', aword which is cognate with numerous other Germanic languages and refers to the fruit of the tree, theacorn, which was regarded as one of the most useful products of the tree, the oak forest being widelyused to fatten swine. Anglo-Saxon laws were in force to protect the trees for this very reason (Prior1879). Britten & Hollland (1886) list 21 variations from 'aac' to 'atchorn' for the more widespreadand familiar 'acorn', plus a further 24 English common names for the tree. The main ancient folklorebeliefs, worship of, and superstitions associated with oak, are summarised in Grigson (1987), Milner(1992) and Vickery (1995), while there are 35 references to oak in various folk modes recounted inFriend (1883).
Threats
There is no immediate conservation threat to native oaks in Fermanagh, but the stock of trees is old andmany individuals in woods and parkland are approaching senescence. An active programme of replacement isurgently required and the best option is to use local acorns. The 2009 arrival in Ireland ofPhytophthora ramosum, the fungal pathogen responsible for so-called 'Sudden Oak Death' in NAmerica, is a worrying development since it is reputed to affect many different woody species. The listof susceptible trees and shrubs includes Fraxinus excelsior (Ash), Betula spp. (Birches),Acer pseudoplatanus (Sycamore), Vaccinium myrtillus (Bilberry) and Viburnum spp.(Viburnum) as well as Larix spp. (Larch), Oaks (especially Q. ilex), fa*gussylvatica (Beech), Aesculus spp. (Horse-chestnut) and Castanea sativa (SweetChestnut). Rhododendron is also seriously affected and it can act as an evergreen carrier and spreaderof the fungus. As a result of this, rampant Rhododendron is being more actively removed from someNational Trust estates than ever before – proving that every cloud has a silver lining!
Native, but also often deliberately planted, common and quite widespread. European temperate.
1882; Barrington, R.M.; around Lough Erne.
Throughout the year.
As there is considerable difficulty with identification of this species due to introgression (gene flowor genetic transfer) with the closely related Q. petraea (Sessile Oak), I have decided toamalgamate the species accounts of our two native oaks and present them both and that of their hybridunder Q. petraea. This is not to deny for a moment that they are separate taxonomic entities, norto suggest despite high degrees of overlap in their biology and behaviour that they are identicalin their ecology and distribution. Rather I believe that the two deciduous oaks native to Britain &Ireland are more similar than different from one another, and the proper degree of distinction is stillunder debate and a matter of active research.
65% of the combined total of deciduous oaks recorded in Fermanagh are listed as Q. robur and it ispresent in 174 tetrads, 33% of those in the VC. While it is common and widespread throughout Fermanagh,Q. robur is still (like Q. petraea) most frequently recorded in woods on the majordemesnes which are situated around the larger, lowland lakes. Many of these woods occupy ground whereoriginal natural forest was clear felled and replanted from the late 18th century onwards, when forseveral reasons outlined by Rushton (1983), Q. robur might well have been the preferred species.Proof of this suggestion, or even whether the planters knew the species they were handling, is difficultto establish. While Pedunculate Oak is over twice as locally widespread as the Sessile Oak in Fermanagh,it is still nowhere near as frequent or widespread as fa*gus sylvatica (Beech), the latter beingthe 11th most frequent tree or shrub in the county.
Threats
Grazing pressure is preventing regeneration and many oaks are reaching a stage of over-maturity whenboughs begin to drop off. As trees die, active management of existing woods, plus a programme ofreplanting using local acorns is urgently needed to maintain the species and its genome at anythingapproaching present levels.
Native, occasional, but possibly under-recorded. European temperate.
8 January 1986; Leach, S.J., Corbett, P. & Dunlop, D.; wood on Gorminish Island, Lough Melvin.
Throughout the year.
There are 60 records from 51 Fermanagh tetrads referring to deciduous oak specimens in woodland whichfield recorders regarded as intermediate and which could not easily be referred to either parentspecies. As such they represent just over 10% of the oak records in the Fermanagh Flora Database – quitea significant proportion. They appear quite widely scattered, but lie chiefly in the northern half ofthe county. Leaf samples examined by Rushton (1983) found that oaks in woods at both the Marble ArchGlen (or Cladagh River Glen) and nearby Rossaa Td (between Mullaghbane and Gortatole) had mixedpopulations of both parents with a high proportion of the hybrid: indeed at Marble Arch almost 50% ofthe trees sampled were hybrids. However several woods also had low proportions of hybrids or none,including Killesher Forest NR, also near Marble Arch Glen, where the samples were exclusively Q.robur. In the Marble Arch wood, ash is the dominant tree on the damp valley floor and the oaksare confined to the higher slopes, where the spacing and even-age of the population suggest a managedplantation origin. The pattern of extensive removal of natural woodland and its replacement byplantations, especially on landed estates, is an all too common one across NI. It reflects alongstanding shortage of timber, major social change and land ownership both ancient and modern(McCracken 1971; Tomlinson 1982).
The limited number of oak populations so far studied in Ireland, and the high levels of introgressionfound in them tend to suggest that the frequent mixed populations of Q. robur, Q. petraeaand their hybrid that occur have most probably (and most often) arisen naturally in ancient times, orthrough plantation of mixed stock, rather than by recent horizontal spread of one or other of the oakspecies (Cousens 1963; Rushton 1983).
Introduced, neophyte, deliberately planted, very rare. Native of eastern N America, widely planted inEurope.
22 August 1986; EHS Habitat Survey Team; Lough Nalughoge, Crom Castle estate, Upper Lough Erne.
A native of NE America introduced to Britain around 1724 and grown for its remarkably attractive autumncolouring, like Q. cerris (Turkey Oak), this large tree has become much more available in thehorticultural trade in recent years. It is now very fashionable for planting in larger gardens and alongdriveways, as well as in public parkland and amenity areas around towns and cities. While it has beenpresent in tree collections in B & I for over 350 years, Q. rubra was not recorded in the'wild' until 1942. Over the last three or four decades it and several other fast growing N American oaksof the 20-strong Red Oak group (mainly Q. coccinea (Scarlet Oak), Q. palustris (Pin Oak)and Q. velutina (Black Oak)) have been increasingly planted across B & I. They are valued fortheir often very large leaves which fairly reliably produce a good bright red or strong yellow autumnleaf colour.
These four oaks are quite variable and phenotypically plastic in leaf form making them difficult todistinguish (Mitchell 1974 & 1996). It is therefore possible that 'Q. rubra' as recorded forthe BSBI New Atlas 2000 survey represents an amalgam of two or more of these rather similarspecies, several of which are regularly confused within the horticultural trade. The two species thatare mainly confused are Q. rubra and the even better colouring tree, Q. coccinea. Theseoaks should perhaps be referred to as Q. rubra s.l. following the example of Clement & Foster(1994).
With this proviso the New Atlas map shows Q. rubra very thinly and widely scattered inIreland, but much more frequent and widespread in Britain. Although Red Oak now appears to seed andnaturalise itself in parts of England (Jones 1959, p. 216), so far the few specimens recorded inFermanagh have all been deliberately planted and they are confined to demesnes or their near vicinity.
There are just eight Fermanagh records, and the details of the other seven are: Corralongford, NE ofColebrooke Park, 1 July 1997, RHN; Castle Coole parkland, October 1998, RHN; Florencecourt House, 7 June2003, RHN; Riversdale Forest, 16 January 2004, RHN & HJN; Derrychara playing fields, 16 May 2008,RHN; N of Rotten Mountain bridge, 9 September 2010, RHN & HJN; track behind old castle, CastleArchdale, 20 September 2010, RHN & HJN.
(ie Preston et al. 2002).
Native, occasional. Eurosiberian boreo-temperate.
1884; Barrington, R.M.; Ely Lodge Forest.
April to September.
Growth form and taxonomic status
Three species of Betula are native in Britain and Ireland. One of them, B. nana L. (DwarfBirch), does not occur in Ireland and in Britain is almost restricted to C and N Scotland. It is adistinctive, very dwarf, deciduous subshrub of high alpine habitats, is easily recognised and normallypresents no taxonomic problems for botanists. The same cannot be said of the two deciduous tree birches,however, B. pendula Roth (Silver Birch) and B. pubescens Ehrh. (Downy Birch). Thetaxonomic status of these two trees has been a matter of long dispute. Linnaeus (1753) treated the treebirches as a single species, B. alba, and some still consider the two forms in Britain andIreland as subspecies of this entity (Tuley 1973). There is a chromosome difference however: B.pendula is a diploid species or subspecies with 2n=28, whereas B. pubescens is atetraploid aggregate (2n=56). Despite the fact that there are only two species and they differgenetically, the tree birches constitute a difficult taxonomic group as a result of common andwidespread intermediate forms that are not readily separable from the parent species. Most intermediatesare fully fertile and therefore they probably backcross with the parents. The taxonomy can really onlybe understood when examined on a world scale using cytogenetic experimental study (Walters 1968).
To deal with the problem of intermediates, Atkinson & Codling (1986) devised a reliable leafdiscriminant coding equation based on three easily measured characters measured on five leaves collectedfrom side branches from the lower crown of the tree and averaged. The methodology is illustrated, but inthe opinion of the current writer, it not adequately explained in Stace (2010). Reference to theoriginal paper in the journal Watsonia is therefore recommended.
'Pure' B. pendula is described as a tree up to 30 m, with slender, pendant, glabrous twigs coveredwith wart-like resin-glands, smooth silvery-white bark and subglabrous, biserrate (doubly toothed)leaves. In addition to the described species identification difficulties, there are as many as ten namedcultivars of B. pendula in horticulture (Schilling 1984; Griffiths 1994).
Hybird birches
Hybrids between the two species occur to complicate matters. B. pubescens in particular isextremely variable, and the hybrid appears to overlap in leaf morphology more with it than with B.pendula (Kennedy & Brown 1983). The overlap of numerous characters occurs to such an extentthat the existence of hybridisation became a matter of dispute (Brown et al. 1982; M.D. Atkinson,in: Rich & Jermy 1998; Stace et al. 2015). The real extent of hybridisation between the twotree birches remains unresolved (Stace et al. 2015). In view of the great difficulty ofrecognising birch hybrids in the field, they have not yet been looked for in Fermanagh at all.
Distinguishing the two Birch tree forms
The Flora of Connemara and the Burren unequivocally states that B. pendula is grosslyover-recorded in Ireland due to the fact that many botanists, particularly visiting, non-nativerecorders, fail to appreciate that in Ireland the twigs of the much more common B. pubescens areoften without a hairy pubescence. This observation renders the possession of glabrous twigs almostuseless as a distinguishing character, and this is particularly the case in the north and west ofIreland. Experience shows Irish material of the two tree birch species can best be distinguished by thelarger, paler, more raised warts (c 1 mm in diameter), present especially on the younger twigs of B.pendula, and secondly by the appearance of its leaf tip, which is much more drawn out into aslender point (ie acuminate) in comparison with the leaf apex of B. pubescens, which is describedas sub-acute to acute (Atkinson 1992; Parnell & Curtis 2012).
B. pendula is regarded as the more 'shapely' of the two species, having fine long-pendulous shootsthat hang from elegantly arched branches borne on a straight trunk. In comparison, B. pubescensis described as having an untidy, shapeless and twiggy crown (Mitchell 1996). Unfortunately, tree shapeagain is not a reliable means of distinguishing the two genetic forms, since pendulous forms of B.pubescens do also occur (Nelson & Walsh 1993).
Preferred habitats
Populations of B. pendula of native Irish status are found mainly on the margins of lowland raisedbogs, or by limestone or in woods and scrub around stony lakeshores. It prefers well illuminated siteson well drained acidic soils, including less heavy clay situations. B. pendula can toleratesomewhat wetter and more shaded conditions as already mentioned, but in such sites B. pubescensis often the more prevalent of the two. In addition, elsewhere throughout Ireland, this colonisingspecies with very light, readily dispersed winged nutlet fruits, originates as occasional to commonnaturalised self-sown seedlings, often derived from nearby planted garden, parkland or demesne trees(Webb 1994; An Irish Flora 1996). Having a small seed, B. pendula is unable to coloniseecologically occupied sites such as grassy swards that might remain open to larger seeded tree species(Worrell & Malcolm 1998). In an unknown number of cases, the gene stock may be derived fromimported, non-native seed. Seedlings are most frequently found on recently cleared ground, especiallywhere vegetation has been burnt, but it can also colonise other forms of open ground, includingroadsides and the gravel drives of houses.
Growth, ecology and longevity
Silver Birch is light-demanding and, while it does not very readily grow in the shade of other trees,occasionally it does. Seedling birches are especially rare under the canopy of young trees of their ownspecies (Kinnaird 1968). Birch therefore tends to be found most often in open areas, including inclearings and on woodland margins. Birches can show very rapid growth during their first 20 years, oftenreaching 20 m in height over this time span. After the initial spurt in development, growth is slow,terminating in a decline that is often rapid due to fungal decay (Kinnaird 1968).
As pioneer colonists, Silver Birch trees are not likely to survive to an advanced age. No dated tree incultivation is known before 1905, but so few are of known age that this means less than it mightotherwise. It is generally assumed that in Britain and Ireland the trees die back, break up, or fallover when less than a hundred years old. In Scottish highland glens, the oldest trees were reckoned tobe around 220 years old (Mitchell 1996).
In birchwoods, it is often observed that apart from the occasional birch seedling, saplings are rarelyobserved under the woodland canopy. This is all the more remarkable as birch can establish itself in theshade of other tree species. Birchwoods consequently regenerate on suitable disturbed or open groundlying adjacent to pre-existing birch stands, rather than within them. Exceptions to this can occur wheregaps appear in deciduous woodland canopy, especially when this is accompanied by soil disturbance or isthe result of a fire. Birch regeneration depends on the vigour of the seedlings, and as with anyspecies, establishment is greatly affected by the local grazing intensity. Seedling vigour is a functionof soil fertility, but on poorer sites it may also depend on the successful establishment of amycorrhizal association with soil fungi (Kinnaird 1968).
Seed production and interactions with birds
Birch fruits are generally produced in abundance, although not in every year by an individual tree. Inother words, there is a sort of birch mast, as in beech and several other species, a good fruiting yearbeing followed by several of low productivity (Kinnaird 1968).
Birch trees are highly attractive to birds. The small, light, winged, single- seeded dry fruits are muchsought after by small finches and tit* including siskins, goldfinches and blue tit*. The foliage inspring and autumn supports large populations of aphids, the first crop of which feed nestling tit* andwarblers. The autumn fruit crop feeds the adult birds, the warblers building up fat reserves to fueltheir migration. Rotting tree bases and stumps of birches are bored for nest holes by willow-tit*(Mitchell 1996).
Fermanagh occurrence
B. pendula, which is also referred to as Pendulous or Warty Birch (the latter name much deplored,being a quite horrible name to give to any plant), is almost a rarity in Fermanagh. It has only beenrecorded from a total of 26 tetrads, 18 of them containing post-1975 records. While the distribution mapshows B. pendula is thinly and widely scattered across the vice-county, to some extent itreflects areas of likely plantation woodland associated with landed estates.
British and Irish occurrence
The New Atlas hectad map shows B. pendula as being common and very widespread throughoutmost of Great Britain, but absent from high ground, and becoming less frequent in NW Scotland. Onaccount of the ability of the two birch tree species (B. pendula and B. pubescens) toreadily colonise felled woodland, Kinnard (1968) speculated that they might well be the commonest treesin Britain. In Ireland, the hectad map indicates that B. pendula is much less frequent than inBritain, and it become rare or absent in the west of the country (Preston et al. 2002). Recently,doubt has been cast on the Scottish distribution of B. pendula, as Worrell & Malcolm (1998)have found anomalies which suggest that it is greatly over-recorded in much of that country, fewrecorders being able to reliably distinguish it from B. pubescens. While no one knows for certainat present, a similar situation might very possibly apply in Ireland, and perhaps even throughout thewhole of the British Isles (Wurzell 1992; Webb 1994).
Names
The genus name 'Betula' is the Classical Roman name for the tree, for example in Pliny (Gilbert-Carter1964). The familiar Latin specific epithet 'pendula', means 'hanging down'. The English common name'Birch' is from the Anglo-Saxon 'birce' or 'beorc', and it has many similar names in other northernIndoeuropean languages, for example, 'berk' in Dutch, 'Birke' in German, 'birk' in Danish, 'beith' inGaelic and Old Irish, 'begh' in Irish. The Indoeuropean root of the name means 'white' or 'shining',probably referring to the bark (Grigson 1974). Many place-names contain the elements 'birk' or 'birch'in England, or 'beagh' and 'behagh' in Irish, meaning 'birch land' (Milner 1992).
Uses
The timber of both tree birches is hard, strong and works and turns well, so it has been used for a widevariety of chiefly indoor household purposes (it is not durable outdoors). Uses include furniture, asbrooms (besoms), tool handles and thread spools (until replaced by plastic). Birch branches also provideexcellent firewood and the jumps for steeplechasing (Milner 1992). The birch (hereafter meaning the twospecies undifferentiated), had the rank of a 'Commoner of the wood', the second grade of four in theeighth century Irish Laws of Neighbourhood. Most likely it achieved this rank on account of itstimber value (Nelson & Walsh 1993).
The bark produces Birch Tar oil, which is, or was in the past, used in the tanning of leather. Thedistinctive smell of Russia leather is due to its use as a dressing. Russia leather is very durable andit was the preferred material for book binding since the tar oil dressing also prevented or discouragedattacks on books by fungi and insects.
Birch sap has had a long history of use in brewing, which continues today in parts of E Europe (Grieve1931; Milner 1992). As a consequence of its long history and many uses, a very well-developed folklorehas grown up around birch, particularly in northern regions of Europe where the trees are mostprevalent, and there are numerous folklore rituals and superstitions attached to it (see Grigson 1987and Milner 1992). Birch sap and birch tar oil have also been used medicinally (see Grieve 1931).
Threats
None.
Native, very common and widespread. Eurosiberian boreo-temperate.
1900; Praeger, R.Ll.; Co Fermanagh.
Throughout the year.
Growth form, identification difficulties and preferred habitats
Although the common and the scientific name both refer to 'downy birch', plants of this species inFermanagh, as in most of Ireland, have twigs which are almost hairless (Webb 1994). B. pubescensis a diploid with chromosome number 2n=28. Birch trees are monoecious and can begin flowering when theyare only five to ten years old. Individual trees are mostly self-incompatible, and as could be expectedin any widespread self-sterile species with a broad geographical and ecological range, there isconsiderable genetic variation and overlap between species, subspecies and forms.
B. pubescens can be subdivided into the tree form, subsp. pubescens and the shrubby uplandform, subsp. tortuosa (Ledeb.) Nyman; the latter is especially frequent in N Britain, but is veryprobably present in N & W Ireland too (New Flora of the BI; M.D. Atkinson, in: Rich &Jermy 1998). No birch subspecies have yet been distinguished in Fermanagh, nor has the hybrid with B.pendula (which is a tetraploid taxon with 2n=56), ever been recorded. However, the wholeproblematic question of the true identity of our birches compared with those in Britain requires furtherstudy.
B. pubescens occupies damp open woods and scrub, frequently but not always in upland areas at ornear the potential tree-line. It also frequents peat bogs (including cut-over examples of these),cliffs, stabilised boulder screes and disturbed and open abandoned ground generally. In most parts ofIreland, birch is most conspicuous when the trees are young and are colonising thickets, or whenwell-spaced individual trees are found invading disturbed open situations on peaty moorland, heath orbog. These colonising situations usually occur after there has been a fire or turf-cutting operationshave taken place, both of which require previous drainage. The other absolute essential for successfulestablishment from seed is the creation of a bare, almost litter-free soil surface (Gimingham 1984).
Variation
Within both species of tree birch that occur in Britain and Ireland (B. pubescens and B.pendula), there is very great phenotypic plasticity (ie environmentally induced variation). Forinstance, when tetraploid trees (therefore supposedly B. pendula) were transplanted into both dryheath and rather wetter bog conditions, Davy & Gill (1984) found that the leaves of those plantedinto the heath soil became more like B. pendula, while those planted into boggy ground resembledB. pubescens! Similarly, a detailed biometric study of transplanted B. pubescens seedlingsgathered from 25 populations which examined and compared no less than 36 variable characters, found thatseedlings with small leaves and drooping branchlets were associated with westerly, wet locations,whereas most seedlings with erect branches and larger leaves originated from drier, more easterlylocations (Pelham et al. 1988).
Glacial history and native status
Both tree birches are native throughout the British Isles and fossil pollen studies prove they were theearliest tall growing woody plants to colonise these islands as the glacial ice finally retreated around10,000 BP. This accords with the position of birch zones on modern tree lines at the margins of both thearctic tundra and the high mountain tundra further south. It appears that before 10,000 BP, tree birchwas well established in much of C and N England, S Scotland and parts of Wales, and tree birches justmanaged to reach two bridgeheads in E Ireland. This pattern very strongly suggests that the spread ofbirch was emanating from the east, probably migrating from the then dry North Sea basin. By 9,500 BP thetree birches had expanded to cover northern Ireland and parts of the Scottish Highlands. Interestingly,expansion was delayed (perhaps for up to 500 years), in areas of the Scottish Highlands which werere-glaciated during the Loch Lomond Stadial (ie a short spell local return of cold, glacial conditionswithin a warm interglacial period). Presumably the delayed birch arrival was caused by the time requiredfor the soil to mature sufficiently after glaciation or periglacial conditions ceased. Periglacialconditions involve near-glacial, freeze-thaw perturbation that effectively turns over and rejuvenatesthe soil. Conditions on the glacial moraine ridges exposed by the thawing ice could also involve soilinstability and 'frost drought', which would seriously affect Betula species since theirseedlings are very sensitive to any form of drought (Birks 1989).
Fermanagh occurrence
Betula pubescens is the locally common and very widespread birch tree and shrub in Fermanagh,present in 396 tetrads, 75% of those in the VC. It is the eighth most frequent woody species in thecounty after Salix cinerea subsp. oleifolia (Common Sallow orRusty Sallow) and the tenth most widely distributed woody plant after Acer pseudoplatanus(Sycamore). In comparison, a Britain-wide forestry survey found birch (hereafter referring toboth tree birch species) was found to be the second most common broad-leaved tree in England andWales and the most common tree in Scotland (Steele & Peterken 1982).
British and Irish occurrence
Our best knowledge of the present day British Isles distribution of the two tree birches suggests thatthey overlap considerably. This is more the case in Britain than in Ireland, where B. pendula isvery much less frequent and widespread than B. pubescens. In general, B. pubescens is moreprevalent in the north and west of both islands, while B. pendula is more common in the south andeast (Gimingham 1984; Preston et al. 2002). Recently, however, doubt has been cast on thesupposed Scottish distribution of B. pendula. Studies by Worrell & Malcolm (1998) foundanomalies which suggest B. pendula is over-recorded in much of Scotland, few recorders being ableto reliably distinguish it from B. pubescens. While no one knows for certain at present, asimilar situation might very possibly apply in Ireland, and maybe even throughout the British Isles(Wurzell 1992; Webb 1994).
The degree of confusion and need for further research is such that some botanists have even called for areturn to the Linnaean single species, Betula alba, demoting B. pubescens and B.pendula to subspecies until their true status can be researched and clarified (Tuley 1973;Nelson & Walsh 1993). Nevertheless, even given the possibility that B. pendula may beover-recorded, the overall trend observed with regard to the distributions of the two tree birches inthe British Isles is believed to be reflected in Continental Europe, ie B. pubescens extendsfurther in the colder north and east, reaching Iceland, while B. pendula stretches further southinto warmer, drier regions than B. pubescens, reaching the mountainous regions of Spain, Italyand Greece (Jalas & Suominen 1976, Maps 269 & 270).
Birch ecology
Tree birches are particularly cold hardy pioneer species, but the northern limit for both species isstill set by exposure to cold and dry NE winds in places like Iceland and Greenland. Even in northernparts of Scotland, seedlings of B. pendula in particular are killed or suffer severe die-back inperiods of very cold, dry, winter weather, while by comparison B. pubescens seedlings suffer lessdamage. This suggests that in at least some parts of Scotland, both birch species are fairly close totheir ecological survival limits with respect to extremely cold winds (Atkinson 1992). The latter authorsuccinctly described the main behavioural characteristics of birch tree ecology (both speciesundifferentiated), in terms of: (a) ability to quickly colonise bare areas, (b) intolerance of shade,(c) lack of affinity for any particular soil type (they can tolerate a very wide range of substrate pH),and (d) ability to grow on nutrient-poor soils.
While this is a generally useful summary, it requires some qualification in that B. pendulaappears more restricted to drier, lighter soils in more sheltered lowland situations than B.pubescens, which can perform perfectly on rather wetter (although seldom permanentlywaterlogged), more peaty, wind exposed situations on upland heaths, moors and bog margins (Gimingham1984). A much rarer and more unusual occurrence is where birch woods or copses are found growing on wetflushes or soaks on bogs, as described by Cross (1987) from two Irish raised and blanket bogs in CosOffaly and Mayo (H18 & H27). It is difficult to imagine the precise factors which allow the ratherstunted, multi-stemmed birches (chiefly, but not exclusively, B. pubescens), to colonise theseparticular bogs. Human interference and disturbance of the bogland in the past may well be involved, eggrazing and burning. Straightforward human neglect of the ground in question may also be partlyresponsible for birch invasion. Whatever anyone thinks of them, these two birch stands on bog soaks aremost unusual, and I am not aware of any Fermanagh parallel to them.
Birch in woodland
Reflecting the broad soil range of both tree birches, the massive yearly output of small, wind-dispersed,single-seeded fruit and their consequent rapid colonising ability, plus their characteristic high lightdemand, rapid initial growth rate and short lifespan (typically 80 years), birches are regularly presentin a wide variety of woodland types throughout Britain and Ireland. In many woodland situations,however, birches are not necessarily very numerous nor very conspicuous. Rather, they simply represent aconstant, minor component, filling gaps in the canopy that occur sporadically within the woodland standdue to the death of larger trees. Alternatively, they may form a marginal exposed fringe to thewoodland, rather than occurring as dominant canopy-forming species within it, or forming an extensivepure birch stand (Gimingham 1984).
In Fermanagh, as in most other parts of Ireland, birch is most conspicuous when young colonising thicketsor well spaced individual trees invade disturbed peaty moorland or heath, say after fire or turf cuttingoperations involving both drainage and, in the case of birch, what is absolutely essential forsuccessful establishment from seed, the creation of bare, almost litter-free soil. Birch also regularlyinvades the drier sloping margins of raised bogs, or abandoned cut-over bogs (which have always beensubjected to drainage operations of some sort). However, as in other situations, this generally requirestemporary severe grazing pressure or some other form of disturbance to create the vegetation gaps andthe bare ground needed for successful seedling colonisation (Gimingham 1984).
Seedling growth and colonising ability
Birches do not form a persistent soil seed bank, and with the seed being so very small, the new seedlingsquickly exhaust the very limited seed food supply, and to survive must rapidly develop the mycorrhizalfungal partnership on which they are so dependant (Milner 1992; Ingram & Robertson 1999). Until themycorrhizal fungal root sheath develops and grows out into the soil, the seedling is very vulnerable toboth nutrient starvation and drying conditions. When invading newly available ground in this manner,birch is sometimes in the vanguard of a vegetation succession responding to major change in theenvironment, and if protected from grazing and other destructive pressures and allowed to develop, wouldeventually build a more diverse woodland dominated by taller, long-lived species more capable ofregeneration within woodland canopy shade than the light-demanding Betula species (Atkinson1992).
This is not to advocate that a completely stable climax vegetation of mixed deciduous woodland dominatedby oak, ash (or beech), would necessarily always develop, since the terrain and its stability might notprove suitable or conducive for such vegetation in the longer term (Tansley 1949). Vegetation historyfrom the Neolithic onwards suggests that man and his activities have played such a dominant determiningrole, that vegetation scarcely ever reaches anything approaching stability or a classical Climax state,but rather it is characterised by reaction to more or less continuous change (Mitchell 1995). Certainlyolder birch may instead be replaced by young birch if human induced disturbance is sufficient tomaintain or recreate open light conditions, thus favouring opportunistic recolonisation by birch seed.As Rackham (1980) so very well puts it, "Throughout the history of birch its supreme colonisingability has been balanced against the greater competitive ability and longer life of other trees."He has also commented, "The reproductive ability of birch is so great as to outweigh even thedisability of its short life-span."
Reputation as soil improvers and nurse trees
In forestry, the pioneer seral role of birch in vegetation change and development is recognised in thatbirches have the reputation of being both soil-improvers and useful nurse trees for longer lived, morevaluable timber crops. In both these aspects, however, this reputation may be challenged. Miller (1984)found that while birch did improve a heath mor podsol, quite rapidly changing it into a less acidicbrown earth with a mull humus and with a greatly increased worm population, it performed this service nobetter than would other tree species of similar growth rate. Likewise, in exposed sites the whippy topsof birch trees are capable of killing the apical buds on leading shoots of adjacent trees, in thesesituations at least, effectively negating any nursing abilities the birch might otherwise possess. Thewhole idea of using 'nurse trees' in forestry is really now a thing of the past due to reductions inavailable manpower, and thus in forest management possibilities.
In reality, the birchwoods most commonly seen in Britain and Ireland appear to have arisensecondarily as a result of felling of woods of all types on a wide variety of soils. Theconditions following felling are obviously highly suitable for colonisation by birch, provided that seedparents are available reasonably close nearby. The relative abundance of lightweight birch fruit and theefficiency of its wind dispersal compared with the seed of other native trees, enables it to rapidlyinvade and exploit the open situation as a pioneer species. In summary, the conditions created byfelling operations which enable birch to invade and compete involve the following; the surfacevegetation is disturbed, the mineral soil is loosened and mixed with the litter and humus to someextent, the soil moisture regime is diversified, soil surface temperatures are elevated by increasedsunlight, and there is an increased rate of decay and mineral release from soil organic matter (Kinnaird1968).
Birch as habitat and food provider
Birch has a very important role as a habitat and food provider supporting other organisms and togetherthe two tree Betula species have an impressive tally. They support at least 334 herbivorousinvertebrate species, more than any other native tree apart from oak and willow (Kennedy & Southwood1984). The presence of so much insect life naturally makes the trees highly attractive to birds. Theleaves in spring and in autumn carry a large population of aphids, the first crop of which supportsnesting tit* and warblers. The autumn aphid crop builds up the fat reserves of adult warblers for theirmigration.
The often abundant seeds are sought by finches and by a wide variety of other small seed-eating birdsincluding blue tit* (Mitchell 1996). Browsing of saplings and seedlings is an important cause of bothmortality and modification of birch tree growth. Trees can respond to mechanical damage by producinglonger shoots, which favour the escape of the growing points to heights above the reach of the browsinganimal. Birch leaves also respond to insect herbivory by increasing their phenol and tannin content,which renders them much less palatable (Atkinson 1992).
Fungal, lichen and bryophyte associates
Apart from the mycorrhizal fungi vital for the establishment and subsequent nutrition of the tree (Masonet al. 1984), the fungal associates of birch include one of the most familiar gall-formingspecies, the witch's broom. These structures, often quite numerous on individual trees, are a tightproliferation of shoots caused by attack on a shoot apical bud by the ascomycete Taphrinabetulina on B. pubescens, and T. turgida on B. pendula (Milner 1992; Ingram& Robertson 1999, p. 103).
Being relatively short-lived and having very acidic bark, the tree birches support just 93 forms oflichen, a total which ranked it eighth among native trees in the British Isles in this respect after thetree willows and hazel (Rose 1974). Subsequently, however, Coppins (1984) listed no less than 235species of lichens and 58 bryophytes (mosses and liverworts), growing as epiphytes on Betula inthe British Isles. The greatest lichen development on birches was found in Scotland for a variety ofreasons, not least of which was the lesser degree of sulphur dioxide pollution (Coppins 1984).
Names
See my Betula pendula account.
Uses
See my Betula pendula account.
Threats
Evidence from Britain suggests that both our native birches may be susceptible to leaf and possibly stemdiseases caused by Phytophthora ramosum. Birches are not at the present moment in anyconservation danger themselves, but as the fruit is so very mobile and invasive, they can become 'woodyweeds' in fens, bogs and heathland. Control is expensive and very difficult to achieve.
Native, very common, widespread and abundant. Eurosiberian temperate, introduced in eastern N America.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Under ideal conditions, A. glutinosa trees can grow large, up to 25 m in height with a girth ofaround 3.5 m. However, in Fermanagh woodland and scrub they usually only reach around 6 to 10 m. Thevast majority of individual trees have several stems, rather than a solitary trunk, and they are notlong-lived (perhaps up to 100 years). As such, they might be considered as large shrubs, rather thanproper trees. The seeds of the tree are chiefly dispersed in flowing water, and thus its prevalence inwaterside habitats. The trees or shrubs often carry dead or dying die-back branches. A.glutinosais readily identified even when minus its leaves, as the distinctive, small, barrel-shaped, woody femalecones are constantly present. In addition, an examination of the bare twigs shows the presence of budswith a purple waxy bloom, borne on distinct stalks (Edlin 1964).
Writing in a British Isles context, Rackham (1980) points out that, "Of all familiar native trees,Alder is the most obviously restricted to particular habitats." Rackham (1980) recognised threetypes of alder woodland in Britain:
(1) Fen alder-woods, extensive on low-lying, level ground in the flood-plains of streams, rivers andaround lakes, almost always containing alder mixed with a range of willows;
(2) Valley alder-woods, forming narrow fringes to streams, often within other types of woodland, notablyash-alder, mixed oak wood, and birch-alder. This category of alder-wood may sometimes climb up flushedslopes, especially in western parts of Britain and Ireland, the dominant tree species changing withaltitude and exposure.;
(3) Plateau alder-woods, in which alder is a constituent of the mosaic of woodland trees on leveluplands, sometimes with slight depressions, and often on a watershed which provides seeping irrigation,at least in winter.
Rackham's first two alder-wood categories are very common in Fermanagh, but there is very little or noneof his plateau woodland involving alder anywhere in the VC. The upland plateau of W Fermanagh is insteadoccupied by modern conifer plantations on ground that previously supported either acidic blanket bog orpeaty moorland pasture.
Fermanagh occurrence
Alder trees help form an important part of Fermanagh's landscape: on almost every streamside, river bankand lakeshore, the dominant species is Alder and it is present in 470 tetrads, 89% of those in the VC.In Fermanagh, A. glutinosa is most often found as a non-woodland tree fringing all types ofwatery habitats, and indeed it is a very good indicator of land liable to flooding. It is characteristicof shrubby woods fringing marshy lakeshores and along river banks. Alder ranks as the fourth mostwidespread woody species in the vice-county, and it is also the tenth most frequently recorded vascularplant in the county survey, which testifies to the damp mild climate and poorly drained soils of thispart of western Ireland.
Alder ecology
Alnus is essentially a pioneer genus with the associated characteristics of heavy seeding, rapidcolonisation of bare ground, fast initial growth, intolerance of shade and a short life span (White& Gibbs 2000). A. glutinosa is very sensitive to shading, the seedling less so, butstill requiring a higher light intensity than seedlings of larger-seeded tree species. This fact aloneexplains why internal regeneration of Alder in woodland is practically impossible (McVean 1953).
Soil reaction
Alder is largely but not totally indifferent to soil reaction. It tends to be restricted to the unstablesoils of stream and lake margins, recent alluvial river soils, flush soils and very wet soils where thewater table is seasonally high, or where drainage is impeded. Alder roots are of great importance inhelping to stabilise waterside soil banks, and the flow of water assists the tree by providing oxygen tothe soil around its roots. Alders do not thrive around stagnant pools or where water flow is slight,although they may persist in a state of suspended animation for a long time after streams have beendiverted (Mitchell 1996). A. glutinosa can grow on deep fen peat, or on acid peat ofMolinia-Myrica bog, but not on blanket or raised bogs as noted above.
Effect of grazing pressure
Moderate grazing of marshes or fens prevents associated tall herb vegetation being kept from smotheringand shading out the trees' seedlings and thus favours the spread of alder. Associated animal tramplingalso helps provide suitable sites for establishment by breaking the surface turf and the leaf litterlayer. Cattle will trim young saplings and limit their growth to bush form if grazing becomes toointensive. Rabbits and hares seldom ring-bark saplings of alder in the way that they do with birch orplantation conifers. McVean (1953) suggested that alder does not establish itself as well in the'natural' aquatic vegetation succession (ie primary hydrosere), as it does when that pattern of habitatdevelopment is disturbed or deflected by the activities of man and his farm animals.
Frost and drought tolerance
Mature alder trees are extremely frost hardy, and once deep taproots have been formed, individuals arealso very drought resistant. It appears therefore that the plant's strong association with wet soilschiefly reflects its seedlings susceptibility to drought and shade.
Beneficial root associates
Mycorrhizal roots and roots bearing nitrogen-fixing nodules are formed; the nodules are of two sizes,many very small and few large, orange-coloured ones like tennis balls, which can live for a decade. Thesymbiotic organism contained in the nodules is an Actinomycete (generally strains of a genus calledFrankia, one of a group of organisms related to filamentous bacteria). These nitrogen-fixingnodules confer considerable nutritional advantages to trees that bear them (Thomas 2000).
Low oxygen in waterlogged soils
Oxygen availability is often a limiting factor for organisms including trees that live in soils subjectto flooding. Some oxygen will dissolve and diffuse down into the upper few cms of waterlogged soil, butbeneath that shallow zone the substrate is devoid of oxygen (anaerobic). Woody roots can surviveanaerobic conditions for a time when the tree or shrub is dormant, but finer roots quickly die, with aconsequent decline in uptake of both water and mineral nutrients, directly affecting growth. Indirecteffects include toxic substances produced in the soil (for instance, hydrogen sulphide) and toxinscreated by the tissues of the tree itself (Thomas 2000). Being able to exist temporarily withoutsomething (oxygen) that is constantly essential for your competitors is one of the great keys toecological survival, the so-called 'anaerobic retreat' (Crawford 1989).
Tolerance of low oxygen
Alders tolerate waterlogged soil extremely well, and some of their roots live permanentlysubmerged in water. Part of the mechanism that permits the tree to survive such conditions involves thepassage of air downwards through the bark and within air-filled sections of the woody xylem tissues ofthe root. The bark- and wood-producing thin cambium layers were once considered impervious to gas, butit is now clear that this is not the case in wetland trees (Thomas 2000). Oxygen diffuses down and leaksout of the root into the surrounding mud, forming an oxygenated envelope around the root, permitting theroot tissues to function normally. By the same route, toxic gases produced by incomplete anaerobicrespiration can escape from the root, although many flood-tolerant trees also have biochemicaladaptations to reduce the production and impact of these toxins.
British and Irish occurrence
Alder is abundant throughout most of Britain & Ireland, but is less common in the far N of Scotlandand in the Central Plain of Ireland due to the extent of extremely wet, cold, acidic Sphagnumpeat (ie raised and blanket bogland) that occurs in these regions, conditions which exclude all treesexcept birch (Betula spp.).
European and world occurrence
Alder extends throughout most of Europe except the Arctic, the Mediterranean basin and the Russiansteppes, and it is also present in Asia Minor, W Siberia and N Africa (McVean 1953).
A new pathogenic disease attacking alders
In 1993, a lethal stem disease affecting A. glutinosa was first noticed and described in Britain.On affected trees the leaves are described as abnormally small, yellow and sparse, especially on thecrown of the plant. The leaves frequently fall prematurely and leave the tree bare. The stem base oftrees that are displaying severe crown symptoms often have black 'tar-like' spots on the bark up to 2 mfrom ground level. The spots indicate that the underlying bark is dead. The pathogen attacks the cambiumlayer of the root and lower trunk so that the tree dies from the base upwards. This new alder diseasehas been recorded throughout Europe and is caused by a fungus that infects trees through the action ofwater-borne zoospores, so it is not surprising that it appears, so far, most frequent and most virulentalong riversides (Rackham 2006).
The causal organism was first considered to be a hybrid between two species of the genusPhytophthora, P. cambivora and P. fragariae, both of which are consideredintroductions to Europe. Neither of the Phytophthora species causes disease in alder on its ownaccount (Brasier et al. 1999). More recent work has indicated that P. × alni was generated by hybridisation between P. uniformis andP. × multiformis (Ioos et al. 2006; Husson et al.2015). The emergence of P. × alni highlights the dangers of sexualhybridisation between Phytophthora species resulting in novel phenotypes/genotypes that can bemore damaging than either of the parental strains (Brasier 2000; Boutet et al. 2010; Érsek &Man, in: 't Veld 2013).
Die-back of alder trees attributed to P. × alni, P. uniformisand/or P. × multiformis has been reported in at least 12 Europeancountries in both forests (11 countries) and in forest nurseries (seven countries) (Jung et al.2016). The presence of the pathogen in forest nurseries implicates the horticultural trade in spreadingthe disease throughout Europe. Since the disease has become widespread in Europe it has focussedresearch throughout the European Community on the importance of alder, to the extent that an ECConcerted Action Plan for alder conservation was approved in 1998. Surveys have shown that diseaseseverity varies in different regions, but it seems to be at its worst in parts of Britain and Francewhere destructive epidemics appear to be developing with thousands of trees dying each year.
In Britain, Webber et al. (2004) estimated the disease had infected some 90,000 alder trees. Today(2017), it is estimated that P. × alni has infected between 16-20% ofthe riverside alder trees in Britain, including introduced exotic alder species. The worst areas for thedisease at present (ie in 2017) are in SE England and along the border with Wales, although the pathogenis now also present along Scottish riverbanks.
The first record of the pathogen P. × multiformis in Ireland wasreported in 1999 in Co Dublin from Alnus bark and from water baits in a riparian zone in thecounty (Clancy & Hamilton 2001). While the distribution of the pathogen in Ireland is not yetunderstood, symptoms of alder die-back and confirmation by isolation has been made in five recentlyplanted alder stands (5-20 years old) in counties Cork (H3-H5) and Kildare (H19) (O'Hanlon et al.2016), as well as in a mature Alnus in a riparian zone near infected recently planted alder treesin Co Cork [VC location not disclosed]. DNA sequencing of these isolates revealed them to be P. × alni and P. uniformis. Fortunately, so far there have beenfew cases reported of the disease in Ireland, although plant pathologists have been alert to thepossibility of infection (A. McCracken, pers. comm., 2001;https://www.teagasc.ie/crops/forestry/advice/management/phytophthora-ramorum/ accessed December 2018).However, because so little is known about the disease, there is no way of predicting the effect on alderpopulations 10 or 20 years from now. Basic research on the nature of the disease is currently beingpursued to discover how the pathogen attack on alder may be arrested, but the P. alni complexappears to be a growing threat to native Alnus stands in both Northern Ireland and the Republicof Ireland (O'Hanlon et al. 2016).
Ecological importance
There is growing concern that loss of significant numbers of the only British Isles native alder todisease may have far-reaching environmental consequences. Apart from providing habitat, food and coverfor a variety of wildlife, aquatic, terrestrial and avian (eg the achenes are a very important foodsupply for finches), alders also play an extremely important role in riverbank stabilisation, erosioncontrol, soil fertility improvement and the regulation of stream water temperature through shading.Alder is particularly important along headland streams where it supports dense populations of trout andsalmon, directly by providing shading, cover and habitat, and indirectly by providing food in the formof insects associated with the canopy, or with leaf litter (Tipping 1999; White & Gibbs 2000).
Uses
Alder can be made into a fine, even-grained charcoal, and is used in the manufacture of top-qualitygunpowder. The wood is noted for its durability under water and, as a consequence, has been used forpilings, sluices, pumps, troughs, boat bottoms and punts. Burr alder veneer is also greatly prized bycabinet-makers for its decorative grain (White & Gibbs 2000). Alder has a long tradition of beingvalued timber in Ireland. In an eighth century Irish document, Bretha Comaithchesa, the Lawsof Neighbourhood, which protected trees and shrubs, alder ranked in the second order ofimportance, Aithig fedo - the commoners of the wood. The fine imposed for cutting a branch off aneighbour's 'commoner', was a sheep. In Irish folklore alder was regarded as unlucky - best avoided on ajourney, and the tree was best left uncut, since the timber, although white when first cut, soon changesto red, like blood (Nelson & Walsh 1993).
Names
The Gaelic Irish name for alder is 'fearnög', 'fearn', or 'fern', which appears in place-names such asFerns, Co Wexford and Ferney, Co Fermanagh. These names have no connection with the English vernacularplant name 'fern', for which the Irish equivalent is 'raithneach' (Nelson & Walsh 1993).
Threats
Importation of infected plant material carrying a newly recognised invasive fungus, Phytophthoraalni, is a recently developed major risk to riverbank Alder populations in Britain.Unfortunately, the disease symptoms are very non-specific, making infection difficult to detect, but acareful eye must be kept on the possibility of the pathogen spreading in Ireland.
Introduction, neophyte, deliberately planted, probably only occasional, but perhaps under-recorded.Circumpolar boreal-montane, but absent as a native from most of W Europe.
1953; MCM & D; Castle Coole estate.
July to August.
Although introduced into cultivation in Britain as long ago as 1780, Grey Alder has only become reallypopular with local county councils in the last couple of decades. It is now recognised as a very hardytree that is easy to grow and which tolerates nutritionally poor, wet soils. As such, it is increasinglyused to plant up roadside cuttings and embankments and alongside rivers.
Alnus incana reproduces in these islands by both seedlings and suckers, so it can readilynaturalise itself. In Fermanagh, it is only occasionally recorded, having been listed in 18 tetrads, 16of which have post-1975 records. In view of the substantial increase of recent amenity planting by localcouncils, the species is very probably under-recorded here, as is often the case elsewhere in B & I(T.D. Dines, in: Preston et al. 2002). Recent records in Fermanagh are from the edges of coniferplantation blocks, where it helps 'soften' the landscape impact, and also from waterside and amenityplantations where it may have been planted as an alternative to A. glutinosa (Alder).
Grey Alder is a native of Eurasia and since it grows very well in the wet soils so common in NI it may beexpected to continue spreading, both with and without the assistance of man.
Threats
So far, A. incana is too rare to present a conservation problem, but in the future it might wellinvade semi-natural fen-carr scrub and then begin ousting native species. A. incana is moreresistant to Phytophthora alni than the native species.
Introduced, neophyte, deliberately planted and rare or very occasional. European temperate, alsocultivated and widely naturalised in Europe.
17 September 1986; McMullin, A.S. & Corbett, P.; Mullynacoagh Td shore, Upper Lough Erne.
January to November.
Growth form and preferred habitats
This is potentially a very large deciduous tree, up to 30 m with a wide, broad crown. Older trees havedeeply fissured bark on the trunk, the longitudinal fissures often spirally curved, allowing the speciesto be easily recognised at any time of year. Ecologically, C. sativa can tolerate mostsoils, although it thrives best on moist, acidic, sandy ones (T.D. Dines, in: Preston et al.2002). In season, the leaf canopy casts a heavy shade, while the deep, persistent leaf-litter creates amor humus, often with a pH of 4 or under (Rackham 1980, p. 332). In Fermanagh, conspicuous, distinctiveand often very large old trees of this species are almost always associated with parkland or treecollections (arboreta) in landed estates such as Castle Coole, Crom or Colebrooke. Unlike thequasi-native chestnut woods in Kent, E Essex and SE Suffolk described by Rackham (1980, pp. 332-9) whichare or were coppiced every 12-14 years for small timber, in Ireland C. sativa never formedwoodland stands. A possible reason for this is the more recent date of introduction and plantation ofthe tree in Ireland, probably sometime before the 17th century – although nobody really knows a definitedate (Nelson & Walsh 1993, p. 105).
Flowering reproduction
Trees flowering in June and July, the minute flowers being borne on long catkins held erect at the tipsof shoots. Individual catkins are 10-20 cm long, with female flowers at their base and more numerousmale flowers on the remainder of the catkin length. Various insects pollinate the flowers and thecharacteristic prickle-covered green chestnuts develop until the autumn. In most of Ireland, the vastmajority of ovules abort and it is doubtful if any of the chestnuts produced in northern counties arefertile and capable of germination as self-sown trees are very rare or totally absent. I expect that thepartially developed chestnuts are eaten by birds and small mammals as they do not seem to accumulate,and in my experience they quickly are taken.
European region of origin
C. sativa is a native species of the more humid areas of eastern Mediterranean countries.However, selected cultivated forms, some of them grafted with superior fruiting scions, have long sincebeen introduced into more northerly regions of Europe, and the tree and its edible nuts and otherproducts has been present in Britain since Roman times (Zohary & Hopf 2000, p. 189). Despite thefamiliar 18th century English common name 'Spanish Chestnut' and the present day frequency of the treein the Pyrenees, the species is unlikely to be native as far west as Spain (Rackham 1980, p. 329). Thenative range of C. sativa is controversial. The editors of Flora Europaea took aconservative view, regarding it as indigenous in the Balkans (Albania, Greece, Jugoslavia) and Turkey,and less probably in SC Europe (Tutin et al. 1993). It is common in Italy as well as in southernFrance and Calabria, where it is extensively coppiced (Rackham 1980, p. 332).
British and Irish occurrence
The New Atlas hectad map demonstrates that C. sativa is much more thinly scattered inIreland than in Britain, and especially so when compared to S England and Wales. The tree flowers andfruits readily, and in suitable, warmer conditions in S England, if not elsewhere at present, the nutsgerminate in the spring after their production. Unlike Beech and Oaks, losses due to seed predationappear rare in England. As it cannot regenerate under its own shade, transport of the large seed intogaps, or to sites under other tree species is necessary, the most likely vectors being rooks (Rackham1980, pp. 334-5).
Fermanagh occurrence
This is a rare or very occasional tree in Fermanagh, having only been recorded in 15 tetrads, 2.8% ofthose in the VC. It is confined to estate parks and woodland plantations, plus a few outliers nearby.Only one or two of the records occur on the margins of the demesnes mentioned above and, although Igreatly doubt it, these might just be self-sown trees. However, in all our years of field recording RHNand I have never found either seedlings or saplings of Sweet Chestnut anywhere in Fermanagh.
In truth, the reproductive biology and local behaviour of the exotic (ie non-native) trees and shrubs inN Ireland have had almost no attention paid to them. In many cases, it is not known whether they cansuccessfully set seed, disperse and establish in this part of the world, as many of them undoubtedly doin S England (T.D. Dines, in: Preston et al. 2002).
A well-known English tree expert, the late Alan Mitchell, suggested that only a few exotic, introducedtrees "plant themselves" in Britain and Ireland. He highlighted one very obvious exception'that proves the rule', as Acer pseudoplatanus L. (Sycamore). Mitchell went on to list SweetChestnut along with several introduced conifers that he believed could produce natural seedlings,although he reckoned these were likely to be "very local" (A. Mitchell, in: Milner 1992, p.158).
In Fermanagh, as elsewhere, in the autumn it is certainly possible to find shiny chestnuts inside theleathery, green or brown, heavily-spined husks which split into four valves beneath the tree. However,it is very doubtful if they contain viable seed in Ireland as far north as Fermanagh. This contrastswith the situation in Essex and Kent where Rackham (1980) found C. sativa regeneratingsuccessfully, and probably even capable of invading woodland of other species. A study of the ability ofexotic, non-native trees to regenerate or not, would be another example of a feasible school project,where a very little study could fill a gap in our knowledge regarding the performance and the localbehaviour of long-introduced plants.
Uses
The tree is valued for its nut crop. For many generations it formed an important part of the traditionaldiet of farming communities, and it was also fed to stock animals.
Names
In S England, Sweet Chestnut survived throughout the Dark Ages, and acquired an Anglo-Saxon name, 'cyst'or 'cisten', the latter form (pronounced 'chisten'), being a derived survival from the Classical Latinname, 'Castanea'. The tree name then made the transition to a word element in at least a few Englishplace names, for instance, 'Cystewde' of 1272, which is today's 'Chest Wood' near Colchester (Rackham1980, p. 330).
Threats
None.
Native, common, widespread and locally dominant. European temperate.
1739; Henry, Rev. W.; Knockninny Hill.
Throughout the year.
Growth form and preferred habitats
Hazel is a very widespread shrub of woods, scrub thickets and hedgerows almost throughout Fermanagh. Itreally is absent or scarce only in areas of extensive peat soils or very high levels of exposure. Hazelis a characteristic species on rocky limestone hills, often forming low, scrubby mixed Ash (Fraxinusexcelsior)-Hazel woodland or pure Hazel scrub either on steep slopes or on ± rocky ground. It ismost abundant on alkaline lime-rich soils, although it does also thrive under Oak (Quercus spp.)in more acidic conditions, provided there is some lime in the substrate or in flushing groundwater.Originally, Hazel would have occupied the more fertile soils available, but Neolithic farmers sooncleared woods from this type of ground (Rackham 2006, p. 382).
Flowering reproduction
The familiar dangling male catkins shed their pollen early in the year, usually in late February or soonafter, before the leaves expand. The development of the hazelnut fruit does not seem to be as reliablenowadays as it was in the distant past, since archaeological evidence shows they were sufficientlyplentiful that they could be collected and stored as winter food. The species does not sucker, althoughit may layer itself in suitable circ*mstances. Thus increase and spread of C. avellana is verydependent upon seed reproduction. The large woody nuts are predated by birds such as rooks, and by smallmammals, especially the squirrel. As with other nut-producing species, it is only the occasional,overlooked, uneaten fruit that survives to germinate and reproduce the plant.
Ecology
Hazel saplings are very light demanding and they do not survive to regenerate the species under closedcanopy of either scrub or woodland dimensions. Thus episodes of wind, fire or heavy browsing whichcreate canopy openings are essential to successful sexual reproduction of both Hazel and native Oaks.Hazel, being a relatively low-growing shrubby plant is really not subject to wind-throw except onextremely steep slopes. Even if it were thrown, it very quickly resprouts from horizontal stems or thestill-rooted portion of the crown.
Fermanagh occurrence
Hazel is the eighth most widespread large woody plant in Fermanagh, ranking just behind Gorse (Ulexeuropaeus), being represented in 409 tetrads, 77.5% of those in the VC. Fermanagh's current mostextensive Hazel wood is probably that on Knockninny Hill, which was mentioned in an appendix in King's(1892) annotated version of Henry's Upper Lough Erne in 1739 (Weir 1987). Here it says of it,"... this hill, being on all sides bordered and adorned with fine stately groves of small woods,planty of heasle nutts, slows and crabs ...". There is physical evidence that the Hazel wood onKnockninny was coppiced, at least on the east side of the hill.
Uses
Despite its unashamedly bushy, multi-stemmed habit and limited stature (often only 4 m or so in height),Hazel was classed by the eighth century Irish Laws of Neighbourhood in the highest rank as a,'Noble of the wood'. This status was given because of the general usefulness of its nuts andstaves or poles (Nelson & Walsh 1993). Pliable Hazel rods were used for making rough, tough basketsfor farm use, eg turf and manure creels, as the ribs of curragh boats, and for wattle fencing amongstmany other uses. Their greatest significance, however, was as 'scollops' – thatching spars, which werethe main product of Hazel coppice for many centuries (Hogan 2001).
It is very likely that all the local estates and the larger farmers with tenants in Fermanagh planted andmaintained some area of Hazel coppice on their land to produce sufficient rods for these purposes. Alocal man (they were almost invariably men), would have had the basketry skills handed down fromgeneration to generation, and would have engaged in the manufacture and trade in baskets in a small,part-time, seasonal manner. Skilled thatchers, on the other hand, were itinerant workers, travellingaround the county to repair or replace worn roofs. The thatchers would purchase some, or perhaps most oftheir materials locally, if they were available.
Names
The genus name 'Corylus' is the Latin name for the hazel, possibly derived from the Greek 'korys' meaninga helmet, the shape of the fruit calyx resembling a helmet (Chicheley Plowden 1972). The Latin specificepithet 'avellana' refers to Avella, a town in Italian Campania where the hazel was largely grown forits nuts (Johnson & Smith 1946). The English common name 'Hazel' is derived from the Anglo-Saxon'hæsl' or 'hæsel', and allowing for dialect, is the same word in all Germanic languages, theinstrumental form of Anglo-Saxon 'hæs', a behest or to give an order. The belief behind this is that ahazel stick was commonly used to enforce orders among slaves and cattle, and therefore was the baton ofthe master (Prior 1879).
Threats
None.
Introduced, neophyte, deliberately planted, locally extinct. Native of N America, an introduced weed inmuch of Europe.
1903; Carrothers, N.; garden plot at Farnaght, SE of Tamlaght.
Fermanagh discovery, growth form and preferred habitats
This hardy summer annual was first noted by Nathaniel Carrothers, father of E. Norman Carrothers, whopublished a short note in The Irish Naturalist in 1903 stating that, "this queerChenopod" had been known from his ancestral farm for over a century. Carrothers (1903) reportedthat the colony always occupied a very limited area in a vegetable patch and although stray plants didoccasionally make it into adjoining fields, they always disappeared after a year or two. This solitaryFermanagh station for the alien was also known to Praeger in the 1930s and to Meikle, Carrothers andco-workers in the late 1940s and early 1950s when they were recording in the VC (Praeger 1939;Typescript Flora; Revised Typescript Flora). The farm was visited by Robert Northridge andmyself in 1989 and the present owner, Mr D. Carrothers was able to point out the exact spot where thespecies had grown up until about 1974, since when it has not been seen (Northridge 1991). There mightwell be seeds surviving dormant in the soil, but for the present the plant remains 'functionallyextinct'.
Appearance and possible origin
C. capitatum is known by the English common name 'Strawberry-blite', or locally in Fermanagh bythe Carrothers' family as 'Strawberry Spinach', on account of its startlingly scarlet globular fruitclusters which resemble Wild Strawberries in both colour and general appearance. Cultivars of C.capitatum with white or red leaves were in use as garden ornamentals in the early 19th century,and it is possible that the Fermanagh plants might have been self-sown survivors of this forgottenhorticultural fashion statement (Reynolds 1994).
Irish occurrence
Elsewhere in Ireland, the only other findings of C. capitatum were from a rubbish tip atStranmillis in S Belfast in 1909 (voucher in DBN), and a recent record by Reynolds (1990), whofound this usually casual alien in September 1988 at Foynes docks in Limerick city on the Shannonestuary. She regarded it as being imported with animal feed. Foynes docks also yielded the first Irishrecords of two other casual alien Chenopodium species in the same year, ie C.glaucum (Oak-leaved Goosefoot) and C. strictum (Striped Goosefoot). The Limerick docksbegan importing commercial animal feed originating from many parts of the world as recently as December1978, and by 1987 they were handling about 25% of the total Irish requirements (Reynolds 1990).
British occurrence
The New Atlas map plots around 45 widely scattered hexad records of C. capitatum inBritain, most frequently occurring across the English Midlands south of a linebetween Liverpool and Hull, but stretching as far north as Inverness. It is considered a neophyteintroduction, most probably a relict of cultivation, or an accidental wool shoddy import, or a bird-seedor esparto alien. The plant tends to be a casual on lowland waste ground and rubbish tips, butoccasionally it may be naturalised on cultivated ground. It appears to have declined in presence in thelast 50 years in both Britain and Ireland.
European and World occurrence
C. capitatum is a native of N America including Alaska and northern Canada. It was formerlycultivated as a garden ornamental, and probably also for its small, edible fruits. These are describedas pulpy, red and strawberry-like. The green shoots provide a spinach-like potherb and the fruit juicecan be used as a red dye. The shoots and leaves can also be eaten as a salad, but they contain toxicoxalates so should only be consumed in small quantities. In truth, the bright red berries are morevisually attractive than tasty. The plant is widely reported as a casual neophyte species in Europe andelsewhere.
Introduction, archaeophyte, a naturalised garden escape, very probably now only casual and locallyextinct. European temperate, widely naturalised in Europe and to a lesser extent in N & S America.
1902; Abraham, J.T. & McCullagh, F.J.; Erne bridge, Belleek.
Growth form and preferred habitats
A more or less ruderal, uniquely perennial Chenopodium species, Good-King-Henry plants are stickyand mealy-leaved when young, and the large, triangular, hastate (with pointed basal side-lobes like aspearhead), entire leaves are often covered with dust when older. The mature plant develops a stouttaproot, and in fertile, nitrogen-rich soil it can produce a large, leafy individual up to 80 cm inheight. From ancient times, C. bonus-henricus was widely and commonly cultivated as a vegetablepot-herb and possibly also used as a medicinal herb or simple. Nowadays, it typically occurs as a relictof cultivation, or a garden escape or discard, in disturbed, dry, nitrogen-rich farmyard and in waysidesituations (always near habitation). On account of its thick taproot which serves as an energy store, itcan be long-persistent in less disturbed wayside sites.
Fermanagh occurrence
There are only two old records in the Fermanagh Flora Database for this species. The first local recordof it, as listed above, dates from 1902, and the second, which is of uncertain date although notmentioned in the May 1951 Typescript Flora and therefore was probably found sometimebetween 1951-7, was observed by Meikle and his co-workers. The later record is mentioned in theirRevised Typescript Flora simply as being, "on waste ground by railway station atBelleek". Belleek today remains a rather small village in the extreme west of Fermanagh, somewhatisolated since 1957 by the loss of its railway connection.
Irish occurrence
Previously, in Ireland, C. bonus-henricus was well naturalised around houses and gardens as an oldcottage garden pot-herb. It was always probably fairly local in its occurrence, but it has not been seenanywhere in Fermanagh for 60 years. Despite the thick taproot and its reputation for persistence, theNew Atlas map depicts its decline to local extinction in many areas in Ireland. Against thisfatalistic view, Reynolds' Cat Alien Pl Ir lists 15 post-1970 records from nine Irish VCs,proving that the plant does survive the intensification of agriculture in a few favoured localitiesscattered around the island. Reynolds still regards the species as a surviving relict of earlierpot-herb cultivation. While this might possibly be true of undisturbed sites near habitation or inderelict ground, nowadays in Ireland the plant is very scarce or rare and sporadically recorded,appearing in dockyards and on roadsides, behaving more like a casual weed. Probably it is occasionallyre-introduced as a contaminant of grain, animal feed, or with garden plants, and generally nowadays itis seldom long-persistent in ruderal sites. A very similar pattern of overall decline and occasionalreappearance as a weed is currently happening in Scandinavian countries (Jonsell et al. 2001).
British occurrence
In Britain, C. bonus-henricus is an archaeophyte introduced vegetable pot-herb dating from theNeolithic-Bronze Age transition (Godwin 1975, p. 157). The New Atlas hectad map shows it is verymuch more frequent and widespread in Britain than is the case in Ireland. Nevertheless, it becomes lesswidespread northwards, and in Scotland becomes more eastern and coastal towards Inverness.
European occurrence
C. bonus-henricus is native to the mountains of C & S Europe, extending into late-lyingsnow-patches on higher ground. It is widely naturalised in temperate areas of both Europe and N America,again being spread by cultivation as an old pot-herb.
Uses
C. bonus-henricus was formerly valued and widely cultivated as a vegetable (ie a pot-herb). Theyoung leaves were boiled and used for food exactly like spinach is today. It is described as beingequally wholesome, but more insipid tasting than spinach, and was fed to both humans and poultry (Grieve1931). C. bonus-henricus was also valued as an herbal medicine, being used to treat both humansand domestic animals, including cattle and sheep. The main uses were as a gentle laxative, or as apoultice or ointment to cleanse and heal chronic sores.
Names
The genus name 'Chenopodium', is a combination of the Greek 'chen' meaning 'goose', and 'pous', 'a foot',a reference to the shape of the lobed leaves (Johnson & Smith 1946). The Latin species epithet'bonus-henricus' translates as 'Good Henry' (see below). The alternative English common name, 'Fat Hen',which also appears in a German equivalent, refers to its previous usefulness, along with allChenopodium species, in fattening poultry for the pot. Grigson (1955, 1987) gives a fullexplanation of the name 'Good King Henry'. The 'king' element apparently is a purely Englishinterpolation. Republicans may prefer the older name, 'Good Henry', used to distinguish the plant fromthe opposed poisonous, 'Bad Henry', which was another common name for Mercurialis perennis, thewoodland Dog's Mercury, which is a very rare species in Ireland. Alternative common names include'Smear-wort', which refers to its use in ointment to treat sores.
Threats
None.
Native, locally rare. Eurosiberian temperate, also native in western N America and widely naturalised, sonow circumpolar temperate.
1934; Praeger, R.Ll.; Trasna Island, Upper Lough Erne.
July to October.
Growth form and preferred habitats
This stout, erect, summer annual grows and becomes conspicuous, mainly in warm summers. It occurs onfertile, nitrogen-rich, bare mud trampled by cattle which becomes exposed when the water level is low inlowland lakes, ponds, ditches and reservoirs. However, it can also be found in more ruderal, wasteground situations where there is sufficient exposure of bare, fertile soil. It is also associated withmanure heaps and farm tracks.
C. rubrum has recently either become much more mobile, or it has begun colonising a new inlandhabitat. The association of Red Goosefoot with nitrogen-rich manure and silage heaps and with manured,trampled ground (Williams 1969) is augmented by an association with discarded heaps of spent mushroomcompost, a relatively new, frequently transported, farm-generated substrate. In August 2003, Ian McNeillfound it at Drumskinny, N of Ederney growing on dumped material that looked like old mushroom compost.
Fermanagh occurrence
As listed above, Praeger (1934c) was the first to find C. rubrum in Fermanagh, but it was not seenagain in the VC until 1989, when six records were made by RHN and by M. Tickner on exposed mudaround Lower Lough Erne. These records were followed by two on White Island in the SE of the Lough in1995, again found by M. Tickner. In 1989, the reddish, grooved stems of mature plants were quitedistinctive and the tall, fruiting plants were photographed by RHN. In October 2001, Red Goosefoot wasfound by RHN in a more ruderal situation, in disturbed open ground at Brockagh Sandpit. Again twice in2003 it appeared on bare earth by the roadside at Ballagh Crossroads and at Tempo Bridge, discovered byRHN & HJN. In 2004, it appeared on a roadside at Teemore and in the old quarry at Brookeboroughwhich is regularly used as an illegal dumping ground. As the tetrad map shows, there are now recordsfrom 15 Fermanagh tetrads, 2.8% of those in the VC.
Irish occurrence
Previously, C. rubrum was a very local and rare plant in Ireland with early records and thoseuntil the 1970s mostly coming from along the E coast between Dublin and the Ards peninsula (CybeleHibernica 1866, 1898; BSBI Atlas 2 (Perring & Walters eds. 1976)). The NewAtlas shows it has increased and spread to a remarkable extent in the 40 years between the twoBSBI surveys, with a doubling of hectad numbers in lowland Britain & Ireland.
British occurrence
C. rubrum is common and widespread in Britain, especially to the south and east of a line drawnbetween Hull to Liverpool. It shows a tendency to become more coastal both further north in NE Englandand E Scotland, and also in southwest England, in the West Country.
Effect of pollution
Recent population increases on exposed mud around ponds, reservoirs and lakes are undoubtedly in partbeing stimulated by eutrophication associated with pollution and enrichment runoff from agriculturalchemicals and sprayed slurry.
Threats
None.
Native, frequent and widespread. Eurasian wide-temperate, but extremely widely naturalised beyond itsnative range to become almost cosmopolitan.
1884; Barrington, R.M.; Co Fermanagh.
June to November.
Growth form and preferred habitats
In Fermanagh, Fat-hen is a frequent summer annual weed of disturbed ground and less frequently of potatofields and root crops and a variety of other open, artificial habitats. It grows on a wide range ofsoils, especially those of moderate pH and mesic to rather dry moisture status in the lowlands. Ittotally avoids strongly acid or wetland substrates. Like C. rubrum (Red Goosefoot), this speciesis essentially a ruderal pioneer colonist of open waste-ground and is, therefore, a poor competitor,unable to persist in closed turf vegetation. It is typically found on heaps of top soil or other formsof loose, recently disturbed surfaces, preferably those that are fertile or at least nitrogen-rich. Itbegins flowering rather late in the season, in July and August, and is particularly obvious in theautumn when it is fruiting (Williams 1963).
Identification
While phenotypically it is very variable in morphology and reproductive output depending upon growingconditions, C. album is usually less robust, more compact and more mealy-surfaced than thesuperficially similar Atriplex patula (Common Orache), in which the lower branches of matureplants tend to be opposite rather than alternate. The fruit is enclosed by five equal sepals rather thantwo large bracts as in oraches.
Stace (1997) details six closely related species that may be confused with C. album, but few ornone of these have ever been recorded in Ireland and I am content to consider all local plants as eitherC. album agg. or C. album s.s.
Fermanagh occurrence
Locally, Fat-hen is quite frequent, having been recorded in 76 Fermanagh tetrads, 14.4% of those in theVC. Considering the collapse to near extinction of arable farming in the area during the last 50 or soyears, this is striking evidence of the species habitat range and ruderal flexibility. The continuingpresence is greatly assisted by the prolonged dormant seed longevity characteristic of the genus.
Seed dispersal
While there is no special seed dispersal mechanism, some fruits remain on the plant until it dies inearly winter, and these may be eaten by animals including birds and the seed transported internally. Theseed contains fat, and it was an important supplementary human food source in ancient times. The lastmeal of the Tollund Man, who was found preserved in a peat bog, was a porridge or gruel containing seedsof Chenopodium and up to 40 other species (Grigson 1955, 1987). Birds (crow, snow bunting andducks) and other animals including pigs and horses, eat the fatty seed, transport it internally anddeposit it with their faeces (Ridley 1930, pp. 359-61). However, the great majority of seed appears tosimply fall around the parent plant, giving rise to patches of the plant in following years inmoderately disturbed habitats.
Seed longevity
Seed of C. album recovered from archaeologically dated soil samples proved viable after burial for1,700 years (Odum 1965).
Population vulnerability
Being strictly a summer annual, C. album has no means of vegetative reproduction, or even ofseedlings over-wintering since the species is frost sensitive. A late spring frost can seriously affectestablishment of the new season's populations, or wipe them out completely. Plants mown, grazed ortrampled in the early stages of growth also have no means of recovery (Grime et al. 1988).
British and Irish occurrence
C. album is widespread throughout Britain & Ireland, although less common in the N & W ofboth islands, and more coastal in both these directions. This is interpreted as due to the wetterclimate and the strongly acid peaty soils prevalent in these regions. The distribution has remainedstable over most parts of these islands, although it has declined in the west due to the move away fromarable agriculture towards pastures (Preston et al. 2002).
European occurrence
While considered indigenous to temperate Europe, C. album has been spread by man and become almostcosmopolitan. It has been transported amongst seed of arable agriculture to the extent that Holm etal. (1977) rank it as the world's worst weed of potatoes and sugar beet. A French botanist hassuggested that C. album is one of the five plants most widely distributed in the world (Coquillat1951).
Names
The genus name 'Chenopodium', is a combination of the Greek 'chen' meaning 'goose', and 'pous', 'a foot',a reference to the shape of the lobed leaves (Johnson & Smith 1946). The Latin species epithet'album' means 'white', presumably a reference to the white, mealy surface of young leaves. The Englishcommon name, 'Fat Hen', which also appears in a German equivalent, refers to the plant's previouslyfrequent usefulness, along with all Chenopodium species, in fattening poultry for the pot.Another common name, 'All good', is also applied to C. bonus-henricus (Good-King-Henry),referring to the usefulness of the plant for both food and medicine.
Grigson (1955, 1987) lists a total of 25 English common names for C. album, underlining thegeneral appreciation given to the plant throughout Britain and Ireland. An interesting one with manyvariants is 'Milds', 'Milder', 'Miles', 'Myles' or 'Meals', 'Melgs', and so on. It can sometimes appearas 'Meldweed', 'Midden mylies', 'Muckweed' or 'Muck-hill weed', referring to one of its significantwaste heap habitats. 'Melde' was the Anglo-Saxon name or word for a food plant that included thisspecies along with C. bonus-henricus and Atriplex patula, all three grown and/or collectedwild for use as leafy green vegetables as already mentioned.
Threats
None.
Native, very rare, but possibly over-looked and under-recorded. Eurosiberian wide-temperate, widelynaturalised beyond its native range including N America, Argentina and New Zealand.
1934; Praeger, R.Ll.; Co Fermanagh.
August.
Growth form, recognition and preferred habitats
Despite its botanical name, A. prostrata is an erect or a near-prostrate (procumbent),much-branched summer annual with large, basal, triangular or hastate (ie lobed, spear-shaped) leaves,upper ones covered with a mealy (farinose) tomentum of hairs beneath (only). The species isdistinguished from the very much more common A. patula (Common Orache), by having paired green ormembranous ovate or triangular bracteoles around the female flower, which are united only near theirbase as opposed to about half way. The prostrate form of the plant is also much more flattened to theground than A. patula, and the leaves are not in the least tapered at the base, but rather, theyare very definitely truncate (ie cut off abruptly at the base) (Taschereau 1985a; Webb et al.1996).
Like A. patula, this leafy annual occupies open, disturbed, damp to moist, unshaded, fertile ornutrient-rich substrates (silt, sand or shingle) of near-neutral soil reaction, either at the coast, orinland in cultivated ground, trampled, muddy margins of ditches, lakes and ponds, or in waste places oron rubbish tips (S.J. Leach, in: Preston et al. 2002). The inland, ruderal, anthropogenic biotypeis often accompanied by A. patula. In common with other ruderal species, the species favoursdisturbed, fertile, productive ground with plenty of patches of bare soil. A. prostrata does nottolerate strong competition from its neighbours and it therefore tends to occur as a pioneer colonist ofopen, moist, unshaded sites, where growth of perennials is delayed or prevented by instability ordisturbance. In freshly disturbed soil, it is often only transient, colonising quickly and soondisappearing. Having said this, plants of A. prostrata are also quickly ousted by grazing ortrampling animals (Grime et al. 2007).
A. prostrata has a particularly prostrate salt-tolerant form that is more or less confined to theupper reaches of coastal salt marshes. Very probably this halophyte is the original form of the species,although in recent decades it has evolved and colonised a wider range of inland habitats in Britain andN Ireland, following the routes of trunk roads salted to reduce accidents in icy conditions. Themajority of roads in the RoI are not salted or de-iced in this way. This type of species migration hasnot yet been observed in Fermanagh, although one very noticeable salt-tolerant species, Cochleriadanica (Danish Scurvygrass ), has been observed advancing inland from Belfast along the M1motorway towards the county.
Variation
Like other members of the family Chenopodiaceae, most characters of A. prostrata arevery variable and plastic in response to its local environment. The A. prostrata group comprisesa number of partially inter-fertile and morphologically similar taxa found on the coasts of W Europe andelsewhere. In Britain and Ireland, it is represented by four species, of which A. prostrata isthe most widespread, and the only one to occur inland from the coast. The other three species in thegroup are A. glabriuscula Edmondson (Babington’s Orache), A. longipes Drejer (Long-stalkedOrache) and A. praecox Hülph. (Early Orache). In the A. prostrata group, leaf size andoutline shape show extreme variation from the base to the apex of the plants. The most constant andcharacteristic leaves to examine are those on the central axis (main stem) in the middle portion of theplant, half way between the base and the terminal inflorescence. These are referred to as the 'lowerleaves' or the 'lower principal leaves', and they often drop off before the bracteoles and seed arefully mature (Taschereau 1977).
Reproduction
A. prostrata reproduces entirely by seed. The seeds are of two types; larger brown seed 1.5-3.0 mmwide and smaller, and in ruderal habitats, more frequent black seed 1.0-2.5 mm. The brown seed are non-or less-dormant than their small, black, shiny counterparts, while the latter are dormant andlong-persistent in the soil seed bank. Seed, of whatever colour germinates in the spring, and the plantsflower from July to September, or until the first frost. The flowers are unisexual, but both sexes areborne separately on the same plant (ie the plant is monoecious). Normally this means the upper flowerson the inflorescence are male and female flowers are arranged towards the base of the spike-likeflowering stem. The number of flowers produced will depend on growing conditions in the particularenvironment, but typically there are over 100 flowers per inflorescence. The flowers arewind-pollinated, but they do also attract insect visitors (syrphid flies) which feed on the pollen andmay assist in its transfer to the female stigma. Like other species in the genus, the flowers aredescribed as facultatively autogamous, meaning they can self-fertilize if other means of pollinationfail to occur. However, the male and female flowers are clustered so closely together in tightglomerules, and they mature and open almost simultaneously, that this greatly reduces the possibility ofcross-pollination (Taschereau 1985a).
Seed dispersal and survival
The seed is about 1.5 mm wide, and when dispersed is enclosed within two 5 mm persistent bracteoles (iesmall, modified leaves). Since these attachments increase the surface area of the fruit propagule, theymust assist its wind-dispersal. They probably also make the fruit more edible, and thus encourage birdsand mammals to ingest and transport them internally. Ridley (1930, p. 361) includes A. prostrata(as A. hastata L.) amongst a selected list of plant species carried internally by domesticanimals in Sweden and subsequently deposited with excreta in a viable condition. The seed islong-persistent in the soil seed bank, a proportion surviving for at least five years (Thompson etal. 1997).
Fermanagh occurrence
Recorded as A. hastata L., it was first found by Praeger in Fermanagh during 1934, but wasnot recorded again until the summer of 1986 by members of the EHS Habitat Survey Team investigating theconservation potential of Upper Lough Erne shores. Known chiefly as a coastal plant and considered rareinland, I personally have never seen this annual during Fermanagh field work. It may possibly have beenoverlooked to some extent by myself and others, though we are not aware of doing so.
There are just four Fermanagh records, confined to two tetrads in the Upper Lough Erne basin. Apart fromthe first record given above, the other record details are: W shore of Corradillar Td, 6 August 1986,L.W. Austin & S.A. Wolfe-Murphy; Sand Lough, Killyclowny Td, 14 August 1986, L.W. Austin & S.A.Wolfe-Murphy; Clonmin Lough, Corsale Td, 19 August 1986, S.A. Wolfe-Murphy.
Hybrids
There are three Atriplex hybrids involving A. prostrata, two of which are occasionalor rare at the coast. The most frequent hybrid is A. prostrata × A.longipes Drejer = Atriplex × gustafssoniana Tascher., whichregularly occurs where the two species overlap in distribution in both coastal and inland sites. Hybridintermediates derived from this combination are also found in sites remote from both parents,particularly on exposed coastal beaches in the far north of Scotland and more occasionally in Shetland.A. longipes differs from A. prostrata most clearly in that its bracteoles are enlarged andfoliaceous (ie green, leaf-like), up to 25 mm and with stalks up to 25 (or, exceptionally, 30) mm,whereas in the latter they do not exceed 6(-8) mm and they are stalkless (ie sessile). There is a rangeof variation within the hybrid forms, however, making determination rather difficult. Having said this,A. × gustafssoniana is a particularly frequent hybrid, and it must still be under-recorded(Stace et al. 2015).
Threats
None.
Native, frequent and locally abundant. Eurosiberian wide-temperate, widely introduced in bothhemispheres.
1900; Praeger, R.Ll.; Co Fermanagh.
June to December.
Growth form and preferred habitats
This very variable, phenotypically plastic, ruderal summer annual is a fairly common pioneer colonist ona variety of lowland, open, artificial habitats (ie created or heavily influenced by man). The largetriangular or rhombic paired vertical bracteoles clasping the fruit on mature plants of A.patula are very obvious and enable its ready distinction from Chenopodium album (Fathen), a related species with which it very often occurs.
Like all members of this family, A. patula prefers loose, nutrient-rich, loamy substrates, whichis why it is so prevalent in gardens and in arable fields. Examples of the latter habitat are now fewand far between in Co Fermanagh. Otherwise, Common Orache crops up in fertile or nutrient-enrichedsituations, including around rubbish tips, manure heaps, margins of farm yards and, in this area,occasionally in disturbed trampled ground or exposed mud around eutrophic lakes and ponds. Otherdisturbed habitats include quarries, sand-pits, roadsides and car parks. Like other Atriplexspecies, it is frequent and sometimes locally abundant in maritime habitats (Grime et al. 2005).
Given a moderate degree of surface disturbance, A. patula occurs on a wide range of substratetextures, from sand and gravel to loam and clay, but it avoids strongly acid peatland. Around Sheffield,Grime et al. (1988) found it ± restricted to soils of pH above 5.0.
Flowering reproduction
A. patula is most conspicuous in late summer and early autumn when it reaches the flowering andfruiting stages (June to November). Unlike Chenopodium species, which have perfect flowers eachcontaining both male and female parts, Atriplex flowers are unisexual. The monoecious plants havedense male and female flower clusters arranged on the same spike. The male Atriplex flowers havea small perianth of five tepals or segments, unlike the female ones which completely lack a perianth,the naked ovary being simply enclosed within a pair of small, leaf-like bracteoles. In A. patula,the margins of these bracteoles are united almost to midway along their length, a feature whichdistinguishes this species from the only other, locally very much rarer Atriplex in Fermanagh,A. prostrata (Spear-leaved Orache). The latter has bracteoles united only at the base.
Being facultatively autogamous, the tiny green flowers are pollinated either commonly by wind, or rarelyby insect, or if these both fail it can self-pollinate, thus ensuring a good seed-set. Again likeChenopodium species, A. patula has the potential to seed quite prolifically, the averageplant producing around 6,000 seeds. The seed is shed still enclosed by the two fleshy bracteoles.
Other biological similarities exist between this species and C. album in that both these weedyruderals produce two types of seed: larger brown non- or less-dormant ones, and small, black, shinyseeds that are long-persistent in the soil seed bank. About 90% of A. patula seeds were of thelatter type, and even after three experimental cultivations of the soil each year for five years, quitea high proportion of the seed remained dormant and viable (ie between 8.6% and 19.2% of three separatesowings) (Roberts & Neilson 1980). It has also been shown that A. patula seed are capable ofsurviving dormant in undisturbed soil for three decades or more (Brenchley 1918).
Native status
The Irish Census Catalogue regards the status of A. patula as "possiblyintroduced" (Scannell & Synnott 1987). There appears to be sufficient British fossil recordsfrom two earlier interglacials, plus evidence of persistence during the last glacial period, to suggestthat a similar pattern of behaviour and survival would be even more likely in Ireland. This would makethe species, despite its current weedy nature, more probably indigenous than an ancient introduction ofBronze Age or later date (Godwin 1975).
Fermanagh occurrence
It is widespread in lowland Fermanagh having been recorded in 139 tetrads, representing 26.3% of those inthe VC. A. patula is about twice as frequent and widespread in the county as the ecologically andbiologically rather similar Chenopodium album (Fat-hen), with which it can readily be confused.
British and Irish occurrence
Common Orache is common and widespread throughout most of Britain & Ireland, becoming less so in NScotland and W Ireland, a distribution pattern often assumed to reflect avoidance of wetter, cooler,more acidic soil conditions (Preston et al. 2002).
European and world occurrence
A. patula is widespread in Europe except the far north. It is also present in N Africa and W Asiaand is naturalised in N America. It has also been introduced in parts of the southern hemisphere,including S America, S Australia and New Zealand (Hülten & Fries 1986, Map 694).
Uses
Common Orache is described by Grigson (1987) as "a poor man's pot-herb", like Chenopodiumbonus-henricus (Good-King-Henry) or C. album, and close enough related to both plants toshare the common names of 'Fat Hen' and 'Lamb's Quarters'. These three plants were collected, boiled,pounded and mixed with butter and served in the same manner as Spinach is to this day.
Names
The genus name 'Atriplex' is the classical Latin name of a plant in Pliny, now applied to this group ofspecies. One suggestion for the derivation (however fanciful, or not), is from the Greek 'a' meaning'no' and 'traphein' meaning 'nourishment', several species of the genus known to be capable of growingin arid, desert soils (Johnson & Smith 1946; Gilbert-Carter 1964). The Latin species name 'patula'means 'spreading', which could apply equally to any of the species in the genus. The English common name'Orach' or 'Orache' again refers to any member of the genus, all of which can be, and were in the past,boiled and used as leafy vegetables in the manner of spinach. The name 'orache' or 'arach' comes throughFrench 'arroche' directly from the Latin genus name 'atriplex' (or 'atriplice') (Prior 1879), and thisin turn is from the Greek 'atraphaxis' in Dioscorides (the father of medicine) (Grigson 1987). It hasalso been suggested that 'orache' is a corruption of 'aurum' meaning 'gold', because the seeds, mixedwith wine, were supposed to cure yellow jaundice (Watts 2000).
Other English common names include 'Lamb's Quarter' or 'Lamb's Quarters', a corruption of 'Lammasquarter', from its supposed blossoming about the first of August, old style, the day of a churchfestival instituted as a thanksgiving for the first fruits of the harvest, when an oblation or offeringwas made of loaves baked with the new season's corn (Prior 1879). Yet another name, 'Hard Iron', isapplied to three quite different unrelated species Centaurea nigra (Common Knapweed),Ranunculus arvensis (Corn Buttercup) and Atriplex patula). In A. patula, theattribution is said to refer to the root, which is described as being as hard as iron, while in theother two species, the name refers to the flower buds or flowerheads (Watts 2000). Personally speaking,I find this supposed root property, of what is after all a summer annual, to be highly fanciful.However, I have no alternative explanation to offer as to the origin of this peculiar common name.
Threats
None.
Native, frequent. European boreo-temperate, native in C & E Asia and N America and thusdiscontinuously circumpolar. Also introduced in both hemispheres.
1882; Praeger, R.Ll.; Inishmacsaint Island, Lower Lough Erne.
Throughout the year.
Growth form and preferred habitats
This small, fleshy, annual or rarely perennial plant is readily overlooked and is very probablyunder-recorded in Fermanagh as also occurs elsewhere. It can grow in a great variety of permanently orseasonally wet situations, changing its habit from elongate, floating stems in streams, to terrestrialforms – either a spreading prostrate mat, or erect or tufted, cushion-like colonies. Blinks grows inwet, muddy hollows in fields, beside ditches, streams and lakeshores, or in flushes and springs onmoorland. A very interesting summary account of the ecology of pioneer plants of exposed mud, includingMontia (as M. chrondrosperma), was penned by Salisbury (1970).
Fermanagh occurrence
M. fontana has been recorded in 158 tetrads, 29.9% of those in the VC. The species is widelyscattered throughout Fermanagh, but is mainly found in the less intensively farmed areas of the county.
Variation and taxonomy
Four subspecies or varieties are recognised by some taxonomists, being distinguished on the basis oftheir black seed coat surface shininess and/or its sculptured texture (Walters 1953; Stace 1991, 1997).However, the different seed types are connected by intermediates (however rare) and thus form acontinuous series of gradual variation. Research in this phenotypically extremely plastic species, whoseform varies with both habitat conditions and vigour of growth, has produced no additional diagnosticcharacter(s) correlated with the seed coat patterns. Therefore, Lekkerkerk et al. (1983)concluded that formal intraspecific subdivision is not a sensible option.
In the recent Flora Nordica, 2, Jonsell et al. (2001) chose to regard one of thevariants, the small erect land form with dull surfaced, tuberculate seeds as a separate species, M.minor C.C. Gmel. (= M. fontana subsp. chondrosperma(Fenzl) Walters). Otherwise, the Nordic authors treat the remaining variation as M. fontana L.,without subdivision. Their taxonomic treatment appears to give heavy weighting to ecologicaldifferences, and it harks back to the situation previous to Walters’s (1953) recognition of the fourseed types.
Irish occurrence
In an Irish context, the authors of An Irish Flora commented that apart from their seed coats,"there appears to be no other difference between these groups in Ireland". All four seed coatvariants of M. fontana were recorded in Fermanagh by Meikle and his co-workers during the summerof 1948. However, no more recent work has been done to distinguish them and we have consistentlyignored them. The records of the subspecies are listed below purely for completeness.
It is clear from the paucity of Irish records of the four 'subspecies' plotted in the BSBI's NewAtlas, that the majority of Irish field botanists are not minded to distinguish Blinks atsubspecific level (Preston et al. 2002).
Looking at the New Atlas hectad map of Montia fontana, the species is widely distributed inthe N and the far S of Ireland, is less well represented on the W and E coasts and only thinly scatteredover inland counties of the Republic of Ireland (Preston et al. 2002).
British occurrence
The New Atlas hectad map shows M. fontana is widely recorded acrossthe entire latitudinal range of Great Britain and its associated isles, the distribution everywheredisplaying a greater and more consistent presence in the N and W. As might be expected from its damp,acidic or neutral soil ecological preferences, it is less well represented in chalk and limestone areas,and from heavily populated, industrialised or intensively farmed areas of S and E England (Preston etal. 2002).
Five Fermanagh records exist in well scattered tetrads. The sites are: N shore of Moorlough Lake;Gortaree Gravel pits, Slieve Rushen; bogs W of Clontymullan Td, near Arney; Cornaleck, Upper Lough Erne;and Lough Ora, SW of Ora More, Ballintempo Forest.
Two records exist, at widely separate sites with no apparent similarity. The sites are: Drumgrenaghanshore near Lackboy, Boa Island, Lower Lough Erne; and Derrynacarbit Lough, Little Dog Forest.
A solitary record exists from the outlet stream of Glencreawan Lough, in the Lough Navar Forest Park.
Montia fontana subsp. chondrosperma (Fenzl) S.M. Walters(= subsp. minor W.R. Hayw.), a form of Blinks
Four records exist, although the first by R.Ll. Praeger dating from 1902 has no specific site. Thedetails of the other three by MCM & D are: Lough Lea, Knocks Td, ENE of Lisnaskea; Mullylusty Td,Lurgan River Glen; and Meenagleragh Lough, NE of Little Dog mountain, in Big Dog Forest.
Names
The genus name 'Montia' is named after Giuseppe Monti, Professor of Botany at Bologna University, Italy(Gilbert-Carter 1964). The Latin specific epithet 'fontana' is from 'fons', 'fontis', a spring orfountain, and thus means 'growing in springs or fountains' (Gilbert-Carter 1964). The English commonname 'Blinks', or 'Blinking-Chickweed', is so called from its half-closed little white flowers peeringfrom the axils of the upper leaves, as if afraid of the light (Prior 1879). It is sometimes also called'Water Chickweed', or 'Water Blinks' (Prior 1879; Britten & Holland 1886), which makes the habitatmore obvious.
Native, rare. Eurosiberian southern-temperate, but widely naturalised beyond its native range.
1901; Praeger, R.Ll.; Co Fermanagh.
January to September.
Growth form and preferred habitats
The favourite habitat in Fermanagh for this apparently rare, but quite insignificant-looking andtherefore possibly somewhat overlooked winter- or more rarely summer-annual is in the open, dry, almostbare or thinly vegetated central strip in gravel tracks in coniferous forestry plantations. Here it isassociated with other small plants including mosses, Arabis hirsuta (Hairy Rock-cress),Eriophila verna (Common Whitlowgrass) and Catapodium rigidum (Fern-grass). A.serpyllifolia is a pioneer colonist of dry, shallow, warm, mainly neutral to calcareous open,lightly disturbed, unproductive bare stony or sandy soils. Usually it occurs in limestone terrain,although elsewhere in parts of England it is known to tolerate a more acidic reaction, down to nearly pH5.0 (Grime et al. 1988, 2007).
Germination, growth and reproductive cycle
Being chiefly a winter-annual, most germination occurs in damp conditions in the autumn, the species thenoverwintering as a seedling or a small plant. A proportion of seed each year remains dormant and becomesincorporated in the soil where it can survive for several years, perhaps up to 30 (Salisbury 1964).Growth of the plantlet resumes in the spring, producing a loose, sprawling and spreading plant up to 25cm in height. Flowering takes place as late as May and continues into August. The small, five-petalled,star-like flowers are mainly self-pollinated and self-fertilised, although small insects may also visitthem for pollen. Seed is set and shed from capsules that split to release them from July onwards.Although there is no obvious dispersal mechanism, the small, light seed appears to be highly mobile(most probably carried by wind), since it occupies a wide range of habitats over a large proportion ofthe country, and it can also be found growing several metres above the ground on the tops of walls(Ridley 1930, p.29).
Fermanagh occurrence
Locally, the species is rare, there being only ten records of A. serpyllifolia in the FermanaghFlora Database scattered over nine tetrads. It is ± confined to disturbed limestone terrain lying westof Lough Erne. The author is confident that the survey records reflect both the insignificant appearanceand the genuine local rarity of this species, since most of the finds have been by RHN.
Apart from the first record already given above, the details are: railway line at Belcoo (nowdismantled), Lower Lough Macnean, 1947, MCM & D; Corraslough Point, Upper Lough Erne, 23 June 1986,RHN & RSF; Lough Nagor, Belmore Mountain, 22 September 1990, RHN; Laghtmacdonnell, 1 km NW of LoughFormal, Big Dog Forest, 23 August 1992, RHN; forest track, Tullinwonny, NE of Lough Formal, 23 August1992, RHN; on gravel roadsides, Killydrum, near Noon's Cave, 4 January 1993, RHN; track at BallintempoForest, 28 August 1993, RHN; Brougher Mountain, 24 April 2000, RHN & HJN; Meenloughabank, 29 May2009, RHN & HJN.
Variation
A. serpyllifolia s.l. or agg. is now known to contain three taxa, either separatesubspecies or occasionally regarded as distinct species. The first two of these are subsp.serpyllifolia and subsp. lloydii (Jord.) Bonnier (subsp. macrocarpa F.H. Perring& P.D. Sell var. lloydii (Jord.) Gutermann & Mennema). Subsp. lloydii is a coastaldune ecotype, confined to shores. The third taxon that previously was named A. serpyllifolia subsp. leptoclados (Rchb.) Nyman, is now given specific status asA. leptocladus (Rchb.) Guss (Stace 2010). These three taxa differ mainly in the size of the plantand in the shape of the fruit capsule.
Irish occurrence
Although this species (in the broad sense, A. serpyllifolia) is widespread in the RoI, especiallySE of a line on the map linking Dundalk, Limerick and Cork, in contrast the majority of N Irish recordsare coastal, from sand dunes, cliffs and walls (New Atlas).
British occurrence
A. serpyllifolia occupies a much wider range of habitats and is very much more widespread inBritain than it is in Ireland. It occurs as a pioneer colonist on bare, open dry ground in areas ofbasic rocks including serpentine in situations that range from upland screes, summits and cliffs tolowland quarries, mine spoil heaps, railway ballast, waysides and margins of arable fields (Grime etal. 1988, 2007; P.S. Lusby, in: Preston et al. 2002). In distribution, it is veryfrequent in lowland situations throughout the country, but while it reaches the coast in the far N ofScotland, it is chiefly and most commonly recorded in the Midlands and SE of England (New Atlas).This distribution is probably explained by the preference the species shows for sunny, dry, south-facingrocky slopes on neutral to calcareous soils (Sinker et al. 1985; Grime et al. 1988, 2007).
European and world occurrence
The native distribution of A. serpyllifolia in the broad sense is centred on middle latitudes ofEurope and Western Asia (Eurosiberian southern-temperate), but it also extends to NW Africa and theCanaries. In Switzerland, A. serpyllifolia ascends to become part of the subalpine flora. Thespecies has been introduced into northernmost Europe and temperate areas of E Asia, N & S America,Australia, New Zealand and the Philippines. The present world distribution is thus discontinuouslycircumpolar (Hultén & Fries 1986).
Names
The genus name 'Arenaria' is derived from the Latin 'arena', meaning 'sand', indicating theplant's preference for sandy places (Johnson & Smith 1946). The Latin author Virgil gave the name'Serpyllus' to the plant we today call Thymus polytrichus (Wild Thyme), and the Latin speciesepithet 'serpyllifolia' means 'thyme-leaved', meaning with leaves like thyme (Gledhill1985). Hence wehave the English common name 'Thyme-leaved Sandwort', which is a perfect example of a so-called 'bookname'. Additional English common names include 'Sandweed', another book name variant of 'Sandwort',applied to all members of the genus Arenaria. 'Chickweed' is a name applied to various smallplants of similar habit, including annual forms of Veronica, Stellaria andArenaria, all of which fit into the category of plants that, "chickens and birds love topick the seed thereof." (Coles 1656, quoted in Britten & Holland 1886 p. 101).
Threats
None.
Native, occasional, but possibly somewhat over-looked. European temperate, also native in parts of W& E Asia and N Africa.
1881-2; Barrington, R.M.; Co Fermanagh.
March to September.
Growth form and preferred habitats
Despite the specific epithet, 'trinervia', the relatively large ovate leaves of this annual Sandwort havethree or five unbranched, almost parallel veins and ciliate margins, which are very obvious anddistinctive when held up to the light. At first glance the species is rather similar in appearance tolarge plants of Stellaria media (Common Chickweed), but its most distinctive habitats in openspots in shady, often damp, glades in deciduous woods and the bottom of hedges, are usually lessdisturbed than the typical conditions occupied by the latter very common weed.
M. trinerva is found in other shaded situations, for instance on cliffs and rock terraces (ledges)and outcrops, and on disturbed mineral soil (Grime et al. 1988, 2007). In coastal regions, it canoccupy heaths, pastures and seashore habitats, very probably always lacking shade. The same may be saidfor a range of man-made, apophytic habitats, including roadsides, lawns, path and field margins,gravel-pits and ruderal ground generally (Jonsell et al. 2001). In Cardiganshire, Wales (VC 46),Chater (2010) recorded M. trinerva growing in shade under Pteridium aquilinum (Bracken)near the coast, and far more surprisingly, on wall tops completely exposed to both light and weather!
While often found tolerating dry situations, the preferred substrate of the species is moist, yetwell-drained, fertile, often nitrogen-rich, weakly acidic to neutral soils that are subject tosufficient disturbance to keep bare ground available for colonisation by seed. It is mostly found invegetation where the environment provides moderate intensities of both stress and disturbance.
Behaviour in woodland
M. trinervia is one of the very few native annuals that we might call a true denizen of woodlandshade, a habitat very much dominated by slow-growing perennials and biennials. The main problem for anannual species living in shade conditions is to grow fast enough to complete its life-cycle, sincewithout a fresh crop of seed it cannot perpetuate itself (Fitter 1987).
Although M. trinervia usually behaves as a summer annual, very occasionally, when growing undershallower depths of shade, individual plants manage to survive the effort of flowering. When thishappens they become relatively short-lived polycarpic perennials, a change that helps the individualovercome the shade survival hurdle for annual species. Despite this flexibility in reproductivestrategy, M. trinervia still relies entirely on seed for its reproduction, there being novegetative means of increase or dispersal available to the plant (Grime et al. 1988).
As with other small, diffusely branching annuals, populations often perform best on warm, sparselyvegetated slopes under trees or shrubs – including sometimes evergreen species like Ilexaquifolium (Holly) (Garrard & Streeter 1983). Another reason for growing on slopes is thelikelihood that, thanks to gravity, leaf litter will not be present, or be deep enough to preventestablishment of a small, weakly competitive annual species. M. trinervia is a poor competitorwith other more vigorous herbs of woodland or shade-tolerant situations, and thus is often confined tomoderately disturbed or unstable ground, such as near rabbit burrows or fallen trees (Sinker etal. 1985; Grime et al. 1988, 2007).
Being something of a nitrophile, M. trinervia sometimes grows under or near nettle patches, orbecomes associated with Galium aparine (Cleavers) (Clapham et al. 1987; Crawley 2005). Thelevel of disturbance in woodland plantations provided by modern mechanical forestry operations favoursthe growth of M. trinervia, and this form of human disturbance probably also serves to assistdispersal of the species. Three-nerved Sandwort has benefitted from logging sites, the construction offorest roads and in general all human activity that disturbs woodland. However, despite this it is notreally a plant that favours human activity, unlike Common Chickweed, with which it is often confused.
Flowering reproduction
The small, solitary flowers produced in May and June bear white petals that are much shorter than thesepals. The flowers are visited by small flies and crawling insects such as beetles, although they willautomatically self-fertilise if not visited. Fruiting occurs and seed is shed in June and July. Thenumber of fruit capsules per plant varies enormously: Salisbury (1942) reckoned the range of capsulesproduced lies between 51 and nearly 6,000, with a mean of about 200 (NB Salisbury calls the speciesArenaria trinervia, p. 227). Seeds per capsule range from four to 17, and therefore the computedmean seed production is around 2,500 per plant (Salisbury 1942).
Seed dispersal
Seeds are shiny, kidney-shaped, black and have an attached highly nutritive white food body – anelaiosome oil appendage – that attracts ants. The ants help disperse seed from around the parent plant,thus minimising seed predation (Clapham et al. 1987). Longer range dispersal of seed in mudattached to animals, including man and his machinery, is undoubtedly also significant.
Seed survival
There are 15 estimates of seed persistence of the species in the survey of soil seed-banks of NW Europe.Five samples indicate that M. trinervia can at best be considered short-term persistent in thesoil, ie they display dormant survival of more than one and less than five years (Thompson et al.1997).
Variation
A modest range of variation occurs within M. trinervia with respect to plant habit, colour andhairiness, most or all of it associated with local environmental conditions and the season of seedgermination (Hind 1985).
Fermanagh occurrence
In Fermanagh, M. trinervia has been occasionally recorded from a total of 35 tetrads, 6.6% ofthose in the VC. As the distribution map indicates, about half its stations are scattered around themoist, wooded shores of both parts of Lough Erne, with the remainder thinly and widely spread across thecounty. It is not a conspicuous plant and is often only present as scattered individuals. It istherefore very probably under-recorded.
British and Irish occurrence
In Britain, the species, as shown in the New Atlas hectad map, is widespread in most lowlandareas, except the English Fens and the Wash. It becomes rare and coastal further N & W in Scotland –a familiar enough distribution pattern. In Ireland, by comparison with Great Britain, the distributionis very much patchier. Whether or not this is an artefact of recorder effort or possible unawareness isnot clear from this evidence, but under-recording remains a definite possibility. The more comprehensiverecording of the species in the three Cork VCs is rather striking, and itself lends support to thiscontention. Again, in NI, there appears to be more records of this Sandwort in the broad basins of LoughNeagh and Lough Erne, perhaps because these two areas have been exhaustively and systematically surveyedin recent years (Preston et al. 2002).
European and world occurrence
More or less restricted to Europe and thinly scattered parts of W Asia and N Africa,although also present in the Canaries. An isolated population in far eastern Asia (Japan), distinguishedas M. platysperma Maxim., is said by Hultén & Fries (1986) to be identical with the Europeantaxon. If this is really so, then it constitutes a major discontinuity in the distribution ofthis species, and probably arose from a case of accidental introduction.
Names
The genus name 'Moehringia' is called after Paul Heinrich Moehring (1720-1805) a physician and naturalistat Jever in Oldenburg, Lower Saxony, N Germany (Gilbert-Carter 1964). The Latin specific epithet'trinerva' means 'three nerved', referring to the three (or up to five) prominent veins observed in theleaves.
Threats
None.
Native, very common and widespread. Eurasian wide-temperate, but naturalised to become circumpolarwide-temperate. It is almost cosmopolitan as a weed, absent from arctic and very arid areas.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
In the temperate zone, Common Chickweed is wintergreen and it can germinate in any month of the year. Itcan therefore behave either as a winter or a summer annual, or as a short-lived perennial. S.media is regarded by many as one of the most common ruderal, sprawling, diffusely branching,mat-forming weeds in Fermanagh, and otherwise as one of the most common plants in Britain & Ireland.It is also notorious world-wide as an economically important weed of arable farmland and gardens. S.media can become a major contaminant in some crops, including soft fruit, potatoes and barley.It sometimes is so abundant, it can overgrow and smother young seedlings of other species andsuccessfully compete for nitrogen in the soil, thus depriving its neighbours of an adequate supply ofthis essential nutrient (Holm et al. 1977; Turkington et al. 1980).
The decumbent or ascending, much-branched plant up to 40 cm in height is distinguished by having asingle row of hairs down its stem internodes and leaf stalks (petioles). It prefers cool, moist,semi-shaded places and quickly colonises almost any piece of disturbed lowland soil and semi-openhabitat offering a damp, preferably fertile or nutrient-enriched substrate (Salisbury 1974). Examples ofwhere it is most abundant include on loose piles of moved earth in gardens or on building sites, onrubbish dumps, around farmyards, and along the junction of roadside pavements and hedges, in neglectedor in young pasture areas, in arable crops and in waste ground (Holm et al. 1977).
The flowering cycle of around five weeks is also very rapid, and in optimal conditions it allows theproduction of three generations per year. It should be stressed, however, that in weedy species likethis, not every individual is capable of behaving in the same way. There now exists a growing body ofinformation revealing habitat-correlated intra-specific genetic variation in life-history traits as wellas a capacity for developmental plasticity (Briggs et al. 1991). The soil seed bank population ofS. media is potentially enormous and buried seed can persist for many years.
These properties, underpinned by the great physiological versatility of the species, often allow CommonChickweed to become a hugely abundant, sometimes dominant weed in suitably fertile, open waste ground orin unmanaged, ± open habitats (Sobey 1981). This is due in part to its abilities to grow and reproduceeven during the winter months whenever temperature rises temporarily, to survive subsequent frosts, andthen to produce very rapid growth in the early spring. S. media is able to continue growing at2°C, when most other plants stop growing below 6°C.
Detailed ecological accounts are given by Sobey (1981) and by Grime et al. (1988, 2007). Ecology,weed behaviour and other very interesting lore is reviewed by Holm et al. (1977), Turkingtonet al. (1980) and Defelice (2004).
Variation
Being an extremely common and widespread, almost ubiquitous weed species, unsurprisingly S.media displays a high degree of phenotypic plasticity, varying in size, habit, hairiness, petallength, stamen number, and number, size and surface details on the exquisite sculptured, tuberculateseeds (Turkington et al. 1980; Jonsell et al. 2001). The variation is so extensive that inthe past 32 varieties or micro-races have been described (Béguinot (1921), cited by Matzke (1932);Peterson (1936)). Also in the past, two subspecies were delimited: S. media subsp. neglecta (Weihe) Gremli and S. media subsp. pallida (Dumort.) Asch. & Graebn., but while these aresimilar to S. media, both are now regarded as separate species (Jonsell et al. 2001; Stace2010).
The second edition of Flora Europaea 1 (Tutin et al. 1993) suggests that S.media sensu stricto contains two additional subspecies: subsp. media (throughout thewhole species range) and subsp. cupaniana (Jordan & Fourr.) Nyman, the latter beingrestricted to C & E Mediterranean regions. There is another form subsp. media described asvar. apetala Gaudin which has petals minute or absent. This latter form has often been confusedwith S. pallida (Lesser Chickweed).
Flowering reproduction
This is essentially by seed only, although prostrate branches can root and spread the individual in loosesoil surfaces, giving the plant additional resources for growth and seed production. The small,star-like flowers are formed either solitary in the axils of upper leaves or few to many borne in loosecyme inflorescences. Each flower normally opens for just one day, and the five petals are so deeplybilobed they look like ten, and are shorter than the sepals. The number of stamens varies from three toeight depending upon the degree of illumination (Salisbury 1964).
Pollination and fertilisation
The flowers are scented and are visited by bees, ichneumon wasps and members of the Diptera, Hymenopteraand Thysanoptera (Turkington et al. 1980). If insects fail to visit, the flowers self-pollinateand self-fertilise (ie they are autogamous). Indeed, autogamy may predominate in this species, and somereports suggest the flowers are cleistogamous, self-pollination occurring while still in bud (Salisbury1974; Jonsell et al. 2001).
Since its sexual reproductive cycle is unfettered by either seasonality (ie day-length) or a need forpollinators, in moist, temperate climates plants of S. media flower and seed right throughout theyear except in very severe weather. During the winter months, flowers tend to be without petals and areentirely autogamous. In general, the influence of day-length, the length of the photoperiod upon thereproductive capacity of flowering plants is to restrict the area of the species distribution. Speciesof wide geographical range that are indifferent to photoperiodic stimulation, and which flower and fruitirrespective of long or short days, include ubiquitous weeds such as S. media and the annualgrass Poa annua (Annual Meadow-grass) (Salisbury 1942, p. 49).
Fruit production
According to Salisbury (1964, p. 191), the average fruit production per Common Chickweed plant is about240 capsules, but one very large plant that he examined produced a total of 1153 capsules! The fruitcapsule is oblong in shape and when ripe, like all members of the genus Stellaria, it splits atthe tip into six triangular valves or teeth to release the seed. The capsule valves open and shutdepending on whether the weather is dry or wet, thus protecting the 5–16 (the mean is ten) containedseeds.
Seed dispersal
The seeds are shaken out of open capsules by wind and appear to be secondarily dispersed in mud attachedto the feet of animals, including man. They are also common in the dung of animals including cattle,horses, pigs and birds. The English common name 'Chickweed' refers to the fact (or the belief) thatbirds peck around and feed on the plant, ingesting the seed as they do so. S. media seeds arealso dispersed as contaminants in seed of many crop plants including cereals, rape, swede, fodder-beet,sugar-beet and kale (Fryer & Makepeace 1977).
Seed dormancy and soil seed bank survival
The seeds are light, disc-like, with a sculptured, knobbly surface, 0.8-1.4 mm in diameter. They vary incolour from yellowish grey to reddish or dark purplish brown.
In temperate and maritime regions, there is no seed dormancy in the population: germination can occurimmediately after release (Peterson 1936; Salisbury 1964). In arctic, subarctic and continentalpopulations, however, germination is delayed (presumably to allow after-ripening of the embryo), withsubsequent rapid development and sparse vegetative growth materialising (Peterson 1936).
While a portion of the seed produced may germinate within a few days of release, the remainder cansurvive in soil for some time. With up to three generations of seed produced in a calendar year, thesoil seed bank population of S. media is potentially enormous, and some buried seed can persistfor years. Seed survival estimates vary, but viable seeds are known to persist for between 25 and 60years or more (Salisbury 1964; Fryer & Makepeace 1977).
Fossil history
Fossil evidence indicates S. media is native in Britain and Europe, seeds having been identifiedin pre-glacial and Mesolithic deposits in Britain. Godwin (1975) regarded it as persistently native inBritain up to and throughout the Weichselian glaciation. In the early post-glacial Flandrian, it waspresent in the middle of what is now the North Sea, but there are no fossil records of it from theBoreal and Atlantic periods when more or less uninterrupted deciduous forest predominated across theland. It reappeared in the fossil record once farmers arrived and cultivation began in the Late Bronzeage and from then onwards it is always present. The numerous interglacial records prove the species isperfectly capable of existing in Britain and Ireland independent of human influence (Godwin 1975, p.149).
Fermanagh occurrence
Despite its widespread reputation for commonness, in terms of record frequency S. media actuallyranks 148th in the Fermanagh Flora Database, showing yet again just how unreliable casualimpressions of relative abundance can be (Crawley 2005). Despite this, Common Chickweed is very commonin Fermanagh, having been recorded in 306 tetrads, 58% of those in the VC.
British and Irish occurrence
A ubiquitous naturalised archaeophyte weed in disturbed, often enriched habitats, the hectad map in theNew Atlas shows S. media is widespread and abundant throughout both islands, except on thehighest mountains and on the driest soils (Preston et al. 2002).
European and world occurrence
Native to Europe and Asia, S. media has been spread everywhere man has introduced cultivation andis now naturalised, cosmopolitan and circumpolar in its geographical distribution. The only ground iteschews are totally inhospitable habitats involving heavily disturbed soils and the high arctic and ariddeserts. In the tropics, it is confined to higher altitudes where more temperate conditions allow it toflourish (Hultén & Fries 1986, Map 724). Fossil records from Greenland show it was present therefrom around 1400 to 1721 AD, before European settlement began (Pedersen 1972). It was first recorded inN America in 1672 in New England (Turkington et al. 1980), and it has spread with agricultureworldwide.
Weediness
S. media was ranked as the 72nd worst weed in the world by Holm et al. (1977) because it isa problem in more than 20 crops in 50 countries. It can also be a serious weed in lawn turf in NAmerica, because its prostrate growth habit and rapid growth rate allow it to survive frequent mowing(Uva et al. 1997). In addition to its direct weediness and power to significantly reduce cropyields, S. media is a host plant for several insect and disease pests of economic crops,including viruses such as Tomato Spotted Wilt Virus (Defelice 2004).
The very rapid growth rate of S. media and its ability to overgrow and smother seedlings and youngplants of other species has led to its use in weed control. In Switzerland, Common Chickweed is used tocombat infestations of Convolvulus arvensis (Field Bindweed) and Calystegia sepium (HedgeBindweed) in vineyards. In such circ*mstances, the Chickweed itself is not a problem weed (Stalder etal. 1973).
Uses and toxicity
Young leaves of the plant when boiled were eaten like Spinach from which it is described by Grieve (1931)as "hard to distinguish". They were also used fresh along with young Dandelion leaves(Taraxacum vulgare agg. (Dandelion)), as salad. However, like other members of the familyCaryophyllaceae, S. media contains saponins, and it accumulates sufficient nitrogen to actuallyrender it toxic if eaten in large quantity. Having said this, there is no evidence of stock animalsbeing seriously poisoned or killed by it in recent years (Cooper & Johnson 1998). There is a longhistory of Chickweed being fed to cage birds and hens. Both wild and cage birds eat the seeds, the youngshoots and the leaves, as do rabbits, cows and horses (Grieve 1931).
In herbal medicine, S. media has long been used for a wide diversity of ailments. The main use hasbeen for a mat of the plant, boiled and applied as a hot, relaxing poultice to treat inflammation,bruises, and reduce swelling. It has been applied in this way for boils, abscesses and ulcers,rheumatism, chilblains and rashes. An ointment made from the plant has also been used for sore eyes, andfor eczema in some parts of Britain. These and many other uses are listed by Allen & Hatfield(2004), along with localities where they are practiced.
Names
The genus name 'Stellaria' is of medieval origin from the Latin 'stella', meaning 'a star', referring tothe shape of the five-petalled flower (Gilbert-Carter 1964). The Latin specific epithet 'media', means'middle-sized' or 'intermediate' and refers to size (Gledhill 1985).
The English common name 'Chickweed' refers to the fact that it has long been used to feed birds. Grigson(1955, 1987) lists no less than 16 variant names for S. media from around Britain and Ireland,amongst which 'Skirt Buttons' from Devon, seems one of the most interesting. Having said this, I cannotfind this name mentioned in any other reference, and have no suggestion as to its origin, even if onetries misspelling either part of the name. Another name is 'Chick Wittles' (ie vituals), whichoriginates from Suffolk, and yet another is 'Tongue-grass' from Ireland (Threlkeld (1727)). German andFrench names that Grigson mentions translate as 'Hen's guts' or 'Hen's bite', again making reference tochicken feed.
Threats
None.
Native, common and widespread. Eurosiberian temperate, introduced in eastern N America and very rarelyand not recently in New Zealand.
1881; Stewart, S.A.; Knockmore Hill.
Throughout the year.
Growth form and preferred habitats
This wintergreen perennial is most conspicuous from April to June when it flowers following a burst ofrapid early spring growth that makes it relatively large (up to 60 cm tall) compared with other typicalcommon hedgerow plants. Other typical habitats include open lightly shaded deciduous woods, scrub,woodland margins, rides and clearings. In Fermanagh, it is most frequently observed along hedgerow andriverside banks and on seldom-mown, rank, grassy roadside verges in the lowlands. It is also wellrepresented on the less fertile, moderately acid to neutral soils in the limestone areas and on the moreupland plateau in the west of the county. In these latter areas, it is associated with clearings or withthe margins of woods and around scrubby ground. Its slender, grass-like stems and leaves can also befound clambering over rocky outcrops and talus slopes, including block screes at the base of cliffs.Around lakes and on wooded isles it occupies the driest available sloping sites, completely avoidingwetland conditions and situations that are liable to flood.
The opposite, acute, linear-lanceolate, grey-green leaves of this familiar and widespread species aredistinctive and recognisable, even when it is not in flower, although it might be easily overlooked whengrowing amongst tall grassy vegetation later in the season. S. holostea plants develop best inmoist and otherwise mesic to infertile soils, but the species can occur over a wide range of soil pH.Plants typically form quite dense patches of ascending stems, which are however, physically weak andbrittle and appear therefore to be supported to some extent by the surrounding, competing mass ofgrasses and other herbs. The ascending stems of S. holostea are not all of the same length, andsome do not flower, so that apart from photosynthesis the secondary role of sterile stems may be to helpsupport the taller, flower-bearing stems (Clapham et al. 1987).
The preferred growing conditions appear 'designed' to limit competition, but Greater Stitchwort isgenerally most conspicuous in springtime towards the top of roadside banks, in obviously well-drained oreven rather dry growing conditions, preferably in full sun or marginal to the still developing summercanopy of an overtopping hedge if there is one present (Clapham et al. 1987; Garrard &Streeter 1983; Sinker et al. 1985; Grime et al. 1988).
Potential vegetative reproduction
S. holostea has a slender creeping rootstock that may root at intervals and allow some degree ofvegetative spread. The overwintering sharply quadrangular, weak, brittle, aerial shoots are supported bysurrounding vegetation until this dies back in winter. When this happens, the rather fragile,wintergreen shoots drop to the soil surface and can sometimes develop adventitious roots, allowing theplant to spread vegetatively and form clonal patches (Grime et al. 1988). A detailed study of thepotential for S. holostea vegetative reproduction by this means remains to be made, a fact thatthrows into focus the paucity of scientific studies in the genus, apart from the huge volume of appliedscience literature available on the arable weed, S. media (Common Chickweed).
Flowering
Flowering takes place from late March to July although the peak lies between April and June. Theinflorescence is large and lax and occupies up to half the length of the shoot. It bears between 3-40white flowers, but usually around ten of them, in a branched cyme. The flowers are insect-pollinated,with nectar and pollen attracting beetles, butterflies, flies and bees for the purpose. Should flowersfail to attract insect visitors, they resort to selfing, the filaments bending forwards to allow thepollen-filled anthers to touch the three stigmas (Proctor & Yeo 1973, p.55).
Seed dispersal
An average of ten seeds are produced centrally in each green, subglobose, dehiscent, fruit capsule, whichis around 9 mm in diameter, about equalling the length of the calyx. The weight of the fruit bends theslender pedicels down towards the ground, and the top of the fruit then opens when ripe by splittinginto six valves to release the seeds.
Ridley (1930, p. 522) reported seeds being found in ant nests despite the absence of an attached foodreward for the insect. He assumed that the seeds, and those of several other common species found in thebig Wood Ant nest (Formica rubra) (eg Luzula campestris (Field Wood-rush) andHypochaeris radicata (Cat's-ear)), were blown there by the wind. Presumably the seeds are shakenout of the dangling capsule and are wind dispersed. Perhaps also, as in S. media, some ripecapsules may be eaten and the seed transported internally by birds (Ridley 1930, p. 457).
Seed size and survival
The seeds are 1.5-2.0 mm across, reddish-brown, and the seed coat is densely covered in papillae. Theseed is at least four times larger and over seven times heavier than that of S. media (Salisbury1942, p. 27). Freshly shed seed are dormant (Grime et al. 1988), and the soil seed bank istransient, ie seeds survive less than one year (Thompson et al. 1997).
Fermanagh occurrence
S. holostea is widespread and fairly common throughout Fermanagh, having been recorded in 199tetrads, 37.7% of those in the VC.
British and Irish occurrence
Avoidance of very wet and strongly acid substrates helps explain the rarity or absence of S.holostea from many parts of W Ireland, most notably Connemara and parts of Clare and Kerry. Thehabitat situation is similar around the English Wash, and also in NW mainland Scotland, the OuterHebrides, Orkney and Shetland (New Atlas).
European and world occurrence
S. holostea is considered native but irregular in occurrence in most of Europe as far north assouthern Finland, but it is rare or absent in S & SE parts of the Mediterranean area. Thedistribution range includes scattered locations in N Africa, and SW and W Central Asia. In Scandinavia,it is regarded as indigenous in the southern part of the region, and occurs most frequently at the coastand at lower altitudes. However, it is also present in parts of Scandinavia as a garden escape, and as agrass-seed alien introduction (Jonsell et al. 2001). S. holostea is listed along withS. media in the Royal Horticultural Society Index of garden plants, S. media being,of course, a notorious weed (Griffiths 1994). S. holostea is introduced, but rare in eastern NAmerica and also in New Zealand, where it has not been seen since 1900 (Hultén & Fries 1986, Map727; Webb et al. 1988, p. 507).
Uses
In both Britain and Ireland, the conspicuous, white, star-like flowers of S. holostea have beenchewed to combat muscular stitches and sharp pain for many years by those who put their faith in thepotency and efficacy of the English common name 'Stitchwort' (Allen & Hatfield 2004). The name issaid to date back to the 13th century, so presumably the tradition goes back that far(Grigson 1955). As mentioned above, S. holostea is occasionally chosen as a garden subject. Thepure white flowers, if present in large numbers in early summer encouraged by good horticultural soil,would undoubtedly be decorative and might be considered by some to be garden worthy. Having said this,Stitchwort is not mentioned by most garden writers, including Graham Stuart Thomas (2004) in hisPerennial garden plants or the Modern Florilegium.
Fossil record
There are a few fossil seed records listed by Godwin (1975), but they are important as they definitelyindicate the long-term native presence of the species. It took time for S. holostea pollen to berecognised at species level, and Godwin therefore can only comment on a few localities where the planthas been recorded. Even so, it is clear that the fossil history of the species stretches well back intopre-history and predates the arrival of Neolithic farming in Britain and Ireland.
Names and folklore
The genus name 'Stellaria' is of medieval origin from the Latin 'stella', meaning 'a star', referring tothe shape of the five-petalled flower (Gilbert-Carter 1964). The Latin specific epithet 'holostea' isthe old generic name of the species, latinised from the Greek meaning 'whole (or 'entire') bone',referring to the fact that it was used in ancient times for healing fractures (Johnson & Smith1946).
The fact that there are a total of 103 English common names listed by Grigson (1955) indicates howwidespread, common and conspicuous this species is throughout most of Britain and Ireland, and how muchfolklore has accumulated around it. Considering this fact, Grigson comments that
'Stitchwort' clearly was believed to be a plant under protection and somewhat feared by both children andsuperstitious folk. They considered that it belonged to or was associated with items of fear, such asthe devil, piskies (also piskey, or pixies?), Jack-a-Lantern (a lantern-carrying elf or goblin of theignis fatuus), and to snakes. In complete contrast, people also believed that the plant hadproperties to banish evil, belonging then, on the other hand, to the Virgin Mary, and associated withWhitsunday and the Star of Bethlehem! There are also rather dubious associations by name with cuckoos,smocks, milkmaids and bachelor's buttons. Some of these latter names may link to the season offlowering, which includes Eastertide and the month of May.
Other names listed by Grigson (1955) are derived from the brittle, easily broken nature of the stems,including 'Brandy Snap', 'Break Jack', 'Dead Man's Bones', 'Jack Snaps', 'Snapcrackers', 'Snapjacks' and'Snappers'. Children in the past liked to squeeze the fruit capsules to make them pop (Vickery 1995),and the names, 'Pop-guns', 'Pop Jack', 'Poppers' and 'Poppy' obviously refer to this practice.
Threats
None.
Native, occasional and locally frequent. Eurasian boreo-temperate, but also widely naturalised, includingeastern N America, Australia and New Zealand.
1884; Barrington, R.M.; Crom Castle Estate.
June to October.
Growth form and preferred habitats
The normally pale, bluish-green or greyish stem and leaves make this a very distinctive locally abundant,rhizomatous, scrambling, patch-forming perennial in Fermanagh. The typical habitat of the species is innutrient-rich, mesotrophic to eutrophic, wet to moist ground, often winter-flooded and swampy. InFermanagh, these enriched, wetland conditions are provided in fens, meadows, pastures and scrub,primarily around the larger lakes, but occasionally also by rivers, streams and ditches, especiallywhere they flood. Although in other parts of Britain and Ireland S. palustris is reputed tocolonise artificial, man-made, disturbed habitats, eg old flooded peat cuttings, it does not do thisanywhere in Fermanagh (P.S. Lusby, in: Preston et al. 2002).
Variation and identification
Apart from the usually distinctive colour of the aerial parts, S. palustris grows up to a heightof around 60 cm and has larger flowers than S. graminea (Lesser Stitchwort) and lacks the ciliatebracts and outer sepals of the latter, the only species with which it might possibly be confused. S.graminea also prefers better drained soils than S. palustris.
We are fortunate that the form of S. palustris present in Fermanagh is so very distinctlygrey-green (glaucous), when in fact the colour of the species elsewhere is very much more variable.Elsewhere, the aerial shoots can be whitish, or a pure or a yellowish green, making such plants muchless immediately recognisable (Jonsell et al. 2001). On finding the green form in Cambridgeshirein 1985, Walters (1986) naturally worried that English field botanists might possibly be overlooking thespecies, since not only does it look grass-like when non-flowering, but it may also display a very muchlater flowering period than the glaucous or pale greyish form, sometimes indeed still being in bloom inearly September.
In the critical Flora Nordica of Finland and Scandinavia (Jonsell et al. 2001, pp. 122-5),no less than six variant forms or 'entities' are discriminated within S. palustris, although theycan only be identified on combinations of several characters, some of which are relative. The entitiesare also linked by intermediates, further complicating their identification. It appears that we aredealing with a polyploid complex with exceptionally high chromosome numbers: the series ranges from10-ploid (2n = 130) up to a 14-ploid with 2n = 182. Little or nothing is known as to how this geneticmakeup relates to the six morphological forms, or how much of the observed variation is environmentallyinduced.
Reproduction
It appears that partial or complete male sterility is common in this species, and unsurprisinglytherefore, seed set is poor. Male sterility adds a third level to the variation within the species, asit is expressed both as floral and vegetative dwarfism (Jonsell et al. 2001). In Britain, theaverage number of seeds per capsule is as low as 13, and the plant does not usually bear many fruits inany case, making its seed output very small (Salisbury 1942, p. 170). Increase in population numbers andlocal dispersal of S. palustris are clearly very much more dependent on vegetative reproductionthan is usual in this genus. Both these aims are achieved by vegetative growth of the slender creepingrhizome, layering of decumbent stem branches, plus stem fragmentation and flotation during floods.
Fermanagh occurrence
In Fermanagh, Marsh Stitchwort has been recorded from 73 tetrads, 13.8% of those in the VC. Nine of thesquares have no post-1975 records, indicating a definite decline. Locally, it occupies a decidedlyrestricted habitat of open areas in winter-flooded marshy grassland, or in base- or lime-rich fens,which in Fermanagh are chiefly found around Upper Lough Erne and along the River Finn. As thedistribution map indicates, Marsh Stitchwort has additionally been found – although very much morerarely – on the shores of Lower Lough Erne and in a few scattered additional sites. However, at severalof the outlying stations recorded in the 1950s or earlier, it seems to have disappeared, as there are nopost-1975 records for them.
British and Irish occurrence
Fermanagh, and in particular Upper Lough Erne, is the northern Irish 'headquarters' of Marsh Stitchwort.Elsewhere in the nine-county province of Ulster there are just three very isolated stations for theplant in VCs E Donegal, Tyrone and Down (H34, H36 and H38) (NI Vascular Plant Database). Apart fromthis, S. palustris has declined to a certain extent elsewhere in Ireland following a trend inwetland habitat losses that dates back to about the 1940s (Walters 1986; P.S. Lusby, in: Preston etal. 2002). In Ireland, Marsh Stitchwort is now thinly and widely scattered around wetlands inthe Central Plain, mainly alongside the long, slow-flowing River Shannon and by numerous tributaries andlakelets off it. Visual inspection of the hectad map for Ireland in the New Atlas indicates that,apart from the four northern VCs already mentioned, it is found in an additional 17 VCs in the Republic.
In Britain the distribution of Marsh Stitchwort is widespread but distinctly thin and patchy, reaching asfar north as Perthshire (VCs 87 & 88). The main areas in England are the Norfolk Broads and theSomerset Levels (Walker et al. 2017). The New Atlas map symbols show that there has been awidespread decline in Britain, especially noticeable in lowland areas of C & E England. The declinedates from before the 1930s and represents more than 30% at the hectad level of discrimination (P.S.Lusby, in: Preston et al. 2002). Surviving populations are now small, having a mean of 17individuals, even in protected sites (Walker et al. 2017).
Reasons for the decline of this wetland species must begin with land drainage, which has undoubtedlyoccurred on a widespread basis and given rise to loss of suitable, open, wetland habitat. However, otherchanges in land use and farming are also significant and cause losses, including under-grazing andeutrophication, which together allow invasion and subsequent dominance by tall, vigorous herbs andgrasses that out-compete S. palustris and quickly replace it.
European and world occurrence
S. palustris is a Eurasian boreo-temperate species and has a widespread distribution in northernparts of Europe and W & C Asia. It extends south as far as the Alps and the Carpathians, but southof these mountain ranges it is very uncommon. In France, it is mainly confined to the N coast, and isonly very thinly scattered elsewhere in the country. It is considered 'endangered' or 'criticallyendangered' in many French Departments. The species is likewise considered endangered in Switzerland,Belgium, the Czech Republic, Hungary, Germany and Norway. In Great Britain, it is regarded as'vulnerable' (Walker et al. 2017). In SW Europe, S. palustris is absent from Spain, mostof Italy and all of the Mediterranean islands except Corsica (Hultén & Fries 1986, Map 729).
One of the most curious and surprising features relating to the distribution of S. palustris isthe fact that the species has been introduced to several regions of the globe, far beyond its native,core area of occurrence. Since it is not a commercial, edible, medicinal or garden plant, it seemsrather inexplicable that S. palustris has been introduced and recorded from eastern Canada,Australia, Tasmania and New Zealand (Hultén & Fries 1986, Map 729). However, the authors of theFlora of New Zealand 4 (Webb et al. 1988), rejected S. palustris as notbeing part of the naturalised flora. One suggestion as to how S. palustris might have travelledacross the world's oceans, is as part of ships' ballast. The fact that some strains of the species cangrow on seashores suggests the possibility that seed, or other plant parts, might have been incidentallytransported in this manner along with shingle, and somehow remained viable until deposited again on afarther shore.
Names
The genus name 'Stellaria' is of medieval origin from the Latin 'stella', meaning 'a star', referring tothe shape of the five-petalled flower (Gilbert-Carter 1964). The Latin specific epithet 'palustris' isfrom 'palus' and 'ŭdis', meaning a swamp of bog, a reference to the typical habitat (Gilbert-Carter1964). The English common name 'Marsh Stitchwort' is a typical, recent, invented, book name of nofolklore significance. The 'Stitchwort' part of the name is examined in my Stellaria holosteaspecies account on this website.
Threats
Habitat loss due to drainage is the principal threat, usually associated with agricultural 'improvement',nutrient enrichment, inadequate grazing, neglect, or general building development. Management usingsuitable cattle breeds or cutting for hay or haylage is often problematic and uneconomic, due to thesmall size of many fragmented sites. Sites that are regularly flooded may also require improvement inpoor water quality and reduction of cultural eutrophication (Walker et al. 2017).
Native, common and widespread. Eurosiberian boreo-temperate, but also widely naturalised, including NAmerica and New Zealand.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This is a very common, widespread and locally abundant weedy perennial of moist to dry, nutrient-poor,moderately acid, stony, sometimes sandy or calcareous grassy habitats. A very variable species, mostfrequent on disturbed grassland, verges and ruderal situations, but elsewhere it can be found on marineforelands, including on grey and fixed (green) coastal dunes (Jonsell et al. 2001). S.graminea is probably one of the most common wildflowers of open ground on acidic loam or sandysoils throughout lowland Britain and Ireland. Generally it appears to prefer lighter, free-draining,acid soils, but while it avoids wet, strongly acid peaty soils, the same cannot be said of it inlime-rich districts, where shallow raw peaty humus accumulates directly over hard crystalline calcareousrock. Lesser Stitchwort is most common in rough grassy wayside places on hedge, river and streamsidebanks, and is frequent in semi-shade in open scrub and woods on rocky talus slopes and aroundlakeshores. In addition, it often forms large stands growing along and supported by wire fences aroundpermanent rough pastures. In the latter situation, it is also frequently associated with taller, rough,nutrient-enriched grass around animal droppings.
Lesser Stitchwort can easily be identified by its narrow, fresh green, grass-like opposite leaves, andits flowers, which are produced from May to August or even later, are smaller than those of S.palustris (Marsh Stitchwort) or S. holostea (Greater Stitchwort), and have very narrow,deeply notched white petals. The species is a hemicryptophyte perennial (ie overwintering buds are borneat ground level, often protected by leaf litter).
Reproduction
Flowering of S. graminea begins in May and can continue until the first frost in October or later,although the peak is in June. The inflorescence is long-stalked, lax and bears around 50 flowers. Theflowers are self-compatible and protandrous, attracting flies, bees and small beetles. Seed set is good.
Vegetative reproduction also occurs, mainly in open habitats where growth of a slender, creeping rhizomedevelops compact, clonal clumps (Jonsell et al. 2001).
Variation
S. graminea is very variable in habit, leaf shape and size, density and length of cilia on bractsand sepals, and number and size of flowers. Much of the variation is environmentally induced, especiallywith respect to light levels and soil moisture. Complete or partial male-sterility is rather common andis associated with dwarfism of flowers, capsules, seeds and vegetative size (Jonsell et al.2001).
Fermanagh occurrence
In Fermanagh, S. graminea is very common and widespread, having been recorded in 319 tetrads,60.4% of those in the VC.
British and Irish occurrence
S. graminea is common and widespread throughout most of lowland Britain & Ireland. However,in comparison with S. holostea (to whose morphology, reproductive biology and general ecology itappears similar, at least at a superficial level of examination), it is less tolerant of shade and morescarce, rare or absent than Greater Stitchwort in the wetter, more boggy, acid peat landscapes of theextreme west of Ireland, NW Scotland and the highlands. In SE England, like S. holostea, itavoids swampy conditions around The Wash (Clapham et al. 1987; Preston et al. 2002).
European and world occurrence
Like S. palustris, Lesser Stitchwort is a Eurosiberian boreo-temperate species and is verycommon and widespread in northern parts of Europe and Asia, well into Russian Siberia. However, it isscarce and thinly scattered south of the European Alps and Pyrenees, and absent from all theMediterranean isles, except Corsica, and the Peloponnese (Hultén & Fries 1986, Map 731). S.graminea has travelled with man well beyond its native range and is now widespread in eastern NAmerica, rarer but present in western N America, and scattered in E Asia and New Zealand.
Names
The genus name 'Stellaria' is of medieval origin from the Latin 'stella', meaning 'a star', referring tothe shape of the five-petalled flower (Gilbert-Carter 1964). The Latin specific epithet 'graminea' means'grass-like' and refers to the leaf shape (Johnson & Smith 1942). The English Common name 'LesserStitchwort' is a typical, recent, invented, book name of no folklore significance. The 'Stitchwort' partof the name is examined in my Stellaria holostea species account on this website.
Threats
None.
Native, common and very widespread. European temperate, but very widely naturalised, including in NAmerica and New Zealand.
1881-2; Barrington, R.M.; Devenish Island, Lower Lough Erne.
March to November.
Growth form and preferred habitats
This slender, delicate, brittle-stemmed, rhizomatous, shallow-rooted, wintergreen perennial formslow-growing mats on wet, open, sometimes disturbed fertile ground of acid to neutral reaction besidestreams, ditches, marshy ground on lakeshores and in flushes by springs in grasslands. Generally, it isa good indicator of moving groundwater. Its creeping clonal mats are often hidden underneath tallervegetation, although it can sometimes grow upright and ascend to 40 cm (Stace 2010). S. alsinealso colonises bare mud in other more frequently disturbed, fairly ruderal, muddy situations, eg aroundfield gates and along wet tracks and rides in woods, where it commonly associates with Juncusarticulatus (Jointed Rush), J. bufonius (Toad Rush) and Montia fontana (Blinks).
Fermanagh occurrence
In Fermanagh, S. alsine is common, with a wide variety of suitable habitat available, so that ithas been recorded in 333 tetrads, 63.1% of those in the VC. In somewhat shaded, strongly eutrophic,peaty, tall fen situations, eg in reed-beds around Upper Lough Erne, it can behave as an emergent onbare mud. Otherwise, in shallow, still or very slow-flowing water it can become a floating semi-aquatic.Bog Stitchwort also occurs in more acid, infertile ground on the slopes of Cuilcagh, the highestmountain, and in mixed upland oakwood at the Correl Glen NR, plus in mixed ashwoods below the limestonecliffs of Knockmore and the sandstone ones of Poulaphouca, overlooking Lower Lough Erne. It is rare, butnot unknown or excluded from the limestone areas of the county.
Reproduction, competition and survival
The small flowers are borne on the upper branches of the plant in lax, few-flowered dichasial cymes. Thepetals are bifid almost to the base and are shorter than the sepals, or sometimes absent. The flowersattract few insect visitors and therefore are mostly self-pollinated. Seeds are rugose-tuberculate, palereddish brown to dark brown. Seed setting is usually sparse, but production in unshaded sites can beprolific. A large, long-persistent population can develop in the soil seed bank and some may germinateafter habitat disturbance (Grime et al. 1988; Thompson et al. 1997; Jonsell et al.2001). Capsules and seeds float, and together with fragments of the brittle stems broken by disturbance,they enable water or mud-borne dispersal and colonisation of suitable moist habitats. This process isoften assisted by man and his stock animals. S. alsine is a weak competitor and is favoured bydisturbance of the soil surface that allows seed germination. Populations are often ephemeral (Jonsellet al. 2001).
Like other members of the genus Stellaria, S. alsine possesses a thin, creeping rootstockfrom which it produces numerous decumbent, ascending stems, not all of which flower (Clapham etal. 1962, 1987). The weak, decumbent or more prostrate stems can layer themselves, particularlyafter trampling, and towards the end of summer by producing adventitious roots. The species may thusdevelop quite extensive creeping clonal mats, hiding underneath taller vegetation which by this stage isbeginning to die down.
Variation
Plants are variable in habit and in the size and shape of leaves. In damp and shady situations, plantsdisplay a more diffuse habit and are less glaucous than those in drier more open sites (Jonsell etal. 2001).
British and Irish occurrence
In Britain, S. alsine is also well represented more or less throughout, but it appears to havedeclined in the S & E of England since 1950. This is probably due to drainage and agriculturalintensification, including a move to more arable farming and the re-seeding of moist pastures. InIreland, it is widespread and common, but more patchily recorded in the C & W of the island (P.S.Lusby, in: Preston et al. 2002).
Names
The genus name 'Stellaria' is of medieval origin from the Latin 'stella', meaning 'a star', referring tothe shape of the five-petalled flower (Gilbert-Carter 1964). The Latin specific epithet 'alsine' isborrowed from a genus name used by Theophrastus for an unknown plant (Gilbert-Carter 1964).Alternatively, 'Alsine' was a name used by Dioscorides for a chickweed-like plant (Gledhill 1985). Theprevious, and until recent specific epithet was 'uliginosa' which means 'growing in marshy places' from'uligo', 'inis', moisture, marshy quality (Gilbert-Carter 1964).
The English Common name 'Bog Stitchwort' is a typical, recent, invented, book name of no folkloresignificance. The 'Stitchwort' part of the name is examined in my Stellaria holostea speciesaccount on this website.
Threats
None.
Native, very rare, but probably a mis-identification. Circumpolar boreo-temperate.
1 June 1995; EHS Habitat Survey Team; Moneendogue ASSI.
Growth form and preferred habitats
This very variable, large-flowered, rhizomatous, loose mat-forming perennial Chickweed is more or lessrestricted to dry, well-drained, calcareous to slightly acid light sandy or gravelly soils, often inopen or semi-closed vegetation in sunny habitats that have been either created or strongly influenced byman. The range of habitats includes short, permanent pastures, margins of arable fields, sand-pits,wayside verges, roadside banks and waste ground. Less frequently, near the coast C. arvense canoccur on sand or sandy gravel in dune grassland. The upper portion of the plant and the inflorescenceare more or less densely covered with short hairs (<1mm), some hairs glandular (but not sticky).Leaves are linear-lanceolate, acute, hairless or sparsely hairy (Garrard & Streeter 1983; Stace2010).
Variation
In Europe, six geographical subspecies have been described, of which only subsp. arvense occursthroughout the species range, including Britain & Ireland (Tutin et al. 1964). Varieties havealso been named in relation to characters such as the degree of plant hairiness and leaf shape. In NAmerica, C. arvense displays a high degree of phenotypic plasticity which is undoubtedlyadaptive, and which helps explain the broad ecological tolerance and the degree of taxonomic variationobserved within the species across its wide geographical range (Wagstaff & Taylor 1988).
Sexual reproduction
Flowering takes place from May to August. The erect inflorescence is few-flowered (3-7), with pure whiteflowers 12-20 mm in diameter, bilobed petals about twice as long as sepals, and a superior ovary withfour or five styles. The flowers are protandrous, nectar is partly concealed and pollination is by smallbees and flies (Proctor & Yeo 1973). The capsule is cylindrical and straight, 6-8 mm, slightlylonger than the sepals. It opens by splitting at the apex to form ten teeth that help release the seed.
Seed production and survival
The dark red-brown seeds are flattened asymmetrically, often wider than long, with acute, narrowtubercules. In Finland and Scandinavia, where C. arvense is a 19th centuryintroduction, seed set is often low or absent, possibly due to high levels of self-incompatibility, plusthe occurrence of purely female clones (Jonsell et al. 2001). On the other hand, elsewhere in itswide geographical and altitudinal range, C. arvense is regarded as an obligate outcrosser(Wagstaff & Taylor 1988).
Survival in the soil seed bank is listed as either transient or short-term persistent, meaning continuingpresence for up to five years (Thompson et al. 1997).
Vegetative reproduction
The plant has a profusely branching rhizome and the lower leafy stems are procumbent and readily root attheir internodes, thus helping to spread the plant and form loose, sprawling clonal mats, or tighter,denser, upright clumps.
Fossil record
The distinctive seeds of C. arvense have been found in all sub-stages of the Weichselian Ice Ageand all zones of the Late Weichselian. However, there is a total absence of any fossil seed in thecurrent interglacial (Flandrian) record, which certainly appears significant. Godwin (1975, p. 147) drewattention to Hultén's description of the plant (Hultén & Fries, 1986) as belonging to the categoryof circumpolar species strongly spread by culture (see European and world occurrence below).
Fermanagh occurrence
There are just two records of C. arvense in the Fermanagh Flora Database, both of which were madeby members of an EHS Habitat Survey Team in separate sites in the far NW of Fermanagh in June 1995. Thedetails of the first record are given above and the second was made on 9 June 1995 at Drumlisaleenmeadow.
Irish occurrence
C. arvense is a very rare and local plant in the N of Ireland, usually occurring on coastal sandbanks and dunes. It has been recorded in one or two sites in Cos Down and Antrim (H38 & H39), anduntil the 1940s and 1950s, also at two inland sites on the shores of Lough Neagh (FNEI 3). JohnHarron (Flora of Lough Neagh) did not manage to refind it in his thorough floral survey of LoughNeagh. Otherwise, C. arvense in Ireland is restricted to dry, calcareous to somewhat acidic sandyor gravelly soils near the coast mid-way along the E & W coasts. It is locally frequent in theBurren, Co Clare (H9) and near Galway city, but only rare and scattered along the Co Dublin coast (H21)(Flora of Connemara and the Burren; Flora of Co Dublin).
While it would not be impossible to find C. arvense in dry limestone grassland in Fermanagh, wereally ought to reject the two 1995 records without supporting vouchers and until such confirmation isprovided we have to regard these as possible mis-identifications. If this species had ever been recordedin Cos Donegal, Sligo or Leitrim (H34 & H35, H28 and H29) we would look on this as a mere extensionof its range, but this is not the case. Having said all that, with its relatively large white flowers itis difficult to imagine which other related species of dry limestone grassland might have beenmistakenly identified as C. arvense. The only candidate is Stellaria holostea (GreaterStitchwort), and it is hard to conceive that one or more members of an experienced EHS Habitat SurveyTeam could mistake such a common plant for anything else. Of course having no vouchers we have no ideaof the condition of the two plant specimens in question, but what is clear is that the field botanistswho listed the C. arvense did not appreciate that it was a significant find in an Irish context.
Unfortunately in these situations quite often the survey workers employed by EHS fail to collect avoucher. In other Fermanagh situations, where these workers found a questionable plant, the EHS fieldcard is sometimes marked with an asterisk or a comment, indicating the recorder knew or felt the recordmight be of special significance. The absence of this type of recognition suggests two possibilities:either the plant wasn't properly examined, or perhaps an imported British botanist was involved whoexpected to find C. arvense in the vegetation being surveyed.
British occurrence and decline
In Britain, C. arvense is widespread, but with a distinct eastern tendency in its distribution. Itis certainly not restricted to coastal situations, but is found across the country on dry pastures,roadsides, wayside banks and field margins (P.S. Lusby, in: Preston et al. 2002).
Within the area of calcareous soils prevalent in E England where the species is principally, butinfrequently recorded, the presence of C. arvense appears to be declining. Particularly at themargins of its range, recording indicates there are losses that probably reflect cultural eutrophicationseeping in from surrounding 'improved' grasslands, together with physical destruction of the dry, oftenrocky grassland fragments in which it is mainly found (Braithwaite et al. 2006).
European and world occurrence
Circumpolar boreo-temperate, native in C & W Europe, although not indigenous in NW Europeincluding Fennoskandia. It is indigenous also in Mongolia, Japan, China, N & S America andGreenland. In the former USSR, the species is distributed in Arctic regions, in the European part, inthe Caucasus, W & E Siberia. It is also native in parts of the Far East and C Asia. As Hultén &Fries (1986) point out, both within and beyond the original native area of the species, which it isimpossible to define exactly, the species sensu stricto and other taxa have been spread considerably byman, eg to S Africa and New Zealand (Hultén & Fries 1986, Map 741).
Names
The genus name 'Cerastium' is from the Greek 'keras' or 'kĕrastēs' meaning a horn or horned respectively,from the fact that some species have curved seed capsules resembling horns as they emerge from the calyx(Johnson & Smith 1946; Stearn 1992). The Latin specific epithet 'arvense' means 'growing on arableland' from 'arval', 'arable land' (Gilbert-Carter 1964). The English common name 'Field Mouse-ear' is abook name, but see the Names section of my Cerastium fontanum species account on this website formore consideration of the name origin.
Threats
C. arvense is regarded as near threatened on the 2005 Red List of plant in the UK, which includesN Ireland and the other three British subdivisions (NBN website, viewed 23 January 2018).
Cerastium fontanum Baumg. (C. holosteoides Fr. = C.vulgatum L.), Common Mouse-ear
Native, very common and widespread. Eurosiberian boreo-temperate, but so widely naturalised as to havebecome circumpolar boreo-temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This very variable, often densely hairy, straggly to loosely tufted, or low creeping, mat-forming,wintergreen perennial (or sometimes annual) weed, grows in a tremendous variety of both natural andman-made, moderately fertile (mesotrophic) habitats. They can vary from moist to relatively dry,unshaded grasslands to numerous types of more or less disturbed ground throughout Britain & Ireland(New Atlas). Typical growing conditions are found mainly but not exclusively in lowland areas,and include moist meadows, pastures, lawns, lake shores, rock outcrops and moderately disturbed groundon roadsides, quarries and rough wayside areas. It does occur, but much less frequently, on soggy, boggyground, both acidic and calcareous, and in also in more shaded situations. C. fontanum can alsoappear in coastal situations on fixed sand dunes and on shingle. It can also invade a range ofartificial habitats, including cultivated ground, waysides, walls and the base of fences.
C. fontanum can be distinguished from the other small-flowered Cerastium species by theabsence of glandular hairs on the sepals, the sepal hairs not overtopping the sepal apex, and the fivestyles being 1.5-2 mm long (Webb et al. 1988).
Fermanagh occurrence
In Fermanagh, C. fontanum is recorded in 482 tetrads, 91.3% of those in the VC. It is absent,however, from three situations: aquatic habitats, strongly acidic bogs and deep or dark, evergreenshade. In the Fermanagh Flora Database, it ranks 40th in terms of recording frequency,falling between Calluna vulgaris (Heather) and Dryopteris dilatata (Broad Buckler-fern).On the other hand, when its presence is measured in terms of tetrad frequency, C. fontanum liesin 12th place in the county and its neighbours on the listing are Hedera helix (CommonIvy) and Taraxacum officinale (Dandelion).
This difference in ranking indicates (albeit in a rather subjective and crude manner) the considerableextent to which semi-natural habitats in Fermanagh were surveyed in detail for their conservationpotential, while comparatively few man-hours have been devoted to disturbed, artificial situations. Thisis very probably always the case with the many local flora surveys that have been carried out over theyears. At least we expect this to be the situation in rural VCs, if not in those with a high proportionof urban areas and associated disturbed, artificial habitats.
Variation
On a world basis, C. fontanum has a very wide distribution and ecological range and is oftenselfed (ie autogamous). This gives rise to marked polymorphism to such an extent that four subspeciesare recognised in Europe (Tutin et al. 1993). Three of the European subspecies are recognised asoccurring in Britain & Ireland, although one of them, subsp. holosteoides (Fr.) Salman, isconsidered a variety in Flora Europaea (Clapham et al. 1987). Two of the Europeansubspecies are recognised in Ireland – the very common form, subsp. vulgare (Hartm.) Greuter& Burdet, and the rarer subsp. holosteoides. The latter form is always associated withrivers, where it grows on grassy riverbanks, marshy flood-plain meadows, or just upstream of thesections of rivers that are under tidal influence. Unfortunately, these two more widespread subspeciesintergrade, and at some sites only intermediates are known (M.B. Wyse Jackson, in: Rich & Jermy1998).
The third taxon, subsp. scoticum Jalas & P.D. Sell, is regarded as a Scottish endemic. Stace(2010) suggested that all these taxa might be better placed in just one subspecies. On the other hand,Sell & Murrell (2018) divide the variation into just two subspecies, subsp. scoticum andsubsp. vulgare, the latter subdivided into four varieties (var. glandulosum (Boenn.) P.D.Sell, var. serpentini Novák, var. lucens (Druce) P.D. Sell and var. vulgare(Hartm.) M.B. Wyse Jacks. The previous species or subsp. holosteoides is subdivided and relocatedmainly in var. lucens (Sell & Murrell 2018).
Both of the subspecies previously listed for Ireland have been very rarely recorded in Fermanagh, but thevast majority of records of this common weed have not been discriminated at subspecific level in thecounty.
Sexual Reproduction
In some situations, Common Mouse-ear can show the rapid relative growth rate typical of weedy ruderalspecies, and under greenhouse conditions it can flower within nine weeks of germination (Peterson 1969).Flowering usually takes place between April and September. Flowering shoots are more erect thanvegetative ones, and are short-lived. The inflorescence typically bears around 20 perfect, hermaphroditeflowers. The flowers have five petals, bilobed to about mid-way (ie emarginate), shorter than, or up to1.3 times as long as the sepals. The flowers are protandrous and are visited mainly by flies. However,they are self-compatible and set seed even if insects fail to pollinate them (Clapham et al.1987). A normal-sized plant produces around 6,500 seeds (Salisbury 1964, p. 159).
Seed dispersal and survival
The curved capsules, 7-18 mm in length, open by splitting at the top to form ten teeth. The brown seedsare rounded triangular in outline and are decorated with tubercles or low ridges. Seeds are presumablydispersed mainly by wind, but they are also carried in mud by man (ie anthropochorus, literally 'carriedby man' as a primary dispersal agent), and other animals. Previously, before efficient industrial seedcleaning, seed of C. fontanum was incidentally transported worldwide mixed with commercial grain(Ridley 1930; Peterson 1969). Seed can survive dormant in the soil for over 40 years.
Vegetative reproduction
While individual flowering stems of C. fontanum may be short-lived, adventitious rooting ofhorizontal/decumbent lower stems and growth of a creeping rootstock constitute vegetative propagation,enabling the formation of persistent, dense clonal mats and lateral spread of the species in permanentgrasslands (Clapham et al. 1987; Grime et al. 1988).
Competition and disturbance
Being an essentially creeping, mat-forming perennial or annual, C. fontanum is not able to competewith more aggressive, taller species. It therefore tends to avoid such competition, tolerating habitatswhere it experiences moderate to severe levels of disturbance in order to survive in the long-term(Grime et al. 1988). Jonsell et al. (2001) mention short, ascending or erect rhizomes (ieswollen stems), in some forms of the species, which may also feature in enabling vegetative propagation.In pastures, C. fontanum can cope with moderate levels of grazing and trampling by cattle, and inlawns it tolerates mowing very well and can persist under such a regime for long periods. This degree ofsurvival ability is largely due to the plant's powers of reproduction and regeneration, which aregoverned by the level of soil fertility present.
Fossil history
Fossil seed of C. fontanum (listed as C. holosteoides Fr.), has been found in Britain andIreland in several interglacials from sub-stage II of the Hoxnian onwards. Four records in the currentFlandrian interglacial are associated with agriculture or settlement and date from 2390 BP through tothe medieval period (4th to 9th century AD). Godwin (1975) comments that itappears to have thrived in periglacial conditions, surviving in open situations naturally throughinterglacials and to have expanded in response to later human clearances.
British and Irish occurrence
C. fontanum, in the broad sense, is very common and widespread throughout the whole of bothBritain and Ireland, including offshore islands. The subspecies are not widely recorded and theirdistribution is not yet clear.
European and world occurrence
This is a Eurosiberian boreo-temperate species, believed to have originated in Europe and W Asia.However, it has been extremely widely distributed as a weed of cultivation, and has become circumpolarboreo-temperate and almost cosmopolitan (Hultén & Fries 1986, Map 747).
Names
The genus name 'Cerastium' is from the Greek 'keras' or 'kĕrastēs' meaning a horn or horned respectively,from the fact that some species have curved seed capsules resembling horns as they emerge from the calyx(Johnson & Smith 1946; Stearn 1992). The Latin specific epithet 'fontanum' is from 'fons, fontis'meaning a spring or fountain, a reference to damp habitats (Gilbert-Carter 1964).
C. fontanum (as C. trivale Link.) is listed with three English common names, 'Chickweed','Mouse-ear Chick-weed' and 'Mouse-ear' by Britten & Holland (1886), who regard it as a generalbook-name for any form of Cerastium, but especially linked with this species. 'Chickweed' is ofcourse shared with Stellaria media and many other small plants of similar habit, and 'Mouse Ear'is likewise shared with Hieraceum pilosella L., in both instances the name is presumably given onaccount of the shape and hairiness of the leaves which resemble a mouse's ear. The name 'Mouse-ear' wasfirst applied to both these species by Henry Lyte (1578) in his translation made from l'Écluse's Frenchversion of Dodoens' Crŭÿdeboeck of 1554, entitled A Niewe Herball.
Threats
None.
Native, common and widespread. European southern-temperate, but so widely naturalised it has becomecircumpolar southern-temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This rather scruffy, slightly sticky, often dust-coated, weedy winter or summer annual is astress-tolerant ruderal that rarely grows anywhere except on disturbed ground (Grime et al.1988). The plant has erect or ascending, flexuous and fragile stems up to about 30 cm in height. Thestems have long, non-glandular hairs below and glandular hairs above. Leaves in the mid-stem areelliptic, 1.3-2.1 times as long as wide and hairy all over. Stickiness also resides in the compactinflorescence of the plant which is densely covered with glandular hairs (Garrard & Streeter 1983).
Common and widespread in lowland areas of both Britain & Ireland, C. glomeratum prefersmoderately fertile or manure-enriched conditions, but it will tolerate a range from moist to dry, lightsandy to clayey soils in sunny to shaded situations. Sticky Mouse-ear is often found in bare areas ofcompacted clayey soil where surface water lies in winter. It is also frequent around gates where animalstale and trample the ground, and at the shady base of damp walls in both urban areas and farmyards.
C. glomeratum sometimes thrives in thinly-grassed patches in lawns and in regularly mown ortrampled grasslands, tracks or soil surfaces, as well as in more open, moderately disturbed wasteground. Previously, before the near extinction of arable agriculture in Fermanagh, it would have been anabundant field weed. It cannot, however, survive regular cultivation in gardens. Likewise, in wellmanaged grassland, it is only a poor competitor with perennial grasses and sooner or later it becomesscarce, rare or almost entirely absent from such situations (Sinker et al. 1985).
Fermanagh occurrence
In Fermanagh, C. glomeratum has been commonly recorded in 287 tetrads, 54.4% of those in the VC.
British and Irish occurrence
Widespread and common throughout lowland Britain & Ireland, it is however scarce in wet, boggy groundin parts of W Ireland and NW Scotland since the species favours or requires moderately fertile soilconditions (Preston et al. 2002)
Flowering reproduction
As with C. fontanum (Common Mouse-ear) and Stellaria media (Common Chickweed), germinationcan occur throughout the year given mild conditions, although there are strong peaks in the early springand autumn. Frost-tolerant overwintering juvenile plants have five to seven small, tightly clusteredleaves, but do not develop a real rosette (Salisbury 1962; Jonsell et al. 2001). The smallflowers appear in compact cymose clusters from April to September or later, even until the occurrence ofthe first autumn frost. The flowers are little visited by insects but are self-compatible and willautomatically self-fertilise. In some populations, flowers are frequently produced without petals andare pollinated and fertilised while still in bud (ie they are cleistogamous, a term literally meaning'closed marriage') (Clapham et al. 1987).
Fruiting plants have a much more lax inflorescence due to elongation of the internodes afterfertilisation, and each 6-10 mm curved, cylindrical capsule contains around 50 seeds (Salisbury 1964).
Seed dispersal
The capsule apex splits to form ten teeth and releases the small, light, tuberculate, yellow-brown seedon the breeze. In common with many other species in the family, the seeds are long-persistent (Thompsonet al. 1997). Apart from wind dispersal, transport of the species is very closely associated withman and his activities. For instance, Sticky Mouse-ear is such a very common weed in horticulturalnurseries and garden centres, customers must often be transporting it home with potted or containerisedplants.
Fossil record
None found. Exactly what should be taken from this negative result is not clear, although the link withman and environmental disturbance due to agriculture and stock animals is obvious. Seeds of other commonrelated species do occur in the fossil record, but sometimes in only a small number of sites, eg C.fontanum and C. cerastoides (L.) Britton (Starwort mouse-ear) (Godwin 1975, pp. 147-8).
European and world occurrence
C. glomeratum has been spread by man from W Europe and the Mediterranean to India, N & SAmerica, S Africa, S Australia and New Zealand, and become circumpolar (Hultén & Fries 1986, Map750). In Norway, C. glomeratum is recognised as being a neophyte introduction imported withballast (Jonsell et al. 2001).
Names
The genus name 'Cerastium' is from the Greek 'keras' or 'kĕrastēs' meaning a horn or horned respectively,from the fact that some species have curved seed capsules resembling horns as they emerge from the calyx(Johnson & Smith 1946; Stearn 1992). The Latin specific epithet 'glomeratum' is derived from'glomus', a ball of yarn and means 'aggregated' or 'gathered into a round mass', clearly referring tothe compact, cymose, cluster of the inflorescence (Gilbert-Carter 1964; Gledhill 1985). The Englishcommon name 'Sticky Mouse-ear' is a recent book name of no folklore significance.
Threats
None.
Native, very rare, possibly a mis-identification. European temperate.
Growth form and preferred habitats
This little, hairy, winter- (or occasionally spring-) annual forms an overwintering basal leaf rosette,dull dark-green in colour (Dark-green Mouse-ear is an alternative English common name for the species,and it used to have the scientific binomial, C. atrovirens Bab.). A plant of light, dry sandy orgravel soils, C. diffusum is a common species of open coastal habitats that rarely, or veryoccasionally, strays inland where it occupies a range of open, disturbed habitats with suitable soils.According to Jonsell et al. (2001), in Fennoscandia it does not prefer saline conditions, butrather, it tolerates sea spray.
In Britain, the inland habitats are generally associated with dry grassland by paths, wall tops, alongroadside verges where salt is regularly scattered for accident prevention in frosty weather, wasteground and rough grassland. Previously, during and in the years after the Second World War, C.diffusum also commonly occurred in Britain on the ballast between railway lines, but followingeconomic contraction of the railway mileage, it has declined in this type of site since then.
Sexual reproduction
The plant flowers from May to July. Diffusely branched flowering shoots are decumbent or ascending, up to30 cm tall and densely covered with short, sticky glandular hairs. The small flowers, 3 to 4 mm acrosshave four (or less frequently five) notched petals, equal in length or shorter than the sepals. Theflowers are rarely visited by insects and automatically self-pollinate (Clapham et al. 1987). Thestraight capsule is 6 mm long and typically contains 30 seeds, although there can be up to 45. The seedsare rounded, 0.7 mm across, pale- or reddish-brown in colour, bearing blunt tubercles (Salisbury 1964,p. 159; Jonsell et al. 2001, p. 151). Seed dispersal is presumably primarily by wind, assisted byhuman activity disturbing sites where it grows.
Fermanagh occurrence
There are two Fermanagh records from the dry limestone grassland at Rahallan Td, S of Belmore Mountain,an area now famous as one of the sites of Neotinea maculata (Dense-flowered Orchid). The plantwas listed in two tetrads at Rahallan on 15 May 2005 by RHN accompanied by Dr D. Cotton, the BSBIrecorder for Cos Sligo and Leitrim (H28 and H29) and Mr F. Carroll. Unfortunately, no voucher wascollected, and these two interesting finds are therefore unacceptable as new county records.
The only other Cerastium listed in the Fermanagh Flora Database for this site is C.fontanum (Common Mouse-ear). We therefore regard the records of C. diffusum as possibleerrors, and the species remains on our list of desiderata.
Fossil history
No information has been obtained on this subject.
British and Irish occurrence
In N Ireland, C. diffusum was previously frequent on the shores of Lough Neagh, but it hasmarkedly declined to rarity (Flora of Lough Neagh; FNEI 3; McNeill 2010). In Britain, SeaMouse-ear is very common on coasts and increasingly scarce inland (New Atlas; Braithwaite etal. 2006).
European and World occurrence
C. diffusum is variable and polymorphic to the extent that in Flora Europaea threesubspecies were recognised, of which only subsp. diffusum is definitely found in Britain &Ireland (Clapham et al. 1987; Tutin et al. 1993; Stace 2010). C. diffusum subsp. diffusum is almost entirely restricted to W, C & SEurope, extending northwards to S Sweden and eastwards to Ukraine. Again, the world map shows it largelyconfined to coastal sites (Hultén & Fries 1986, Map 753), the oceanic distribution of this Europeantemperate species suggesting that it is more frost-sensitive than warmth demanding (Jonsell etal. 2001). Hultén & Fries (1986) recognise that the distribution is incompletely known, andthere exists a small number of scattered introductions in N America. It is very puzzling to work out howand by what agency this annual species managed to cross the Atlantic Ocean, but it cannot be doubtedthat it succeeded in doing so on a number of occasions. The answer must somehow involve accidental orincidental human transport.
Names
The genus name 'Cerastium' is from the Greek 'keras' or 'kĕrastēs' meaning a horn or horned respectively,from the fact that some species have curved seed capsules resembling horns as they emerge from the calyx(Johnson & Smith 1946; Stearn 1992). The Latin specific epithet 'diffusum' means loosely spreadingor diffuse (Gilbert-Carter 1964). The English common name 'Sea Mouse-ear' is a recent book name of nofolklore significance.
Native, occasional to locally frequent and quite widely scattered. Eurosiberian boreo-temperate, but alsopresent in N America.
1860; Smith, T.O.; Co Fermanagh.
April to October.
Growth form and preferred habitats
Rosette forming, diffusely tufted perennial, taller than other Pearlworts with stems, ascending orprocumbent, usually 10-15 cm tall. Plants sometimes flower in their first year of growth. The stemleaves of S. nodosa are linear-subulate as in other Pearlworts, with the leaf length diminishingupwards from 10-15 mm to 1-2 mm. The plant gets its characteristic 'knotted' appearance from thepresence of bunches of shorter leaves (or undeveloped shoots) in the upper stem leaf-axils. The flowers,0.5-1.0 cm in diameter, are not always present in great numbers, but are most attractive. They have purewhite, entire petals nearly twice as long as the sepals, making the plant rather more showy andconspicuous than other Sagina species.
Knotted Pearlwort is a widespread, locally common, stress-tolerant, perennial colonist of open ordisturbed, slightly damp to wet, muddy to gravelly, moderately fertile, base-rich lake shores, shortfens, flushes, runnels and forest tracks. It is particularly associated with calcareous fens and moistto wet limestone grassland, including upland peat or clay areas flushed by seepage from preferablybase-rich springs. S. nodosa also frequents open, moderately acid to calcareous, sandy ground indune grassland and dune-slacks near the coast. In addition it is reported from ledges on coastal cliffs,less commonly in moist, open woods, and infrequently as a weed of arable cereal crops (Clapham etal. 1987).
Since it requires open habitats, S. nodosa tolerates, or possibly even demands, a moderate degreeof grazing and trampling to keep the ground sufficiently open, through limiting and discouraging morevigorous, taller competitors (Sinker et al. 1985; Grime et al. 1988).
Variation and Taxonomy
The species is polymorphic to a considerable degree and varies widely in hairiness, habit (ie prostrateor erect), length of shoots and leaves, degree of leaf succulence, flower size and number of flowers andbulbils (Jonsell et al. 2001). A form, described as var. moniliformis (G.F.W.Meyer) Lange, has stems more or less procumbent and the upper axillary buds, with their characteristicbunches of dwarf stems and leaves readily become detached, forming bulbils that can disperse andvegetatively propagate the plant (Clapham et al. 1987; Sell & Murrell 2018).
The great variability of the plant is reflected in the difficulty taxonomists have faced in assigning thespecies to a genus: in the past Sagina nodosa has been listed and named within the generaAlsine, Arenaria, Moehringia and Spergularia (Jalas & Suominen 1983).
Reproduction
Flowers are produced from July to September. They are protandrous (male first), and S.nodosa has the highest rate of cross-fertilisation in the genus. Larger flowers are bisexual,but smaller ones which are rare, may be entirely female. Few insects visit the flowers which areautomatically self-pollinated. The fruit capsule is short, ovoid, 4 mm and splits to the base to formfour or five valves. Seeds are 0.4 mm, dark brown and tubercled. The seed is long-term persistent in thesoil (ie survives more than five years) (Thompson et al. 1997).
Flowers are often sparse or absent, and when this occurs bulbils tend to be more numerous. In some areas,eg in seashore habitats, vegetative reproduction predominates (Jonsell et al. 2001).
Fermanagh occurrence
In Fermanagh, S. nodosa is occasional to locally frequent and has been recorded in 102 scatteredtetrads, 19.3% of those in the VC. In nine of these tetrads, there are pre-1975 records only, whichsuggests there has been some local loss of suitable habitats. It is principally found around the UpperLough Erne basin, which is partially fed by lime-rich waters arising in the hills to the SE. However, itis widely and more thinly scattered in the upland limestones of the Western Plateau, and also morerarely along damp forest tracks.
Irish occurrence
It is rather strongly and obviously associated with the wet, gravelly shores of the larger lakes in thewest and centre of Ireland which are geologically underlain by limestone basem*nt rocks. Elsewhere inIreland, S. nodosa has a definite N, W & C pattern of occurrence at the hectad scale. It isfrequent in open areas of sandy and gravelly seashores and in coastal grasslands, especially along thewestern Atlantic coastline.
British occurrence
In Britain, the species is also widespread, but in a more patchy and attenuated manner than in Ireland,displaying a definite, strong northern and rather weaker western predominance in its distribution(New Atlas). The current distribution of S. nodosa, in Britain at least, is partiallyimposed by major losses of suitable open, calcareous habitats, especially in S & E England. Analysisof the New Atlas Flora Database indicates that the decline affecting Knotted Pearlwort hasoccurred since 1950 (P.S. Lusby, in: Preston et al. 2002), pointing to changes in land use andagricultural intensification being most probably responsible.
European and world occurrence
Native in N & C Europe from Iceland to mainly coastal areas of N Spain and Portugal andstretching north-eastwards to northernmost Norway and C Russia. Not present in the Mediterranean basin,nor in Macaronesia (ie Canary Isles & Madeira) (Jalas & Suominen 1983, Map 910). S.nodosa also occurs in Asia, although the map provided by Hultén & Fries (1986, Map 757)shows it very thinly scattered there. The latter authors comment that the species is probably rare inAsia, and admit that their map of its Asian distribution may be incomplete. S. nodosa is alsowidespread in N America and therefore belongs to the amphi-Atlantic group of northern European plantsthat span the ocean (Hultén 1958, Map 105).
Names
The generic name 'Sagina' is Latin, meaning 'food crop' or 'fodder'. Spergula arvensis (CornSpurrey), which was cultivated and much valued in Flanders as a fodder crop for dairy cows, or as humanfamine food in the 15th century, was previously called Sagina spergula (Gilbert-Carter1964; Thirsk 1997, p. 17). The Latin specific epithet 'nodosa' means 'knotty' or 'many jointed' and isderived from 'nodus' meaning 'a knot'. In this instance, 'nodosa' refers to the characteristic bundlesor fascicles of dwarf branches in the upper stem leaf axils.
The English common name 'Knotted Pearlwort' is a modern book name, but the 'Pearlwort' portion is of17th century origin, a reference by John Ray (1660) either to the small fruit capsule, or tothe pearl-like unopened flower (Grigson 1974).
Threats
Cultural eutrophication from intensive farming and nitrogen in road traffic pollution stimulatingcompetition from taller, more vigorous species.
Native, common and very widespread. Eurosiberian boreo-temperate, but very widely naturalised worldwide.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
S. procumbens is a very common and widespread, locally abundant, wintergreen, rosette,mat-forming, or more rarely tufted, moss-like perennial, or sometimes a ruderal, annual. The plant isoften, but not always, tightly appressed to the substrate on which it grows, especially when growing insoil compacted by regular trampling. It colonises a wide range of semi-natural, open habitats, includingdamp, shaded cliff ledges, rocky areas or bare, sometimes marshy ground on lakeshores and river banks,and in damp, flushed areas on moors. It is also common in moist, moderately fertile, shaded artificialhabitats, for instance in short, heavily-grazed pastures, lawns, by paths, ditches, crevices inpavements and at the base of walls. It is indifferent to lime and really avoids only the most acid,waterlogged, or submerged aquatic situations (Grime et al. 1988).
An all too familiar garden and plant-pot weed, at a glance the tuft of linear leaves of the sterilecentral rosette, and the slender spreading branches can make it look quite moss-like, especially whenclones of it infest short mown grass in lawns. Numerous creeping stems arise from the sides of theflower-less, tufted central leaf rosette, and these root at intervals and can spread and interweave toform dense mats in suitable growing conditions.
As one might expect, the longevity of the individual plant is very much determined by the pattern andseverity of disturbance in the particular ground it occupies, and in many artificial, anthropogenicsituations, the species is almost entirely ruderal and annual. In more stable situations, it remains apersistent polycarpic perennial, and for instance has been observed surviving in a lawn for around eightyears (Grime et al. 1988).
Flowering reproduction
S. procumbens has a long flowering period stretching from April to September. Plants produceshort, erect, flower stalks bearing tiny, solitary, regularly or automatically self-pollinating flowers,which are often without petals or else have four (or occasionally five), minute white or greenish ones(Clapham et al. 1962). The flowers often remain closed, only opening when it is both sunny andwarm. It is, therefore, probably mainly autogamous, and seed-set is good (Jonsell et al. 2001). Acurious feature of S. procumbens is the way the pedicel of the developing fruit capsule flexesand allows it to droop and dangle, hook-like for a time, yet when ripe it straightens up and standserect to release the seed!
Capsule contents vary considerably, the number of seeds ranging from 20-135 with a mean of 74. The seedis extremely lightweight, almost dust-like, and despite being released from a very low point of origin,they are very readily lifted and dispersed by the slightest breeze that crosses the open mouth of thefruit capsule (Salisbury 1942, 1964). The seed is also adhesive in mud and is very probably transportedby man and other animals in this way. In common with many other members of the family, under suitableconditions this species persists for at least five years in the soil seed bank (Thompson et al.1997).
Variation
In Scandinavia there is considerable variation in ciliation (short hairiness) of the leaves, number ofsepals and petals, presence and length of the petals, and length of the mucro point on the leaf tip(Jonsell et al. 2001). In Britain & Ireland, there is considerable variation in the length ofthe leaf blade (0.5-1.2 cm), degree of leaf ciliation (including its absence), flower diameter (2-4 mm),sepal number (4, occasionally 5) and capsule size (2-3 mm) (Sell & Murrell 2018).
Hybrids
While selfing is the norm in S. procumbens, the fact that occasional or rare hybrids with otherSagina species are known, indicates that some out-breeding can and does occur (Stace 1975; Staceet al. 2015). No hybrid Sagina has ever been found in Ireland.
Vegetative reproduction
In areas of mild climate, established plants of S. procumbens can continue to grow almost all yearround, and when cut or grazed, small stem fragments may root and regenerate the plant in open sites withdamp, disturbed soil. Prostrate stems are also capable of rooting, and will form new plantlets ifdetached (Grime et al. 1988).
Competitive ability and survival strategy
On account of its appressed habit and low growth form, Grime et al. (1988) concluded that S.procumbens is a stress tolerant herb, unable to compete with the majority of taller andbroader-leaved pasture species. However, it is flexible in both its growth and reproductive strategies,and is sufficiently variable in terms of both genetic and environmental responses, to be capable ofcolonising artificial and freshly available semi-natural habitats (Jonsell et al. 2001). It isable to survive and compete in vegetation that is regularly and frequently mown, trampled or otherwiseconsiderably disturbed. It also avoids most competitors by growing in sites that provide severelylimited growing conditions, such as cracks in pavement.
Fermanagh occurrence
S. procumbens has been recorded in 370 Fermanagh tetrads, 70.1% of those in the VC. ProcumbentPearlwort is abundant in damp, shady situations at all altitudes in Fermanagh, on a wide variety ofsoils.
British and Irish occurrence
It is common throughout the whole of Britain and Ireland, and is especially frequent in man-made,artificial habitats.
European and world occurrence
Eurosiberian boreo-temperate, native in most of Europe (including Iceland, the far north of Scandinaviaand the Macaronesian islands), although rarer in the Mediterranean basin and eastern areas of thecontinent (Jalas & Suominen 1983, Map 917). Also present in N Africa and the mountains of W Asia.Introduced by man in many parts of the world, including N America, Australia, New Zealand (arrived c1873), the South African Cape Region (arrived c 1845), and parts of S America (Kloot 1983; Hultén &Fries 1986, Map 762; Webb et al. 1988).
Names and folklore
The generic name 'Sagina' is Latin, meaning 'food crop' or 'fodder'. Spergula arvensis (CornSpurrey), which was cultivated and much valued in Flanders as a fodder crop for dairy cows, or as humanfamine food in the 15th century, was previously called Sagina spergula (Gilbert-Carter1964; Thirsk 1997, p. 17). The Latin specific epithet 'procumbens' means 'extend', 'spread', 'creepingforward' or 'lying along the ground', ie prostrate (Gilbert-Carter 1964; Gledhill 1985), all of whichdescribes the typical habit of the species.
The English common name 'Pearlwort' is given to all Sagina species and was first used by John Rayin 1660. It refers to the small fruit capsule, or to the unopened flower. 'Procumbent Pearlwort' is amere book name translation of the botanical name. Alternative names listed for this very common plantare 'Beads' (Wiltshire), 'Bird's Eye' (Sussex), 'Little Chickweed' (Somerset) and 'Poverty' (Norfolk)(Grigson 1955, 1987).
In folklore, the Pearlwort plant has special magical powers associated with a tradition that believedChrist stood on it when he first came to earth or when he arose from the dead, making it a good-luckplant (Grigson 1955, 1987). In Scotland, it was one of the plants fixed over doors for good luck, or tokeep away the fairies who might spirit away the inhabitants (Grigson 1955, 1987; Vickery 1995). Grigsonlists several other myths associated with cows, milk and kissing!
Threats
None.
Native, occasional. European southern-temperate, but widely naturalised.
1900; Praeger, R.Ll.; Co Fermanagh.
May to September.
Growth form and preferred habitats
This small, stress-tolerant winter and summer annual is a pioneer colonist of dry or well-drained, sunny,lowland, open artificial or disturbed habitats where there is a virtual absence of biologicalcompetition. It frequently occurs in a wide range of such habitats, including, eg on the tops of walls,in cracks in weathered concrete, between paving stones, on or along forestry tracks and in bare orsparsely vegetated areas of sandy, clayey, and gravelly or otherwise stony soils. The most naturalhabitats it occupies are gravelly or sandy soils on heaths, sea cliffs and by paths on commons (Sinkeret al. 1985; Stace 2010).
Compared with the generally perennial, prostrate, appressed, spreading habit of S. procumbens(Procumbent Pearlwort), the typical S. apetala plant generally has a more diffuse, more uprightappearance, growing up to 15 cm tall. Both these species are very tolerant of trampling and they cancolonise heavily compacted soil on and near pathways. In such circ*mstances, they are often extremelydwarfed in height and flattened in appearance.
Flowering reproduction
The erect inflorescence bears 2-7 flowers from June to September. Although the Latin specificepithet 'apetala' translates as 'without petals', S. apetala may sometimes possess minute greenpetals within the ring of (usually four) sepals. While small crawling insects may visit the flowers,most pollination and fertilisation is probably the result of automatic selfing (Clapham et al.1962). Seed is produced in considerable numbers, about 50 per capsule. The light and dust-like seed isvery readily wind dispersed. Since the plant most frequently occupies disturbed, often trampled ground,seeds may also be carried in mud or hair by animals, or footwear and clothing by man (Salisbury 1964).Estimates of dormant longevity in the soil seed vary from transient up to viable for five years(Thompson et al. 1997).
British and Irish occurrence
S. apetala is common and widespread, especially in urban settings in lowland Britain &Ireland, but it becomes more scattered and scarce or absent in wetter, more acid, peaty conditions,especially on higher ground in the N & W of both islands (New Atlas). There is room for doubtabout the accuracy of the distribution of this and related forms, due to confusion regarding taxonomyand identification in recent years, and reassignment and renaming of subspecies (P.S. Lusby, in: Prestonet al. 2002). This is especially the case with regard to confusion with S. filicaulis (seeVariation below).
Fermanagh occurrence
In Fermanagh, S. apetala in the broad sense has been occasionally recorded in 59 tetrads,11.2% of those in the VC. In Fermanagh, outside urban and village areas, it is especially associatedwith ground in and around quarries.
Variation
Previously two subspecies were recognised, S. apetala subsp.apetala and subsp. erecta. However, S. apetala subsp.erecta F. Herm. and var. filicaulis have recently been subsumed into a new species, S.filicaulis Jord. (Slender Pearlwort), which is regarded as containing no less than four namedvarieties (Sell & Murrell 2018). As is shown below, this new combination has not beenrecorded in Fermanagh, where identification to subspecific level is only rarely achieved.
European and world occurrence
The total distribution is poorly known due to confusion of S. apetala with S. filicaulis.S. apetala is probably native in the whole of C Europe from Britain and Ireland to Poland andRomania, and southwards to Switzerland and the Pyrenees (Jonsell et al. 2001). Earlier treatmentby Hultén & Fries (1986, Map 763) suggests a much wider occurrence in Europe extending east toGreece and south to the Mediterranean islands, and possibly to N Africa. A very similar range is plottedin Jalas & Suominen (1983, Map 919), although they they show it as essentially a W European species,near absent in all of Scandinavia. A. apetala is also widely introduced to remote regions aroundthe world including S Africa, S Australia, Tasmania, New Zealand, E & W coast USA and S America.
Names
The generic name 'Sagina' is Latin, meaning 'food crop' or 'fodder'. Spergula arvensis (CornSpurrey), which was cultivated and much valued in Flanders as a fodder crop for dairy cows, or as humanfamine food in the 15th century, was previously called Sagina spergula (Gilbert-Carter1964; Thirsk 1997, p. 17). The Latin specific epithet 'apetala' refers to the petals being absent (oralmost so – see Flowering reproduction above). The English common name is a typical book name ofno folklore significance.
Threats
None.
1976; Dawson, Miss N.; Brockagh Point, Boa Island, Lower Lough Erne.
April to October.
In common with elsewhere in N Ireland, and indeed probably throughout the rest of Ireland, this is theusual form of this weedy annual that is found in Fermanagh (New Atlas). The Fermanagh FloraDatabase contains frequent records of subsp. apetala from eight tetrads, representing a presencein 1.5% of the VC. As mentioned in the species account above, it is only frequent in urban areas(including the larger villages), being very occasional and scattered elsewhere in the county.
Threats
None.
Native, very rare, but possibly over-looked and under-recorded.
1958; Mackechnie, R.; railway tracks near Pettigoe village.
This form of the small annual has been recorded only once in Fermanagh as listed above. Sadly the railwaytracks have long ago been lifted. The respected Scottish botanist, R. Mackechnie, who was visiting thecounty regularly at the time, recorded it. Notice of the record first appeared in the RevisedTypescript Flora.
Generally, elsewhere, this subspecies occurs in similar or identical habitats to the common form which wehave in Fermanagh, subsp. apetala, ie in dry, sunny, bare or open, man-made, disturbed orcompacted ground. A widespread smattering of both old and recent records of Fringed Pearlwort areplotted throughout Ireland in the New Atlas, a fact which strongly suggests that we, along withmany other botanical field workers, are probably guilty of overlooking this variant. We may partially beexcused by past name confusion created by the taxonomists, but all the identification points were madeclear in New Flora of the BI (1991, 1997).
Perhaps unfortunately, in terms of remembering the distinguishing identification feature, the developingfruit capsule of subsp. erecta has patent sepals arranged around it, NOT erect to erecto-patentones as in subsp. apetala. The sepals are blunt at their apex, whereas those of subsp.apetala are acute.
As noted above, this taxon has subsequently been combined with other forms and included in a new species,S. filicaulis Jord. (Sell & Murrell 2018).
;
Introduction, archaeophyte, occasional, but locally abundant. Eurosiberian wide-temperate, but widelynaturalised and now both circumpolar and well represented in the southern hemisphere.
1881; Stewart, S.A.; Co Fermanagh.
June to December.
Growth form and preferred habitats
A mainly summer annual, S. arvensis is an ascending or erect, glandular-hairy, somewhat stickyplant, often unbranched, or with 1-8 long branches arising from near the base. Stems bear slender,linear, fleshy leaves that are opposite but often appear whorled due to the presence of leaf clusters orfascicles in the stem leaf axils. Previously a common arable weed of as many as 25 different crops, andparticularly associated with cereals and flax, S. arvensis is now a declining ruderal species offertile, disturbed habitats, including sand and gravel quarries and more or less sandy, moist lakeforeshores (Holm et al. 1977; Grime et al. 1988; Rich & Woodruff 1996). S.arvensis appears to prefer light, sandy, acidic, moderately fertile, lowland soils (pH 4.6-5.5),and it shows a slight (or sometimes more definite) calcifuge tendency.
Although S. arvensis is said to be intolerant of trampling (New 1961), one of the most predictablehabitats we find it in is around field gateways where the soil is poached and manured by concentrationsof cattle. S. arvensis never occurs as a member of a stable, closed vegetation community. Rather,it is confined to open or disturbed situations where bare ground is available for colonisation. It isfrequently found in abundance in recently sown fields or garden lawns, but plants quickly disappearafter the first year as the closed turf gradually develops. Another situation where S. arvensiscan become conspicuously abundant is on the exposed shores of lakes after a period of hot, dry, summerweather.
Flowering reproduction
S. arvensis reproduction and survival relies entirely on seed. The plant grows rapidly, flowerswithin eight weeks of germination and seeds a fortnight later (New 1961). On account of its rapid growthand reproductive cycle, two generations readily fit within a growing season. The tiny white or slightlypinkish flowers are 97% selfed, although a range of insect visitors including flies and bees have beenrecorded (Knuth 1906-9). Sometimes flowers are pollinated and fertilised in bud (ie cleistogamy occurs)(New 1961). Flowers do not open until about mid-day, and they close again in the late afternoon. In coolor dull conditions, they remain shut. Depending upon source, an unpleasant odour is emitted by the openflowers (New 1961), or they are slightly fragrant (Jonsell et al. 2001)!
Fruit capsules are ovoid and up to twice the length of the sepals. They split to form five broad teethand release the seed. Seeds are lenticular, rugose, with or without club-shaped papillae, narrowlywinged and black, brownish, or grey in colour. The number of seeds produced per plant is between300-7,500, depending upon the number of capsules produced. This in turn reflects plant size and thedegree of branching. During experimental studies in India using local seed, Trivedi & Tripathi(1982) found seed production varied from 540-3,155 per plant, highlighting the great degree ofplasticity found in this cosmopolitan species.
Fossil record and status
S. arvensis is a cosmopolitan species of unknown origin which Holm et al. (1977) recognisedas one of the world's worst weeds, particularly of cereals. Until very recently, S. arvensis wasgenerally assumed to be native in Britain & Ireland (eg Stace 1997). However, Godwin (1975) pointedout that fossil finds are almost exclusively from archaeological sites from the Neolithic onwards, soearly introduction could not be ruled out. The editors of the New Atlas have now recognised it asan ancient introduction (Preston et al. 2002).
Variation
S. arvensis is very variable with respect to habit, stamen number and even seed surface texture.Some of this variation is environmentally induced, including the degree of branching (and hence flowerand seed production), degree of glandular hairiness and presence or absence of papillae on the seed coat(New 1961). A cline in the frequency of the two seed forms (papillate and non-papillate) exists acrossBritain & Ireland, running in a north to north-west direction. The papillate form (var.arvensis) is predominant in the south, but as one goes north and westwards it is graduallyreplaced by the smooth form (var. sativa). A matching cline with respect to hairiness exists, thedensely hairy form being predominant in the north and west of Britain & Ireland. These genotypicdifferences relate to general climatic factors: smooth seeds germinate better than papillate at lowertemperatures, but they proved more sensitive in this respect to higher moisture tensions (ie drierconditions) than the papillate ones (New & Herriot 1981).
Germination
S. arvensis germinates in the spring (usually in April) from a very large, persistent seed bank.It cannot easily be classified as a summer or winter annual, since although its seeds germinate inspring and summer, which is characteristic of a summer annual, dormancy seems to be broken by hightemperatures, which is typical of winter annuals (Karssen et al. 1988). Corn Spurrey appears todisplay a dormancy cycle varying seasonally from conditional dormancy to non-dormancy (Milberg &Andersson 1998). Bouwmeester & Karssen (1993) found that dormancy was broken in spring andre-induced in autumn with rising and falling temperatures respectively. They also showed, however, thatexposure to light, addition of nitrate and a desiccation treatment of seeds prior to the germinationtest strongly stimulated germination. Furthermore, a combination of these factors allowed S.arvensis to germinate in all seasons. Milberg & Andersson (1998) found that seed buriedoutdoors at the end of November and exhumed monthly for testing, germinated from March to December undertwo light treatments, but was greatest in the autumn months.
Seed dispersal
Seed transport is achieved through fruit capsules ingested and voided by grazing animals, such ascattle, sheep and birds (Holm et al. 1977). Seed is also unintentionally transferred in mud onfarm vehicles and boots. Long-distance seed dispersal is again accidental, chiefly as a contaminant ofa*gricultural seed (New 1961).
Seed longevity
In arable fields, S. arvensis can develop a huge, long-lived, soil seed bank of up to 23 millionseeds per hectare (New 1961). Seed longevity is variously reported, maxima ranging from around 50 years(Chippendale & Milton 1934), or "at least 50 years" (New 1961), to less than 1% viabilityafter 9.7 years (Conn & Deck 1995). There is also a fascinating report of ancient seed of S.arvensis recovered from archaeologically dated soil samples proving viable after burial for1,700 years (Odum 1965).
Fermanagh occurrence
S. arvensis is still a widespread species in N Ireland, but Fermanagh is the one area where thereare gaps in the hectad map. Locally, it has been recorded in 52 tetrads, 9.9% of those in the VC. It isoccasional and as the distribution map shows, it is thinly scattered across the county, being mostfrequently found in the better farmland east of Lough Erne. Corn Spurrey appears particularly frequentaround the Tempo area, E of Enniskillen, although with the exception of around exposed lakeshores, evenhere it is rarely abundant. It should also be looked out for on piles of topsoil and around recentlydisturbed roadsides, or in newly sown grasslands.
British and Irish occurrence
Common and widespread on suitable soils throughout. It mainly avoids basic calcareous and waterloggedsites. S. arvensis has declined markedly due to industrial seed cleaning and the widespread useof herbicides (New Atlas). It really has become more of an occasional species, often found onlyin small numbers.
European and world occurrence
Area of origin unknown, but the species common in C & E Europe, thinning eastwards in theMediterranean basin. S. arvensis becomes much more rare in eastern Europe. Jalas & Suominen(1983, Map 990) show it stretching from the far north of Scandinavia (70°N) to the south of Spain, butbecoming rare in Greece and Crete. It is introduced almost worldwide, including commonly in N America,especially in eastern states (Hultén & Fries 1986, Map 770). It is almost cosmopolitan, from thearctic to the tropics, although confined to high ground in the latter (New 1961).
Nutrition and Uses
Despite its small size and very narrow leaves, the species is very nutritious and is avidly eaten by farmand other animals, being rich in phosphate, magnesium and sodium (Wilman & Riley 1993). In the past,it has also been cultivated and eaten as food for humans, not only in times of famine. It was part ofthe last meal of Tollund Man (one of the archaeological bog bodies). Corn Spurrey was eaten by humans inShetland in historic times, being ground into meal and used for bread (Grigson 1987).
As grasses can be deficient in some elements for animal production purposes (including N, P, Mg, Na andsometimes, Ca), comparative digestibility trials using sheep were carried out by Derrick et al.(1993) who found that the voluntary intake of S. arvensis measured in terms of weight was 22%higher than that of Perennial Rye-grass (Lolium perenne). Fream (1900) reported S.arvensis being sown for grazing and silage in the past. The live-weight gain of lambs fed onCorn Spurrey has been found to be higher than that on a Rye-grass diet, and a combination ofpalatability, a lower proportion of cell wal and ease of physical breakdown were the significantdigestibility features compared with grass (Derrick et al. 1993). On the other hand, the lowerconcentration of water-soluble carbohydrate in Corn Spurrey and five other dicotyledonous speciescompared with Rye-grass, would put them at a disadvantage from the point of view of silage making(Wilman & Derrick 1994). The overall conclusion of these experimental measurements was that CornSpurrey and other common dicotyledonous grassland weed species such as Common Chickweed (Stellariamedia), Broad-leaved Dock (Rumex obtusifolius), Ribwort Plantain (Plantagolanceolata) and Dandelion (Taraxacum officinale), extend the range of concentration ofsome major elements in herbage beyond that normally found in temperate grassland.
There is little evidence of medicinal use of the herb, although Allen & Hatfield (2004) quote JohnParkinson (1640) who claimed that country folk used the sap of the bruised plant for healing cuts.
Names
First recorded in Great Britain by l'Obel (De l'Obel & Pena 1571), Grigson (1974) follows Lyte (1578)in suggesting the name 'Spurrey' or 'Spurrie' comes from the Dutch 'spurie' (West Frisian, 'sparje'),from which l'Obel (himself Flemish), seems to have coined the Latinised name 'Spergula'. However, thereare alternative explanations of its origin. One interesting suggestion is that spurrey derives from thearrangement of the leaves, "On the stalk are set ... small narrow leaves, waving or bending inmanner of a star or spur rowel of many points" (Britten & Holland 1886). Prior (1870) believedit more likely that Spurrey came from Spergula, and suggested that the latter was a contraction of'Asparagula', a presumed diminutive of 'Asparagus', "a plant which the Spurrey somewhatresembles".
In England, an alternative name for Spurrey was 'Francke' (Lyte 1578), 'Franke Spurrey' (Gerarde 1633),'Franck Spurry' or 'Francking Spurnewort' (Parkinson 1629), meaning a fattening herb for cattle. Thename 'Francke' given by Lyte, is said to be derived from an obsolete word for a fattening pen or sty(Prior 1870; Britten & Holland 1886; Grigson 1974).
In addition to the above, Grigson (1987) lists a total of 17 further local common names for the plant,some of which are regularly applied to alternative species (eg Dodder (Cuscuta spp.)), or whichdenote a vaguely similar appearance (eg Toadflax, referring to Linum catharticum, Fairy -,Mountain - or Purging Flax).
Threats
None.
Native, common and very widespread. Eurosiberian temperate, but widely naturalised including in eastern NAmerica and New Zealand.
1881; Stewart, S.A.; Co Fermanagh.
April to January.
Growth form and preferred habitats
This colourful, attractive perennial grows erect and up to 75 cm tall from a slender, short to fairlylong, rhizome-like, branching rootstock. L. flos-cuculi is a very typical and conspicuous plantof wet meadows and rushy, spring-fed pastures where it can frequently occur in considerable quantity.Ragged-Robin grows well on both mineral and peaty soils, including those of widely differing nutrientstatus and it is especially plentiful, thriving and beautiful where there is some degree of winterflooding. It is also found in marshes, swampy tall-herb fens, ditches, stream-sides and to a lesserextent in the wetter, more open parts of woods. In suitably moist ground, L. flos-cuculireally only avoids the most acidic, most nutrient-starved situations. This is such a well-markedcharacteristic of the species that it can even be seen reflected in the British & Irish distributionat the hectad scale in the New Atlas.
Since wet lakeshore pastures, fens and even fen-carr alder and willow stands in Fermanagh are regularlygrazed by cattle, the plant obviously tolerates herbivory and the associated manuring and tramplingdisturbance.
Variation
A dwarf coastal form, var. congesta, exists in parts of England and Scotland fromKent to Shetland. Stace (2010) transferred Lychnis to the genus Silene and called thisspecies Silene flos-cuculi (L.) Clairv. Sell & Murrell (2018) however, leave it as L.flos-cuculi L.
Fermanagh occurrence
L. flos-cuculi is very widespread throughout Fermanagh, occurring in 317 tetrads, 60% of those inthe VC. It is most frequent in the seasonally flooded, base- and nutrient-rich peaty and mineral soilsaround Upper Lough Erne and by much smaller, more upland lakes on the Western Plateau.
Flowering reproduction
The flowering period is long, stretching from May to August. The narrow, tubular flowers up to 6 mm long,are borne in long-stalked dichasial cymes that come close to forming a flat-topped corymb-likepresentation. The bright, rose-pink petals are deeply cleft in four, making the flower very distinctive.They are protandrous, contain well-concealed nectar and are visited and pollinated by long-tonguedinsects, including bees, butterflies, moths and hoverflies (Proctor & Yeo 1973; Fitter 1987).Although plant size and the number of flowers produced both vary between wide limits, Salisbury (1942,p. 167) reckoned an average sized plant would produce around 40 ovoid capsules, and estimated the meanseed output as 4,500 per plant. This is a relatively high figure for this flowering plant family, andSalisbury reckoned that Ragged-Robin is a rather short-lived perennial which tends to colonise groundwhere competition for light is temporarily diminished. It is thus an opportunistic colonist ofintermittently available habitats, a fact probably linked to the temporarily high water tableexperienced in many of its sites, which also accounts for the high seed production observed.
Seed dispersal and longevity
The seed capsules at the top of the tall, flexible stem, release the seed on the breeze in a typicalswaying censer mechanism (Fitter 1987). Seed are 0.5-0.7 mm across, blackish brown and tubercled(Clapham et al. 1987; Jonsell et al. 2001). Dormant seed is long-persistent, ie survivingmore than five years in the soil (Thompson et al. 1997).
Vegetative reproduction
The plant also reproduces vegetatively by means of offsets, ie basal shoots that produce leaf-rosettes(Salisbury 1942, p. 167). This is especially efficient in wet habitats (Jonsell et al. 2001).
Fossil record
Seeds and pollen have been identified from mild sub-stages of the Hoxnian and Ipswichian interglacials,and from the end of the Late Weichselian through to the Flandrian. The historical pattern is similar tothat of its relative, Silene dioica (Red Campion), and Godwin (1975) says it may therefore beregarded as a long persistent native. However, it differs from S. dioica in showing response tohuman settlement from the Iron Age to the Mediaeval period.
British and Irish occurrence
L. flos-cuculi is a common and widespread species of damp to wet habitats throughout Britain &Ireland at altitudes up to 750 m. Harron (Flora of Lough Neagh) regarded Ragged-Robin as adeclining species around Lough Neagh and the FNEI 3 also mentions its decline in the Magilliganarea of Co Londonderry (H40), marginal to Lough Foyle. We have no evidence of any corresponding lossesin Fermanagh, although there are four tetrads with pre-1975 records only.
The decline of L. flos-cuculi commonly reported elsewhere in local Floras in Britain & Irelandin recent years is presumably caused by drainage and pasture 'improvement' operations (ie ploughing,re-seeding and the spraying of fertilisers and herbicides). However, the extent of agriculturalintensification in Fermanagh has not greatly impinged on its local populations as yet (Crawley 2005; NIVascular Plant Database 2006).
European and world occurrence
L. flos-cuculi is native in Europe and stretches from Iceland to Siberia and the Caucasus. It ispresent in Corsica, Sardinia and Sicily but absent from all other Mediterranean isles (Jalas &Suominen 1986, Map 1015; Hultén & Fries 1986, Map 775). A separate form, subsp. subintegraHayek, is endemic to the Balkan area (Jalas & Suominen 1986, Map 1016).
The beautiful and unusual rose-pink, occasionally white and especially the lovely double flower have beencultivated in gardens, probably for a long period. In this manner, it was introduced to N America aroundthe middle of the 19th century, first appearing in the wild around 1867 in Canada (Cody &Frankton 1971). It is now quite widespread across northern states of the US, coast to coast. It alsoreached New Zealand in 1871 (Webb et al. 1988).
Names
The genus name 'Lychnis' is derived from the Greek 'lychnōs' , meaning 'a lamp', as the hairy leaves of arelated species Coronaria tomentosa were used as lamp wicks (Gilbert-Carter 1964; Stearn 1972).The Latin specific epithet 'flos-cuculi' means 'cuckoo flower' and is a reference to the early floweringof the plant at the season the cuckoo calls (late-May) (Gilbert-Carter 1964).
The plant has numerous English common names, Grigson (1987) listing no less than 23. Many of them includereferences to the early flowering (Cuckoo and Cuckoo-flower), ragged petals (Rag-a-tag, Ragged Jack,Ragged Willie, Ragged Robin and Shaggy Jacks) and Robin (Red Robin, Robin Hood and Rough Robin). Theflowers were used for garlands, but not for food or medicine (Grigson 1987).
Threats
It is not directly threatened, but its moist meadow habitat could be subject to agricultural'improvement'.
Introduction, archaeophyte grain contaminant, now a very rare casual. Native range uncertain due tospread with early arable agriculture, possibly E Mediterranean, but very widely naturalised and nowcircumpolar boreo-temperate and widespread in the southern hemisphere.
1902; Abraham, J.T. & McCullagh, F.J.; Gubbaroe Point, Lower Lough Erne.
Growth form and preferred habitats
This rather handsome, purple/pink-flowered, ancient introduced annual grows erect, 30-100 cm, sparinglybranched from a stout taproot. The narrow leaves and flowering stems are covered with white appressedhairs. The British and Irish distribution suggests it prefers lime-rich loam or light sandy sites, butit can grow in any soil. In earlier days, it was an important, widespread weed of cereals and otherarable crops, especially wheat and rye. As a consequence of this, it often grew on field margins andappeared as a ruderal on disturbed, open, wayside habitats. Nowadays, it sometimes appears on steep,sandy slopes where erosion continually keeps the habitat open (Sinker et al. 1985).
Weed populations of this markedly anthropochorous species declined very rapidly after the introduction ofimproved seed cleaning methods and increased use of herbicides in the 1930-60 period. Cornco*ckle is aweak competitor, not growing well in dense fields of fertilized crops (Jonsell et al. 2001). Inthe last 100 years in Britain & Ireland, it has been reduced to a rare, casual ruderal, confined toopen wayside situations or gardens, where it is increasingly found as a component of sown wild-flowerseed mixtures.
Flowering reproduction
The plant flowers from June to August and has pointed sepals that project appreciably beyond the fivepetals of the 2-5 cm diameter corolla. The flower is protandrous, and while visited by butterflies, itis largely automatically self-pollinated (Clapham et al. 1987; Jonsell et al. 2001). Thecapsule is ovoid and protrudes slightly from the calyx tube. It opens by splitting into five valves. Theseeds are large, oval and flattened, about 3 mm in length. They are brownish-black or black in colourand are covered with wart-like protuberances. Each fruit capsule contains around 25 seeds.
Seed survival
The survey of soil seed banks in NW Europe provided seven estimates for A. githago, six of whichconsidered dormant seed transient and one reckoned it short-term persistent, ie it may survive more thanone year but less than five (Thompson et al. 1997). Thus there is no persistent soil seed bank,germination generally occurring within a few weeks of release from the plant. There are reports,however, that A. githago reappears from time to time when soil is disturbed, and in former arablefields (Sinker et al. 1985). This suggests a small proportion of seed may be long-term persistentin the soil seed bank.
Variation and taxonomy
There is not much variation over its large geographic range. The species has been moved to and from twoalternative genera, Lychnis in the past, and Silene more recently. Jonsell et al.(2001), Stace (2010) and Sell & Murrell (2018) all keep it in Agrostemma L.
Fossil record
Records in Britain & Ireland are almost all of seed and date from the Roman period onwards. Onepollen record, from Old Buckenham, is from levels considered to be Iron Age, Anglo-Saxon and Normanages, along with abundant evidence of arable cultivation. There is no evidence of natural status: theabundant fossil seeds are always associated with a local collection of weeds of cultivation (Godwin1975).
Toxicity
In the past, Cornco*ckle was an important cause of animal and human poisoning. Previous to its decline,when A. githago was a plentiful weed of cereal crops, its seeds used to be threshed out withcereal grains and either sown with them for the next crop, or ground with them for flour. Unfortunately,the Cornco*ckle seed contains high levels of toxic colloidal glycosides with the properties of saponins.These saponins are not inactivated by heat, and bread made from flour contaminated by A. githagoseed can cause poisoning. The flour is discoloured a greyish hue, has a bitter taste and an unpleasantodour (Cooper & Johnson 1998). Although nowadays A. githago is of much rarer occurrence, inPoland there were 30 outbreaks of poisoning by it between 1951 and 1963, with high mortality among pigs,cattle, horses, fowl, geese and ducks (Cooper & Johnson 1998, p. 54).
In humans, a chronic type of poisoning called 'githagism' can occur after consuming saponin-contaminatedflour. This involves weight loss, lassitude and gastrointestinal disturbances, including ulceration andfrothy diarrhoea. Death can occur if the contaminated diet is continued (Cooper & Johnson 1998). Ithas been suggested that the high saponin content of contaminated flour may also have causedsusceptibility to leprosy (Godwin 1975, p. 146).
Uses
Despite the severe toxicity of its irritant saponin poison, according to Grieve (1931) in the past A.githago was used in herbal medicine as a supposed cure for dropsy and jaundice. Its use byallopathic herbalists has been discontinued for obvious reasons, but in hom*oeopathy a trituration of theseeds has been used in treating paralysis and gastritis (Grieve 1931, p. 223). On the other hand, Allen& Hatfield (2004) dismissed as a probable error the very suggestion that A. githago ever hada role in herbal or folk medicine anywhere in the British Isles.
Fermanagh occurrence
There are just two records of A. githago in the Fermanagh Flora Database. The original early findby Abraham and his co-worker McCullagh on the shore of Lower Lough Erne as given above, and secondly ina recent patch of roadside sown with a 'wild flower seed mix' at Drumawill in Enniskillen, on the SligoRoad, which was found by RHN and HJN on 14 September 2003. Cornco*ckle forms a regular component of suchseed mixes and has recently appeared in roadside plantings in other Irish counties, eg within Birr town,Co Offaly in 2017.
Irish occurrence
In the 16th century, A. githago was regarded as a menace to husbandry and one of theworst weeds (Salisbury 1964). It was still a frequent and widespread cornfield weed in Ireland in thelate 19th century, especially in light, sandy soils (Praeger 1901). It was suggested it was"usually introduced with vetches, in which crop it appears to be more abundant than in corn"(Colgan & Scully 1898). The noticeable decline in the population was noted in The Botanist inIreland where the author commented on the distribution in the country being "rarer in thecentre" (Praeger 1934).
A. githago is now considered more or less extinct in the wild in Ireland, but prior to 1950 itstill was a relatively common, or at least a familiar weed of grain fields in all but eight VCs acrossthe island (Cen Cat Fl Ir 2). The distinctly patchy display of old records plotted for Ireland inthe New Atlas suggests it probably was never all that frequent or abundant, apart that is, fromin the major cereal growing areas of the country, in SE of the island and in the Lecale peninsula of CoDown.
Apart from the two Fermanagh records already mentioned, the New Atlas map records just sevenpost-1987 hectads elsewhere in Ireland. The closest stations to Fermanagh were a few plants on disturbedground at Belladrihid Bridge, Co Sligo (H28), found in 1999, and in a disused limestone quarry atAnnahaia, in Co Monaghan (H32), dated post-1994, regarded by the finder, Ian McNeill, as probablydiscarded garden material (Cat Alien Pl Ir).
British occurrence
The New Atlas map shows A. githago surviving scattered throughout most of lowland Britain,but with a greater frequency in areas of limestone geology and the region of the map south of a linebetween Liverpool and Hull. It has been suggested that A. githago has some unknown biological orecological link with the Rye crop, and that the rapid decline of the weed in the late 19thcentury, which preceded the intensification of agriculture, might be due to the move away from plantingRye that definitely occurred (Godwin 1975). Nowadays, A. githago is either a rare casual ofwaysides, or it is found in deliberately sown wild flower seed mixtures in gardens and municipalplantings. In England, it is also maintained in special agricultural Nature Reserves (Firbank 1988).
European and world occurrence
The centre of origin of A. githago is unknown, but it is presumed likely that it arose somewherein the E Mediterranean region, possibly derived from its near relative A. gracile Boiss. In WEurope, Cornco*ckle was one of a number of contaminants of crop seed spread by agriculture in ancienttimes which has in the last 60-100 years declined to rarity or extinction. The European distribution mappublished by Jalas & Suominen (1986, Map 1022) is particularly useful, since it indicates where thespecies is now regarded as extinct, or probably extinct, or not recorded since 1930. This shows A.githago extinct or almost so in Scandinavia, most of Britain & Ireland, N Germany, Holland,Belgium and in N, W & S France, although surviving and widespread in Spain, Portugal, Italy andEuropean areas further east.
Hultén & Fries (1986, Map 779) describe A. githago as indigenous in Eurasia but also note thisrapid decline in distribution. Their map shows A. githago as very widespread throughout C, W& E Europe but thinning northwards into Scandinavia. The map also shows the species widespread in CAsia, where presumably the limit of its native range lies. Elsewhere the species has been spread byarable agriculture to N & S America, the Far East of Asia, including Japan and Korea, C & SAfrica, New Zealand and the Falkland Isles.
Names
The genus name 'Agrostemma' is a combination of two Greek words 'agros', meaning 'field', and 'stĕmma','crown', 'garland' or 'wreath', a reference to the beauty of the flowers − considered suited forgarlands of wild flowers (Johnson & Smith 1946; Gilbert-Carter 1964; Stearn 1982). The Latinspecific epithet 'githago' is an old genus name of the plant and is probably a combination of two words,'git' or 'gith', a name given by Pliny to Nigella sativa (Black Cumin), and '-āgō', a commonsuffix in plant names. The word combination 'githago' is intended to indicate the similarity of theblack seeds of the two plants, both of which were used in herbal medicine (Gilbert-Carter 1964).
A wide range of English common names exist for A. githago, Grigson (1955, 1987) listing 20 fromaround Britain & Ireland. The name 'Corn co*ckle' or 'co*ckle-weed' is thought to date from the18th century, the earlier form being simply 'co*ckle', from the Anglo-Saxon or Old English'Coccel', perhaps connected with co*ck, the bird, but later applied to weeds generally, and especially to'Corn Pink', another common name for A. githago (Prior 1879). There are 21 English common namesgiven by Britten & Holland (1886) for Lychnis githago (one of several synonyms forAgrostemma githago). These include 'Gith', 'Gye' and 'Field Nigella', which recall the Plinyconnection with the black seed mentioned above.
Threats
No threat to A. githago itself, but if the planting of so-called 'wild-flower seed-mixes'containing it continues and becomes more fashionable, this poisonous species could escape cultivation,increase and become a nuisance.
Native, very rare. Eurasian southern-temperate, but naturalised in N America so now circumpolar, alsointroduced in S America, S Australia and New Zealand.
1901; Praeger, R.Ll.; Co Fermanagh.
Growth form and preferred habitats
This is a very variable, long-lived, ± hairless, many-branched perennial arising from a stout, deeplypenetrating taproot. At ground level, the plant produces a number of persistent, short, stout, woodystem bases from which erect or decumbent, flowering shoots are formed. Flowering stems up to 60 cm, bearoval, pointed leaves that vary greatly in length (up to 10 cm) and width, either broad or narrow(0.7-2.5 cm). In some strains, the leaves are slightly waxy or smooth, but others have leaves coveredwith short hairs (Salisbury 1964). The leaves are often blue-green (glaucous) in colour.
Bladder Campion is a lowland plant and weed of the margins of cultivated fields, gravelly hedge banks,roadsides, quarries and other occasionally or regularly disturbed, rough, grassy places. Although itoccurs in a wide range of soils, it shows a preference for moderately fertile, open, calcareous, sandyor gravelly conditions with some humus present. It entirely avoids very wet, strongly acidic, peatysubstrates.
S. vulgaris tolerates partial shade and can occur in woodland glades and edges, or on the banks ofhedges and ditches (Sinker et al. 1985; Jonsell et al. 2001; P.S. Lusby, in: Preston etal. 2002). The range of disturbed wayside habitats indicates a definite anthropochorous linkwith man and his activities, which clearly assists seed dispersal and facilitates colonisation bycreating suitable open habitat for the species.
Flowering reproduction
Flowering takes place from June to August. The flowers on any one plant of S.vulgaris are either unisexual (ie male or female) or perfect (ie hermaphrodite, male andfemale), but occasionally some may bear perfect and unisexual flowers. Male flowers are very rare, buthave been reported in C Europe (Clapham et al. 1987). The flowers are produced on a branchedcymose inflorescence, usually bearing around 30, rarely fewer than five, or up to 50 nodding flowers.The white flowers (rarely pink or greenish) have five free petals, with a tube around 10 mm deep; thelimbs that spread at right angles at the calyx mouth are very deeply bi-lobed. The flowers are also veryvariable in their proportions, some bearing an urn-shaped, inflated sepal tube, while others have a morecylindrical, fused calyx. The number of styles is also variable, usually three, but there can be four orfive on female and perfect bisexual flowers.
Pollination
The flowers produce fragrance and nectar at the base of the stamens only in the evening, and areprincipally pollinated by night-flying moths and flies. The anthers release their pollen at around 20.00hours, and little remains the next morning. A large number of insect species are known to visit S.vulgaris flowers, including solitary bees, flies and moths. The long-tongued bees mop up anyremaining nectar during daylight hours. Night-flying moths of the genus Hadena not only pollinatethe flowers, but also lay their eggs in them, acting as 'parasitic pollinators' (Pettersson 1991).
Fruit and seed
The fruit capsule remains enclosed by the rather inflated calyx. The capsule is subglobose and becomesdry and somewhat leathery as it ages. When ripe, it splits at the top to form six (sometimes eight orten) spreading teeth, and the seeds are shaken out by the wind. The kidney-shaped seeds, 1-1.6 mm, aregreyish or black in colour (white when immature). They are covered with conical tubercles (projections,bumps) that vary considerably in height, ie low, rounded, high, acute, or even like flat plates,although the latter type is rare (Clapham et al. 1987; Jonsell et al. 2001; Stace 2010).Germination occurs in May and June, at least eleven months after seed production, and the germinationsuccess rate can be as high as 90%. The seed is also long-term persistent, a proportion surviving morethan five years burial in the soil (Thompson et al. 1997).
Variation
It is absolutely clear from the content of the previous two sections, that S. vulgaris is anextremely variable species, varying considerably or even widely in almost every character one examines.There is a large body of evidence of this wide-ranging variation from two in-depth studies of S.vulgaris made by Marsden-Jones & Turrill (1957) and Aeschimann (1985). Bladder Campionvaries, for instance, in leaf shape, hairiness and habit. It has been suggested that in Scandinavia thisvariation may be the result of repeated immigration from various geographical sources (Jonsell etal. 2001).
In Britain, hairy-leaved plants appear to be more prevalent on drier, more calcareous soils, suggesting apossible association with a physiological difference. However, in general in would seem that we havehere an example of diverse genetic patterns, all of which have similar survival value, and thusdifferent forms can persist side by side (Salisbury 1964, p. 186). In Scandinavia, on the other hand,local populations are often found to be hom*ogenous with regard to seed wall sculpture (ie high or lowtubercules), yet no correlation between seed form and ecology or geography has ever been discoveredthere (Jonsell et al. 2001).
In Flora Europaea 1 (2nd edition), the variation in S. vulgaris issubdivided into five subspecies and the closely related maritime species, S. uniflora Roth, intoa further four subspecies. Only two of the S. vulgaris subspecies occur in Britain, thewidespread subsp. vulgaris, and the introduced and long established subsp. macrocarpaTurrill, which is tightly restricted to Plymouth Hoe in Devon (Stace 2010).
The variability within the S. vulgaris species aggregate is so great that in the past taxonomistshave struggled to describe, delimit and classify the various subordinate entities. As many as fouralternative genera have been proposed, for the taxon, namely Behen, Behenantha,Cucubalus and Oberna (Jalas & Suominen 1986, p. 55).
Fossil record and Status
Seed of S. vulgaris was "extraordinarily abundant in full Weichselian time where it wasrepeatedly found in geological conditions and faunistic evidence indicative of fresh soils, openhabitats, salinity and severe periglacial climate" (Godwin 1975, p. 143). Seeds have also beenfound from the Wolstonian glacial stage. In contrast, there are no Flandrian fossil records at all (iefrom the current warm interglacial period), except for those associated with Roman times, and from oneMediaeval site at Shrewsbury. The fossil evidence clearly indicates that S. vulgaris was wellable to live through glacial stages, but almost certainly had much more difficulty contending with plantcompetition during interglacial woodland conditions (Godwin 1975).
When one considers the near absence of interglacial fossils for S. vulgaris, its absoluterequirement for open habitats, and its definite link with sites created or strongly influenced by humanactivity (ie artificial and disturbed habitats), it is perfectly possible to conclude that BladderCampion might well represent another archaeophyte to add to our growing list of ancient introducedspecies, now thoroughly naturalised in Britain and Ireland.
Fermanagh occurrence
S. vulgaris has always been a very rare species in Fermanagh. Robert Northridge and the currentwriter, RSF (the joint Botanical Society of Britain & Ireland Recorders for the county) have neverseen it here, and there are just four records in the Fermanagh Flora Database. Additional to the firstrecord listed above, the other three are: roadside at Cranbrooke near Fivemiletown, 1951, MCM & D;Clonmackan quarry, near the border and Clones, 1951, MCM & D; and Tattynuckle Td, near Fivemiletown,1987-99, I. & D. McNeill.
Northern Ireland occurrence
Around Lough Neagh, Harron (1986) regarded S. vulgaris as, "very sparingly distributed,generally rare and a (non-persistent) colonist". In fact he listed just four records there, all ofthem Victorian! Similarly, the FNEI 3 lists a total of 45 records for the three counties, butonly 13 of them are post-1970 and the majority (25 of them) are 19th century or earlier.
A careful examination RSF made of the 91 records for both S. vulgaris and S. vulgaris subsp. vulgaris in the N Ireland Vascular Plant Database coveringall six counties in N Ireland, suggested that, despite an incomplete computerisation of early recordsand allowing for an increased recorder effort in recent decades, S. vulgaris remains scarce, butis not showing major decline in the other five VCs in NI.
British & Irish occurrence
Previously in Ireland, this was a more common and widespread perennial weed of cultivated fields,gravelly hedge banks and roadsides than is the case today (Mackay 1836; Cybele Hibernica 1866).In their day, Colgan & Scully (1898) suggested it was more frequent near the sea, but there probablywas some confusion with S. uniflora Roth. (= S. maritima With.) (Sea Campion), althoughthese authors did distinguish the latter.
S. vulgaris has always been much less frequent towards the N & W in both Britain &Ireland, disappearing completely or never recorded in much of Scotland and N & W Ireland. However,comparison of the two BSBI atlases shows there has been a rather surprising, appreciable decline since1962 (BSBI Atlas 2; New Atlas).
A subsequent 'Local Change' re-survey in Britain of the 1987-8 'Monitoring Scheme' sample hectads andtetrads has shown that while S. vulgaris is not a strict calcicole, it is now frequentonly on calcareous substrates. Furthermore, it appears that grassy field margins where itpreviously grew have now become too nutrient-enriched to provide the open vegetation structure thespecies requires, and plant competition has become too severe for it to survive there (Braithwaite etal. 2006).
The species is strikingly variable, both genetically and in response to environment. Plants are large andvigorous, dormant seed is long-persistent in soil, and when germination occurs, it enjoys a high levelof success. Populations also occupy a wide range of disturbed and artificial habitats, so S.vulgaris is intimately associated with human activities. The observed gradual decline, and itsvery widespread extent, are therefore more than a little unexpected.
European and world occurrence: S. vulgaris is very widespread on the European continent,stretching in an almost continuous manner from the far north (70°N in Scandinavia), to the Mediterraneanislands, and from the western coastline well into the eastern continental heartlands where it appears togradually thin out (Jalas & Suominen 1986, Map 1109). According to Hultén & Fries (1987, Map788) "This polymorphic, partly anthropochorous species is indigenous in Europe and large parts ofAsia." The latter authors do acknowledge that S. vulgaris is widely spread by man into N& S America, S Australia, Tasmania, New Zealand and elsewhere. With its present distribution, thespecies (shown as S. cucubalus Wibel. in Hultén 1974, Map 214) belongs to the circumpolar plants.
Uses
Although we generally think of Bladder Campion as a weed of disturbed places, nevertheless in pastures inpast times it was regarded as a good fodder plant. Young shoots, which appear in March or early April,were previously rated as a vegetable suitable for human consumption (Salisbury 1964). It is listed byMabey (1972, p. 92) in his book Food for free, as a substitute for Chickweed (Stellariamedia), itself now regarded as a substitute for Spinach.
Names
The origin of genus name 'Silene' is obscure (Gilbert-Carter 1964) but might possibly be derivedfrom the Greek 'sialon' meaning 'saliva', referring to the gummy exude from the stem which wards offinsects (Johnson & Smith 1946). Another suggestion is that 'Silene' is Theophrastus' name foranother plant (Viscaria), a different Catchfly (Gledhill 1985; Stearn 1992). The Latin speciesepithet 'vulgaris' simply translates as 'common'.
There are a plethora of English common names, Grigson (1987) listing no less than 33 varied names, somemore fanciful and unique than others. Quite a few names are shared with related species, particularlywith Ragged-Robin (Lychnis flos-cuculi), Red Campion (Silene dioica) and White Campion(Silene latifolia). In addition to 'Bladder Campion', there are variants that include 'Bladder'such as 'Bladder bottle', 'Bladders of lard', 'Bladderweed' and 'Bletherweed'. Other names like 'PopGuns', 'Poppers', 'Poppy', 'Corn pop', 'Snappers' and possibly 'Clapweed' and 'Cow-cracker', refer tothe fact that the inflated calyx can be popped or snapped when young before the contained fruit capsuleripens. 'Rattle-bags', 'Rattleweed' and 'Cow-rattle', refer to the sound of the seed rattling in theshaken ripe capsule. 'Cow-rattle' might also suggest the calyx is shaped like a cowbell. The Scottishname 'Cowmack' refers to a suggestion or belief that the plant helped make the cow want the attentionsof the bull. Six of the names Grigson lists refer to the white, night pollinated flowers, eg 'Whiteco*ckle', 'White Hood' and 'White co*ck Robin' (Grigson 1987, p. 81-2).
Threats
Cultural eutrophication, caused by nitrogen and phosphate slurry and fertiliser drainage runoff fromfarmland, has increased the competition S. vulgaris faces from more vigorous plant species,leading to its displacement.
Introduction, archaeophyte, extremely rare and almost certainly extinct. Eurosiberian southern-temperate,but widely naturalised and now circumpolar, also introduced in Tasmania and New Zealand.
1947; MCM & D; rubbish tip near Enniskillen.
Growth form and preferred habitats
In good growing conditions this conspicuous, short-lived wintergreen perennial, or occasionally annual orbiennial develops a taproot more than 40 cm deep and flowering branches up to 100 cm tall. It is notcompletely frost hardy in Britain & Ireland (Salisbury 1964), but it is extremely resistant todrought (Baker 1947). Like S. vulgaris (Bladder Campion), it develops a thick, almost woodyrootstock from which a number of short, non-flowering shoots and taller, erect, robust, flowering stemsarise. Shoots are covered with soft hairs, long ones below, short and mostly glandular, but not stickyabove (Baker 1947).
It is a weed of arable fields and their margins that also occurs on other man-made, dry, well-drained, ±exposed ruderal habitats, including roadside verges, hedge banks, disturbed ground in waste places andthe base of walls. It is especially associated with harbours and ports. S. latifolia toleratesdry, light, sandy soil conditions, but thrives best in deep, well-drained, near-neutral pH, lowlandcalcareous substrates, in sunny, open situations (Salisbury 1964). Nowadays in many sites it typicallyoccurs solitary, or as a few scattered individuals, and it only rarely forms small patches (Baker 1947;Clapham et al. 1987; Jonsell et al. 2001).
Cattle, horses and sheep graze the plant, but rabbits do not. Plants usually regenerate rapidly afterbrowsing. S. latifolia stems are easily broken by trampling at the flowering stage, but leafrosettes survive intermittent trampling (Baker 1947).
Flowering and vegetative reproduction
White Campion is a long-day plant, and has a prolonged flowering season. Flowering extends from April orMay in the south, until October, or November in sheltered situations (Baker 1947). Peak flowering isfrom June to August. The inflorescence of S. latifolia is lax with spreading branches.Technically, it is a dichasial cyme, but with monochasial upper parts! The species is dioecious, havingseparate male and female flowers. Male flowers bloom earlier than female, and they also persist later.Unlike S. vulgaris (Bladder Campion), perfect, bisexual flowers are very rare in S.latifolia. The flowers are large, about 30 mm across. Female flowers have an urn-shaped orsubglobose and inflated calyx, while in male flowers the calyx is campanulate or ± cylindrical. The maleplants bear more numerous, slightly nodding flowers than females. In both sexes, the calyx is coveredwith downy, slightly sticky hairs.
The white petals are deeply two-lobed, and flowers release a slight perfume and provide nectar at nightto attract noctuid moths as pollinators (Jonsell et al. 2001). Crawling insects are excluded by aconstriction at the mouth of the corolla tube. As in S. vulgaris, long-tongued bees visit indaytime, collecting any remaining nectar, and they may help pollinate the flowers. Short-tongued beesregularly steal nectar by making a hole near the base of the flower tube (Baker 1947). Some noctuidmoths of the genus Hadena are serious parasites, laying eggs in the female flowers and causingsevere reduction in seed production in affected flowers (Baker 1947; Pettersson 1991).
The fruit capsule is cylindrical to broadly ovate and it becomes leathery as it ages. In summer, thecapsule ripens around four to five weeks after flowering. It splits open, forming ten slightly spreadingteeth. The dry seeds are shaken out by wind, the capsule behaving like a censer. In wet weather, thecapsule teeth tend to bend inwards, keeping the seed dry and preventing them from getting wet andsticking together (Baker 1947). The typical plant produces between 16-20 fruits, but a large plant couldset close to 100 capsules, each containing between 50-400 seeds (Baker 1947; Salisbury 1964).
The fact that S. latifolia, like some plants of S. vulgaris, bears unisexual flowers only,very probably reduces the amount of seed set when the plant is scarce in a geographical area.Inevitably, low seed production will lead to, or hasten, local extinction of the species. This isparticularly the case with a plant like White Campion that has no specialised structures adapted forvegetative reproduction. Having said this, partially uprooted plants of S. latifolia, eg as aresult of ploughing, are able to produce elongated leafy stolons that can re-establish the plant inunder a fortnight given suitable growing conditions (Baker 1947). Only the buds on the root crown permitvegetative regeneration (McNeill 1977), and in the normal habitat situation, vegetative reproduction ofany description is too slow for anything more than establishment of existing colonists (Baker 1947).
Seed production
Seed set is generally good and in England, an average sized female plant of S. latifolia willoften produce around 5,000 seeds. Larger plants can manage up to perhaps three times this figure(Salisbury 1964). Seed output estimates in N America vary widely, but at best they are more than doubleSalisbury's estimate (McNeill 1977). Seed are grey or black, bluntly tuberculate and about 1.5 mm long(Baker 1947). Seed shed in early summer can germinate the same season, or they may do so the followingspring, or later.
Seed survival
As with many other members of this family, S. latifolia seed can survive dormant in undisturbedsoil for up to 20 years (Lewis 1973). Despite this fecundity, S. latifolia is essentially aspecies of sunny conditions, well adapted to dry soils and is extremely drought resistant. It does notcope well with damp, humid conditions, however, since attack by Fusarium species cause seeds torot and seedlings to damp off (Baker 1947). This alone, or in combination with other factors, includinga marked decline in arable agriculture and improved seed hygiene, helps explain why White Campion is soseldom met in northern and western parts of these islands.
Variation and taxonomy
There is a cline in variation from E to W in Europe with respect to a range of characters, including seedornamentation (Jonsell et al. 2001). Variation within the Silene vulgaris aggregate and inrelated species including S. latifolia has occasioned numerous changes in classification andnomenclature over the years. Silene latifolia has been in two other genera, namely Lychnisand Melandrium.
Fossil record and centre of origin
The fossil record in Britain stretches back only to the Early Bronze Age (Godwin 1975). The species mostprobably arose from an ancestor more closely resembling S. dioica (Red Campion) byeco-geographical divergence. It has dispersed from a centre of origin somewhere in the Middle East, theMediterranean and C Europe, where it grows in dry, sunny, natural habitats. S. latifolia verylikely arrived in Britain & Ireland along with Neolithic arable farmers, and probably it was thenmost common in S England (Baker 1947; Godwin 1975; Clapham et al. 1987).
Fermanagh occurrence
White Campion has only been seen once in Fermanagh – at the municipal rubbish tip just outsideEnniskillen by Meikle and his co-workers in 1947.
Irish occurrence
The Cen Cat Fl Ir 2 lists S. latifolia (as S. alba) occurring in 28 of the 40 IrishVCs, and for quite some time this weed of cultivated land, or casual of waste ground, dockyards androadsides has been regarded as a probable introduction in Ireland. In her Cat Alien Pl Ir,Reynolds reckoned White Campion is fairly common across the island in disturbed and sandy ground,eg on roadsides and especially near ports. She points out that Praeger and others in the late19th century realised that it was being occasionally imported with foreign grain.
Reynolds (Cat Alien Pl Ir) may be over-estimating the Irish presence of S. latifolia, atleast in a N Ireland (NI) context, since it is really only a casual colonist here (Flora of LoughNeagh; FNEI 3). Apart from the solitary Fermanagh record the NI Vascular Plant Databasecontains 40 records for the other five VCs in NI. Almost exclusively, these date from 1976 onwards, sowe assume earlier records have been neglected. Computerisation of early NI plant records is not yetcomplete. The Flora of Co Tyrone lists eleven recent records, all of them post-1981(McNeill 2010). Nevertheless, despite the incomplete NI picture the number of records does suggest thatseed of S. latifolia is still being occasionally imported, and the plant appears sporadically forshort periods in suitably open ground.
British status and occurrence
Previously in Britain, botanists traditionally assumed S. latifoia was an indigenous species.Stace (1997) vainly clung to this belief, long after evidence otherwise was clearly presented byBaker (1947). Received opinion has been recently revised (Preston et al. 2004), and S.latifolia is now recognised as an early introduction (ie an archaeophyte) in both Britain &Ireland. White Campion has always been much more scarce and local in Ireland in comparison with Britain.It remains fairly common and widespread in Britain today, although with a definite eastern tendency inits distribution, and the distribution becomes more coastal northwards (Preston et al. 2002). TheNew Atlas map suggests there have been losses of this species at the western margins of itsBritish & Irish distribution.
Locally in Fermanagh, we can attribute loss of weeds like this to the decline in arable agriculture tonear absence. Elsewhere in Britain & Ireland, the decline in arable weeds is certainly due tochanges in agricultural practices, including the use of cleaner crop seed, widespread spraying ofherbicides and the use of artificial fertilisers and slurry to increase crop productivity. There havebeen reports, however, that S. latifolia is resistant to commonly used herbicides, such as 2,4-Dand 2,4,5-T (Salisbury 1964; McNeill 1977).
The findings of the recent 'Local Change' re-survey of BSBI 'Monitoring Scheme' hexads in Britain,emphasises the fact that S. latifolia is struggling to maintain its distribution, especially inthe more pastoral west of the island (Braithwaite et al. 2006).
European and world occurrence
S. latifolia is widespread as a native weed throughout Europe northwards to 68°N in Scandinavia,ranging east to Mongolia and SE Siberia and south to N Africa (Jalas & Suominen 1986, Map 1183). Italso occurs as a neophyte or archaeophyte introduction in more northern areas of Scandinavia, Iceland,Greenland, Ireland, Great Britain, parts of India, Australia, Tasmania and New Zealand. The presentdistribution is amphi-atlantic and almost circumpolar (Hultén & Fries 1986, Map 793). Preston &Hill (1997) classified it as Eurosiberian southern-temperate, but widely introduced. In Scandinavia, itis believed to have been introduced in the 19th century along with ley seed and cereals, andperhaps also with ship ballast and along with foreign troops (Jonsell et al. 2001).
In North America, White Campion was introduced from Europe sometime in the early 19th centuryprobably as a crop seed contaminant. In Canada, it remains an important weed of field crops such asalfalfa (Medicago sativa), clover (Trifolium spp.) and small grain and legume grain-foragerotation crops. In 1965, S. latifolia was reported as one of the five worst weeds in pastures andhayfields in six states from New Hampshire and Connecticut to Minnesota, and at that time infestationwas still increasing (McNeill 1977).
Names
The origin of genus name 'Silene' is obscure (Gilbert-Carter 1964) but might possibly be derivedfrom the Greek 'sialon' meaning 'saliva', referring to the gummy exude from the stem which wards offinsects (Johnson & Smith 1946). Another suggestion is that 'Silene' is Theophrastus' name foranother plant (Viscaria), a different Catchfly (Gledhill 1985; Stearn 1992). The Latin speciesepithet 'latifolia' translates as 'broad-leaved'.
Threats
In Britain & Ireland, cleaner crop seed, the increased use of herbicides and a gradual shift awayfrom arable agriculture are probably jointly responsible for the decline of this species.
Silene dioica (L.) Clairv., Red Campion
European Boreo-temperate
1953; MCMD (1957, Typescript Flora); Belcoo, Lough Macnean
Throughout the year.
Growth form and preferred habitats
A very attractive and conspicuous plant when bearing its red or pink, few-flowered cymes, this fairlyshort-lived perennial is locally uncommon, occasional and rarely abundant in semi-shaded or more open,north-facing ecological conditions, mainly in a variety of semi-natural, sometimes only intermittentlyavailable habitats (Salisbury 1942). The softly pubescent, broad, basal, wintergreen leaves are quitedistinctive and the species can therefore be easily recognised throughout the year.
S. dioica is most regularly found in sheltered, moist, open situations at the base of cliffs, orforming loose patches in woodland clearings, or along paths within and on the margins of woods, or onhedged river banks, stream sides or ditches, or in tall-herb roadside banks and verges. S. dioicarequires damper soils than S. latifolia (White Campion) does, and in comparison it is much less,or not at all, drought resistant.
While Red Campion tolerates a broad range of substrate conditions, it prefers fertile, nitrogen-rich,moderately acid to neutral soils in sites where the vigour of competing species is limited either byshade, or by occasional disturbance (eg slippage on unstable slopes or temporary flooding). It tends tobe less prevalent on lime or base-rich soils, except when these are in more open, rocky, steep sites, orin deep, narrow limestone crevices (called grykes), that protect the plant from grazing pressure bytheir sheer inaccessibility (Clapham et al. 1987; Grime et al. 1988). Like the closelyrelated S. latifolia, Red Campion is not fully frost hardy (although rarely killed by frost inBritain & Ireland), and it also avoids waterlogged and strongly acidic substrates (Baker 1947; Grimeet al. 1988).
S. dioica has a taproot that grows horizontally near the surface of the soil for a distance beforeturning down vertically, but it never reaches the 40 cm soil depths that S. alba roots achieve.Red Campion performs best in light shade and it fails to flower and seed when growing under very heavyleaf canopy. Being essentially a woodland mesophyte, it cannot compete in very exposed, windy conditions(Baker 1947).
In the Sheffield area, Grime et al. (1988) concluded that S. dioica showed a,"pronounced inability to exploit conditions of extreme stress or disturbance", and the lifestrategy of the established plant was described as 'C-S-R', ie intermediate between the primarystrategies of Competitor, Stress-tolerator and Ruderal. Species of similar ecological behaviour in thisrespect include Hypochaeris radicata (Cat's-ear) and Rumex acetosa (Common Sorrel).
Variation
Genetically, this is a very variable species both within and between populations. The range of plantvariation is related to both geographic location and habitat environment and several named varietiesexist (Jonsell et al. 2001). Thick-leaved coastal populations in Shetland were once consideredsufficiently different to constitute a separate variety (var. zetlandicum Compton), or even asubspecies. Dwarf variants occur on mountains, exposed rocks and sand dunes (Tutin et al. 1993).However, subsequent studies in Britain and Europe concluded that S. dioica variation consists oflocal, weakly differentiated races that intergrade at their geographic margins, and thus there is nopoint in recognising the variation taxonomically (Prentice 1980).
Variation within the Silene vulgaris aggregate and in related species including S.latifolia and S. dioica has occasioned numerous changes in classification andnomenclature over the years. Silene dioica has been in two other genera, namely Lychnisand Melandrium.
Hybrids
A fully fertile hybrid with S. latifolia occurs wherever they overlap.
Fermanagh occurrence
For some reason, probably explained by the aforementioned species preferences, locally S. dioicais most frequently met in Fermanagh around the Boho region in the mid-SW of the vice-county, but it isrecorded in a total of 44 of our tetrads, thinly scattered around the wooded shore areas of Lower LoughErne, and even more widely and sparsely elsewhere in the county.
Reproduction, vegetative and flowering
Red Campion tends to have a loose rhizome system or branching secondary roots near the soil surface,which allows a minor degree of vegetative spread (Dalpra 1965). Apart from this, S. dioicareproduction and dispersal is entirely achieved by seed. As the scientific name indicates, Red Campionis a dioecious species, and in this instance, the frequency of male plants to some extent predominatesover females. Male plants produce more leaves and flowers than females (Cox 1981; Kay & Stevens1986). Although germination and growth can be quite rapid in a suitable environment, up to 80% of plantsdo not flower until their second season of growth (Baker 1947).
The number of flowers per plant is seldom large, many bearing less than 20. The flowering season runsfrom April to September, or until the first frost. In females, the calyx is urn-shaped and inflated,becoming rounded in fruit, while in males it is more cylindrical or campanulate. Petals are red orbright pink, rarely white and the flowers are (18)20-25(30) mm in diameter. The flowers are open indaytime, and while they produce no perfume, they attract mainly bumble-bees and butterflies aspollinators. The female flowers are subject to the same attacks as S. latifolia by egg-layingmoths of the genus Harmodia. The insect larvae parasitize the developing ovules, causing a majorreduction in seed production (Baker 1947).
Seed production, germination and seed predation
Seed production is extremely variable, being dependent upon local environment and individual plant sizeand vigour, a phenotypic reflection of the former factor. The number of seed per capsule varies from10-233 (Baker 1947), while Salisbury (1942) found the range in his samples in S England stretched from41-288, with a mean of 215 seeds per capsule. The distance of seed dispersal by wind (censer mechanism)is rarely more than 2.5 m, and germination takes place in any month except August, but it chiefly occursin spring (Knight 1978 a, b).
Fungal pathogens, such as Smut, Rust and Damping-off, limit both flowering success and seedlingestablishment. In a damp, humid oceanic climate like ours, these combined stresses are so forceful, theyeffectively tip the balance against S. dioica survival when indigenous populations are low tobegin with (Baker 1947).
Dormant seed survival
Dormant seed persistence in the soil is also extremely variable. The survey of seed survival in NWEuropean soils lists 21 estimates: seven transient, five short-term persistent, two long-term, and sevenuncommitted in this respect (Thompson et al. 1997).
Fossil history
Seed of S. dioica have been found in early glacial and interglacial periods and in all zones ofthe Late Weichselian, and many of the current Flandrian interglacial. The presence of this species intwo glacial and two interglacial stages contrasts strongly with the absence of S. latifolia. Thepresent day ecology of S. dioica and its distribution so far north in Europe, suggests it couldbe a periglacial survivor, having perhaps been present continuously in or near these isles (Godwin1975).
Irish occurrence
The New Atlas hexad map indicates that S. dioica is considerably more widely (and perhapsmore frequently) recorded in Northern Ireland than in the Republic of Ireland, where it is only verythinly and widely scattered (Preston et al. 2002). However, even within N Ireland, Red Campion isnot evenly widespread, but rather it is absent from much of Cos Armagh and Down (H37 & H38), adistribution feature which yet remains to be explained (NI Flora Website 2006).
British occurrence
S. dioica is locally abundant throughout almost all of lowland Britain, but rare in the dry sandyBreckland of East Anglia and the intensively farmed area of Cambridgeshire and its immediate surrounds.
Overall, S. dioica has a very curious distribution in Britain and Ireland, the New Atlashectad map looking at first glance, as if the species had a decidedly United Kingdom (UK) flavour to it.In many of the 34 vice-counties in the Republic of Ireland, no resident member of the BSBI recordingcommunity exists and the designated BSBI Recorder in numerous VCs may live a hundred or more kms outsideits boundaries. The number of recording hours and coverage they can achieve is hugely reduced incomparison to British VCs. Thus the representation of species distribution we see in the publishedNew Atlas maps is necessarily an artefact, since the data over the whole area involved are notuniform in quality and are not comparing like with like.
The Balkans as a possible species origin
The centre of origin of S. dioica remains unknown, but it most probably lies in the BalkanPeninsula, or nearby. A study of S. dioica and its close relatives, S. latifolia andMelandrium nemorale, suggests that the latter, a woodland plant of Rumania, Bulgaria and Greece,has characters very similar to those of both S. dioica and S. latifolia. M.nemorale may resemble the supposed or expected ancestor of all three species. The geographicaldistribution of M. nemorale also appears to represent the point of divergence of these threespecies and of all of their recognised subspecies (Dalpra 1965). The Balkan region represents the areawhere the C European and Mediterranean climatic regimes meet. Its mountains and valleys provide anenvironment where climate varies greatly over a relative small area. Species mingle here in unusualproximity, and ecotypes can readily arise and diverge.
The Balkans is also one of several southern refugia areas to which European plant life retreated duringrecent ice ages/glacial stages. Another important factor, is that change is particularly encouraged whenman disturbs vegetation and creates new, artificial habitats, some of them associated with theintroduction of tillage and grazing agriculture. Agriculture has favoured the spread of S.latifolia, which thrives as a crop weed, while the destruction of woodland to create fields hasbeen to the detriment of S. dioica populations. S. dioica has fewer weedy characteristics,and appears much less able to compete with cereal roots than S. latifolia (Dalpra 1965).
European and world occurrence
S. dioica occurs throughout most of Europe northwards to the Faeroes and Spitsbergen, but it isabsent from parts of the Mediterranean, including all of the islands. Although indigenous in Europe, thespecies is very closely associated with man (ie it is anthropochorous). In many lowland areas ofScandinavia, it occurs entirely in man-made habitats and is considered a recent immigrant, imported withley and hay seed. This is especially so in Finland, where S. dioica is considered native only incoastal sites (Jalas & Suominen 1986, Map 1189). Further north in Scandinavia, S. dioicabecomes rare and casual (Jonsell et al. 2001). The distribution extends eastwards to C Asia asfar as Altai, and it is also present in N Africa and Greenland. In Iceland, S. dioica is anestablished garden escape. It is introduced in eastern N America and in New Zealand where it isrecognised as a garden escape (Hultén & Fries 1986, Map 795; Webb et al. 1988, p. 499).
Uses
The flowers are brightly coloured and the plant very easily cultivated making it a popular decorativegarden subject. As many as eight garden varieties are listed by Griffiths (1994).
Names
The origin of genus name 'Silene' is obscure (Gilbert-Carter 1964) but might possibly be derivedfrom the Greek 'sialon' meaning 'saliva', referring to the gummy exude from the stem which wards offinsects (Johnson & Smith 1946). Another suggestion is that 'Silene' is Theophrastus' name foranother plant (Viscaria), a different Catchfly (Gledhill 1985; Stearn 1992). The Latinisedspecific epithet 'dioica' in Greek means 'two houses', or 'double houses', a reference to the separateplants, each with flowers of one sex only.
'Red Campion' has a very long list of English common names, Grigson (1987) listing no fewer than 63variants. The names have snake, devil, goblin and 'death if picked' folklore connotations. A number ofthe names are shared with two other May-flowering, woodland species, Bluebells (Hyacinthoidesnon-scripta) and the Early-purple Orchid (Orchis mascula), eg 'Cuckoo-flower'. 'Campion',itself is a 16th century Elizabethan book name that has never been convincingly explained. Itremains a bit of a mystery. It first appeared in 1576 and was soon taken up by poets as well asbotanists. 'Campion' may be equal to, or be another version of 'champion', ie a champion flower of thesummer garden (Grigson 1974).
Threats
None.
Introduction, archaeophyte, a rare or very scarce garden escape or discard.
1884; Barrington, R.M.; beside the avenue of Crom Castle, near Lough Nalughoge.
April to December.
Growth form and preferred habitats
A rarely met decorative garden escape or throw out, usually found in small colonies, this attractive,moderately tall, robust perennial has the potential to spread vegetatively very efficiently by the dualmeans of a creeping rhizome and long stolons. If it manages to establish in the wild, Soapwort canbecome very long-persistent. It tends to grow in occasionally mown grassy vegetation on the moderatelyfertile soils of roadside verges and banks, or in rather drier, barer, more open ground, on walls, inquarries or on waste ground. It is very rarely met far from houses, and in some sites it may be theresult of fly-tipping of garden waste.
Reproduction
The large red, pink or white flowers, produced from July to September, are self-compatible butnevertheless strongly protandrous, favouring cross-pollination. An intense, sweet perfume is emitted inthe evening attracting night-flying insects. A study in C Europe found the frequency of night-flyingvisitors was surprisingly low, whereas bumble-bees and syrphid hoverflies carried out pollination duringdaylight hours. Seed set is apparently rather poor (Jürgens et al. 1996). However, vegetativereproduction by runners is efficient and helps establish colonies. Dispersal is most probably achievedby transport of rhizome fragments in soil and garden refuse.
Variation
Previously, this species was included in the genus Silene and referred to as Silenesaponaria Fries.
Fermanagh occurrence
In Fermanagh, there are 13 records from ten tetrads, mainly situated around the shores of both parts ofLough Erne, a fact undoubtedly reflecting the human settlement pattern. Most of the Fermanagh Soapwortshave 'double' flowers lacking functional stamens, the anthers being replaced by small petaloid lobes.
Apart from the first record above, the local record details are: roadside, near a derelict house,Woodhill, 1 km N of Lower Lough Erne, August 1982, RHN; on wall at entrance to Castle Hume estate, nearCoagh Quarry, 4 km N of Enniskillen, August 1982, RHN; Crossmurrin NR, Marlbank Loop, 1 December 1989,RHN; Coolnamarrow Lough, Mount Darby Td, 22 June 1990, RHN; disused quarry, S of loughshore jetty nearKnockninny, 25 August 1995, RHN & HJN; roadside, N of Blenalung Bay, Lower Lough Erne, 20 April1995, RHN; Culliondoo, 5 km S of Lisnaskea, 19 April 1996, RHN; Bigwood Td, W of Boa Island, Lower LoughErne, 13 August 1996, RHN, RSF & D. Cotton; Bosallagh Bridge, 2.5 km E of Mount Darby, 6 September1997, RHN & RSF; roadside, near Drumbad House, S shore Lower Lough Erne, 15 September 2002, RHN;Knockninny Quarry, August 2004, RHN.
Irish occurrence
S. officinalis is an archaeophyte and is thinly but widely scattered throughout Ireland. TheIrish Census Catalogue has it recorded from 38 of the 40 Irish VCs, the missing two being E Mayo(H26) and Monaghan (H32) (Scannell & Synnott 1987). The New Atlas hectad map shows that theplant has now been recorded in Co Monaghan, but there is no symbol indicating past presence in anotherVC, namely W Galway (H16) (Preston et al. 2002).
British occurrence
The same New Atlas map shows that Soapwort is very much more prevalent in the southern half ofBritain, becoming scarce and rather coastal northwards into Scotland. Previously, it was regarded aspossibly native in parts of Britain (Devon, Cornwall and N Wales), but now it is recognised as anestablished alien archaeophyte throughout (Clapham et al. 1987; Preston et al. 2004).
European and world occurrence
This widely cultivated species is claimed to be indigenous in C & S Europe, but in C Europe thenative and secondary introduced ranges are hardly separable due to widespread garden planting. It is notnative in most of Denmark and all the more northerly countries of Scandinavia, although it reaches 55°N(Jalas & Suominen 1986, Map 1312). The natural habitat of the species is on streamsides and dampwoods on alluvial soils (Clapham et al. 1987). S. officinalis is also a widespreadintroduced species in N America and occurs also in S America (Hultén & Fries 1986, Map 802).
Uses
Formerly cultivated as a drug, for making a soap-like lather for washing fabrics, and still in use as anornamental garden plant.
The dried root is the important commercial part of the plant, being a good source of saponin, resin, gum,mucilage and fibre. In herbal medicine, a decoction was used to cure an itch. It was also used to treatjaundice and other visceral obstructions (the 'wood' of the root being a pale yellow colour, this wasanother example of the 'doctrine of signs' herbalists believed in). Soapwort was also regarded as a goodcure for 'old venereal complaints, especially where mercury has failed'. It was also regarded as atonic, diaphoretic, and a valuable remedy for rheumatism or cutaneous troubles resulting from any formof syphilis. Grieve (1931, p. 748), who provides this information, warned that Soapwort should be verycautiously used owing to its saponin content.
As a detergent, rubbing a leaf between the fingers will produce a slight slippery froth. Boiled in water,the plant produces a green lather that previously was used to lift grease and dirt, especially fromfabrics and from animal wool. The presence of saponins provides the desired detergent effect which, likeinorganic soap, has the power to lubricate and absorb oil and dirt particles. Since vegetable saponinsare much gentler than soaps, they have recently been used for washing ancient delicate tapestries (Mabey1997).
As a decorative garden plant, S. officinalis has at least four named garden cultivars, featuringdouble flowered forms and also a variegated plant (Griffiths 1994).
Names
The genus name 'Saponaria' is from the Latin 'sāpo', 'ōnis', meaning 'soap', from the laundry propertiesof the contained, frothy saponin (Gilbert-Carter 1964). The Latin specific epithet 'officinalis' means'of the shop', referring to the druggist's shop where it would be available for purchase.
Apart from 'Soapwort' or 'Soap-wort', which refer to the saponin lather that is mentioned above under'Uses', there are a number of other English common names to examine. 'Fullers' Herb' (HerbaFullonum) and 'Fuller's Grasse' are names that also refer to the removing of stains andimpurities from woollen cloth and the thickening of it. 'Fulling', also known as 'tucking' and 'walking'(or 'waulking' in Scotland), was a trade that goes back to ancient times. Other laundry-associated namesinclude 'Latherwort', 'Scourwort', 'Buryt', 'Borith' or 'Borit', and 'Saponary', the last four obtainedfrom the early English herbal with the highest reputation, The Grete Herball of 1526 (Arber1938). Two other names listed for this species in the same herbal were 'Crowsope' and 'Herbe Phylyp'(Philip?), but the derivation, meaning and association appear unknown (Grigson 1987).
'Bruisewort' is also associated with the plant, a name it shares with Bellis perennis (Daisy), fortheir supposed efficacy in treating bruises (Prior 1879). 'Mock Gilliflower' suggests the herb may havebeen used as a cheap substitute for clove or Dianthus (Pink), its spicy odour being used toflavour wine. 'Gill-run-by-the-street' seems to be a rather peculiar local derivative of this name.'Hedge Pink' might refer to the same idea of use in wine-flavouring, or to one of its habitats, and'Farewell Summer' to the plants' ability to flower in August and September (Britten & Holland 1886).
Soapwort was taken to New England by early settlers who valued it for treating the skin rash caused bytouching Toxicodendron radicans (Poison Ivy), a climbing member of the cashew and pistachiofamily, the Anacardiaceae. S. officinalis is naturalised in the USA and is known there by twoimported English west country names, 'Bouncing Bett' and 'Lady-by-the-gate' (Grigson 1987). 'BouncingBett' might refer to the irrepressible vigour of the plant in the garden setting, which tends to lead toits wayside dumping. This name is also applied to Centranthus ruber (Red Valerian), which alsoshows plenty of vigour, often taking possession of old walls (Grigson 1974).
Threats
None.
Introduced, neophyte, a rare garden escape.
12 August 1996; RHN, RSF & Cotton, D.; in a small quarry at Mullanacross, 5 km E of Garrison.
March to October.
Growth form and preferred habitats
This is a vigorous, colonising, rhizomatous garden perennial, native of the Himalaya and W China. P.campanulata is valued by the gardener for its long decorative display of many, small, clustered,dense panicles of usually pink, rarely red or white bell-flowers from midsummer until well into October.Leaves are 2-4 cm wide and have a dense, pinkish-brown tomentum of hairs beneath. Lesser Knotweed likesmoist or cool soil, and in the garden it looks best when grown in part-shade (Thomas 2004).
P. campanulata was introduced to gardens in Britain and Ireland around 1909 and was first reportedin the wild in 1933, since which time it has slowly but continually expanded across these isles. Thespecies can tolerate a wide variety of growing conditions however when found discarded on roadsides in WIreland. It appears to thrive in the suitably wet, cool soils provided by our mild oceanic climate.
Much of the increase of P. campanulata in Britain and Ireland, which has been very well describedby Conolly (1977), has been in scattered W & SW coastal areas of both islands, rather than in inlandor east coast regions. This distribution suggests the species rather surprisingly might not be all thatfrost hardy. The New Atlas hectad map supports this notion to some extent, but the true positionis not clear since a minority of inland records do exist both in the English Midlands, and in theclimatically more continental SE region of the country. It is possible that the represented stations liewithin urban areas that provide higher night-time temperatures, but we do not know if this is the caseor not.
Reproduction
The small, pink or red, bell-shaped (campanulate) flowers are heterostylous, with long- and short-styledflowers borne on separate plants. This feature may help account for the surprising lack of reports ofseed-set in these islands (Lousley & Kent 1981). Lesser Knotweed, which despite its English commonname is often tall, some annual stems reaching up to 60-90 cm in height. Plants spread and maintainthemselves by means of extensive prostrate stems and both above-ground stolons and buried rhizomes. Inwinter they die down to just a few basal leaves.
Taxonomy and nomenclature
Classification of this family is such that many members have changed genus and species names time aftertime. Synonyms for this particular species include: Aconogonum campanulatum (Hook. f.) H. Hara;A. lichiangense (W. Smith) Soják; Polygonum campanulatum Hook. f. and Polygonumcampanulatum var. lichiangense (W. Smith) Stewart.
There are several garden cultivars of this plant in use. The darker rose-pink flowered form is called'Rosenrot' or 'Roseum', and there are white-flowered forms called 'Album' and 'Southcombe White'(Griffiths 1994).
Fermanagh occurrence
P. campanulata is rather rare in Fermanagh, there being just eleven records, all post-dating 1995.It was first discovered, dumped with garden rubbish, in a small roadside quarry in 1996 as listed above,and since then ten additional finds, all involving RHN, have been made in seven further tetrads,scattered mainly in the northern half of the county. The plant is generally found in damp ground onroadsides, rarely far from houses.
Apart from the first record above, the local record details are: roadside at Gubbakip Td, 13 August 1996,RHN & RSF; opposite abandoned cottage, Mountdrum Td, 19 October 1997, RHN; Clonelly, NW of Kesh, 17March 1999, RHN & HJN; gateway at Lough Skale, 22 September 2000, RHN; roadside, E end of BoaIsland, 6 September 2001, RHN; roadside, Castle Caldwell, 7 September 2002, RHN & HJN; roadsideLackboy, 7 September 2002, RHN & HJN; Coolbuck Td, 11 September 2002, RHN & HJN; gateway, 100 mN of Slisgarrow Td, 16 August 2006, RHN; between road and shore, Bleenalung Bay, Lower Lough Erne, 4October 2010, RHN & HJN.
British and Irish occurrence
In England & Wales, Lousley & Kent (1981) regarded P. campanulata as only semi-naturalisedin wet places at or near the point of planted introduction. They contrasted this with the behaviour ofthe species in Scotland & Ireland, where they considered it fully naturalised in remote areas ofboth countries. The New Atlas hectad map shows the species is only thinly and very widelyscattered across both Britain and Ireland, although it covers the whole range of latitude from Jersey toShetland. The New Atlas map also indicates the quite marked westerly trend in distribution foundin both Britain and Ireland.
Apart from the expected 'garden escapes', in Ireland, P. campanulata is sometimes recorded as arelic of previous cultivation in landed demesnes and parklands, eg Castleward in Co Down (H38) andCastle Dobbs in Co Antrim (H39) (Cat Alien Pl Ir). Some of the stands in ground remote fromhabitation may have arisen as the result of fly-tipping of garden refuse and outcasts, a reprehensibleand all too common practice nowadays throughout these islands. The most extensive naturalised stand ofthe plant, reported by Conolly (1977), was along the bank of the River Camp in south Co Kerry (H1),where it stretched for at least 1 km. It appears unusual for P. campanulata to spread greatlyfrom the area where it is either planted or discarded, but damp, linear habitats, such as riverbanks,probably represent sites where the species is best able to colonise and spread, assisted by water flow.
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The Latin specific epithet'campanulata' is a diminutive of 'campana', meaning 'bell', and thus means 'small bell', in this case avery good description of the individual flower (Gilbert-Carter 1964).
Introduced, neophyte, occasional.
1947; MCM&D; Mountdrum Td, N of Lisbellaw.
May to November.
Growth form and preferred habitats
This 2 m tall, large-leaved, vigorous rhizomatous, decorative garden perennial forms dense clumps, and isa native of the Himalaya, the Indian subcontinent and Afghanistan. It is similar to the more notoriousJapanese Knotweed (Fallopia japonica), but the leaves are longer and more pointed. The exact yearof introduction of this large, dense, spreading, dominant perennial to gardens in Britain and Ireland isunknown, but it must lie around the very end of the 19th century. The first report beyond the gardenwall was in 1917 in N Devon (Conolly 1977).
Most records of this giant plant in Britain and Ireland are from roadsides, hedge banks, railway banks,streamsides, waste ground and old quarry settings. From the evidence of the associated species, theyclearly derive from discarded garden rubbish, presumably containing pieces of the rhizome. Very fewreports occur anywhere in Britain and Ireland of situations where the plant might have spread beyond theconfines of a garden by natural vegetative extension, ie under its own power of growth, without humanassistance to transport it.
Taxonomic synonymy and variation
P. wallichii has been previously classified in three other genera, Reynoutria,Aconogonum and Polygonum. Its most recent synonym is Polygonum polystachyum(Wallich ex Meissner). A more hairy stemmed variant form exists and has been around since 1917 (Conolly1977). It has been given the name Polygonum polystachyum var. pubescens Meissner (Clement& Foster 1994). So far, this form has not been recorded in Ireland (Conolly 1977; Reynolds 2002).
Fermanagh occurrence
The Flora Database records suggest this garden escape or discard has been rapidly increasing in the VC inrecent years. Previously there was just the one old record listed above from the Meikle era (1947-53),the plant having escaped (presumably to a roadside location) from a nearby cottage garden outsideLisbellaw. Nowadays, however, there are 30 additional records dating from 1988 onwards. As theFlora map indicates, these are scattered across a further 21 tetrads, mainly in the NE and E ofthe VC. Most of the Fermanagh records are from roadside, waste ground and old quarry settings, and fromthe evidence of the associated species, derive entirely from discarded garden material.
Reproduction
Himalayan Knotweed rarely if ever sets seed in Britain and Ireland, and its propagation and dispersal isachieved by rhizome fragments transported by man in soil, mud, or amongst garden rubbish. Very fewreports occur of situations in Britain and Ireland where the plant has or might have spread beyondgarden confines by natural vegetative extension, ie under its own power of growth, without humanassistance.
In recent years, we have also noticed Himalayan Knotweed becoming quite a common roadside plant in WDonegal (H35), where it again forms well-established, often dense pure stands, and indeed where it wasfirst reported outside a garden setting in Ireland as long ago as 1928 (Conolly 1977).
British & Irish occurrence
While the New Atlas map shows there is a continuing increase in records throughout these islands,it is in SW Britain and in NW Ireland, where the two first 'escapes' occurred (or were recorded), thatP. wallichii still maintains its greatest, most invasive presence (Preston et al. 2002).The great English horticultural expert Graham Stuart Thomas (2004) regarded P. wallichii as,"a tremendous spreader and only fit for landscape planting in moist ground, where it will smothereverything and provide a wonderful display of fragrant plumes in October".
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The Latin specific epithet'wallichii' is the genitive meaning 'of Wallich', the surname of a taxonomist botanist associated withthe genus.
Threats
None as yet in Fermanagh, particularly since it does not seem to set any seed. If this situation shouldchange, perhaps encouraged by rising temperatures, secondary dispersal from existing established clumpscould then pose a major risk to semi-natural vegetation. This has already happened with other invasivealien species in Britain and Ireland. As it is, the plant produces very dense clumps with which nativespecies cannot compete. It has no natural enemies, and once established will be practically impossible,or extremely expensive, to eradicate.
Introduced neophyte, a rare garden escape. Eurasian boreo-temperate, but so widely naturalised it hasbecome circumpolar.
1884; Barrington, R.M.; field near shore W of Ely Lodge House.
April to August.
Growth form and preferred habitats
This tall, pink-flowered, rhizomatous garden perennial is definitely a neophyte, introduced alien speciesin Ireland. In Britain, it is now recognised that the native range is obscured by additional introducedgarden forms across the whole country (A.R. Akeroyd, in: Preston et al. 2002). P. bistortaprefers damp, mildly acidic, humus-rich soils, often near water. The most natural habitats it occupiesare base-poor, damp soils on river banks, tall-herb communities in river valleys, and mountain ledges.It also occurs in more artificial, man-made habitats, including pastures, hay meadows and roadsideverges.
Reproduction
P. bistorta can reproduce and disperse vegetatively, by means of its stout rhizome, and byflowering. The latter takes place from May or June to August, cross-pollination of the perfumed whiteflowers being carried out by insect visitors, including sawflies (Proctor et al. 1996). Seeds arescattered by birds feeding on the fruiting heads (Grieve 1931). As a result of these reproductivemethods, P. bistorta often forms dense clonal clumps that can prove long-persistent.
Fermanagh occurrence
P. bistorta is rarely found in 'the wild' in Fermanagh, having been recorded in just elevenscattered lowland tetrads in the VC. It is most often found in or near demesnes such as Ely Lodge,Castle Coole and Florencecourt, from the gardens and grounds of which it tends to 'escape'. In morerecent years, it has also been found by the sides of roads and forest tracks, and at least once near aruined cottage where it certainly is a relic of cultivation. As a local instance of its persistence,P. bistorta still occurs at, or very close to, Barrington's original 1884 meadow site at ElyLodge on the shore of Lower Lough Erne.
Fossil record
Fossil pollen of P. bistorta and P. vivipara (L.) Ronse Decr. (= Polygonumviviparum L.) (Viviparous Bistort or Alpine Bistort) cannot be distinguished, but P.bistorta seed is recognisable. It has been recorded once, in E Yorkshire, from zone IV of theFlandrian, the current interglacial period we are living in. This is an early post-glacial zone, calledthe pre-boreal, approximately dating to just after 10,000 BP (Godwin 1975). If this solitary record iscorrect, it indicates the presence of P. bistorta well before human immigration and settlement.The species is thus regarded as indigenous in NE England, but elsewhere, and especially in SE England,it is possibly or probably a garden escape.
Irish occurrence
Common Bistort was first recorded in the wild in Ireland in 1746 in Co Waterford (H6). Since the mid-19thcentury onwards it has always been recognised as an alien of garden origin, "formerly muchcultivated", probably as a pot and medicinal herb (Cybele Hibernica 1866). By the end of the19th century, the plant was being considered an increasingly rare, casual relict of cultivation in muchof what is now the RoI (Colgan & Scully 1898; Irish Topographical Botany). Interestingly, theNew Atlas map indicates that to this day, over a century later, P. bistorta is very muchmore frequently found in Ireland north of a line drawn between Sligo and Dundalk. Nevertheless, recentlythe species has been recorded at two sites in Co Waterford in 1997 and 2001 (Green 2008), and the NewAtlas indicates it has been found in a post-1987 hectad in E Cork (H5). These appear to be theonly Common Bistort sites discovered in S Ireland for very many years (New Atlas; Cat Alien PlIr).
British occurrence
Bistort is always local in Britain and Ireland, but is most frequently found in NW England and in SScotland. Elsewhere in Britain, it is widespread but very local, with the exceptions of the S and EEnglish Midlands and the Scottish highlands and islands, in both of which it is scarce or absent. Withregards the species status, P. bistorta is generally considered native in Cumbria and S Scotlandwhere it is most plentiful, although in truth as time goes on it appears increasingly difficult, if notnigh impossible, to distinguish introduced populations of garden origin from native ones throughout theisland (Lousley & Kent 1981; J.R. Akeroyd, in: Preston et al. 2002).
This becomes especially the case when one realises that in the past Bistort was widely cultivated aroundhabitation, not like today for its decorative flower heads, but rather for use as an edible pot herb. Itwas used as a spring green vegetable, valued for the supposed nutritional quality of its leaves thatwere reputed to cleanse the blood. It may have had additional herbal medicinal properties (Grieve 1931;Grigson 1987; and see below).
European and world occurrence
P. bistorta is widely distributed and considered native in boreal Europe and W Asia, but in SEurope it becomes mainly confined to the mountains. It is absent as a native in Ireland and in most ofFennoscandia (Jalas & Suominen 1979, Map 414; Tutin et al. 1993). A form known as P.bistorta subsp. plumosum (Small) Hult. occurs in E Siberia andnorthwestern N America (Hultén & Fries 1986, Map 653). The Siberian distribution remainsinsufficiently known. The European form of the species has been introduced widely beyond its nativerange, including to N America, making it now a circumpolar boreo-temperate species.
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The Latin specific epithet'bistorta' is the Mediaeval or 16th century name of the plant, and it was also previously its genericname (Grigson 1974). It is a combination of two words, 'bis' and 'torta'; it translates as 'twicetwisted', a reference to the plant's characteristic stout, contorted rhizome which is often 'S'-shaped(Grieve 1931; Clapham et al. 1962; Gilbert-Carter 1964).
This feature of the plant and its creeping nature appears to be the origin of some of the alternativeEnglish common names, which refer to coiled snakes or adders. Early English herbalists including Gerardand Lyte called the plant 'Adderwort' and 'Snakeweed', and by the Doctrine of Signatures, or theprinciple of sympathetic magic, they regarded it as a remedy for snake bite or poison (Grigson 1987).
Seven of the sixteen English common names from around Britain and Ireland listed by Grigson (1987) referto some aspect of Easter (eg Easter Ledger, Easter Ledges, Easter Mangiants), or Passion (Passion-tidebeing Easter), (eg Passion Dock and Patience Dock). The reason for this appears to be that P.bistorta was an ingredient of a food called 'Easter Ledger Pudding' (Grigson 1987).
Uses
In the English Lake District, where today the species is most frequent and abundant, the young springtimeleaves are still eaten as a vegetable like spinach and they are used as one ingredient of a traditionalEaster pudding with fertility and blood cleansing connotations. Grigson (1955, 1987) provides a detailedand plausible explanation of the mysterious beliefs surrounding this matter, and also details afascinating symbolic link with the marvellously beautiful 'Captured unicorn' French wedding tapestry of1514. This seasonal folk use led to the local cultivation of the plant in Cumberland and Westmorland,and it is known there as 'Easter Giants', 'Easter Mangiants', 'Easter Ledges', Easter Ledger', 'PassionDock' and 'Patience Dock' amongst other names, from which Grigson (1955, 1987) draws meaning.
Additional medicinal folklore and Easter pudding recipes are provided by Vickery (1995) and Mabey (1996).Grieve (1931), and Allen & Hatfield (2004) list medicinal uses of the plant, which is one of thestrongest astringents around. It is also highly styptic (ie it stops bleeding), and has been used bothas famine food and for the treatment of numerous complaints, including wounds, headaches, worms, andbowel and urinary problems.
Threats
None.
Introduced, neophyte, a very rare garden escape or discard.
14 June 1985; Northridge, R.H.; Castle Caldwell FNR.
June to August.
Growth form and introduction
This attractive and distinctive, up to 1 m tall, tufted, mat-forming, garden perennial, is a native ofthe Himalaya, and ranges from Afghanistan to SW China. It has a stout woody rhizome and large cordateleaves. In the wild in Britain and Ireland, the plant is found naturalised on roadsides, waysidethickets, quarries and stream banks, often in sites where garden refuse is deposited.
Red Bistort was first introduced to garden cultivation in Britain and Ireland in 1826 for its decorativedeep red flower spikes. It was first recorded in the wild in Britain around 1908. In Ireland, by 1920,it was said to be naturalised on the Hillsborough demesne, Co Down (H38) (J.R. Akeroyd, in: Prestonet al. 2002; Cat Alien Pl Ir).
Reproduction
Red Bistort plants develop deep purple-red, claret-coloured or white flowers borne densely packed inlong, slender, cylindrical spikes up to 8 cm long on naked flowering stems. The main flowering period isin late summer, from August to October, and it continues until the first frost. While Red Bistort rarelyor never sets seed here, it possesses a vigorous branched rhizome or woody rootstock which spreadsaggressively to the extent that it may become an invasive weedy nuisance, especially in smaller gardens(Lousley & Kent 1981). Fragments of the plant transported in soil, or with discarded garden waste,frequently enable it to 'escape' from gardens.
Variation and taxonomic nomenclature
Like other Knotweeds, this species has been renamed several times and has been placed in other genera asBistorta amplexicaulis (D. Don) E. Greene and Polygonum amplexicaule D. Don. Seven gardencultivars are listed in the RHS Index of Garden Plants (Griffiths 1994), varying in flowercolour, habit and size (eg 'Inverleigh' is a dwarf form).
Fermanagh occurrence
In Fermanagh, Red Bistort is apparently well naturalised just to the west of the Old Castle Caldwell. Theonly other recorded site in the VC is amongst tipped garden refuse on the moss-covered rocky floor ofthe disused roadside quarry, just opposite the turning for Brookeborough village.
Irish occurrence
The New Atlas map shows this species is fairly rare or scarce in Ireland, widely but thinlyscattered throughout, but having been recorded in 19 of the 40 VCs. Elsewhere in N Ireland (NI), thereare one or two scattered records in each of counties Tyrone, Down and Antrim (H36, H38 & H39). Atone of these sites, at least, the plant persisted for over 20 years, yet Paul Hackney (in the NIVascular Plant Database 2002) did not consider the plant fully naturalised anywhere, presumably becauseit does not reproduce sexually.
British occurrence
In Britain, records are concentrated in the S & SW of England, but an attenuated, increasinglydisjunct presence extends northwards, mirroring a decreasing human population. P. amplexicauliseventually reaches a remote outlier in North Aberdeenshire (VC 93). The majority of its stations arealong roadsides, or in other no-man's-land where fly-tipping of rubbish and excess garden material isall too frequent. Regrettably, dumping of garden material also occurs at the coast where relatively openconditions on sand, loose gravel and shingle are especially conducive to the plant's establishment.
The increasing frequency of records in the post-1987 period displayed in both the New Atlas and inReynold's Cat Alien Pl Ir, strongly suggests that P. amplexicaulis is still activelyspreading.
European occurrence
Apart from its occurrence in Britain and Ireland, the only other record mapped for Europe by Jalas &Suominen (1979, Map 413) is from a single site in Czechoslovakia.
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The Latin species epithet'amplexicaulis' is a combination of 'amplexus' meaning 'encircling' or 'embracing' and 'caulis' meaning'stem' referring to the leaf stalk (petiole) or leaf base partially clasping the stem, which is the casein this species. The English common name 'Red Bistort' is a recent book name that merely recognises thedecorative deep red flower spikes.
Threats
None.
Native, frequent but rather local. Circumpolar boreo-temperate, but also widely naturalised.
1900; Praeger, R.Ll.; Co Fermanagh.
April to October.
Growth form and preferred habitats: A very variable, truly amphibious,rhizomatous perennial, which in water is a very distinctive and beautiful stand-forming perennial withpink or reddish, dense flower spikes. This species also has a much more easily overlooked terrestrialgrowth-form of very different appearance – a weed of damp waterside soils or somewhat drier, rough ordisturbed ground. In Fermanagh, as elsewhere in Britain and Ireland, the terrestrial form appears tovery rarely flower, a fact which together with its occurrence amidst other weedy species is sufficientto account for it being somewhat under-recorded.
The aquatic form of P. amphibia is a plant of still or very slow moving, nutrient-rich, shallowwaters, roughly between 50 and 200 cm deep. It occurs in a wide variety of generally muddy, silty orpeat floored, lowland wetland habitats, including ditches, rivers and lakeshores (Preston & Croft1997). Studies elsewhere show it can occur within the pH range 3.0 to 8.0, although it is extremely rareat pH levels below 4.5 (Grime et al. 1988; Partridge 2001).
The plant is remarkably tolerant of pollution and increasing eutrophication and it is one of the lastaquatic macrophytes to succumb during this form of often rapid environmental change (Partridge 2001).The aquatic form is able to tolerate water turbidity caused by algal blooms or sediment disturbance,since submerged leaves that might become dependent in such circ*mstances, are entirely absent (Preston& Croft 1997). It is also very well adapted to fluctuating water levels, but does not handle shadewell (Partridge 2001).
Identification
The vegetative terrestrial form of P. amphibia can readily be misidentified as the very commonweedy annual, Redshank (Persicaria maculosa), and the aquatic form, when non-flowering, maysimilarly be taken for Broad-leaved Pondweed (Potamogeton natans) (although it differs in theparallel leaf veins) (Partridge 2001). The aquatic and terrestrial forms of P. amphibia are sovery dissimilar, they could easily be mistaken for separate species by the uninitiated (Lousley& Kent 1981). The two growth-forms were nevertheless recognised as belonging to the same speciesback in the early 18thcentury (Ray 1724).
With the exception of the stalk of the inflorescence the aquatic form is entirely hairless and hasstalked, waxy, floating leaves with petioles 2-4 cm long, while the terrestrial plant is erect anddistinctly and generally hairy, at least to a degree, and its leaves are stalkless, or almost so (iesessile, or with a very short petiole). Immature or early season leaves of the terrestrial form oftenbear a dark chevron-shaped blotch. To add to the general confusion, a transitional form of the plant canoccur at water margins (Partridge 2001).
Reproduction
Despite aquatic populations flowering freely, reproduction appears to be mainly vegetative – by extensiongrowth of its slender, creeping rhizome and through the ready rooting of detached plant fragments (Grimeet al. 1988; Partridge 2001). Seed production appears uncertain and infrequent, due to flowersbeing self-incompatible and some clones being male-sterile and requiring transported pollen.
Bright red, pink, or white flowers are produced from July to September. They produce scent and nectar andare visited by flies and butterflies. Seed on male-fertile plants is said to be infrequent, there rarelybeing more than 5-10 per inflorescence. This suggests there is also a high degree of ovum infertility(Partridge 2001).
Various patterns of reproductive strategy have been suggested, but Partridge (2001) concluded that seedis associated only with survival in extreme situations, although it is vital for long-range dispersal(ie jump-dispersal) between water systems. As with aquatic plant fragments, seeds float for 7-10 daysafter release from the parent plant and are readily transported by water within a drainage system.However, transport of seed by birds between water bodies or from adjacent moist terrestrial ground,possibly adhering in mud, would be highly significant, even if it only very rarely occurred. Mammals andbirds (especially water-fowl), eat the plants and seeds, but there does not appear to be any record ofthem affecting dispersal (Partridge 2001). More work is required to elucidate the true situationregarding dispersal of this species.
Fossil record
Seeds and pollen of P. amphibia appear in British sediments back to the Cromer Forest Bed series,but fossils become more frequent later on in all zones of the Late Weichselian. Fossil records persistinto the beginning of the current interglacial, ie the early Flandrian, but then there is an absenceuntil records reappear in later zones (Flandrian viib & viii), and in Roman times. We can say,however, that P. amphibia has persisted in Britain through from at least the middle of the lastinterglacial, and thus is a definite native species (Godwin 1975, p. 232).
Fermanagh occurrence: Amphibious Bistort has been recorded from a totalof 125 tetrads, 23.7% of those in the VC. Around 90 of the tetrads are represented by the shores of bothparts of Lough Erne. The species is almost omnipresent and often abundant in wetlands around the frettedshores of mesotrophic to distinctly eutrophic Upper Lough Erne. It is also present, but to a lesserdegree, on the much more open, wave-exposed, more calcareous shores of Lower Lough Erne and around othersmaller, lowland water bodies, including rivers and ditches in the VC.
The terrestrial form is much less abundant in marshy, or drained but damp lowland grassland areasadjacent to the lakes, limestone turloughs and slow flowing waters in rivers and ditches scatteredaround the county, but it does also rarely occur on roadside verges, for instance near Carry Bridge.
British and Irish occurrence: P. amphibia is frequent andwidespread throughout lowland Britain and Ireland, absent only in upland areas lacking suitably fertile,wetland habitats (Preston et al. 2002). The New Atlas hectad map suggests that in WIreland P. amphibia becomes almost confined to coastal or near-coastal wetlands, while in NIreland it is closely associated with the larger freshwater lakes (Lough Neagh & Lough Erne).However, this appears to be less the case in Co Down (H38) – a VC which also happens to have avery long coastline. In contrast, P. amphibia appears more or less absent in large inland swathesof Co Tyrone and W Donegal (H35 & H36), where thanks to their regular and heavy annual rainfall, onemight conjecture that plenty of suitable marshy ground is available, although this may prove too acidicor too leached for Amphibious Bistort to successfully colonise.
P. amphibia is extremely frost-sensitive and is the first wetland species to die-back in autumn.This accounts for the entirely lowland distribution of the species, and also for its increasingconfinement to maritime regions as one travels northwards on these islands. The species reaches itsmaximum altitude in Britain of 570 m at Blind Tarn in Westmorland (vc 69) (J.R. Akeroyd, in: Prestonet al. 2002).
European and world occurrence
P. amphibia is widespread throughout lowland regions in temperate latitudes of Europe. As is thecase in Britain, it becomes more prevalent in coastal districts further north in Scandinavia, and it israre and more scattered towards the Mediterranean area (Jalas & Suominen 1979, Map 408). Beyond itsEuropean area, the native range of P. amphibia extends east into C Asia. In the European Alps, itextends to about 1,200 m and in the Himalayas to above 2,700 m (Hultén 1974). It is also an introducedweed in many countries in Africa and in N & S America (Hultén & Fries 1986, Map 652).
Hultén (1974, Map 116) recognises an E Asian form as subsp. amurense (Korsh.) Hult., and he alsodistinguishes the native form in N America as subsp. laevimarginatum Hult. There must be genetransfer where the introduced European form (subsp. amphibia) and these other subspecies meet.All subspecific taxa of the species have both aquatic and terrestrial forms that differ significantly ingeneral appearance and degree of pubescence. P. amphibia s.l. belongs to the circumpolarboreo-temperate geographic element of plants (Hultén 1974; Hultén & Fries 1986; Preston & Hill1997).
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The Latin specific epithet'amphibia' is derived from the two Greek words 'amphi' meaning 'both', and 'bios' meaning 'life', thusmeaning 'living a double life', ie living both on land and in water (Gilbert-Carter 1964). The Englishcommon name is a simple translation of the Scientific name.
Threats: None.
Native, common and widespread. Eurasian temperate, but extensively naturalised, and thus circumpolar andwidespread in both hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
April to December.
Growth form and preferred habitats
Throughout almost all of lowland Britain and Ireland, this strict summer annual is one of the most commonand abundant weeds colonising open, sunny, lowland, recently disturbed ground. P. maculosa is avery variable erect to sprawling decumbent, occasionally dwarfed, prostrate plant, 3-80 cm tall, butgenerally it grows erect, up to 30 cm in height. The stems are often suffused with red (hence theEnglish common name), and leaves are often black-blotched on the upper side.
Typical habitats include damp soil heaps, cultivated but unsown surfaces in fallow ground in gardens, asa weed among broad-leaved crops in fields and gardens, and on open or disturbed roadside verges,especially where there has been recent road-works. It is also found around building sites, on wasteground and in manured, heavily trampled or otherwise disturbed ground where competition is greatlyreduced. Redshank also grows on damp or moist muddy ground exposed near lowered water bodies, asituation where it may meet the terrestrial growth form of P. amphibia (Amphibious Bistort),although the two species do not hybridise (Stace 1997).
P. maculosa can tolerate an extremely wide range of soil conditions. However, like many otherruderal weed followers of man and species of intermittent habitats, it is a species characterised by arapid growth rate, high levels of seed production and long-persistence in the soil seed bank. Like otherweeds, Redshank generally grows most abundantly and luxuriantly on damp, bare, naturally fertile ornutrient-enriched conditions.
Redshank is generally absent, if not absolutely so, from strongly acidic soils on bogs and moorland. Inthe English Midlands, Grime et al. (1988) reckoned P. maculosa was most frequent in the pHrange 5.0-7.0, and it avoided acidic conditions below pH 4.5. It also avoids closed turf vegetation,aquatic or permanently waterlogged conditions and shade. It is, however, often associated with winter-or seasonally-flooded ground near water bodies. Whatever else varies, P. maculosa remainsconfined to sites that have been recently subjected to some form of disturbance. This reflects theessential lack of competitive ability typical of ruderal species, but they are also often characterisedby extreme variability, both genetic and plastic with respect to their immediate environment. P.maculosa fits this pattern of ruderal characteristics and behaviour very closely.
Fermanagh occurrence
In Fermanagh, it is very widely distributed throughout the lowlands, less frequent but still present atintermediate levels, including on the Western Plateau. It has been recorded in 285 Fermanagh tetrads,54% of those in the VC. Since there has been very little arable agriculture in the county since the endof World War II, it occurs most frequently on trampled and manured, cattle-grazed pastures around lakeshores, on roadside verges, open waste ground, and in more fertile, disturbed urban sites.
Reproduction
Plants flower after about six to eight weeks' growth, usually doing so from May to September. P.maculosa (sometimes known as 'Pink Persicaria' or 'Red Persicaria'), produces small, bright pink(or occasionally white) flowers, around 50 of which are borne on a short, rather dense, cylindricalspike inflorescence. The flowers are not perfumed but do contain a little nectar. They eitherautomatically self (stamens incurving to touch the stigma), or are insect-pollinated. The fruit is asingle-seeded achene or nut, and they are of two kinds. Most fruits are triangular with three hollowfaces, about 2.5-3.0 mm long, blackish brown and shining. Usually only a small proportion of the achenesare of the secondary compressed, lenticular or biconvex type, but the relative number of the two kindsproduced by an individual plant, and between plants, is very variable (Simmonds 1945). The achenes, whenreleased are ± contained within the dry, withered perianth, which in water assists flotation anddispersal (Lousley & Kent 1981; Grime et al. 1988). A typical sized plant produces between200 and 1,200 achenes (Salisbury 1964).
Fruit dispersal
Dispersal is achieved by birds, horses, cattle and other animals feeding on the plant and internallytransporting the still viable seed (Ridley 1930). Man has also helped transport the species – locally inmud on clothes and machinery, and in past generations, around the globe with his agriculture, throughcrop seed impurities of cereal, flax, and particularly with clover for which there was huge agriculturaltrade (Salisbury 1964). In this manner, P. maculosa has reached near-cosmopolitan weed status intemperate and tropical areas around the world (Simmonds 1945; Hultén & Fries 1986, Map 650).
Germination
Seed germinates in spring after winter chilling, and depending upon seasonal conditions this may beginhappening as early as mid-March, or as late as mid-April. Most germination takes place between April andJune (Salisbury 1964), but if there should be a dry period in spring, the process can be prolonged, eveninto mid-July (Witts 1960). However, a situation like this would be exceptionally unlikely to occur inFermanagh's damp, mild, oceanic climate.
Variation
Reflecting the wide range of disturbed habitats P. maculosa can occupy, there is a wealth ofgenetic variability within the species. A number of varieties have been described (eg var.agreste Meisn., var. ruderale Meisn., and var. prostratum Bréb.), but the feelingis that the variation is plastic and environmentally induced, and therefore does not easily lend itselfto or deserve taxonomic treatment (Simmonds 1945).
The newly published critical Flora of Great Britain & Ireland Volume 1 has taken a leapover this hurdle and recognises two subspecies, the native one containing three varieties. Subsp.hirticaulis (Danser) S. Ekman & T. Knutsson is a very rare Asian introduction. Thedistinctions centre on the presence or absence of appressed or spreading hairs on stems and peduncles(the stalk of the inflorescence), and whether or not glandular hairs or short-stalked glands are presenton the peduncles (Sell & Murrell 2018).
Fossil history
Fossil nuts have been recorded from the Cromer Forest Bed series onward to the early Flandrian. Godwin(1975) felt it reasonable to say the species has been persistently present since the opening of the LateWeichselian to the beginning of the archaeological record in the Iron Age, and again later in Roman,Norman and Medieval times. Thus there is no doubt that P. maculosa is a native species. Thenative habitat is more difficult to identify, but clearly it must have involved some circ*mstance, likeregular disturbance, that limited competition from co-habiting plants.
British and Irish occurrence
The New Atlas hectad map and species account indicate that P. maculosa is very widespreadand stable in its distribution throughout Britain and Ireland, becoming less common, or absent only onhigh ground, strongly acidic bogland, and permanent wetlands (J.R. Akeroyd, in: Preston et al.2002).
European and world occurrence
Common and widespread throughout temperate Europe (including Iceland) and W & C Asia. In Scandinavia,it is common only in the south, although it is scattered along the W coast and extends to 70°N. Thedistribution thins considerably towards the Mediterranean, although it is present on most of the westernisles (Jalas & Suominen 1979, Map 406). It has been introduced very widely around the globe withagricultural seed (including N & S America, Australia, New Zealand, scattered parts of E Asia andAfrica) (Hultén & Fries 1986, Map 650). It has become naturalised in many areas and is now almostcosmopolitan. The development of more efficient agricultural seed-cleaning means further introductionsare much less likely, and we might expect gradual reductions in the presence and range of this weed.
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The current Latin specificepithet 'maculosa' means 'spotted', a reference to the black blotched leaf marking typical of thespecies (Gilbert-Carter 1964).
The list of English common names runs to 24 in Grigson (1987), of which five include 'red' in referenceto stem colour (Redshank, Red Legs, Red Joints, Red Knees and Redweed). The dark leaf spot features inseveral Easter-related legends, suggesting drops of blood from the cross marking the plant and givingrise to such names as 'Pinch-weed' and 'Virgin Mary's Pinch' or 'Devil's Pinch' or 'Useless' because theVirgin (or the Devil) pulled up the plant, left the mark on the leaves and discarded it as useless, ielacking the peppery taste of P. hydropiper (Water-pepper) (Grigson 1987, p. 232).
Threats
None.
Native or possibly introduced, occasional, perhaps only casual. Circumpolar southern-temperate, but alsowidely naturalised in both hemispheres.
1900; Praeger, R.Ll.; Co Fermanagh.
July to October.
Growth form and preferred habitats
Overall this genetically very variable and phenotypically extremely plastic annual knotweed looks rathersimilar to the extremely common ruderal weed, P. maculosa (Redshank). It is larger thanthe latter, sometimes reaching 1.2 m in height, although more often it is less than half this. The stemsare usually very pale green, tinged pink, and the flowers typically have a greenish-white, ratheranaemic appearance (and thus its English common name, 'Pale Persicaria'). Flower colour is veryvariable, however, and it can also be a dingy pale or a deeper pink. Also, the peach-like leaves areblotched with black (often), or sometimes not; the yellow glands on the peduncle are densely present andcan often be seen with the naked eye (Lousley & Kent 1981). Flower colour, habit and pubescence areall very variable, as Stace (1997) very helpfully points out!
Like many other Persicaria species (and especially like P. maculosa), ecologically it is aweedy pioneer colonist of a wide range of open or recently disturbed habitats, including cultivatedland, manure heaps, waste ground and moist soils marginal to water. P. lapathifolia occurs onvarious types and textures of soil, from river gravel to silt, through sand and clay, but it appearsmost often, and can sometimes form large, dense stands in mildly acid, nutrient-rich, peaty fenconditions, avoiding strongly acidic and nutrient-poor terrain. The species is a weak competitor andvery stress tolerant. The chief habitat requirement therefore is sufficient disturbance to reducecompetition from associated, often weedy species (Simmonds 1945; Sinker et al. 1985; Mitich1998).
Variation
In addition to the above remarks, P. lapathifolia is extremely variable genetically, and isphenotypically very plastic with respect to local environments. Two varieties are recognised in Britainand Ireland by Sell & Murrell (2018), var. lapathifolia, and var. salicifolia, thelatter having leaves that are whitish and densely arachnoid-hairy beneath (ie cobweb-like). InScandinavia, two subspecies are described, subsp. lapathifolia and subsp. pallida (With.)S. Elkman & T. Knutsson, the latter being a crop weed that can be further divided into twovarieties, var. incana and var. linicola, the latter being a weed specific to flaxcrops.
Worldwide, P. lapathifolia s.l. is an extremely complicated and variable complex in which a greatnumber of species, subspecies, varieties and forms have been described. In the boreal belt of the Nhemisphere, the variation is grouped around two main types, P. tomentosum Schrank and P.nodosum Pers. However, the two types are not everywhere distinctly separate (eg in Scandinavia),and also their ranges overlap (Hultén 1974, pp. 244 & 390, Map 236).
Hybrids
Hybrids of P. lapathifolia are known with P. foliosa (H. Lindb.) Kitag., P.hydropiper (Water-pepper) and P. maculosa, which underlines the fact that reproduction inthis species does not entirely involve selfing (Jonsell et al. 2000).
Reproduction
Being annual, P. lapathifolia reproduces entirely by seed. Decumbent lower branches may root atnodes touching the ground, although there is no suggestion of vegetative reproduction. The plant dies inthe autumn.
Flowering occurs around four to six weeks after germination, generally between June and September (Mitich1998). The flowers are bisexual (perfect) but pollen production is very low (about 30 pollen grains peranther). The low ratio of pollen to ovule production is characteristic of inbreeding, and indeedfertilisation is almost or entirely autogamous (selfing), although small insects (thrips) have beenrecorded visiting, possibly assisting pollination (Simmonds 1945; Jonsell et al. 2000).
Fruiting takes place from mid-July onwards. The fruit is single-seeded, dry (ie a nut-like achene), andit is contained within the persistent perianth. The achene is lens-shaped (lenticular) or rarelytriangular in section (trigonous), brown or black in colour, and either glossy or dull. The number ofseed produced is extremely variable, ranging from 10 to about 1,500 per plant (Simmonds 1945). A weedseed trial found an individual plant could yield up to 19,300 seeds when grown without competition(Stevens 1957).
Seed dispersal
There is no specific seed dispersal mechanism, the mature fruit simply fall or are knocked off the parentplant. As with P. maculosa, birds and herbivorous mammals (cattle, horses and deer) may eat theplants and transport seed, depositing it with their faeces (Ridley 1930). There is no informationregarding seed viability after passing through the alimentary canal, although this was reported for theclosely related P. maculata.
In the past, fruits were known to occur as impurities of agricultural clover seed and livestock feed, andthe species has undoubtedly been spread by man in this manner (Simmonds 1945). Seed cleaning processesare very much more efficient nowadays, and dispersal as a crop seed impurity has probably been verygreatly curtailed, if not totally eliminated everywhere.
Seed germination and longevity
P. lapathifolia and P. maculosa not only look alike, an experimental study found theirseeds also behave in a very similar manner when cultivated and disturbed three times per year. Resultsshowed a relatively high percentage of Pale Persicaria seed germinated in the April and May of the yearafter production, and in subsequent years seed numbers progressively declined, although a smallpercentage persisted and remained viable after five years (Roberts & Neilson 1980). This is entirelyconsistent with the ecology of other weedy members of this family, all of which appear to demonstrateprolonged survival in the soil seed bank.
Fossil record
All records are for achenes which have been found in small numbers from the Cromer Forest Beds onwards inboth glacial and interglacial stages, proving ancient native status and suggesting possible periglacialsurvival. In the current Flandrian, records are numerous and extend from the Bronze Age to the Mediaevalperiod in archaeological sites. The evidence suggests the fruits were previously part of the prehistorichuman diet (Godwin 1975).
Fermanagh occurrence
P. lapathifolia is only occasional in Fermanagh, but it is widespread mainly around Lough Erne andin the lower-lying, rather more fertile, less wet farmland of the east of the county. Prior to thecurrent flora survey (Forbes & Northridge 2012) there were just four records. The original Praegerrecord of 1900 was completely unspecific with respect to site and there were three additional recordsmade by Meikle and co-workers in the 1946-54 period. A total of 34 subsequent records were made from1986 onwards by several recorders, although the majority were by RHN. The records of this annual are nowspread across 32 Fermanagh tetrads, 6.1% of those in the VC. It is mostly found around Lower Lough Erne,but is also scattered thinly to the east of the county.
It occurs on cultivated and disturbed ground, including waterside habitats, roadsides and in oldquarries.
Status in Ireland
In the Connemara and the Burren regions of western Ireland, Webb & Scannell (1983) regarded P.lapathifolia as being, "perhaps little more than casual" in its appearances.Furthermore, the Irish Census Catalogue regarded its status as, "possibly introduced"(Scannell & Synnott (1987).
British and Irish occurrence
While P. lapathifolia is physically, biologically and ecologically most similar to P.maculosa, and the two species often occur together, it is nothing like as common as the latterin Britain and Ireland. The difference in presence is no doubt due to Redshank possessing widerecological tolerances, rather than it being more competitive.
The New Atlas hectad map shows P. lapathifolia is not quite as widespread and omnipresentin lowland England and Wales as Redshank. In Scotland, it becomes much less prevalent than further southin England, being scattered and increasingly eastern and coastal northwards. The species map for Irelandlikewise shows that P. lapathifolia is very much less common and widespread than it is in Englandand Wales. The Irish distribution of P. lapathifolia is also very much more eastern than thealmost ubiquitous P. maculosa (Preston et al. 2002).
Although the New Atlas species account suggests that the distribution of P. lapathifoliahas not changed appreciably in the last 40 years (calculated Change Index = -0.04), other work suggeststhe species has been, or is increasing in Britain and Ireland, both in the longer term and also morerecently. The authors of the The Flora of County Dublin commented on the positive change thepresence of Pale Persicaria has undergone between 1904 and their late 20th century survey: "Verymuch commoner now than in Colgan's time." (Doogue et al. 1998). The BSBI 'LocalChange' project which repeated the 1985-6 'Monitoring Scheme' survey in 811 pre-selectedtetrad squares in Britain, also suggests that P. lapathifolia is increasing in Britain(Braithwaite et al. 2006).
Neither of these reports proposes reasons why P. lapathifolia may be experiencing increase, but itmay be due to a combination of nitrogen deposition enriching the soil, increased levels of disturbance,and better taxonomic treatment in published Floras, giving rise to more confident field recording.
European and world occurrence
The taxonomy and nomenclature of this taxon has been so confused in the past that Jalas & Suominen(1979) list twelve synonyms. The European range extends from the far north of Scandinavia at 70°N, tothe Mediterranean and Macronesian islands in the south (Jalas & Suominen 1979, Map 407). In northernScandinavia, it becomes rare, casual or ephemeral, and is considered a recent incomer (clearly of cropseed weed origin) (Jonsell et al. 2000). However, taking this polymorphic species in the broad,sensu lato sense, it has spread with agricultural crop seed from a widespread European boreo-temperateorigin to become a circumpolar, near-cosmopolitan southern-temperate naturalised weed of arable anddisturbed soils. It now extends SE to Iraq and E to the Himalaya region, C China, S Japan and SE Asia,Australia, N America, Mexico, Chile, Argentina, tropical and S Africa. It reached New Zealand as longago as 1904 (Hultén & Fries 1986, Map 651; Webb et al. 1988).
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The previous generic name'Polygonum', in use until recently and occasionally revived, is from the Greek 'poly' meaning 'many',and 'gony' meaning 'knee', a reference to the characteristic swollen stem nodes found throughout thegenus. The Latin specific epithet 'lapathifolia' means 'leaf like a sorrel', or 'leaf dock-like'(Gilbert-Carter 1964).
The English common name 'Pale Persicaria' is a mere book-name, but at least it indicates the differencein colour of flowers and stems between this species and Redshank (Persicaria maculosa).Alternative common names include 'Pale Smartweed' and 'Pale Knotweed' (Mitich 1998).
Threats
None.
Native, common and widespread. Circumpolar temperate, but also widely naturalised in both hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
June to November.
Growth form and preferred habitats
The very hot, peppery taste of the 3-8 cm long lanceolate leaves is a very distinctive feature of thisstrictly summer annual. P. hydropiper is a hairless (ie glabrous) plant up to 75 cm tall, withgreen stems, often with a red ring beneath each node. As the plant ages the stems become increasinglyred. It produces long, slender, nodding inflorescences of greenish-white flowers from July to September.The perianth flower parts are covered with yellow glands, and the number of styles is usually two, morerarely three. The peppery taste of the leaves arises from glandular dots on their surfaces. There is noodour off the plant (Timson 1966; Clapham et al. 1987).
P. hydropiper is confined to a quite limited range of habitats and phenotypic variation in thisspecies is slight. Nevertheless, it grows in a variety of wet, often muddy, open, waterside, but alwaysterrestrial habitats. Indeed, it colonises any bare or open muddy ground where shallow water tends tostand in winter and dry out in summer. Occasionally this includes the plant appearing in ± temporarypuddles, created for instance by vehicle tracks, hoof-prints or simply the lie of the land in damp,patchy meadows, on woodland rides, or along roadside verges.
P. hydropiper colonises a wide range of soil textures from clay to peat, but while Praeger (1934)regarded it as a strongly calcifuge (lime avoiding) species in Ireland, and in their Flora ofConnemara and the Burren Webb & Scannell (1983) more or less concur ("distinctlycalcifuge in its general distribution"), the latter authors allow that P. hydropiper doesoccur in abundance in some calcareous habitats, especially in the eastern inland portion of the Burren,Co Clare (H9). Lousley & Kent (1981) hedged their bets and described the species as, "usuallycalcifuge". In Fermanagh, plants are most often growing in moderately acid to near-neutral, base-or lime-poor to rich soil conditions. As it occupies such a wide range of soils, it is not a usefulspecies indicator of marked substrate acidity, or of lime- or base-avoiding nature.
Being essentially a pioneer colonist of open, bare, muddy ground, Water-pepper is probably only weaklycompetitive, but it has most of the properties of a successful ruderal and it behaves in a similarmanner, being tolerant of stress in the form of trampling and winter flooding. It cannot withstandeither drought or frost and therefore is mainly confined to lowland sites, typically near water (Timson1966).
Reproduction
Like other small-flowered Knotweeds which have long, lax, terminal inflorescences and small subsidiaryinflorescences in the axils of leaves, P. hydropiper is rarely visited by insects and is almostentirely inbreeding (ie it is autogamous and automatically self-fertilises). Furthermore, the flowers inthe small inflorescences formed in the axils of leaves on lower stem nodes are cleistogamous, meaningthey self-fertilize while still in the unopened bud stage of development (Timson 1966). The fruit ofeach individual flower is a single seeded nut-like achene. The achenes are c 3 × 2 mm, biconvex ortrigonous (three-sided), depending on whether the flowers that formed them had two or three stigmas. Theachenes produced by cleistogamous flowers are irregularly shaped, and all achenes are dark brown toblack in colour and have a matt surface.
P. hydropiper flowers in all its many habitats, but it produces fewer seeds in drier or poorersoils. The typical plant produces around 300-400 seeds, but there is great variation in quantity, inresponse to habitat conditions (Timson 1966). Like other members of the genus, the single-seedednut-like, achene fruits are capable of long-term survival in the soil seed bank (more than five years),although the record on this is quite variable, several European studies suggesting they are ephemeral(Thompson et al. 1997).
In view of the degree of selfing shown, the small amount of Persicaria pollen produced and itssticky nature, Timson (1965b, 1966) felt that the virtual absence of cross-pollination ruled out anylikelihood of previously claimed hybrids involving P. hydropiper, or else at least it renderedthem extremely rare reproductive anomalies. Also, the putative parents all being annual species curtailsthe time available for the genetic readjustment necessary for some of the possible hybrids to survive.The absence of any form of asexual reproduction or apomixis certainly prevents the survival of triploidsor any other genetically unbalanced forms that might arise (Timson 1965b).
Hybrids
A study at Lough Neagh of P. hydropiper and its close relatives, P. mitis (= P.laxiflora Weihe) Opiz.) (Tasteless Water-pepper) and P. minor (Small Water-pepper),showed that while the latter two species did cross with one another, P. hydropiper did nothybridise with any other species (Parnell & Simpson 1988). Later study appears to have accepted thatP. hydropiper can rarely form hybrids with other annual species P. maculosa, P.lapathifolia, P. minor and P. mitis (Jonsell et al. 2000; Anonymous 2002,p. 230). Stace et al. (2015) examined hybrid claims in Britain and Ireland involving crossesbetween P. hydropiper and P. lapathifolia, P. mitis and P. minor. Theyconcluded that all of the hybrid claims made in both islands still require confirmation.
Dispersal
As one would expect, the strong acrid flavour of the plant sap is extremely effective in protecting theherb from grazing animals, which soon learn to avoid it. However there is a downside to this protectivefeature, since it rules out an efficient means of dispersal – ie the internal animal transport ofingested achenes (Timson 1966). However, despite lacking any obvious means of species dispersal, apartthat is from accidental transport in mud, transient flotation in water, or as a contaminant ofa*gricultural crop seed (especially amongst clover seed), P. hydropiper is a common and widespreadspecies in Britain, Ireland and beyond. Inefficient seed dispersal also results in Water-pepper tendingto occur in dense patches, either large or small (Timson 1966).
Fossil history
Fruits have been recorded from the Pastorian interglacial and the late Flandrian, and also from twointervening glacial stages – the late Anglian and tentatively, from the Middle Weichselian. From thelate Flandrian the fossil record stretches through the Bronze Age more or less continuously to theMediaeval Period. Many of the finds are from archaeological sites, but the record certainly representsthat of a native species (Godwin 1975).
Fermanagh occurrence
Water-pepper is the most frequently recorded member of the genus Persicaria in Fermanagh. It beatsP. amphibia (Amphibious Bistort) into second place by nearly 100 records. While it isconsiderably more common than the third ranking species P. maculosa (Redshank), however, it isnot as widely distributed as the latter, being recorded in 221 tetrads, 41.9% of those in the VC,compared with Redshank's 53.8% tetrad presence. This situation probably reflects Water-pepper's lack ofanimal transport. A small minority of people cannot detect the acrid taste, however, and this may alsohappen with some herbivores!
Nevertheless, P. hydropiper is the most common and the second most widespread member of the genusin our survey, and as the tetrad map shows, it is widely scattered throughout Fermanagh, although itappears to be most heavily represented around the shores and water-meadows of Upper Lough Erne. Thisdistribution is undoubtedly achieved and maintained by achene flotation. However, as mentionedpreviously, it is possible that the extremely detailed botanical survey of the Upper Lough carried outby DOENI field survey staff in the mid-1980s (the forerunners of the EHS Habitat Survey Team), may skewour species distribution, over-representing the presence of the plant in comparative terms with the restof the VC, which has not received the same level of recorder man-hours.
British and Irish occurrence
The New Atlas describes and shows that P. hydropiper is widespread in both Britain andIreland, although it is rather rare in C Ireland (previously the main area for raised bogs, and withsoil underlain by a rock skeleton of Carboniferous limestone). In Britain, the species has a distinctlywestern trend in its distribution as one goes northwards from the English Midlands and on up intoScotland, where it gradually thins out and disappears to the east and then the north (Preston etal. 2002).
European and world occurrence
A native of Eurasia, P. hydropiper is widely distributed in middle latitudes across Europe,thinning noticeably towards both the north and the Mediterranean. It is not uncommon in southern partsof Norway, Sweden and Finland and it penetrates to a couple of sites within the Arctic Circle (Timson1966; Jalas & Suominen 1979, Map 404). The distribution extends east into C Asia and it is thinlyscattered as a naturalised alien through many areas of China, Japan, SE Asia, S Australia and NewZealand, presumably introduced as an impurity with crop seed (especially with clover) (Hultén &Fries 1986, Map 649).
The status of the species in N America is uncertain (Timson 1966). Hultén (1974, Map 209) discussed thequestion and decided it needed a monograph study to resolve it. Hultén & Fries (1986) suggest thereis a mixture of native and introduced forms of the species spread across N America, but the verywidespread alien occurrence of P. hydropiper elsewhere in the world, renders this ideaunconvincing.
Uses
As its peppery taste indicates, P. hydropiper contains a sharply acrid, acidic sap that causesirritation and inflammation to the skin, and to the gut if it is ingested. However, there are no recentreports of poisoning from this source (Cooper & Johnson 1998). In the past, P. hydropiper wasa valued plant used in herbal medicine (Grieve 1931), and recently published biochemical studies, mainlyfrom the Far East, supports the notion that it has useful healing properties.
European herbalists regarded it as a stimulant, a diuretic (induces increased urine), diaphoretic(induces sweating), emmenagogue (induces menstral flow) and it arrests bleeding. In Scotland, it wasalso used to treat dysentery, flatulence and piles as well as gravel in the kidneys or bladder. It wasused for treating minor cuts and bruises and ailments such as colds, coughs, toothache and herpesulcers, as well as for more serious complaints such as epilepsy, gout and gangrene (Launert 1981; Darwin1996). In Bangladesh, leaves of the plant are traditionally used in the treatment of rheumatic pain,gout, and for skin diseases such as ringworm, scabies, boils, abscesses, carbuncles and the bites ofsnakes, dogs and insects.
The juice of the plant is a common remedy against body lice in cattle and sheep, and it is also used asan insect repellent. Yellow and gold-coloured dye can also be obtained from the chopped plant (Darwin1996).
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The Latin specific epithet'hydropiper' is a combination of the Greek 'hydor', 'water', and Latin 'piper' meaning 'pepper' andhence the most commonly used English common name, 'Water-pepper', is a straight translation of thebotanical species name.
Alternative common names include 'Marsh-pepper', 'Pepper Plant', 'Biting Persicaria', 'Bity Tongue','Smart-weed', 'Smartass', 'Arcmart', 'Hot Arsmart', 'Arsesmart', 'Red Knees', 'Bloodwort', 'Ciderage'and 'Culrage' (Grieve 1931, p. 743; Darwin 1996). As Grigson (1987) quotes John Minsheu (1617) Ductorin Linguas, it is 'Arsesmart' "because if it touch the taile or other bare skinne, itmaketh it smart, as often it doth, being laid unto the bed greene to kill fleas".
Threats
None.
Native or possibly a quite recent neophyte; rare, but readily subject to mis-identification. Europeantemperate.
1988; NI Lakes Survey; Derrykerrib Lough, Upper Lough Erne.
July to August.
Growth form, identification and preferred habitats
A summer annual very similar to P. hydropiper (Water-pepper), but leaves not wavy margined andwithout the sharp (but often slowly developing) taste of that species, and the inflorescence nearlyerect or slightly nodding (not drooping), usually redder (pink to purplish-pink) than P.hydropiper rarely greenish white. Glands on the flower perianth and the peduncle (ie theinflorescence stalk) are smaller, flatter and rather sparse (only c twelve per flower) in comparisonwith P. hydropiper. Other distinctive features that separate the two species are bristles at thetip of the stipules >3mm (not <3 mm), and the achenes are 3-4.5 mm, biconvex, ± shiny (not matt).Ripe fruit are almost essential in order to successfully separate out the riparian Persicariaspecies (Clapham et al. 1987; Parnell & Simpson 1988; Rich & Jermy 1998; Stace 2010; Sell& Murrell 2018).
In genetic terms, P. mitis (2n=40) is an autopolyploid derivative of P. hydropiper (2n=20),and therefore unsurprisingly they are very similar in their ecology. P. mitis is a plant of wetor damp, nutrient-rich ground near lakes, ponds, streams and ditches and in wet hollows in pastures(Parnell & Simpson 1988). It also appears as a pioneer colonist of nutrient-rich or manure-pollutedmud or fen peat left exposed in dry summers when water levels in ditches, ponds and lakeshores is drawndown. It does not appear to display any particular preference in terms of soil reaction (J.O. Mountford,in: Stewart et al. 1994).
By themselves, the habitat conditions do not account for the rarity of this species, which seems to bemore concerned with correct determination of critical plant characters. At the present time (April2018), P. mitis remains rather poorly and unreliably recorded, due to its close similarity toP. hydropiper.
Fermanagh occurrence
All 20 records for this annual species in the Fermanagh Flora Database are from the NI Lakes Survey(1988-91), but since P. mitis (= P. laxiflora (Weihe) Opiz = P. mite Schrank) isregarded as a very rare species in Ireland, and it is readily confused with otherPolygonum/Persicaria species, the identifications of all of the records really do still requireexpert determination. The Fermanagh records for P. mitis lie in a total of 15 tetrads, themajority of which are from the shores of both parts of Lough Erne. The remaining five lake sites areKeenaghan Lough, Parabaun (or Finnauan) Lough, Ross Lough near Carr Bridge, Lough Bresk near Lisnarrickand Watsons Lough.
It appears that only two voucher specimens were deposited in BEL to support the Fermanagh recordsof P. mitis, and Paul Hackney, when herbarium curator, reckoned one of them was incorrectlyidentified and required re-determination. Although considerable doubt is thus cast on all the NI LakesSurvey identifications of this taxon, in a case like this the neglect is tempered by the fact that thestudy was carried out by competent scientists who would (or should) have been aware of the relevant andimportant work carried out in NI towards distinguishing this species from its relatives. The relevantstudies were published by Webb (1984) and by Parnell & Simpson (1988).
Irish occurrence
In his review of all the available Irish herbarium material, Webb (1984) defined several distinguishingcharacters and examined how it was that mistakes had been made by the experienced botanists concerned.Webb concluded with regret that all the pre-1969 Irish records were unreliable, and that most of themwere wrong. The plants in question instead proved to be P. minus, P. hydropiper or P.persicaria.
The reasons for the mis-identifications were undoubtedly associated with poor choice of reputedlydiagnostic characters by taxonomists of a pervious era, plus inadequate description in many or all ofthe published Floras of the time, together with the usual taxonomic problems and confusions surroundingdifficult or critical plant groupings. P. mitis, P. minus and P. hydropiper allshow a wide range of phenotypic variation in response to environmental factors, and P. maculosa(Redshank) and P. lapathifolia (Pale Persicaria) can also be difficult to separate from theothers, particularly from P. mitis. Webb (1984) suggested that P. mitis was most probablyintroduced to Ireland fairly recently, perhaps during World War II (although this could hardly beproven). He concluded that the earliest reliable records for the species in Ireland were John Harron'spost-1968 finds in the Lough Neagh basin (Kertland & Lambert 1972; Harron 1986).
In their Lough Neagh study, Parnell & Simpson (1988) found that P. maculosa and P.lapathifolia were in fact readily distinguishable from the three other species; the denser spikeand small black patches on the leaves of P. maculosa, and the possession of both these featuresplus pedicel glands in P. lapathifolia, were perfectly distinctive. Parnell & Simpson (1988)made a detailed numerical analysis of 17 morphological characters to compare P. mitis, P. minusand P. hydropiper from Lough Neagh, and they devised a table of the nine characters which bestseparate the three species.
Previous work by Webb (1984) and by Lousley & Kent (1981) had indicated that the possession of glandson the perianth of P. hydropiper is one of the key characters allowing its discrimination fromboth P. mitis and P. minus. However almost all the P. mitis plants Parnell &Simpson examined also possessed perianth glands, which on close examination proved to be smaller, fewerin number and almost flat in comparison with the more numerous and more prominent raised glands on P.hydropiper.
The same situation was found with respect to perianth glands when British and European P. mitisplants were examined, a feature ignored in standard works on the genus. As is often the case, acombination of characters needs to be compared when attempting to separate these three similar species(Parnell & Simpson 1988).
Hybrids
To add to identification difficulties there are intermediate hybrids formed with P. maculosa(Redshank), P. hydropiper and P. minor (Small Water-pepper), although they all are eitherrare, very rare, absent or in need of confirmation in Britain and Ireland (Stace et al. 2015;Sell & Murrell 2018).
British occurrence
Stace (1997, 2010) and Sell & Murrell (2018) regard P. mitis as native in Britain, where theplant is described as being, "rare and very scattered over England, Wales and NE Ireland".Stace (1997, 2010) is of the opinion that it is over-recorded. The New Atlas treatment addslittle or nothing to the above, but recognises that it is a scarce species.
European and world occurrence
P. mitis is a native of temperate areas of W & C Europe, and from the publishedmap in Jalas & Suominen (1979), Map 403, appears to be mainly confined between 40° and 55°N, thedistribution thinning markedly both towards the Mediterranean and the Baltic. P. mitis does notfeature at all in Jonsell et al. (2000). P. mitis is shown as an Eurasiatic species byHultén & Fries (1986, Map 648), although they describe their map as tentative because ofdifficulties in identification of the species. Clapham et al. (1987) also mention P. mitisoccurring in W Asia.
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The Latin specific epithet'mitis' means 'mild', 'mellow' or 'bland' (Gilbert-Carter 1964), and clearly refers to the absence ofthe peppery taste of P. hydropiper, the more common species with which it is most likely to beconfused.
The English common name 'Tasteless Water-pepper' is an excellent example of an invented, so-called 'bookname', given to a plant with absolutely no folk associations attached to it.
Threats
None.
Native, occasional or locally frequent. Eurasian temperate, introduced in N & S America and otherareas.
1884; Barrington, R.M.; SE of Muckros Point, near Kesh Water Foot, Lower Lough Erne.
June to October.
Growth form and preferred habitats
P. minor is a rather variable wetland summer annual that resembles a smaller, straggling, morediffuse, more branched, rather decumbent form of P. maculosa (Redshank), with leaves that neverhave black blotches. In common with most other Persicaria species, P. minor is a weaklycompetitive summer annual, confined to ± sunny, open, somewhat disturbed, or man-made, damp, ± enrichedhabitats in lowland areas. The typical site in our area is a lake shoreline, trampled, poached andmanured by periodically visiting thirsty cattle. In Ireland, Scotland and N England, this generallyscarce or rather rare, phenotypically variable yet reasonably distinctive little annual occurs on open,gravelly or sandy marshy ground liable to intermittent flooding, ie on ground that lies above thenormal water levels of lakes, ponds and ditches. Around lakes that are used as reservoirs, it canadditionally occur in the draw-down zone, ie in intermittently exposed ground usually belownormal water level (Lousley & Kent 1981; J.O. Mountford, in: Stewart et al. 1994).
Several closely related Persicaria species have overlapping riparian (waterside) habitattolerances. In the survey results entitled Scarce plants in Britain, J.O. Mountford remarked howin S England and in Wales P. minor is confined to the more muddy, drawn-down water level type ofterrain. This riparian habitat is sometimes colonised by three closely related Persicaria species(P. hydopiper (Water-pepper), P. minor (Small Water-pepper) and P. mitis (TastelessWater-pepper)), and rarely by up to six of them (ie additionally P. amphibia (AmphibiousBistort), P. maculosa and P. aviculare (Knotgrass)). Mountfield also noticed that in thearea he studied, P. minor tends to occur in slightly more acidic and more nitrogen-richsubstrates than P. mitis (J.O. Mountford, in: Stewart et al. 1994). However, in Ireland,P. mitis is too rare a species, and its local ecology has been too little studied to allow validspecies habitat predictions and comparisons of this type to be made. The species habitat comparisonsoffered in Stewart et al. (1994) tend to reflect a southern English perspective, and until scarcespecies are further investigated locally, they should probably be applied with reservations by botanistsworking in Ireland and Scotland.
Ideally for accurate identification, waterside Persicaria plants should be checked carefullyagainst the nine characters delimited by Parnell & Simpson (1988), particularly to see if anyspecimens can be referred to the much rarer P. mitis (= P. mite = P. laxiflora)(Tasteless Water-pepper). This involves looking at the fruit (a single seeded nut or achene) for length,breadth and surface shininess; perianth glands (shape (raised or flat) and number); perianth length;flower colour; the extent to which the inflorescence spike nods or does not; and the length of teeth onmid-stem ochrea (J.R. Akeroyd & T.C.G. Rich, in Rich & Jermy (1988); Parnell & Simpson1988). A useful distinguishing point for P. minor is that the ratio of leaf length:breadth is upto 8.5, whereas the ratio for P. hydropiper and P. mitis is up to 4.5 to 4.8 times as longas broad (Parnell & Simpson 1988). Having said all this, Lousley & Kent (1981) regarded P.minor as "a distinct little plant not easily confused with other species."
Reproduction
P. minor reproduces entirely by seed, flowering taking place from July until October. The slender,spike-like inflorescence is ± erect in comparison with P. hydropiper and P. mitis,both of which have definitely lax, nodding flowering branches and the flowers are typicallyreddish (sometimes described as crimson), although flower colour is very variable and it can be white(Lousley & Kent 1981). Styles are usually two or occasionally three and the species isself-fertilising (autogamous), requiring no pollinator. The nut-like fruit is a single-seeded achene orseed which is lenticular (biconvex) or rarely trigonous (three-sided) in shape, with a shiny (not matt)surface, varying from brownish-black to pure black in colour (Sell & Murrell 2018).
Variation
In common with all Persicaria species, there is a wide range of phenotypic variation inP. minor, with respect to leaf width (0.2-2.3 cm), flower colour (red, pink or sometimeswhite), and nut (achene) size (1.5-2.7 mm). In the view of Jonsell et al.(2000), most of this variation is probably genetically determined.
Two distinctive variant forms of P. minor are taxonomically recognised. Var. minor isan elegant, decumbent plant with stems up to 25 cm, long-branched from the base and withleaves mostly narrowly linear; the other form is now referred to as var. latifolia (A. Braun)Akeroyd. It is taller (up to 60 cm) and more erect, with short branches up the stem and broader leavesthat are described as narrowly elliptical or linear lanceolate (Sell & Murrell 2018). This is theform that is most likely to be confused with P. mitis.
Hybrids
Although P. minor habitually selfs, and its pollen like that of other Persicaria species is'sticky' and produced in small quantity, nevertheless very rarely it can produce intermediate hybridsoccasionally with P. maculosa, rarely with P. mitis (five samples from Lough Neagh), andeven more rarely with P. hydropiper. In the latter case (P. ×subglandulosa (Borbás) Soják), confirmation of the hybrid in Britain and Ireland is required,although there are a few records of it from mainland Europe (Parnell & Simpson 1988; Stace etal. 2015).
Fermanagh occurrence
In Fermanagh, we regard P. minor as an occasional, or even a locally frequent species, since ithas been recorded over a hundred times in as many as 50 tetrads, 9.5% of those in the VC. It is chieflyfound in either muddy fen peat, shingle or sand around both parts of Lough Erne, but additionally it hasbeen recorded on the shores of eleven smaller lakes scattered both N and S of the main lake basin. Inthe past, these included one or more limestone turloughs (ie the 'Green Loughs' near Fardrum). P.minor has also been recorded just once in a sandy quarry adjacent to one of these minor lakes(Lough Keenaghan). It is interesting that only nine of the records in the Fermanagh Flora Database arepre-1975.
Irish occurrence
In N Ireland, as far as we know, P. minor occurs largely around the two major lake systems ofLough Neagh and Lough Erne, but it is also present on the gravel, sandy or muddy shores of some smallerlakes and ditches in the N Down area (NI Vascular Plant Database 2006). In Ireland as a whole, P.minor has been recorded at least once from 23 of the 40 VCs. The New Atlas map showsSmall Water-pepper is mainly found in N Ireland (Fermanagh, Armagh, Down & Antrim, H33 & H38-40)and thinly scattered down the west of the island to Cos Clare and Cork (H9 & H4-5) (Preston etal. 2002).
British occurrence
P. minor is widely, but thinly scattered throughout most of England and Wales, with a greater frequencyin Cumbria. In Scotland, it is well represented in the SW in Dumfries, Kirkcudbright, Wigtown & Ayr(VCs 72-75), and quickly thins out northwards, although it reaches outliers in Main Argyll (VC 98) andEast Perth (VC 89) (Preston et al. 2002). Although the taxonomic problems associated withPersicaria species have made recording of this species group difficult in the past, there isevidence that P. minor has declined in recent years in S England. This follows a decline in thenumber of suitable farm ponds and ditches and greater regulation of water levels, which together havereduced the extent of the wet mud habitat the plant colonises (J.O. Mountford, in: Stewart et al.1994).
European and world occurrence
P. minor is widespread in Europe between 45°N and 65°N, but absent from the arctic and from mostof the Mediterranean region – although it does reach Tuscany and N Greece (Jalas & Suominen 1979,Map 401). It extends eastwards into E Asia and is a rare introduction in N & S America, S Africa andSri Lanka (Hultén & Fries 1986, Map 646).
Names
The genus name 'Persicaria' is from the Latin 'persicum' meaning peach, and translates as either'peach-leaved' (Gilbert-Carter 1964), or 'peach-like' (Gledhill 1985). The Latin specific epithet'minor' means 'smaller', which is reflected in the English common name 'Small Water-pepper'. The latteris another perfect example of a so-called 'book name', as it is purely invented for convenience andbears no burden of historic folk lore usage.
Threats
None.
Introduced, neophyte, a rare casual, or a relict of 19th century cultivation. Locally very rare, butpossibly a mis-identification. Eurasian temperate.
17 July 1988; Northridge, R.H.; Gublusk Bay, Lower Lough Erne.
Growth form, origin and preferred habitats
Buckwheat is a tall or medium-sized (20-60 cm in height), erect, little branched, hollow-stemmed, pink orwhite flowered annual. It is a native of temperate E Asia, and wild forms are found in SW China (Yunnan)and Siberia. As a non-grass cereal crop, it was introduced into Europe from China in the Middle Ages,reaching Holland and Germany early in the 15th century. From there it spread with agriculture and wascultivated for several centuries in England, France, Spain, Italy and Russia. Apart from China, it alsofound a place in the agriculture of Japan, India, Africa, Brazil and Australia, and was one of the firstcrop plants introduced to the American Colonies where it continues to be grown (C.G. Campbell, in:Simmonds 1976; Vaughan and Geissler 1997).
Buckwheat was quite widely sown in Britain and Ireland for poultry and game bird fodder from the 16thuntil the early 20th century. Nowadays, F. esculentum is much more commonly grown in other partsof the world, particularly in Russia (50% of the world crop), China (17%) and in N America wherebuckwheat flour is still used to make pancakes (Allison & Day 1997).
The similarity of the blackish brown or grey nut-like achene to the much larger beech nut, both fruitsbeing sharply triangular in section, and its use for bread flour, gives rise to the German name,'Buchweisen', which translates as 'Beechwheat'. The English common name 'Buckwheat' is a corruption ofthis translation (Salisbury 1964).
Since the crop is a seed and it can be used to make gluten-free flour, Buckwheat is described as apseudo-cereal crop. The discovery of gluten allergy in many people in western society in recent yearshas enhanced interest in and growth of the crop as a suitable dietary replacement for sufferers of thisdebilitating condition. F. esculentum grows well in nutrient-poor, light or sandy, acid soils oflow fertility, always provided they are well drained. In hot climates, it can only be sown late in theseason so that it blooms in cooler weather. It produces most seed when pollinating insects areplentiful. These properties, together with the high nutritional value of the crop (protein content 18%),made it popular and widely grown around the world for centuries.
Buckwheat grows quickly, has a growing period of just 10-12 weeks, and apart from its use as apseudo-cereal, can be used as a cover crop to overgrow and smother weeds, eg in vineyards, or toconserve soil in areas subject to erosion. It can also be used as a green manure, being ploughed intothe soil to improve fertility and moisture retention. Another use for which Buckwheat is deliberatelysown is to provide protective cover for pheasants and other game birds, since the plants in addition toshelter also provide food in the form of seed later in the growing season (Stace & Crawley 2015).
Reproduction
The inflorescence consists of spike-like, cymose clusters of 4-6 small flowers borne on peduncle stalksin the axils of the upper leaves. The flowers, which become more congested towards the top of the stem,are heterostylous, like the Primrose (Primula vulgaris), having 'pin' and 'thrum' types,depending upon the length of the style and the position of the stamens. An unusual and rather curiousfeature is that in F. esculentum, the thrum-eyed flowers are noticeably larger than the pin-eyed(Jonsell et al. 2000). Heterostyly is unusual in annual plants, and doubly so in a seed cropplant (Stace & Crawley 2015). Heterostyly is an obvious outbreeding mechanism, the flowers beingtotally self-incompatible. The necessary pollinating insects (which include honey bees) are attracted bythe greenish-white, pink or rarely bright red or deep pink flower perianth and nectar from globularnectaries arranged between the inner and outer rings (whorls) of the eight stamens. The nut-like fruitis 5-6 mm long, 2-3 times longer than the perianth (Clapham et al. 1987; Sell & Murrell2018).
Some years ago plant breeders managed to create hom*ostylous, highly self-compatible, diploid strains ofBuckwheat which obviously have the potential to greatly increase seed yields of the crop (C.G. Campbell,in: Simmonds 1976; Stace & Crawley 2015).
Variation
There are rare variants with dark pink or bright red flowers. Other forms have fruits with silvery greysurfaces (Jonsell et al. 2000).
Fermanagh occurrence
There is just one record in the Fermanagh Flora Database made by RHN, but since there is no voucherspecimen to support it, some doubt must attach to the identification.
Irish occurrence
The New Atlas map shows just three post-1987 records in Irish hectads in N Tipperary, Fermanaghand Tyrone (H10, H33 & H36). The only other Irish records mapped in any of the BSBI plant atlasesare in three hectads near Belfast, all pre-dating 1930 (Perring & Walters 1962, 1976; Preston etal. 2002).
British occurrence
Buckwheat seed is still imported into Britain and Ireland and is used as feed for poultry and pheasants.In parts of Britain, a little Buckwheat may still be sown as a green manure and as a bird- andbee-fodder plant (J.R. Akeroyd, in: Preston et al. 2002). Cultivation is only on a very smallscale in Britain and there does not appear to be any at all in Ireland. Buckwheat still occurs, however,as a component of wild bird seed mixtures that are increasingly used in gardens to attract avianvisitors, especially during the winter months.
F. esculentum is occasionally found in the wild in Britain on waste ground, especially in themidlands of England south of a line between Liverpool and Hull (Preston et al. 2002). Nowadays,it only appears either as a casual from bird seed in garden borders or parks, or as a relict ofcultivation on field margins (Reynolds 2002). In either situation, its occurrence is erratic and itusually does not persist long (Clement & Foster 1994).
Toxicity
Buckwheat contains a pigment called 'fa*gopyrin' that is present in both fresh and dried plants. When fedto farm animals, and particularly chickens, fa*gopyrin is believed to induce photosensitivity tosunlight. When the black seeds are dehusked, however, they are considered harmless (Cooper & Johnson1998).
Uses
The protein of Buckwheat seed and flour is of excellent quality and it is also rich iniron, zinc, selenium, vitamin B and dietary fibre. Apart from the above mentioned uses as animal fodderand flour for pancakes, dehulled Buckwheat achenes are cooked as porridge and the flour is used inbiscuits, noodles, cereals and as a substitute for other grains (C.G. Campbell, in: Simmonds 1976).
Names
The genus name 'fa*gopyrum' is a combination of the Latin 'fa*gus' meaning 'beech' and Greek 'pyros','wheat'. The Latin specific epithet 'esculentum' means 'edible' or 'good to eat' (Stearn 1992). TheEnglish common name 'Buckwheat' is a mistranslation of the German name 'Buchweisen' or the Dutch'Boekweit' (Gilbert-Carter 1964). Alternative names include 'Common Buckwheat', 'Japanese Buckwheat' and'Silverhull Buckwheat'.
Threats
Apart from the toxicity of the whole plant (fresh or dried), rather than the dehusked seed, there is noreal threat from this species. However, people who suffer from food allergies would be wise to avoid it.
Introduction, archaeophyte, locally frequent. Eurasian wide-temperate, but also widely naturalised.
1937; Praeger, R.Ll.; Co Fermanagh.
June to December.
Growth form, identification and preferred habitats
This small stress-tolerant ruderal summer annual is similar to P. aviculare s.s. (Knotgrass), butit is always prostrate or procumbent, rather than erect. It has a strong, deep taproot sometimesreaching down to around 40 cm, and its wiry stem is often much branched, allowing it to form a densesurface-spreading mat (Salisbury 1964). P. arenastrum has evenly-sized leaves, whereas those ofP. aviculare s.s. noticeably decrease in size towards the tip of the shoot. Both knotweed specieshave a long flowering period stretching from July to November and both are mobile, well-adapted, pioneercolonists of cultivated and disturbed open habitats, occupying a wide range of moderate soils of acid toneutral reaction (pH usually above 5.0 according to Grime et al. 1988). They both show preferencefor moderately fertile, damp to dry, often winter-wet sites and full sun situations. Unlike many otherannuals, both of them have the ability to recover after early season damage, including grazing pressureand trampling.
P. arenastrum is typically found in dry or well-drained, often sandy, gravelly or stony soils inwayside and waste ground habitats which are open and disturbed. It can survive, and indeed even thrive,in very compacted, heavily trampled surfaces, eg on paths, around the sides of yards, in gateways,between paving stones and in cracks in concrete or asphalt, often in droughted, dwarfing conditionswhich P. aviculare s.s. simply could not tolerate (Lousley & Kent 1981; Sell & Murrell2018).
Both Knotgrass species also commonly occur as successful weeds of arable agriculture and horticulture,preferring fertile, damp, loose, recently disturbed soils, newly sown grassland, or disturbed orovergrazed meadows and pastures where patches of bare earth are present. They may also be presenttogether in much more ruderal, artificial, man-made habitats, eg in waste ground, rubbish tips and alongthe kerbsides of busy roads, tolerating the spray and slipstream buffeting provided by passing roadtraffic. Presence of these two species in coastal situations and along roadside kerbs immediatelyadjacent to metalled surfaces is strong evidence of their salt-tolerance (Grime et al. 1988).
Reproduction
Polygonum species are typically inbreeding, although in Scandinavian countries the flowers areoften wide open, and at least the larger flowered species are occasionally visited by flies and othersmall insects so could from time to time be cross-pollinated (Jonsell et al. 2000). Normally,however, seed production in all species of the genus is assured by habitual self-fertilization, and arobust plant can produce up to 1,000 relatively large seed. The seeds (often referred to as nuts orachenes), are dispersed by both man and animals, eg in mud, with harvested crops, as an impurity ofa*gricultural seed, or they may be ingested and transported internally by both domestic stock and wildbirds (Ridley 1930).
The flowers of P. arenastrum are borne in 3-7 flowered cymes on the upper parts of stems andbranches from July to September. The perianth is 2-4 mm, the five tepals laterally overlapping, butforced apart as the fruit grows. The tepals are green with wide white margins and they usually containthe nut fully, although the tip may be visible. The three-sided or biconvex nuts, 2.5-3.0 mm, areblackish brown, the faces almost smooth or coarsely papillose.
Variation
The high degree of inbreeding found in this genus leads to the formation of numerous infraspecific racesand ecotypes within it. In taxonomic terms, the number of useful morphological identification charactersis limited, and when this is combined with polyploidy and great plasticity, the taxonomic limits betweenforms are often obscured (Jonsell et al. 2000).
Fermanagh occurrence
In Fermanagh, P. arenastrum is widely but thinly scattered throughout lowland areas, having beenrecorded in 81 tetrads, 15.3% of those in the VC.
Irish occurrence
The New Atlas hectad map shows P. arenastrum is widespread throughout most of Ireland,particularly in the N and in the region around Dublin plus in the extreme S, but more patchily dispersedelsewhere.
Status in Britain and Ireland
The editors of the New Atlas regard P. arenastrum as an ancient introduction (anarchaeophyte), while they take the ecologically very similar P. aviculare s.s. to be indigenous.It is not obvious why they make a status distinction between the two. The fossil diagram and associatedtext for P. aviculare s.l. in Godwin (1975), indicates that a gap occurs in the fossil record inthe current interglacial period (called the Flandrian in Britain and the Littletonian in Ireland), untila sudden expansion of fossils (both fruits and pollen) occurs in close association with archaeologicalsituations in the Bronze Age. In a paper subsequent to publication of the New Atlas, Prestonet al. (2004) remark that the authors of most accounts of the fossil record do not distinguishbetween the segregates of P. aviculare (ie palaeoecologists like Godwin treat it as sensu lato).Nevertheless, Preston et al. (2004) single out P. arenastrum and label it an ancientintroduction. Both these species, and P. aviculare s.s. in particular, have a long history ascommon cornfield and broadleaf crop weeds (Styles 1962). The current author (Ralph Forbes) considersthem as equally probable ancient archaeophytes which accidentally and repeatedly have been transportedto these islands amongst crop seed by farmers from the Neolithic period or the Bronze Age, onwards.
For comparison, in the critical account of these taxa given in Flora Nordica, Jonsell etal. (2000) treat these two segregates of P. aviculare s.l. as subspecies. Subsp.aviculare is regarded as indigenous on the coasts of Denmark, S Norway and W & S Sweden,while inland it is closely associated with man and is considered a probable archaeophyte, except in thenorth. The status of P. arenastrum (as subsp. microspermum (Jord. ex Boreau) Berher) isgiven as, "anthropochorus in Norden" (ie associated with man throughout the whole surveyarea), and "probably archaeophytic, except in the north." (Jonsell et al. 2000).
European and world occurrence
The distribution is imperfectly known, but P. arenastrum is regarded as probably common throughoutcontinental Europe, extending well into northern regions of Scandinavia where it is an archaeophyte(Jalas & Suominen 1979, but not mapped; Jonsell et al. 2000). It is fairly common as anestablished alien in N America. Hultén & Fries (1986) map the Polygonum aviculare complex ofaround 50 taxa as circumpolar and very widely introduced worldwide (Map 645).
Names
The genus name 'Polygonum' is a combination of two Greek words, 'polys', many or much, and either'gonos', meaning offspring or seed, an allusion to the numerous seeds characteristic of the plants, or'gony', knee-joint, an allusion to the swollen joints of the stems. The Latin specific epithet'arenastrum', means 'resembling Arenaria', from 'arena', 'sand', an allusion to sandy placeswhere many of these species prefer to grow (Stearn 1992).
Threats
None.
Probable introduction, archaeophye, common and widespread. Circumpolar wide-temperate, but very widelynaturalised in both hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
June to November.
Growth form and preferred habitats
The P. aviculare species agg., or P. aviculare s.l. is genetically very variable andphenotypically extremely plastic with respect to environmental growing conditions. British material ofthis weedy ruderal annual was split into six separately named, geographically and/or ecologicallydistinct taxa by Styles (1962), four of which are accepted today. Along with the species aggregate theseare separately mapped in the New Atlas. Only two of these segregated species have proven to becommon and widespread throughout most of Britain and Ireland and they alone are known to be present in NIreland, ie P. aviculare s.s. and P. arenastrum (Equal-leaved Knotgrass).
Fermanagh occurrence
The majority of recorders operating in Fermanagh since Styles's study have regularly distinguished P.arenastrum from P. aviculare agg. However, with just one solitary exception (a NILakes Survey record), Irish field botanists (including ourselves), for reasons that are not immediatelyobvious, have continued to field record this weedy little annual in the broad sense, listing it as,'P. aviculare agg.' or 's.l.', rather than 'P. aviculare s.s.'.
The aggregate or broad species taxon thus appears to combine records of both these weedy summer annualsand as a result the broad combination is more commonly and more widely recorded in Fermanagh than theonly distinguished constituent species, P. arenastrum. At the time of writing (December 2010),the Fermanagh Flora Database incorporates records of the broad taxon, Polygonum aviculareagg. or s.l., from a total of 127 tetrads (24.1%) widely scattered in the lowlands, while forP. arenastrum there are records listed from 81 tetrads.
Reproduction
Although by definition P. aviculare s.s. and P. arenastrum are distinguishable, they arerather similar in both their form and biology. Furthermore, while they differ slightly but significantlyin their ecological preferences and tendencies, they share an essentially ruderal, stress-tolerant,pioneer colonist nature. Consequently, and unsurprisingly, their distributions often overlap, formingmixed populations in open, disturbed more or less fertile ground. Despite this overlap and closephysical proximity, hybrids do not occur between the two forms. The tiny bisexual flowers of both ofthese small, weedy annuals fail to attract insect visitors and they therefore habitually self-fertilise.
British and Irish occurrence
The New Atlas hectad distributions for P. aviculare s.s. and P. arenastrum inBritain are remarkably similar in both frequency and pattern. In Ireland, P. aviculare s.s. isless well distinguished and recorded than could be expected in comparison with P. arenastrum.Thus P. aviculare s.s. appears patchy and sparse or absent in large areas of C Ireland and inparts of the west (notably the Burren, Connemara and Donegal). No obvious climatic or soil factor seemsto explain such a pattern, so it might reflect some degree of under-recording.
The New Atlas editors map not only the four segregated British species, but also include P.aviculare agg. This is shown to be virtually ubiquitous in lowland Britain and Ireland, withjust a few missing hectads scattered in C & W Ireland and in NW Scotland.
European and world occurrence
Since the species aggregate includes forms like P. aviculare s.s. which are significant arableweeds of both cornfield and broad-leaved crops, it is not surprising that it has travelled around theglobe with man into far distant lands, including New Zealand, the South Sea Islands, S America and C& S Africa (Hultén & Fries 1986, Map 645). The spread of the plant might also be partially dueto its use and reputation in herbal medicine, since the fibrous roots were used as a quinine substitutein N & C Africa, and the seeds are emetic and cathartic (Mitich 1998).
Names
The genus name 'Polygonum' is a combination of two Greek words, 'polys', many or much, and either'gonos', meaning offspring or seed, an allusion to the numerous seeds characteristic of the plants, or'gony', knee-joint, an allusion to the swollen joints of the stems. The Latin specific epithet'aviculare' means 'of or pertaining to small birds' or 'eaten by small birds' from ' avicula',diminutive of 'avis', bird, as Sparrows and finches feed on the fruits (Gilbert-Carter 1964; Gledhill1985).
The English common name 'Knotgrass' was coined in the 16th century by Turner (1538) who alsocalled it 'Swyne grys' (ie Swine's grass) as it was used in herbal medicine for ulcers and other sores,and it was, according to Gerard (1597), fed to pigs "when they are sicke, and will not eat theirmeate." Grigson (1987) lists no less than 30 English common names from around Britain and Irelandfor this familiar plant.
Threats
None.
Introduction, neophyte, a frequent, widespread and locally dominant garden escape or discard. Native ofJapan and the Far East, widely introduced and established in Europe and N America.
12 June 1974; Hackney, P.; Lower Lough Macnean, near Belcoo.
April to October.
Growth form and preferred habitats
Of far-eastern origin (ie Taiwan & N China), F. japonica is a tall, stout, erect,clump-forming dioecious perennial first introduced to gardens in Britain and Ireland in 1825, from whereit first 'escaped' into the wild around 1886. Apart from a rarer compact form (see below), it appearsthat all the F. japonica plants grown in European gardens derive from a single Dutch import fromJapan made in the 1820s. A detailed history of the introduction and spread of F. japonica, F.sachalinensis and their hybrid F. × bohemica has been provided by Bailey &Conolly (2000). F. japonica is designated a noxious weed and it is an offence to allow it toescape into the wild.
Japanese Knotweed is now regarded by many as the most aggressive introduced herbaceous plant in the floraof Britain and Ireland. It is widespread and well established throughout these islands in lowland,man-made, disturbed or linear habitats, especially by flowing water (Grime et al. 1988; Charter1997; Hollingsworth & Bailey 2000; J.R. Akeroyd, in: Preston et al. 2002). The Fermanaghsites closely fit this pattern of occurrence in neglected ground on wayside, waterside and woodlandmarginal areas, including in landed estates. It also grows on many types of artificial man-made habitatsand generally appears on unmanaged, disturbed ground including beside railway tracks and in oldquarries. The species is described as light-demanding and salt-tolerant (Jonsell et al. 2000). Itis, however, sensitive to both frost and drought.
The overwintering organs of F. japonica lie below ground level in the form of a thick rhizome androots. The fleshy, bamboo-like aerial parts of the plant can reach over 2.0 m in height, but they diedown and disappear each year, apart from a minority of standing dead stems. Nevertheless, whenintroduced to new ground the plant very quickly establishes and develops a laterally creeping, deeplypenetrating, thick, perennating rhizome. In spring, this underground foodstore enables very rapid growthof both root and stem, producing vigorous, spreading, dense leafy clumps. A large dense canopy developswhich provides the good nutrient stocking that underpins the high competitive growth strategy that isstrongly characteristic of the species (Grime 1979; Schnitzler & Muller 1998).
Reproduction
Creamy-white female flowers with sterile staminodes are borne in tassel-like panicles up to 15 cm long inthe axils of upper leaves in late summer and early autumn. Male-fertile plants of var. japonicaare unknown in the British Isles (Beerling et al. 1994), and thus the plant spreads entirely byvegetative means. The dispersal of Japanese Knotweed is closely associated with linear landscapefeatures, eg waterways and railways (the latter now largely defunct in much of rural Ireland), and morerecently along roadsides to which it has spread through the all too common unauthorised hedgerow dumpingof rhizome-infested garden waste (ie 'fly-tipping'). The legitimate trade in transported topsoil is alsoknown to have spread the species, some loads being contaminated with the plant. Tiny fragments ofrhizome as little as 8 gm fresh weight are sufficient to allow regeneration, and under suitableconditions cut stems also can produce adventitious roots within a few days.
Hybrid Knotweed
Populations of F. japonica generally contain both male-sterile and hermaphrodite individuals, asis also the case with its much larger-leaved close relative, F. sachalinensis (Giant Knotweed).When these two knotweeds occur together they frequently hybridise, and their intermediate product (F.× bohemica) forms viable seed. Although F.sachalinensis is decidedly rare in Ireland, it is very possible that some of the many standsrecorded as F. japonica really belong to this hybrid, but are mistakenly identified as the morefrequent parent species.
Variation and taxonomic nomenclature
In its native area of East Asia (Japan, China, Taiwan and Korea), F. japonica is very variable andis often a pioneer colonist of bare volcanic soils. In Europe, however, very little of this variation isrepresented in the introduced material (Jonsell et al. 2000).
Two varieties of F. japonica are described (var. japonica and a much smaller dwarf, var.compacta), but the two have not been distinguished in Fermanagh. The dwarf form is the preferredgarden subject, but it is also capable of becoming so very well established it may offer a threat to theground it occupies.
Apart from the type race var. japonica and var. compacta there is a further very tallvariant in W Norway with stems up to 2.5 m and leaves 18 cm long that are darker and thinner than thetype. This variant is referred by some to Reynoutria japonica var. teminalis Honda.However, Jonsell et al. (2000) reckon that further studies are needed to confirm the identity ofthis form of the plant.
The taxonomic revision of this member of the Polygonaceae has also given rise to numerous changes,reflected in its naming. Fallopia japonica is also known as Reynoutria japonica Houtt.,and previously it was referred to Polygonum cuspidatum Siebold & Zucc.
Fermanagh occurrence
There seems to have been just one possible record of Japanese Knotweed prior to 1974, when Meikle andco-workers recorded either this species or F. sachalinensis (Giant Knotweed) in 1953 on aroadside between Tempo and Brougher Mountain. F. japonica is now very widespread throughout thecounty and has been recorded in 102 tetrads, 19.3% of those in the VC.
F. sachalinesis is rare in Fermanagh and F. × bohemica has notyet been recorded, although the latter has been very rarely recorded near Lough Neagh in Co Antrim(Hackney et al. 1992). In confirmation of this, a plant of F. ×bohemica was recorded at the mouth of the River Six Mile Water, Antrim town in summer 2009 (P.Hackney, pers. comm. November, 2010).
British and Irish occurrence
F. japonica is common, widespread and produces very persistent, well-established dense colonies insuitable lowland ground as described above throughout both Britain and Ireland. It appears to still beincreasing (Clement & Foster 1994). Conolly (1977) first documented the history of the spread ofF. japonica and three other members of the family Polygonaceae in the British Isles. Herecognised three phases of invasion (also observed for many other alien plants): a pioneer stage ofscattered occurrences accumulating relatively slowly, followed by rapid secondary spread from primaryfoci, and a final era of consolidation resulting in more or less total coverage of all suitablehabitats.
As is typical of other alien introductions, Japanese Knotweed is host to relatively few herbivorousinsects, and it is not attacked by either nematodes or fungi. Its distribution in the British Isles islimited by late frosts and by summer drought, neither of which figure greatly in Fermanagh or in otherwestern areas of Britain and Ireland. On the other hand, these two limiting factors do help explain whyit is essentially a lowland species.
Gilbert (1989, 1994) has suggested that Japanese Knotweed is not as serious a problem in Britainas it is often considered to be. His observations of its behaviour around Sheffield indicate that it isshowing signs of adapting to British conditions. Furthermore the indigenous flora and fauna are alsoadapting to it. Not everyone is quite so sanguine about this vigorous competitor as Dr Gilbert is,however.
Since 2013, in Britain it is illegal to sell property without checking the ground to see if JapaneseKnotweed is affecting it. Where it is, the seller is required to provide a management plan for itseradication from a professional company. Japanese Knotweed isclassed as 'controlled waste' under the Environmental Protection Act 1990. Thisrequires its disposal at licensed landfillsites.
Toxicity
Aerial parts of F. japonica are grazed by sheep, cattle and horses, but the plant containglycosides and is unpalatable. The rhizome at least is known to have killed a goat in Britain which ateit (Cooper & Johnson 1998, p. 180).
European and world occurrence
On the European continent, F. japonica is widespread being represented in over 40% of theterritory. It is particularly frequent in W & C Europe. In Scandinavia, it is mainly coastal (Balticand N Atlantic), while southwards in Europe it peters out north of the Alps, Pyrenees and other mountainranges (Jalas & Suominen 1979, Map 422). F. japonica is also an introduced, invasive alien inN America (including Canada) and New Zealand.
Eradication or control
Cutting and burning are useless in controlling the plant (Beerling et al. 1994). Eradication isdifficult and expensive, requiring application of systemic herbicide carefully timed to coincide withthe near exhaustion of stored photosynthetic reserves in the late spring or early summer. The NationalRivers Authority advises that eradication by cutting can take up to ten years, but other evidence wouldsuggest this form of extirpation (or even attempted control) is quite impossible (see Beerling etal. 1994, p. 967). The only real alternative to this is the use of chemicals (ie glyphosate or2,4-D amine), which is permissible under licence where no runoff into watercourses is anticipated.Repeated spraying for up to four years may be required. Spraying is most effective when the plant is atthe flowering stage in late summer or early autumn.
Uses
The dried rhizome of F. japonica is used in traditional Chinese and Japanese herbal medicineagainst a variety of complaints including dermatitis, gonorrhoea and athlete's foot (Beerling etal. 1994)! Despite what is said above regarding its toxicity, young stems are considered edibleas a spring vegetable, although they have a flavour similar to extremely sour Rhubarb. The flowers arean important source of nectar for honeybees, and in NE USA it gives rise to a monofloral honey oftenreferred to as 'Bamboo honey' (https://en.wikipedia.org/wiki/Fallopia_japonica (consulted on 16 June2018)).
Names
The genus name 'Fallopia' was given in honour of Gabriele Fallopi (1523-1562), the Italian anatomist,after whom the Fallopian tube was also named. The alternative genus name 'Reynoutria' was given inhonour of Reynoutre, a 16th century French naturalist who was an acquaintance of Lobel (Stearn 1992).The Latin specific epithet 'japonica' simply refers to the geographic origin of the species in E Asia.
English common names include Japanese Knotweed, Asian Knotweed, Fleeceflower, Himalayan Fleece Vine,Monkeyweed, Monkey Fungus, Hanco*ck's Curse, Elephant Ears, Pea Shooters, Donkey Rhubarb, Sally Rhubarb(although it is not a rhubarb), Japanese Bamboo, American Bamboo and Mexican Bamboo (although it is nota bamboo) (https://en.wikipedia.org/wiki/Fallopia_japonica(consulted 12 June 2018)). Some of these names refer to the hollow, bamboo-like stems, others are moremysterious.
Threats
A noxious weed, threatening woodland, hedgerows and waterside vegetation. Eradication is very difficultand needs to be sustained over several years.
Fallopia sachalinensis (F. Schmidt) Ronse Decr., Giant Knotweed
Introduction, neophyte, a rare garden escape or discard.
1953; MCM & D; road between Tempo and Brougher Mountain.
April to December.
Growth form, introduction and preferred habitats
This very large rhizomatous, herbaceous perennial develops annual stems that can grow up to 3, 4 or even5 m tall. Like F. japonica (Japanese Knotweed), it has a vigorous, thick, spreading rhizome thatenables it to develop dense clonal colonies, described as "coarse thickets" by Lousley &Kent (1981), but considered less dense than F. japonica clumps by Jonsell et al. (2000).The heart-shaped leaves are of coarse texture and are the largest in the family Polygonaceae, growing upto 40 cm long and 28 cm wide. The species is rather similar to Japanese Knotweed, but is distinguishedby its greater height and its leaves having a heart-shaped (not straight) base and a crenate margin. Itis gynodioecious, the majority of plants being functionally female, although bisexual (hermaphrodite)plants also occur.
Like F. japonica, F. sachalinensis is a native of NE Asia that was introduced to farms andgardens in Britain and Ireland in the mid-19th century (possibly as early as around 1861) (Conolly1977). In this case, the species came from both Sakhalin Island and N Japan. Originally young shoots ofthe plant were considered good forage for horses and cows, especially since it grew well in times ofdrought. Giant Knotweed was experimentally planted for this purpose and was said to produce 80-120 tonsof green fodder per acre. You need a metric conversion as well inbrackets. I have calculated this to be (rounding) 30-44 tonnes/ha but please check Later itwas realised that it was not as productive nor useful as predicted, and its fodder cultivation dwindledand eventually ceased or was abandoned.
Despite its dingy white flower panicles, the plant was grown as a decorative garden subject. It waspromoted by Veitch's nursery in the 1880s and 1890s, one of the most important suppliers of exotic treesand shrubs to the Victorian and Edwardian gardener well into the first quarter of the 20th century.F. sachalinensis plants also appeared in the catalogue of the Daisy Hill Nursery in Newry in1891, offered at one shilling each (Bailey & Conolly 2000).
On account of its larger scale, growing to around twice the height of F. japonica, cultivation ofF. sachalinensis was naturally avoided by most gardeners with more modest-sized plots orproperties, and only the owners of estates and very large gardens would have been likely to introduceit, perhaps intending it as cover for game. In some cases it may have been planted to provide rapidgrowing fodder for horses and cattle, although this usage was probably confined to areas of the Europeancontinent. Otherwise it is easy to see the species being a fashionable horticultural status symbol atthe end of the 19th century.
By 1896 Giant Knotweed was reported growing unplanted in the wild, on waste ground near the Lagan Canalat Lisburn, Co Antrim (H39). The subsequent spread of the species, however, has not matched that ofF. japonica, most reports of it representing primary escapes or cast-outs of garden material(Conolly 1977).
The species is somewhat more shade tolerant than F. japonica and its typical habitat in Britainand Ireland consists of open areas in estate woods, on damp areas of riverbanks, lakeshores, roadsidesand waste ground, where it develops large, persistent clones that die down in winter leaving a tangle ofdecaying branches. In its native Japan, characteristic habitats include unstable, moist soils on riverbanks, volcanic detritus and on basaltic lava flows. It can also appear there on stabilised scree belowcoastal cliffs (Conolly 1977).
Variation
Unlike its near relative F. japonica, F. sachalinensis displays considerable variationacross Europe. This reflects the origin of the plant, probably collected from numerous sites in itsnative territory in the 19th century, and possibly its introduction to Europe as seed, as well asrhizome fragments (Bailey & Conolly 2000).
Reproduction
Both F. japonica and F. sachalinensis are gynodioecious, with female and bisexual(hermaphrodite) plants occurring. Most plants of F. japonica in Britain and Ireland are var.japonica which only has female plants. The dwarf form, F. japonica var. compacta,has both male and female plants here. Most plants of F. sachalinensis are female, but male plantsare not regarded as all that rare (Sell & Murrell 2018). The inflorescence of F.sachalinensis is shorter and on stouter branches than in F. japonica and the flowers aregreenish white, the 8 cm panicles drooping. Giant Knotweed plants occasionally bear mature-lookingfruits and may set viable seed. Despite this there are no definite reports of establishment andnaturalisation of F. sachalinensis from seedlings in the wild anywhere in these islands(Conolly 1977; Hollingsworth & Bailey 2000).
Evidence of genetic diversity in clones examined along rivers in England and Scotland, however, doespoint to the possibility of seed reproduction and dispersal occurring, at least in some areas of SEEngland where the species is most prevalent. The pattern of spread along riverbanks, however, could alsobe achieved by floating plant fragments re-rooting and establishing.
Despite the vigour, size and shading capacity of the annual shoots, and the potential for both sexual andasexual reproduction, F. sachalinensis does not appear to be nearly as invasive as the smaller,much more widespread F. japonica, although reproduction of the latter is entirely vegetative.Analysis of the dates of published records of F. sachalinensis in non-garden sites clearly showsthat it does not spread rapidly in these islands (Conolly 1977; Bailey & Conolly 2000).
It thus appears that the reproduction and spread of F. sachalinensis may well be limited, at leastin part, by climatic conditions. Perhaps our winters are too mild and wet for seeds to chillsufficiently, break dormancy and germinate in spring. Instead, the seeds may simply die, rot anddisappear. Maps of the species ex-garden occurrence in Britain and Ireland very probably simply reflectthe incidence of independent primary garden escapes near habitation, or discards established on open,disturbed or waste ground (Conolly 1977; Preston et al. 2002).
Hybrid Knotweed
Populations of F. sachalinensis, like those of its smaller relative F. japonica, generallycontain both male-sterile and hermaphrodite individuals. When these two knotweeds occur together theyfrequently hybridise, and their intermediate product (F. × bohemica(Chrtek & Chrtková) J.P. Bailey (Bohemian Knotweed)) forms viable seed in Britain, though reportedlyit does not do so in Scandinavia (Jonsell et al. 2000). The hybrids are describedas "variably fertile" and back-crossing can occur (Stace et al. 2015). While F.sachalinensis is decidedly scarce in Ireland, it is very possible that some of the very manystands recorded as F. japonica, really belong to the hybrid form, but are mistakenly identifiedas the more frequent parent species.
Fermanagh occurrence
In Fermanagh, there are records of this very large rhizomatous perennial from a total of just tentetrads, eight with post-1975 dates. As the distribution map indicates, its ex-garden Fermanaghdistribution is centred on the Irvinestown-Ballinamallard area NW of Enniskillen and it is chieflyrepresented on the Necarne estate and at Riversdale Forest. The latter is now a conifer plantation, butpreviously it was a subsidiary portion of the nearby Castle Archdale estate. Either of the two estateswhere the species persists might have been the primary point of Giant Knotweed's local introduction.
The first Fermanagh record, as listed above, dates from 1953 and was made by Meikle and his co-workers,except that they remarked that they might have mistaken it for F. japonica (JapaneseKnotweed), and that it needed checking! The site and habitat given for their find, on the roadsidebetween Tempo and Brougher Mountain, certainly suggests to us that F. japonica is by far morelikely the correct identification of their plant. Therefore the first definite record for F.sachalinensis is Paul Hackney's find in Enniskillen Town in September 1974, a voucher for whichexists in BEL. Subsequent records are as follows: Glen Lodge, 3 km N of Ederny, 1986, D. McNeill;by riverside track through conifer plantation, Riversdale Forest, Ballycassidy, 27 December 1987 and 8July 2000, RHN; old railway ground to NE of Ballinamallard, 1991, I. McNeill; Belleisle, Upper LoughErne, 27 June 1992, RHN; Necarne estate near Irvinestown, 19 September 1993 and 3 July 1995, RHN; S. ofDrumsloe, 12 April 1995, RHN; Jamestown House, Magharacross, 2000, I. McNeill.
So far, there are no records of the hybrid, F. × bohemica fromFermanagh, but it grows in similar damp wayside habitats as Japanese Knotweed, and it is equallyinvasive (Stace et al. 2015).
British and Irish occurrence
Giant Knotweed had escaped from cultivation and was naturalised in the north of Ireland by1896 and in Britain by 1903. It has subsequently spread and become widespread in both islands, but it isnot as frequent, abundant, or as rapidly invasive as its smaller relative, F. japonica.Nevertheless, in Britain it stretches from S Cornwall to Shetland. The frequency of F.sachalinensis is greatest in the south of England and around the larger conurbations, itspresence thinning noticeably northwards from Liverpool. In Ireland, while extending from SW Kerry (H1)to N Antrim (H39), it is much more thinly spread than in Britain and the distribution displays a slightnorthern and western tendency.
In 1981, the British parliament passed the Wildlife and Countryside Act which proscribed GiantKnotweed and Japanese Knotweed. It is now an offence to introduce these species into the wild anywherein the country.
European and world occurrence
F. sachalinensis is quite widely naturalised in NW and C Europe, although itis nothing like as frequent or omnipresent as F. japonica (Jalas & Suominen 1979, Map 423).The species has been introduced and become naturalised in N America, and it is also present to a lesserextent in S Africa, India, Australia and New Zealand (https://www.cabi.org/isc/datasheet/107744(consulted on 18 June 2018)).
Control or eradication
Both Knotweed species are well known to be remarkable difficult to eradicate. Once a clone has outgrownthe space the gardener notionally allotted it, attempts to limit further spread, and to reduce oreliminate the problem it becomes, would naturally tend to involve uprooting and discarding of portionsof the plant. However, digging, cutting and burning have proved useless in controlling the plant. Thethick and extremely tough ramifying rhizome of F. sachalinensis can easily survive even the mostsevere physical mistreatment, and even a very tiny weight of rhizome tissue can regenerate the plantafter its upheaval, transport and refuse deposition. Real, effective, professional eradication isdifficult and expensive, requiring application of systemic herbicide carefully timed to coincide withthe near exhaustion of stored photosynthetic reserves in the late spring or early summer. Eradicationoften requires repeated spraying over several growing seasons.
Names
The genus name 'Fallopia' was given in honour of Gabriele Fallopi (1523-1562), the Italiananatomist, after whom the Fallopian tube was also named. The alternative genus name 'Reynoutria' wasgiven in honour of Reynoutre, a 16th century French naturalist who was an acquaintance of Lobel (Stearn1992). The Latin specific epithet 'sachalinensis' simply refers to the geographic origin of the speciesin E Asia.
Threats
Although this is a very large, dominant patch-forming alien, it has not proved as vigorous and mobile asF. japonica. Since it remains rare and is confined to demesnes, one could not yet call itinvasive in Fermanagh. Nevertheless, it is a noxious weed that is a potential threat to woodland andwaterside vegetation, and since it produces a vigorous, fertile, invasive hybrid with F.japonica, it is all the more threatening to native vegetation. Eradication is very difficult,involves spraying with expensive herbicides, and to be effective it must be sustained over severalyears.
Introduction, archaeophyte, a rare, casual. Eurosiberian wide-temperate, but widely naturalised to becomecircumpolar wide-temperate.
1884; Barrington, R.M.; Lower Lough Erne.
July to November.
Growth form and preferred habitats
Black-bindweed is a tiny-flowered, summer annual possessing a taproot, an efficient nutrient-gatheringfibrous root system, and a sprawling, rapidly increasing, smothering, vine-like stem that develops atwining climbing habit when tall supporting plants are available.
It is a pioneer colonist of disturbed, open, lowland, unshaded bare ground. Previously an annual weed ofarable cultivation, it now appears mainly as a garden weed or as scattered individual plants onroadsides, rubbish tips and in disturbed waste ground or on rubble in building sites (Lousley & Kent1981; J.R. Akeroyd, in: Preston et al. 2002).
Reproduction
As an annual, F. convolvulus reproduces entirely by seed. It flowers from July to October,producing cymose clusters of greenish-white flowers, many congested at the top of the stem andindividual cymes of up to 8 flowers borne in leaf axils lower down. The flowers are bisexual and areinsect- or self-pollinated, sometimes cleistogamous (ie fertilised while still in bud). Seeds areprolifically produced and are long persistent in the soil seed bank (Grime et al. 1988).
Fermanagh occurrence
There are records of F. convolvulus in the Fermanagh Flora Database from a total of 20 tetrads,but only 14 of them have post-1975 finds. The pattern of Fermanagh records provides slightcirc*mstantial evidence of a species decline, but it forms part of a wider trend within N England andScotland whereby F. convolvulus has gradually disappeared from marginal areas where cultivationhas been abandoned (J.R. Akeroyd, in: Preston et al. 2002).
In Ireland, the change from a mixed arable and pastoral pattern of farming towards almost exclusivelypastoral agriculture began immediately after the potato famines of the 1840s and the associated humande-population. Apart from brief reversions during both world wars, this process has continued to thepresent day and arable cultivation has almost reached extinction in Fermanagh. Consequent to this,Black-bindweed is rare in Fermanagh but is most frequently found on suitable open ground around thequarries in the Derrylin area. Since it possesses a long-persistent seed bank, the species also crops upsporadically on heaps of disturbed topsoil in other parts of the VC, eg around building sites.
Irish occurrence
The Cen Cat Fl Ir 2 lists Black-bindweed as having been recorded in every Irish VC, butwhile still widespread, it has been in decline for a long period, perhaps for 180 years. Nowadays,Fermanagh is near the north-westerly inland Irish limit of this weedy annual, and indeed in Donegal(H34, H35) it appears largely coastal (BSBI Atlas 2; New Atlas).
British occurrence
Although elsewhere within its overall distribution F. convolvulus occurs very much further north(for instance at 70N in Greenland), within Britain and Ireland the species shows a definite thinningtowards both N & W throughout these islands.
Species status
Previously accepted as a native species without question by most British and Irish authorities (eg Stace1997), Webb (1985) listed F. convolvulus along with 40 other species as being, in his opinion,probably or almost certainly introduced by man. It is odd that the introduced status of this common weedshould only recently have been recognised and acknowledged (Preston et al. 2002; Preston etal. 2004). F. convolvulus has been very closely associated with human agriculturalactivities since recorded history began, and its seeds or achenes are a notorious, almost cosmopolitancrop seed contaminant, especially of cereals. In many parts of the world, it is regarded as the numberone weed of a wide variety of arable crops (Holm et al. 1977). It is odd that Sell & Murrell(2018) revert to regarding F. convolvulus a native species.
Fossil record
While there are fossil records of F. convolvulus from earlier interglacial periods, in the currentwarm period there are no pre-Neolithic fossils which would indicate its presence before farming gotunderway (Godwin 1975). The discovery of large accumulations of seed in some archaeological digsstrongly suggests that the seed was an important source of food or fodder (Hume et al. 1983).
European and world occurrence
F. convolvulus is common and widespread across the whole of Europe, although the distributionthins towards the south as it approaches the Mediterranean coast (Jalas & Suominen 1979, Map 420).Black-bindweed spreads widely with agriculture as an arable crop seed contaminant, and it is nowvirtually cosmopolitan. In many parts of the world, it is regarded as the number one weed of a widevariety of arable crops, and in other areas including Argentina, Canada and the United States it is inthe top three cereal weeds. It has increased in seriousness due to its prolific seed production, deepdormancy and long seed persistence in soil, its habit of germinating throughout the growing season, andits resistance to herbicides. As other weeds more susceptible to herbicides are removed from the field,competition is effectively reduced, allowing F. convolvulus to flourish (Holm et al.1977).
Names
The genus name 'Fallopia' was given in honour of Gabriele Fallopi (1523-1562), the Italiananatomist, after whom the Fallopian tube was also named. The Latin specific epithet 'convolvulus' isderived from 'convolvo' meaning 'roll around' or 'interweave' (Gilbert-Carter 1964). Synonyms includePolygonum convolvulus L., Bilderdykia convolvulus (L.) Dumort, fa*gopyrumconvolvulus (L.) H. Gross, fa*gopyrum carinatum Moench, Helxine convolvulus (L.)Raf., Reynoutria convolvulus (L.) Shinners and Tiniaria convolvulus (L.) Webb & Moq.Other English common names include Bearbind, Bindcorn, Climbing Bindweed, Climbing Buckwheat, Cornbind,Corn Bindweed, Devil's Tether and Wild Buckwheat.
Threats
The continuing decline in arable agriculture limits the availability of open sites suitable for thisspecies.
Rumex acetosella L., Sheep's Sorrel
Native, very frequent and widely scattered throughout. Eurosiberian wide-temperate, but widelynaturalised around the world and now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This often small but rapidly growing, 4-30 cm tall, creeping, clump and patch-forming rhizomatousperennial is a characteristic early, but rather persistent, coloniser of unshaded, open or disturbedground, both lowland and upland. It is typically found on infertile acidic, relatively dry, sandy, stonyand peaty soils, including those on overgrazed pastures, heaths, bogs and grassy moorland. Thelanceolate, hastate (spear-shaped) leaves with their small, spreading basal lobes are very distinctiveand their strong, acid taste confirms identification.
It is an occasional to frequent weed in urban situations, on roadsides and in waste ground, sand-pits andquarries. Nowadays, it is also very often found growing in the peaty composts used in garden centreplant pots. Being low-growing, it is a poor competitor with taller herbs and grasses and therefore it isconfined to very infertile, disturbed, or physically limited conditions for growth. Sheep's Sorrel isalso an early coloniser of freshly available bare soil and of vacant rock crevices. The plant isespecially linked with recently burnt ground in peaty areas, eg on cleared ground in forestryplantations, around cut-over areas of bogs and in fired areas of heathland. R. acetosella alsosurvives in narrow rock crevices on cliffs and outcrops, in situations where competition is much reducedor absent.
Variation
R. acetosella is a polymorphic and very widespread circumpolar species within which there is anintricate and taxonomically not fully worked out range of variation (Hultén & Fries 1986). Thechromosome base number is x=7 and three levels of ploidy are known to occur (2x, 4x and 6x). However,study shows there appears to be only minor correspondence between ploidy level and either morphology(including size characters and leaf shape) or distribution. Taxonomic distinctions are based onmorphologically defined variants, and several ploidy levels may co-exist within each described taxon(Jonsell et al. 2000).
Stace (2010) recognises two subspecies in B & I, although their distribution is not yet adequatelyresearched. The more widespread form is named subsp. acetosella (R. tenuifolius (Wallr.)Á. Löve), which very likely covers both islands. A small form of this plant, with narrow, linear leavesthat is ± confined to very dry coastal sands previously described as R. tenuifolius (Wallr.) Á.Löve, is treated as a mere variety by Stace (2010), namely var. tenuifolius Wallr. The lesswidespread of two subspecies he recognises is subsp. pyrenaicus (Pourr.) Akeroyd (subsp.angiocarpus auct. non (Murb.) Murb., R. angiocarpus auct. non Murb.). This form probablyis mainly located in the S of Britain and appears to be absent in the N, although the distribution andecology of the two subspecies very likely overlap as they do in continental Europe.
In their recent, critical consideration, Sell & Murrell (2018) relocate R. acetosella to agenus on its own, naming it Acetosella vulgaris (Meisn.) Fourr. They also recognise threesubspecies within it: subsp. vulgaris, subsp. tenuifolia(Wallr.) P.D. Sell and subsp. pyrenaica (Pourr. ex Lapeyr.) P.D. Sell. Subsp. pyrenaica isdistinguished by having its inner perianth segments fused to the achene, while subsp. tenuifolia,as its name suggests, has narrow basal leaf lobes, usually 5–15 times as long as wide, usually curvedforwards, separating it from subsp. vulgaris (basal leaf lobes wider, usually 2-7 times as longas wide and usually patent, directed backward, or occasionally absent (Sell & Murrell 2018, pp.512-3).
Fermanagh occurrence
In Fermanagh, R. acetosella has been recorded in 147 tetrads, 27.8% of those in the VC. Apart fromthe previously mentioned habitat types, it is also very common in many rocky upland areas, including inthe area on the thin peaty soils developed over limestone on the Knockmore cliffs and the stabilisedscree above Doagh Lough, as well as on the acidic rocks and slopes of Cuilcagh, the highest mountain.Having said that, Sheep's Sorrel is very definitely a calcifuge species; seedlings grow very poorly andthe plant cannot compete for long in lime-rich growing conditions. Frequency and abundance measureselsewhere show R. acetosella strongly prefers acidic soils within the pH range 3.5-5.5 (Grimeet al. 1988).
Although most frequent and abundant in disturbed, bare, peaty soils, Sheep's Sorrel can also be longpersistent in short-turf, acidic, unproductive grassland, even when this type of ground is heavilygrazed by sheep. The reason for this is at least partially accounted for by the sharp, bitter-tastingoxalates present in its tissues which deter some types of browser.
Toxicity
As the English common name (Sheep's Sorrel) suggests, livestock will graze on the plant, but since it issmall in size they seldom consume large quantities. The species contains bitter-tasting oxalates thathelp deter browsers, and sometimes they also provide sufficiently high concentrations of nitrates thatcan cause poisoning. A third potential toxin is tentatively identified and called rumicin (Cooper &Johnson 1998).
Phenology and vegetative colony production
After seed germination in spring, the young plant develops a stout, deeply penetrating taproot early inits growth. This quickly gives rise to a spreading network of branching horizontal underground stems andshallowly running fibrous roots. Adventitious shoots are prolifically developed from the more shallowparts of the root system, which together with the rhizomatous growth, allows the individual plant cloneto rapidly colonise and carpet the available open ground (Salisbury 1942, 1964).
Clonal colony development is very readily observed whenever R. acetosella first invades loose,sandy, bare or burnt soils to form a living carpet. A study in Canada found that a clone there couldeasily cover an area of 4 m2 after just two growing seasons (Vezina et al. 1986).
Sexual reproduction
R. acetosella is a dioecious perennial with unisexual flowers on separate male and female plants.Plants flower from May to September and are wind-pollinated. Perhaps on account of its clonal growth andseparate sexes, fruit production is sometimes poor. However, most plants produce some seed, and plantsin favourable sites and seasons generate an abundance of it.
Seed dispersal
The seed or achene (a single-seeded dry fruit) is extremely lightweight, even in comparison with those ofother species of open habitats. The seeds are often released surrounded by the persistent perianth and,despite the often small stature of the plant, they are dispersed readily enough by wind (Salisbury1942). The fact that R. acetosella grows on cliffs and in rock crevices is further evidence ofits excellent powers of dispersal, although in addition to carriage by wind this might be achievedthrough animal ingestion and internal transport. Regarding the latter dispersal mechanism, Ridley (1930)reported viable seed being excreted by birds, pigs, horses, cattle and goats.
Seed survival
The survey of soil seed banks of NW Europe uncovered no less than 52 estimates for this species, of whicheleven recorded buried survival for at least five years (Thompson et al. 1997).
Fossil record
Fruit and pollen of R. acetosella s.l. are common in B & I throughout the interglacial andglacial fossil record from the Cromer Forest Bed series onwards. R. acetosella was prevalent(along with R. acetosa (Common Sorrel)) in all zones of the Late Weichselian, but it diminishedgreatly in zone IV of the Flandrian, probably in response to a closing of the herbaceous vegetationcover as the forest canopy developed. It does not reappear in the fossil record until there is evidenceof deforestation brought about by human interference in the Bronze Age (Godwin 1975).
Fossil nutlets of R. acetosella agg. have been found in a full-glacial freshwater deposit ofMiddle Midlandian age, radio-carbon dated to 30,500 BP, discovered at Derryvree, near Maguiresbridge, CoFermanagh (Colhoun et al. 1972). The flora and fauna of the deposit indicated that open tundravegetation and a periglacial climate prevailed at the time when it was laid down.
British and Irish occurrence
R. acetosella s.l. is almost ubiquitous throughout these islands from the coast tothe uplands. However, the two or three subspecies that have been taxonomically described in recent yearsare not yet sufficiently recorded to reveal the true pattern and extent of their occurrence (Prestonet al. 2002). Subspecies or variety tenuifolia is described by Lousley & Kent (1981)as locally abundant on dry, humus-deficient, sandy soils and occurring commonly near the S and E coastsof England and Scotland from Dorset to Sutherland. It is also said to be present on inland heaths inSurrey (VC 17), Berkshire (VC 22) and the Breckland (VCs 25-28), with widespread scattered localitieselsewhere, although the distribution is imperfectly known. This remains the case.
European and world occurrence
R. acetosella s.l. is common and widely distributed as a native species throughout Europe from theMediterranean to the far N of Scandinavia (Jalas & Suominen 1979; Map 427). Having said this, fromEurope it has been very widely dispersed as a weed in association with human movement and activities andhas been introduced to N America and indeed almost worldwide (Hultén & Fries 1986, Map 658). R.acetosella s.l. is now boreal circumpolar and has also spread into the southern hemisphere,including most temperate and some tropical areas (Jonsell et al. 2000).
Names
The genus name 'Rumex' is an old Latin name for Sorrel from Pliny derived from the Latin 'rumo' tosuck, from the Roman habit of sucking Sorrel leaves to allay thirst (Johnson & Smith 1946). TheLatin specific epithet 'acetosella' is derived from 'acetum' meaning vinegar, from the acid taste of theplant. 'Acetosa' and 'Acetosella' were both pre-Linnaean names for Sorrel and any other plant withacid-tasting leaves (Gilbert-Carter 1964; Stearn 1992).
The English common name 'Sheep's Sorrel' or just 'Sheep Sorrel' is from the 14th century French 'surelle'or 'sorele', a diminutive from the Low German 'suur' meaning 'sour' or 'acid', a reference to theacidity of the leaves. As such it might be translated as either 'little acid plant' or 'little sourplant', both of which are appropriate (Prior 1879; Grigson 1974).
Threats
None.
Native. Extremely common and widespread, locally abundant. Eurosiberian boreo-temperate, but widelynaturalised in both hemispheres and now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This is a very common, variable, phenotypically plastic, sometimes abundant wintergreen, tufted,semi-rosette perennial of established, unimproved grasslands on a wide range of moist to dry soils in ahuge variety of habitat situations. Like the smaller R. acetosella, leaves are characterised by asour, acid taste, in spring sometimes described as refreshing and therefore long used in diet and folkmedicine (see below). In Fermanagh, as in most of the rest of B & I, Common Sorrel is almostubiquitous in old, established grassy places with mildly acid to neutral, impoverished to moderatelyfertile, base-poor to base-rich soils.
It is successful on account of having both wide, easily met ecological tolerances, plus physiological andbiological characteristics that allow it to cohabit with grasses and many herb species in sub-optimalmeadow and pasture environments. These conditions allow moderate levels of competitive interaction todevelop, yet they prevent sustained, vigorous plant growth by any plant species (Grime etal. 1988). Given sufficient time and continuing, but moderate levels of varying disturbance,such conditions encourage the eventual development of species-rich grassland vegetation. Apart fromevading strong competition in fertile soils, R. acetosa chiefly avoids environmental extremessuch as aquatic, regularly flooded, strongly acid or deeply shaded conditions.
Like other Dock species, it contains toxic levels of bitter-tasting oxalates which make livestock avoidit, and it can thus survive fairly high levels of grazing. R. acetosa is morphologically flexibleand tolerates mowing very well. Its rootstock can also survive moderate degrees of trampling, fire andother forms of disturbance which limit the plants' growth, reproduction and competition.
In addition to agricultural grasslands, R. acetosa is commonly found on track-sides, openings inwoods, open areas on roadside verges, waste ground and cliffs.
Variation
R. acetosa has wide ecological tolerances and a matching wide geographical amplitude based on anextensive range of both phenotypic and genetic variation. Much of the phenotypic response to theenvironment is continuous variation and it is therefore unsurprising that taxonomic treatment of it hasvaried greatly over the years (Tutin et al. 1993; Jonsell et al. 2000). R. acetosais regarded as a polymorphic species, and in Flora Europaea, Tutin et al. (1993) supportthree European subspecies. However, in their recent critical Flora of Great Britain and Ireland,Sell & Murrell (2018) transfer R. acetosa to a genus on its own, naming it Acetosapratensis Mill., within which they recognise four subspecies.
The additional form in the latter is subsp. biformis (Lange) P.D. Sell, which is small (up to 20(or exceptionally 30) cm), and has rather thick, succulent leaves and papillae ± confined to the basalmargin of leaves. It is said to be native on sea cliffs in Cornwall, Cardiganshire and Co Clare (Sell& Murrell 2018). The authors of Flora Europaea believed subsp. biformis could not besatisfactorily separated from robust plants of subsp. acetosa, and it is very similar indescription to subsp. hibernica (Rech. f.) P.D. Sell, except the latter has papillae and veryshort hairs on all vegetative parts.
Hybrids
Unlike the Dock species of the subgenus Rumex which cross frequently, there are no hybridsinvolving either R. acetosa or R. acetosella (Stace et al. 2015).
Fermanagh occurrence
In Fermanagh, by 2010, Common Sorrel was recorded well over 1,150 times (more than 300 records ahead ofthe next Rumex species, R. obtusifolius (Broad-leaved Dock)). It has been recorded in 467Fermanagh tetrads, 88.5% of those in the VC, thus making it the 20th most widespread species in tetradterms in the 35-year survey (1975-2010) carried out by Robert Northridge and the current author. Despitethe high frequency of R. acetosa finds in Fermanagh, we feel that if we searched long enough itmight well be possible to record it in every single tetrad in the county!
Reproduction
R. acetosa might be even more common if it were not for the fact that individual plants arenot long-lived. Also, unlike R. acetosella, this species has only limited powers ofvegetative reproduction, plants only producing daughter rosettes for recovery purposes after theysurvive moderate levels of disturbance.
Although it has separate male and female plants (ie it is dioecious), seed (achenes) are copiously set bythe large female inflorescences. Flowering occurs from May to June, pollination is by wind and seed isset from June to September.
Natural dispersal is both by wind and through birds and other animals ingesting the shiny achenes (Ridley1930). Human transport is probably even more significant, however, Sorrel seed being present in hay,silage or as a contaminant of crop seed (Grime et al. 1988).
Germination and establishment
R. acetosa is characterized by early autumn germination and it exhibits only a transient type ofseedbank (ie surviving less than one year). In water meadows or other forms of wet ground, the death offlood-intolerant achenes results in a lack of multiple post-flood germination cohorts. Taken togetherthese seed properties limit R. acetosa in river and lakeshore flood-plains to rarely floodedgrasslands, where the plants face a relatively high level of competitive interactions. This contrastsstrongly with the flood-tolerant properties of the related wetland species, R. crispus (CurledDock) (Voesenek & Blom 1992).
British and Irish occurrence
Analysis of the 2002 New Atlas hexad survey results for the whole of B & I indicates thatR. acetosa is the most widely recorded Dock in these islands, again leading R.obtusifolius, the next most widespread Dock by 52 hexads (Preston et al. 2002).
European and world occurrence
R. acetosa is widespread throughout most of Europe (although rare further south), and Eurasianforms are widely naturalized across N America so that the species sens. lat. has become circumpolar.R. acetosa has also been accidently introduced by agriculture in widely scattered places aroundthe southern hemisphere including S America, S Africa, S Australia and New Zealand (Hulten & Fries1986, Maps 660, 661).
Uses
The sour taste (or the "grateful acidity", as Grieve 1931 has it) of R. acetosa sap isproduced by the oxalate content of the plant which is mildly toxic. The flavour is much less sharp andpungent in springtime and does not really reach its maximum until the plant is in full flower in Juneand July. Sorrel has an ancient pedigree as a medicine for cleaning or purging the blood, and thusclearing up spots on skin. In the past, the plant was cultivated for its medicinal and edible qualities,its leaves eaten raw as a salad appetiser or digestive, cooked as a pot herb in ragouts, fricassees andsoups, or treated as spinach and eaten, for instance, with stewed lamb or veal (Grieve 1931).
The plant was also used much as we use lemons: as a cure for scurvy and in the making of sauces for foodsuch as fish (Grigson 1987). Leaf infusions were drunk to assist the kidneys and to cool fever. Leaveswere also used to heal sores, boils, bruises and burns, presumably being made into poultices. They werealso used to staunch bleeding and to treat a range of ailments from jaundice to heart trouble (Grieve1931; Allen & Hatfield 2004).
Names
The genus name 'Rumex' is an old Latin name for Sorrel from Pliny derived from the Latin 'rumo' tosuck, from the Roman habit of sucking Sorrel leaves to allay thirst (Johnson & Smith 1946). TheLatin specific epithet 'acetosa' is derived from 'acetum' meaning vinegar, from the acid taste of theplant. 'Acetosa' and 'Acetosella' were both pre-Linnaean names for Sorrel and any other plant withacid-tasting leaves (Gilbert-Carter 1964; Stearn 1992).
The English common name 'Common Sorrel', or just 'Sorrel', is from the 14th century French 'surelle' or'sorele', a diminutive from the Low German 'suur' meaning 'sour' or 'acid', a reference to the acidityof the leaves. As such it might be translated as either 'acid plant' or 'sour plant', both of which areappropriate (Prior 1879; Grigson 1974).
No less than 37 English common names are listed by Grigson (1987) and 17 of them include the word element'sour'. Of the remaining names, several refer to the edible nature of the species, such as'Bread-and-cheese', 'Donkey's Oats' and 'Green Sauce', while others refer to the spring season then theplant is most edible, eg 'Cuckoo's Meat', 'Cuckoo's Sorrel' or 'Cuckoo's Sorrow'.
In his Typescript Flora of County Fermanagh, Meikle noted that R. acetosa was called"Clock sorrel" by a farmer in Meenagleeragh, because the leaves are similar in shape to acertain style of "Wag at the wall" pendulum clock.
Threats
None.
Native, locally frequent and abundant. European temperate, locally spread by man.
1836; Mackay, J.T.; an unspecified site on Lough Erne.
April to December.
Growth form and preferred habitats
This large, stout, tufted, rhizomatous perennial Dock produces large stands by rhizome growth up to 2 mtall, and its very long (up to 1 m in length), lanceolate leaves held vertically are absolutelyunmistakable in emergent lakeshore habitats. R. hydrolapathum is a species of shallow (up toabout 30 cm depth), slow-moving, lowland, base-rich, eutrophic waters (Preston & Croft 1997). Thusthe margins of Lough Erne and its feeder streams and rivers provide an ideal habitat for the plant.
Fermanagh occurrence
The British & Irish hectad distribution maps in Preston & Croft (1997) and in the NewAtlas (Preston et al. 2002) suggests that with modern records occurring in 14 hectads,Fermanagh is the Northern Ireland headquarters of R. hydrolapathum. On the same crude basis,however, Co Down (H38) may be a close second in terms of frequency of the largest Irish 'Docken'species. In Fermanagh, Water Dock has been commonly recorded in 93 tetrads, 17.6% of those in the VC. Itis an almost constant component of the Upper Lough Erne shoreline vegetation. The species is alsoprevalent in shoreline marshes at the SE end of Lower Lough Erne, but apart from these two areas, it isotherwise rare, extremely local and only very thinly scattered in ten or twelve outlying wetlandstations beyond the Erne basin.
The discovery of some of the outlying Fermanagh sites dates from the late Praeger and the Meikle eras (iebetween1933 and 1953). On the other hand, six new sites were discovered during the very comprehensive NILakes Survey of 1988-90, suggesting that the species was either previously under-recorded in Fermanaghor that it might even be expanding.
Irish occurrence
Elsewhere in Ireland, Water Dock is widespread and well represented throughout the whole of the RiverShannon catchment in the Republic of Ireland. Closer to Fermanagh, Flora of NE Ireland 3describes R. hydrolapathum as, "local but fairly well distributed in lakes andmarshes", and it is said to be, "still abundant about Downpatrick and in [the] Lecale[peninsula] generally" (ie in SE Co Down, H38).
British occurrence and status
The New Atlas map shows that in Britain, while Water Dock is scattered ± throughout, it is verymuch more frequent and widespread south of a line between Preston and Bridlington, and that furthernorth and into Scotland it becomes much more thinly scattered (Preston et al. 2002).
In many areas of its wider distribution, R. hydrolapathum is regarded as a follower of man (ieanthropochorous), being spread in the past in ship ballast, for example (Jonsell et al. 2000), orplanted for garden decoration (Lousley & Kent 1981). In Northumberland (at least VCs 67 and 68),Water Dock is regarded as a naturalised garden escape (Swan 1993), the species having not been recordedfor the county in the Victorian Flora of Northumberland and Durham, of Baker & Tate (1868).At that time, it was then known in Co Durham and was described by them there as, "notuncommon". The New Atlas map shows these N England records in red indicating their suspectedintroduced status, and a few other northern records, recent and not so recent, are likewise shown aspossible introductions.
No such doubt attends the native status of the species in Fermanagh, the first record having been made aslong ago as 1805 by Prof. Robert Scott. In England, particularly in the south, loss of aquatic habitatshas reduced the frequency of Water Dock, but it is still widespread in suitable situations in this sameregion (Preston & Croft 1997). In contrast, in Angus in E Scotland (VC 90), Ingram & Noltie(1981) have suggested that R. hydrolapathum is actually increasing.
Reproduction
The balance of successful establishment and increase in this species is probably towards vegetativereproduction by water transport of its stout, black rhizome or of the whole plant. This is probably notuncommon (Jonsell et al. 2000). The species is not strongly competitive and even established fenstands can be ousted by more vigorous competing tall emergent perennials such as Phragmitesaustralis (Common Reed), Typha latifolia (Bulrush) and Iris pseudacorus (YellowIris, Yellow Flag). The plant does not flower in its first year. From the second year on, floweringtakes place from June to September, the plants producing one or more tall inflorescences. Theinflorescence arrangement is a branched raceme or panicle. The reddish, bisexual (perfect) flowers areborne in lax whorls and they are wind pollinated.
Seed production and dispersal
An average sized plant of R. hydrolapathum in England produces about 16,000 achenes, which are3.5-4.0 mm long, pale brown and trigonous with sharp edges (Lousley & Kent 1981; Sell & Murrell2018). Ridley (1930) reports their dispersal by both wind and water floatation. Since birds and otheranimals regularly consume the achenes of other Dock species, it is reasonable to suppose that Water Dockseed, which float for a long time, might be eaten and be internally transported further, and indeedbetween water bodies, in this manner.
Germination and establishment
A field and laboratory experimental study made in England at Woodwalten Fen by Wells (1966) found highlevels of seed viability and reasonable germination success (ie 37% of sown seed germinated after 1.5months) on freshly bared peat slopes around ditches. This was followed by high seedling mortality,however, due to a combination of insect damage and the drying-out of the peat at a critical stage ofseedling development. As a result of this, only 7% of the small plants survived into September. Clearlylocal conditions suitable for seedling establishment are very critical for this Dock, as indeed they arefor any species dependent upon seed reproduction.
Hybrids
Being wind pollinated, Dock species of the subgenus Rumex cross freely whenever species grow inproximity and their flowering periods coincide. R. hydrolapathum can form hybrids with five otherDock species, R. aquaticus (Scottish Dock), R. crispus (Curled Dock), R.conglomeratus (Clustered Dock), R. longifolius (Northern Dock) and R. obtusifolius(Broad-leaved Dock) (Jonsell et al. 2000; Stace et al. 2015). Only two of thesepotential species crosses has ever been reported in Ireland − R. hydrolapathum × R. crispus, recorded from a solitary hectad, and R. hydrolapathum× R. obtusifolius, which has been recorded from six well scatteredIrish hectads (Stace et al. 2015).
European and world occurrence
Beyond our shores, R. hydrolapathum is very much confined to temperate areas of Europe betweenlatitudes 40N and 63N, from southern Scandinavia to Sardinia, Italy and the Balkans and stretching asfar east as Iran. It becomes rare towards its southern European limit. Beyond this it is known only fromNW Anatolia and one isolated station in the Caucasus, where it is also considered indigenous, althoughlocally spread by man (Jalas & Suominen 1979, Map 454; Hultén & Fries 1986, Map 666). It isintroduced in both N & S America (Sell & Murrell 2018).
Uses
As a decorative garden subject, the very large size and potentially invasive behaviour of R.hydrolapathum limits the suitability of the species to wetland gardens and shallow pools inlarger premises. Alternatively, specimens could be planted in containers submerged in shallow water torestrict their spread.
Names
The genus name 'Rumex' is an old Latin name for Sorrel from Pliny derived from the Latin 'rumo' tosuck, from the Roman habit of sucking Sorrel leaves to allay thirst (Johnson & Smith 1946). TheLatin specific epithet 'hydrolapathum' is derived from the Greek 'hydor', water, and 'lapathi' meaning'sorrel-like' or 'dock-like'. The name originated in Pliny, referring to a Dock growing in water(Johnson & Smith 1946; Gilbert-Carter 1964; Gledhill 1985). The English common names 'Water Dock','Great Water Dock' and 'Greater Water Dock' are modern book-names with no local folk connectionsanywhere.
Threats
None.
Native, common, widespread and locally abundant. Eurosiberian southern-temperate, but very widelyintroduced worldwide and now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
R. crispus is an extremely variable short-lived perennial, or rarely an annual, which flowers inits first year after germination. Perennial forms develop a thick, fleshy, yellow taproot that can reachto a soil depth of 150 cm. Autumn germinated seedlings overwinter as small rosettes which grow rapidlyin the spring to produce tall flowering stems up to 2 m in height, but often reach only half thisheight. In comparison with R. obtusifolius (Broad-leaved Dock), the other most competitive Dockspecies, the basal leaves are narrower, more parallel-sided and their margins are more wavy (undulate),crenate (with small, blunt, rounded teeth) and crisped (tightly undulate). The leaf base is wedge-shapedor only slightly indented at the stalk, which is long (sometimes almost as long as the blade). Anotherdistinguishing feature observed in the ripe fruit is that the three inner perianth segments (tepals),which become enlarged to form valves surrounding the fruit (a solitary nutlet in each flower), havemargins almost entire, and each may or may not bear one smooth, oblong tubercle (Lousley & Kent1981).
Like the other very common weedy dock, Broad-leaved Dock (R. obtusifolius), R. crispus is avery troublesome plant for farmers, being a particularly serious weed in arable fields, a habitatnowadays almost completely absent in Fermanagh (Foster 1989). Unfortunately, Curled Dock is also all tooabundant in pastures. It is believed to be one of the twelve most widespread and successfulnon-cultivated, colonising species in the world (Allard 1965; Holm et al. 1977). TypicalFermanagh habitats are damp to wet lowland lakeshore pastures, marshy grasslands in general, plus gardenlawns and beds, in parkland grassed areas, roadsides, old quarries and waste ground. In other countieswith coastal habitats, subspecies of it are also common on shoreline shingle, dune and estuarine saltmarsh habitats.
Variation
Five subspecies and a "numerous" range of minor forms, all coming true from seed, arerecognised in Britain & Ireland (Sell & Murrell 2018). Probably the only native forms of R.crispus occur on coastal shingle, dunes, salt-marshes and muddy river estuaries (Sell &Murrell 2018). The other more inland forms commonly found in cultivated, rough and waste ground areregarded as weeds introduced at an early date by man and his domestic animals, namely, subsp.crispus, subsp. robustus (Rech.) Pestova and subsp.strictissimus (Rech.) Pestova. The most likely native form is subsp. littoralis (Hardy)Akeroyd, followed subsp. uliginosus (Le Gall) Akeroyd, which appears to be restricted to a numberof muddy estuaries in the S of England and Wales, and is probably also mostly native (Sell & Murrell2018).
Fermanagh occurrence
While Curled Dock is generally regarded as the commonest native Dock in Britain & Ireland as a whole(Lousley & Kent 1981), it is less common than R. obtusifolius in Fermanagh. Nevertheless, itis common and widespread, being recorded in 315 of the tetrads, 59.7% of those in the VC. It isparticularly widely scattered across lowland areas of the county. In comparison, R. obtusifoliusis represented in the Fermanagh Flora Database by records in 456 tetrads, 86.4% of the total area.
Reproduction
R. crispus reproduces mainly by seed (nutlets). The inflorescence is a lax to dense panicle30-60 cm long, ± simple, or with branches held upright at an angle of no more than 30° to the main stem.Flowers are borne in whorls of 10-30. The species has a very long flowering period extending from Juneto October. One plant can produce from 100 up to 60,000 seeds per year. Buried seed longevity isimpressive, some remaining viable for up to 80 years. The plant can also regenerate vegetatively, butonly the uppermost 4 cm of rootstock is involved. The deep taproot enables the species to survive bothvery dry seasons and severe freezing. Vegetative regeneration is most successful when it occurs in theearly part of the growing season (Hudson 1955). Regeneration from root fragments, together withprolonged seed survival, makes eradication of the plant infested ground almost, or practically,impossible (Holm et al. 1977).
Seed dispersal
Seed is lightweight, has an attached wing and can readily be dispersed by wind. Seed also floats and canbe transmitted in flowing water or transported in mud attached to feet and coats of animals, includingman. It may also be transported in mud on machinery. Viable seed can also pass through the digestivetracts of birds and cattle. Despite these dispersal possibilities, in pastures and in open, neglectedground R. crispus often forms patches. This reflects the fact that in very many species,including Docks, the majority of seeds produced fall near the parent plant and germinate there (Holmet al. 1977). R. crispus can establish quickly from seed, colonising pasture openingscaused by trampling, gouging, fire or dung patches.
Germination
Seeds can only germinate on open ground as the presence of a leaf canopy reduces red light wavelengths(in absolute terms), and increases far-red radiation (relatively), which 'switches off' germination(Foster 1989). Temperature fluctuations reaching 15°C at the upper limit, promotes germination of buriedseed, which helps explain the observed flush of spring germination (Roberts & Totterdell 1981).Until a tap root develops, seedlings are vulnerable to competition from other more established plants,and to disturbance, such as further cultivation.
Polymorphic variation exists in seed germination between plants in different habitats, and even withindifferent parts of the same plant (Cavers & Harper 1966). Differences in germination between seedsof separate plants was greater than between habitats. Interplant differences were greater thandifferences between the germination of seeds borne on different parts of the same plant, plants ofdifferent age, or different degrees of ripeness (Cavers & Harper 1966). Once well established,plants of both R. crispus and R. obtusifolius can easily withstand even quite severetrampling, although flowering may be inhibited. In growth experiments in pots, R. crispus hadmuch higher rates of germination and growth rate in freely drained and partly waterlogged conditionsthan under fully waterlogged conditions (Cavers & Harper 1964). The species appears to be unaffectedby severe drought or frost, and while occasionally it can survive periods of very waterloggedconditions, it is not very successful there (Cavers & Harper 1964).
Weed ecology and control measures
Curled Dock is a primary coloniser of disturbed ground and is regarded as the most 'weedy' species in thegenus Rumex due to its ability to establish quickly from seed, flower in the first year ofgrowth, and because its dormant seed can persist for many years buried in soil until another favourableenvironment arises (Cavers & Harper 1964). Weed infestation of pasture tends to be encouraged by: 1.spreading of slurry and farmyard manure, 2. cutting for silage, 3. high nitrogen levels, 4. soildisturbances, 5. previous damage to the sward (eg direct drilling), and 6. selective grazing by horses(Haggar 1980). The same author also found that Docks are discouraged by flooding, sheep grazing andcutting for hay, but poaching of soil and grazing by cattle have apparently no effect on the abundanceof Docks.
Mowing of grassland containing Docks is often advantageous to the establishment of both R. crispusand R. obtusifolius, since after mowing new shoots are quickly sent up from ground level andthese two species can very quickly flower and set seed before winter. Both these Dock species producelarge numbers of persistent seed and they also show considerable powers of regrowth from vegetativefragments left in the soil (Foster 1989). Apparent elimination of established Dock plants only givesshort-term control due to their very large and long persistent seed bank.
Practical methods for the non-chemical control of Docks include pulling by hand (ie 'docking' them) ingrowing crops or on waste ground. Pulled Docks should be burned in case they should still go to seed.Hoeing of ploughed or dug ground, especially in the autumn, is effective in removing seedlings. Frenchet al. (1986) worked on natural flavour-related chemicals which might be used to stimulate thegermination of R. crispus seed at a time unfavourable to successful growth of the weed. They alsosuggested the premature stimulation of seed germination, before shedding from the mother plant, could bedeveloped as a control method against the species.
European and world occurrence
R. crispus is a very obvious follower of man (ie an apophyte), having spread with agriculture andbecome a serious weed in every continent and across a wide range of crops. It is considered one of thefive most widely distributed plant species in the world, its spread completely masking its area oforigin (Hughes 1938). On the other hand, Hultén & Fries (1986, Map 667) consider it originated inEurasia. R. crispus occurs throughout Europe up to a latitude of 69°N in both Norway and Russia.It is present in Iceland and the Faeroes, but interestingly not in Greenland (Cavers & Harper 1964).Despite existing discontinuities, mainly in C Asia, Hultén & Fries (1986) regard it as circumpolar,and it is widely introduced in the southern hemisphere, including remote areas such as the FalklandIsles, Réunion and Tasmania.
Toxicity
R. crispus plants have a pleasantly sharp taste and are apparently palatable to farm animals,which often eat them without showing any obvious ill effects (Cooper & Johnson 1998). However,Curled Dock is also known to have caused fatal poisoning in both sheep (Panciera et al. 1990) andhumans (Reig et al. 1990). The active toxic substances include oxalic acid, oxalates andanthraquinone glycosides. Leaf and 'tops' samples of R. crispus measured by Panciera etal. (1990) contained from 6.6 to 11.1% oxalic acid calculated on a dry-weight basis.
The quantity of oxalate required to induce acute poisoning in ruminants is found to be quite variable(0.1-0.5% of body weight), and the active dose depends on several factors. Ruminal gut microflora canreadily detoxify a proportion of ingested oxalate, so that mature animals that slowly ingest potentiallytoxic concentrations do not become affected. In addition, because the population of oxalate-degradingmicroflora in ruminants increases with gradual exposure to higher concentrations of oxalate for a fewdays, such animals are able to consume considerably greater quantities of toxic plant material than areanimals that rapidly consume a single comparable dose (James & Butcher 1972).
Most reported cases of Dock poisoning in Britain and in the USA have involved sheep. It has also beennoted that well-fed sheep can tolerate considerably greater doses of ingested oxalate than can animalssuffering from restricted feed and water intake (Cooper & Johnson 1998).
Uses
R. crispus has a long history of use in herbal medicine. It produces an acid called rumicin, andthe root contains chrysarobin (chrysophanolanthranol) and other substances of similar composition. Thelarge leaves of both common 'Dock' or 'Dockens' species, ie R. crispus or R. obtusifolius,are very generally reached for in many communities when anyone suffers a nettle sting, since the leaf orits sap, provides a welcome cooling effect when rubbed on the affected skin (see the Urticadioica (Stinging Nettle) species account on this website). Root extracts of R. crispushave been used as laxatives, astringents in piles, rheumatic tonics and for blood diseases, chronic skindiseases and jaundice. A homeopathic tincture made from unopened inflorescences has also been used totreat sore throats and coughs (Grieve 1931).
Names
The genus name 'Rumex' is an old Latin name for Sorrel from Pliny derived from the Latin 'rumo' tosuck, from the Roman habit of sucking Sorrel leaves to allay thirst (Johnson & Smith 1946). TheLatin specific epithet 'crispus' means 'curly', a reference to the undulate, crisped leaf margin of theplant (Gilbert-Carter 1964). The English common name 'Curled Dock' is a typical book name based on thescientific name and the general appearance of the leaf. As such, it has no folklore significance, andBritten & Holland (1886) do not list any other English names specifically attached to R.crispus. However, the commonly used names 'Dock' and 'Docken' apply equally to thisspecies and to its very common relative, R. obtusifolius (Grigson 1987; Allen & Hatfield2004), their most frequent use being as a coolant for nettle stings, burns and scalds.
Threats
None.
Native, locally frequent to common, widespread. Eurosiberian southern-temperate, but widely naturalisedin both hemispheres and now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
May to November.
Growth form and preferred habitats
Docks in general are a difficult group to identify accurately, really requiring well-grown plants withripe fruits to be certain of naming them correctly. To make matters worse, P.D. Sell carried out adetailed study around Cambridge and found that early flowering stems often did not produce viable fruitand seeds, even when the plant otherwise appeared a good specimen of the species. The same ticket-markedplant would then form ripe fruit and viable seed on fresh stems later in the year. Thus the same plant,whether species or hybrid, could be sterile or fertile at different times of year according tocirc*mstances (Sell & Murrell 2018).
R. conglomeratus is a very variable, short-lived, tap-rooted perennial that typically flowers fromJuly to October. Along the several inflorescence branches, leafy bracts occur on the basal two-thirds tothree-quarters or even more of their length in this species. In comparison, R. sanguineus (WoodDock) has its inflorescence bracts confined to the basal one-third to one-half of each flowering branch(Stace et al. 2015). The greater extent of bracts in the lower part of the Clustered Dockinflorescence, its more open, wider-spreading flowering branches, and the fruiting sepals (or tepals)with three pronounced swollen, oblong, corky tubercles per flower surrounding the solitary nut orachene, together help to distinguish R. conglomeratus from the rather similar R.sanguineus, which in Fermanagh is the more widespread species of the two (Lousley & Kent1981; Garrard & Streeter 1983).
Essentially, R. conglomeratus is a pioneer colonist, most frequently associated with muddy,somewhat disturbed, open, unshaded, low-lying ground around eutrophic lakeshores, stream-sides and baresoil or patchy vegetation near ditches, paths and gateways. Typically, these areas are subject toflooding in winter, or occasionally they temporarily flood after heavy or prolonged rainfall at othertimes of year. The occurrence of the species is associated with the high water mark at which floatingachenes (nutlets) become stranded amongst other drift materials and where, after winter exposure, theymay germinate in the spring and establish (Vergnano 1966; Lousley & Kent 1981; Roberts &Boddrell 1985).
R. conglomeratus is also very frequently associated with wet, muddy, artificial, man-madehabitats, or those heavily influenced by man and/or his stock animals (ie like R. crispus (CurledDock), it is a definite apophyte). Sometimes it also behaves as a ruderal on wet or waterlogged wasteground, eg in abandoned quarries, waste tips and gardens. It also more rarely appears on damp to wet,grassy roadside verges.
Buried achenes of R. conglomeratus can survive burial in soil for more than one year but less thanfive (Thompson et al. 1997). In comparison, R. sanguineus displays a rather wider range ofecological tolerances than R. conglomeratus, including growing in shade and in quite a lot drierground, eg on wood margins, waysides and in other ruderal situations.
Apart from its special soil-water relations, Clustered Dock requires or is confined to open, ± unshaded,moderately fertile, mildly acid to calcareous situations. Its presence in meadows, pastures and roadsideverges speaks of its tolerance of occasional mowing and/or some degree of grazing pressure, but it isdefinitely commonest in the more open, pioneer communities of intermittently disturbed habitats whereaggressive competition from vigorous accompanying species is absent or severely limited.
Fermanagh occurrence
Locally, R. conglomeratus has been recorded in 128 tetrads (24.2%), while the comparable figurefor R. sanguineus is 204 tetrads (38.6%). Clustered Dock, nevertheless, is a locally frequent oreven common plant in Fermanagh, being quite typical of rather wet, marshy, lowland habitats, especiallyaround the margins of lakes and ponds. It is frequent about both parts of Lough Erne and along the banksof the River Finn and its lakelets. However, as the tetrad distribution map indicates, it is only thinlyscattered elsewhere in the VC.
Hybrids
Hybrids are formed with eight other Docks which share wet ground with R. conglomeratus, includingseveral that are common or locally abundant species in our area, R. sanguineus (Wood Dock), R.hydrolapathum (Water Dock), R. crispus (Curled Dock) and R. obtusifolius(Broad-leaved Dock) (Lousley & Kent 1981). However, none of these Dock hybrids haveyet been identified in Fermanagh. The hybrid with R. sanguineus (Rumex × ruhmeri Hausskn.) is the most likely one to occur but it isdifficult to recognise since the parent species and their variability are very similar (Stace etal. 2015).
Hybrids are said to resemble either parent, or they may be intermediate between the two in manycharacters. The hybrid is also fertile which compounds the identification problem (Sell & Murrell2018). The number of fruiting tubercles can be one, two or three but it is most often three (which isthe usual number in R. conglomeratus), although in the hybrid one of the three is larger. Thetubercle shape in the hybrid is typically elongate (not ± spherical as is usual in the single tubercleof R. sanguineus) (Stace et al. 2015).
Since the pattern of variation shown by the hybrid tends to overlap that of each parent, itsidentification is only convincing when it shows not only reduced fertility but also a mosaic of parentalcharacters or several intermediate characters. In Stace et al. (2015), there are a totalof just 15 widely scattered hectads across Ireland with records of this particular hybrid.
Toxins
Like other Docks, R. conglomeratus contains oxalates and other toxins, which might deter some ifnot all kinds of herbivores.
Irish occurrence
The New Atlas shows R. conglomeratus is quite frequent and widespread in Ireland, ifsomewhat patchy in its occurrence. However, there are areas of the N & W (ie Donegal (VCs H34, H35)& Connemara (H16)), where it is absent or unrecorded. This might be partly explained by theprevalence of strongly acid, nutrient-impoverished soils in these areas, together with a high proportionof upland terrain which would also be unsuitable for this species.
British occurrence
The New Atlas map shows that in Britain Clustered Dock is widespread in lowland England and Walesand rare or absent in mountain areas. It becomes increasingly scarce and more obviously apophytic as onemoves north into Scotland (Preston et al. 2002). In NE Scotland (eg near Inverness), the nativestatus of the species was regarded as 'doubtful' by Mary McCallum Webster (1978), and she rejected aserrors many older records in the N & NE of the country.
European and world occurrence
R. conglomeratus is widespread throughout W, C and S Europe and N Africa, in the north reachingDenmark and the southern tip of Sweden, but only an ephemeral alien elsewhere in Scandinavia (Jalas& Suominen 1979, Map 460; Jonsell et al. 2000). It has been spread, presumably by agriculture,widely beyond its native range into the Azores, S Africa, N Turkey, the Caucasus, N Iran, C and E Asia,N and S America, S Australia and New Zealand (Hulten & Fries 1986, Map 668).
Names
The genus name 'Rumex' is an old Latin name for Sorrel from Pliny derived from the Latin 'rumo' tosuck, from the Roman habit of sucking Sorrel leaves to allay thirst (Johnson & Smith 1946). TheLatin specific epithet 'conglomeratus' means 'crowded together' and refers to the clustered flowers andpossibly the three tubercules on the inner tepals surrounding the nut or achene fruit (Gilbert-Carter1964). The English common name 'Clustered Dock' is a mere modern book name of no folklore significancebeing a straightforward translation of the specific scientific name.
Threats
None.
Native, frequent and widespread. European temperate, introduced in N America, S Africa and Australia.
1900; Praeger, R.Ll.; Co Fermanagh.
April to February.
Growth form and identification
In the past, there has been some confusion between this short-lived, rosette-forming perennial and theclosely related R. conglomeratus (Clustered Dock), but the identification differences are nowmuch clearer and better known (Lousley & Kent 1981; Stace 2010; Sell & Murrell 2018). R.sanguineus normally grows 30-60 cm tall and produces a few upright flowering stems that arisedirectly from the rootstock. Basal rosette leaves are 15-30 cm, oblong lanceolate, and in one variety ofthe plant (var. sanguineus), they develop a strongly marked blood-red or purplish coloration inthe midrib and veins (Sell & Murrell 2018). The stems and inflorescence branches of this variety arealso often suffused with purplish-red, and together with the leaves this makes the plant sufficientlyattractive to induce a small minority gardeners to actively choose to cultivate it (Griffiths 1994). Thecommon, wild form of the plant is var. viridis Sibth., which does not have this strong degree of red orpurple coloration.
At the flowering stage, R. sanguineus can be separated from R. conglomeratus (ClusteredDock) by the panicle being less diffuse on account of its branches being more acutely and morevertically arranged. The inflorescences of both species have sparse, widely separated flowering cymesscattered along their length. The inflorescences of R. sanguineus are leafy only at their extremebase. In addition, the three inner, tongue-shaped sepals (or tepals) of each newly opened flower areconspicuously unequal in size, and at the fruiting stage, usually only one of them bears awell-developed, swollen, almost spherical tubercule, c. 1 mm in diameter, which is as wide as, or widerthan the tepal that bears it (Lousley & Kent, 1981; Webb et al. 1996).
Variation
The characteristic feature that probably is most regularly relied upon to distinguish R.sanguineus from R. conglomeratus is the solitary swollen tubercule on the inner tepal ofthe fruiting flower. However, sometimes this character can be unreliable: in addition to the largetubercule on one tepal, occasionally the other two tepals in the fruiting flower surrounding the nut mayeach display minor, vestigial tubercule swellings, or very rarely all three tepals surrounding thesingle, trigonous, nut-like fruit bear large, globose tubercles like those of R. conglomeratus.Variants with a well-developed tubercule on all inner tepals have been collected in several places in SSweden (Jonsell et al. 2000).
As is always the case, a minimum of two characters should be tested to distinguish between similarspecies; in this case, the very unequal size of the three inner tepals of the R. sanguineusflower prior to fruiting would provide the necessary confirmation.
Three varieties are recognised by Sell & Murrell (2018), namely: var. sanguineus mentionedabove, with parts suffused blood-red; var. sanguinalis (Moss) P.D. Sell, which has leaves, stemsand inflorescences ± suffused with rusty red; and var. viridis Sibth., the common native formwith leaves, stems and inflorescences ± green.
Comparison of the preferred habitats of these two Dock species
The difference between these two rather similar Dock species, R. sanguineus and R.conglomeratus, can readily be related to their habitat requirements and tolerances: R.sanguineus occurs in lowland areas in both half-shade and full sun conditions on a range of soiltypes (although probably more frequently on calcareous substrates). In terms of habitat, R.sanguineus ranges from the margins of damp deciduous woods and hedgerows to ± dry roadsideverges. More rarely it can appear as a ruderal on waste ground. On the other hand, R.conglomeratus is more ecologically confined, being usually found in much wetter or damperconditions than R. sanguineus – essentially it is frequent only in low-lying marshy ground onlakeshores, stream-sides and ditches, including winter-wet meadows and pastures, although it too canrather more rarely than R. sanguineus behave as a ruderal on disturbed ground in quarries, wastetips and in gardens (Lousley & Kent 1981).
Hybrids
R. sanguineus forms hybrids with R. crispus, R. conglomeratus (see that speciesaccount on this website), R. obtusifolius (Broad-leaved Dock) and R. pulcher (Fiddle Dock)(Lousley & Kent 1981). Of these, the hybrid with R. obtusifolius (Rumex × dufftii Hausskn.) is the most frequently recorded in Ireland, havingbeen found in a total of 18 hectads widely scattered across the island (Stace et al. 2015).
Fermanagh occurrence
Wood Dock has been recorded slightly less often than Clustered Dock in Fermanagh but it is considerablythe more widespread of the two species, occurring in 204 of the tetrads, 38.6% of those in the VC. LikeR. conglomeratus, Wood Dock is most commonly found in the Fermanagh lowlands, but in this case itis also scattered across the Western Plateau at moderate altitudes.
British and Irish occurrence
R. sanguineus is a widespread and quite frequent plant in S England, Wales and throughout most ofIreland (although here in many areas it is rare or absent down the W and SW coasts). Further north inBritain, however, it becomes scarce, and in Scotland beyond the River Clyde it becomes progressivelyrarer and more coastal in its occurrence (Preston et al. 2002).
European and world occurrence
Wood Dock is widespread in W & C Europe, but it does not penetrate beyond 60N in Scandinavia,confining it to the southern shores of that large peninsula landmass. To the south of Europe, it extendsonly thinly into the Iberian peninsula, Italy, the Balkans, Sicily, Corsica and Sardinia. Beyond this,as a native species it reaches Turkey, the Caucasus and N Iran. It is an introduction in E North Americaand in scattered stations in temperate regions of the S Hemisphere, including S Africa and Australia(Jalas & Suominen 1979, Map 461; Hulten & Fries 1986, Map 669; Jonsell et al. 2000).
The record of occurrences of R. sanguineus in New Zealand in the 1930s and later has been rejectedby Webb et al. (1988, p. 983), due to the absence of vouchers, and the possibility of confusionwith R. conglomeratus, which is also present as an alien throughout both islands.
Names
The genus name 'Rumex' is an old Latin name for Sorrel from Pliny derived from the Latin 'rumo' tosuck, from the Roman habit of sucking Sorrel leaves to allay thirst (Johnson & Smith 1946). TheLatin specific epithet 'sanguineus' translates as 'blood-red' from 'sanguis', 'inis', 'blood'(Gilbert-Carter 1964), a reference to the colour that generally develops in the leaf mid-rib and sideveins. The English common name 'Wood Dock' is a typical book name that merely indicates one of the mainhabitats of the plant.
Threats
None.
Native, very common and widespread. European temperate, but widely naturalised in both hemispheresincluding N America, although not circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
A very variable, large-leaved, rosette-forming, indigenous perennial with a stout and often deeplypenetrating taproot, capable of producing a flowering stem up to 150 cm tall, R. obtusifoliusoccurs scattered through lowland grassland, field edges and gateways, lakeshores, streamsides,riverbanks, wood margins, as well as in a range of ruderal, disturbed and waste ground habitats. Theabsence of the species from upland areas and mountain tops in Britain & Ireland is more associatedwith soil conditions (ie the avoidance of peat) than climatic factors, such as length of the growingseason or winter cold, since it grows inside the Arctic Circle in coastal Norway (Cavers & Harper1964; Jonsell et al. 2000). R. obtusifolius is a very common, troublesome perennial weedof fields cut for silage, and it is listed as one of the world's worst weeds, being important in no lessthan 37 countries (Holm et al. 1977).
Despite its great frequency across these islands and Europe, Broad-leaved Dock has a low competitiveability as a seedling and it cannot become established in closed vegetation plant communities. Once itproduces its deep, branching taproot, however, it becomes very difficult to eradicate. The large leavesand tall stem give the species a great advantage over shallower-rooted grasses and herbs, particularlyin drier soils. Nevertheless, it does not persist in well-managed permanent grassland and it is regardedas somewhat easier to eradicate than the closely related R. crispus (Curled Dock).
Variation
R. obtusifolius is a highly variable species and Stace (1997, 2010) lists three varietiesoccurring in the British flora based on the shape and dentation of the tepals (var. obtusifolius,var. microcarpus Dierb. (= subsp. sylvestris (Wallr.) Čelak.), and an intermediate betweenthese, var. transiens (Simonk.) Kubát.). Stace states that only the first of these is native. Onthe other hand, in Scandinavia, Jonsell et al. (2000) raise these three forms to subspecies, andthe authors state that, "only one of them (subsp. sylvestris) is probably native in lessdisturbed habitats". They go on to comment that the distributions of the three forms has becomewidened by increased trade and travelling, and that "their distinctness is now much obscured byhybridization in secondary habitats".
Sell & Murrell (2018) have gone deeper into the variation of the species in Britain & Ireland andhave described three subspecies and no less than five varieties of subsp. obtusifolius. Of allthese varied forms, they regard only var. obtusifolius as probably native in Britain &Ireland.
However, neither subspecies nor varieties were described or distinguished in any of the eight editions ofWebb's Irish Flora (Webb 1977; Webb et al. 1996; Parnell & Curtis 2012), and to dateno species subdivisions have been recorded in Co Fermanagh.
Flowering reproduction
R. obtusifolius plants tend to flower freely from the second year of growth onwards andindividuals may continue doing so for several years. Large individuals can have over 1,000 self-fertileflowers per inflorescence and they may flower and fruit twice during a season, in early and again inlate summer. The number of seed produced varies per plant from less than 100 to more than 60,000 in asingle year (Cavers & Harper 1964). The tendency for plants to die after seeding is not as marked asit is in R. crispus. Achenes (nutlets) are shed continuously from late summer to winter and theseed can germinate in any month of the year.
Seed dispersal
The lightweight fruits are mainly dispersed by wind, but they have also been found in cattle dung whichsuggests, if not proves, that they have been eaten and can be transported internally (Salisbury 1964).Spines on the perianth segments additionally assist dispersal by attaching the nut-like fruitsexternally to the coats of animals, including man, purely by chance. Seeds may also cling to mud orbecome embedded in it and be accidently transported (Cavers & Harper 1964).
Abundant R. obtusifolius populations have been associated with intensive silage production and theapplication of organic manures (slurry) to grassland. Courtney (1985) found that seed remained viableafter storage in cattle slurry for 12 weeks. The application of such slurry may not only act as a meansof seed dispersal but, even more significantly, concentrated deposits of slurry may also cause localiseddestruction of the grass sward, introducing gaps, reducing competition and favouring the colonisationand establishment of seedling Docks in grassland (Humphreys et al. 1997).
Germination and dormancy
Seed is extremely variable (polymorphic) with respect to germination and dormancy, but it can persist for40 years or more and may form a very large seed bank. Cideciyan & Malloch (1982) found that thegermination rate of seeds of R. obtusifolius increased as seed size decreased. They also showedthat although initial growth was slower from smaller seed, no difference in plant size (biomass) wasdetectable at the end of the growing season.
Vegetative reproduction
Fragments of underground stem and root can reproduce the species after ploughing. Pino et al.(1995) found that R. obtusifolius had a 'phalanx' type of invasive clonal vegetative growth ingrassland, the main structure involved being the stem. Regeneration can occur at depths up to 15 cm, butgrowth and recruitment is prevented in wet soils (ie those maintained at water-logging or fieldcapacity). This is very significant in a Fermanagh context, since local soils are often this wet.Despite the possibility of vegetative spread, the major means of R. obtusifolius reproduction anddispersal is undoubtedly by seed.
Toxicity
Like other Docks, R. obtusifolius contains oxalates and other toxins which are sometimes said togive it a pleasantly sharp taste, a fact which apparently allows it to remain palatable to some forms ofstock (Cooper & Johnson 1998). Mabey (1972) reported that R. obtusifolius (like R.crispus) has been used as 'wild food' in human communities, the leaves being collected veryyoung, cooked and eaten with bacon or ham and a little vinegar. The leaves of both species are bittertasting, R. obtusifolius being the worst of the two in this respect. In 1990, a Spanish familysuffered acute oxalate poisoning after such culinary use of R. crispus, resulting in one fatality(Reig et al. 1990), so its use in the human diet is very definitely not recommended.
Other agricultural research on fodder plants suggests that R. obtusifolius is quite nutritious,being high in protein, low in crude fibre and rich in Mg (Fairbairn & Thomas 1959; Wilman &Riley 1993), and it can be acceptable to dairy cows (Courtney & Johnston 1978). As with otherspecies of Rumex, the conditions under which oxalate poisoning occurs are not clearly defined(Cooper & Johnson 1998). It is generally refused by cattle, sheep and by even more so by chooseyanimals such as horses. Derrick et al. (1993) found that while the voluntary intake of dried Dockby sheep was satisfactory in their experiments, fresh Dock, especially if chopped, was avoided. Theoff-putting factor was presumed to be either the taste or the smell of the fresh Dock, which appears toprovide the plant with an effective defence against at least some herbivores.
Fermanagh occurrence
R. obtusifolius is the second most widespread and abundant Dock species in Fermanagh,occurring in 459 tetrads, 86.9% of those in the VC. The most abundant Dock in Fermanagh is R.acetosa (Common Sorrel).
R. obtusifolius is widespread and very common throughout Fermanagh, except on the highest, wettestand most acidic ground.
British and Irish occurrence
The New Atlas map shows R. obtusifolius recorded in 95.5% of hectads across Britain &Ireland. It is present everywhere except on the highest ground and in the wettest, most acid peatlandsof Scotland and Ireland (Preston et al. 2002).
European and world occurrence
R. obtusifolius occurs in 90% of European territory and has been accidently introduced to everycontinent (Grime et al. 1988). The European distribution mapped by Jalas & Suominen (1979,Map 465) shows R. obtusifolius widespread in W & C Europe and on the W coast stretching fromS Spain to beyond the Arctic Circle in Norway. The distribution thins gradually southwards andeastwards, becoming increasingly scattered in the Iberian, Italian and Balkan peninsulas. Althoughplotted as native and indigenous throughout most of W, S & C Europe, the accompanying text makes itclear that the species is not native in some areas including Finland, the Azores, "and perhapselsewhere". R. obtusifolius is shown as introduced on the Macaronesian islands (Madeira, theCanaries and Azores), and to the north in Finland, and up the Baltic coastline of Sweden (Jalas &Suominen 1979, Map 465).
Beyond Europe, R. obtusifolius is very widely introduced in N America and is especially frequentin eastern states of US and Canada. It is also reported from S America, Cuba, S Africa, Japan,Australia, Tasmania, New Zealand and many remote island groups including the Falklands Isles (Hultén& Fries 1986, Map 670).
Possible biological control mechanisms
Studies of grazing by the chrysomelid beetle, Gastrophysa viridula De Geer on Broad-leaved Dock,showed that it resulted in a reduction in both seed number and seed weight (Bentley et al. 1980).In combination with similar debilitating effects on infection by the rust fungus, Uromycesrumicis (Schubiger et al. 1986), there may just be the possibility of a biologicalcontrol mechanism being developed sometime for this troublesome and very common weed (Hatcher etal. 1994).
Uses
R. obtusifolius is sufficiently common for the astringent properties of the leaf to be frequentlyused by both children and adults to ease the pain of a nettle sting (see the Urtica dioica(Stinging Nettle) species account on this website). Thus, together with the other common 'Dock' or'Docken' species, R. crispus (Curled Dock), from which R. obtusifolius is very oftenundistinguished, the latter has earned its reputation as a generally useful medicinal herb. In this wayit has accumulated a good supply of folk names (see below). Although the leaf alone is often applied toa nettle or an insect sting, traditionally a far more effective poultice was made from the roots, bycleaning, peeling and crushing them to a pulp to make a paste (Darwin 1996).
Apart from stings, Dock is used in herbal medicine to treat the pain of burns and scalds, to relieverashes and to staunch bleeding from a cut (regarded as the second best herb to Plantain (Plantagomajor or P. lanceolata), which is greatly and universally preferred for that purpose)(Allen & Hatfield 2004). Other medicinal uses are to relieve pain, include leaves for headaches andrheumatism. Boiled seeds have also been used to draw the pus from a wound, to cure boils, to treat allkinds of cough, and for colds and bronchitis. A liquid or tea processed from Dock roots has beenconsidered a curative tonic for cleansing the blood, for keeping scurvy at bay and for liver trouble,including jaundice. The leaves have been used to remove warts and corns (Grieve 1931; Allen &Hatfield 2004).
Names
The genus name 'Rumex' is an old Latin name for Sorrel from Pliny derived from the Latin 'rumo' tosuck, from the Roman habit of sucking Sorrel leaves to allay thirst (Johnson & Smith 1946). TheLatin specific epithet 'obtusifolius' translates as 'blunt-leafed' (Stearn 1992).
The English common name 'Broad-leaved Dock' is a modern book name based on the Latin scientific specificepithet and as such has no folklore connections. A total of 16 interesting alternative English commonnames from around Britain & Ireland are listed by Grigson (1987). Apart from 'Dock' and 'Docken'mentioned above, the names 'Butter Dock', 'Butter Docken' and 'Smari Dock' (a Scottish name) refer tothe previous convenient practice, in the absence of waxed paper, of wrapping lumps of butter in thelarge, broad leaf of the plant.
Threats
None.
Native, very rare. European temperate, also reported from Senegal, W Africa.
29 August 1991; NI Lakes Survey; Upper Lough Macnean, N of Inishteige Island.
Growth form and preferred habitats
This tiny, inconspicuous, prostrate or low-growing, opposite-leaved waterweed has its Irish strongholdsin the far SW of the island and in Co Down (H38), with occasional other scattered stations, especiallyalong the west coast. On intermittently exposed bare, soft sandy or peaty mud, silt, or fine gravel,Six-stamened Waterwort is an opportunistic colonising, minute, green, aquatic annual with fragile,creeping stems. It typically occurs in or on the margin of bodies of still or sheltered, shallow,mesotrophic water up to 50 cm deep. When exposed to the air or when just covered with water, the slenderstems creep across the mud, rooting at every node. Individual plants, or several growing closelyintermingled, can form temporarily dominant, but ephemeral, patches on the mud.
Much more rarely, E. hexandra grows permanently submerged in deeper, open water situations, oftenin sheltered bays of larger water bodies. In these deeper waters, the plant assumes a more stragglygrowth, as stems that are only barely anchored to the lake bed, reach up towards the water surface andlight (Mitchell 1983). In the deep water situation, the species behaves as a more-or-less persistentperennial, although it usually fails to flower and is forced to rely on vegetative growth for anyincrease or spread (Preston & Croft 1997). When submerged in shallower water, E. hexandraflowers and fruits each year, although like Callitriche stagnalis (Common Water-starwort),individual plants probably are not long-lived (Salisbury 1967; Hawkins 1982).
While E. hexandra appears to prefer moderately nutrient-rich, mesotrophic conditions, inherentvariation enables it to tolerate a much wider range of water chemistry than this implies. Occasionallyit can occur on peat at the edges of moorland lakes (A.J. Silverside, in: Stewart et al. 1994).In reality, E. hexandra eschews only extremely calcareous hard water situations (Salisbury 1967;Preston & Croft 1997).
Elsewhere in Britain & Ireland, E. hexandra and its near relative E. hydropiper(Eight-stamened Waterwort), generally associate with Littorella uniflora (Shoreweed),Eleocharis acicularis (Needle Spike-rush) and Baldellia ranunculoides (LesserWater-plantain), all of which occur quite frequently in Fermanagh. Like these other smallspecies, both Elatine species cannot compete with tall perennial colonists of marginal muddyhabitats, such as reeds and the larger sedges. Rather they are restricted to disturbed areas of exposedmud, where the larger, vigorous, secondary colonists are prevented from establishing by the instabilityof the habitat, due to, for instance, wave action, grazing stock animals or boating activity.
The physical extent of the water-marginal, bare mud habitat is determined by local rainfall andconsequent water-levels in lakes and pools. Therefore population numbers of E. hexandra are knownto fluctuate enormously from year to year (Salisbury 1967; A. J. Silverside, in: Stewart et al.1994).
Although mainly a lowland species, Six-stamened Waterwort does occur in sites up to 440 m in N Kerry(H2), and to altitudes of almost 500 m in the Scottish Highlands (Preston & Croft 1997).
The plant is so small and so very easily overlooked (or mistaken for young Callitriche specimens),a careful eye needs to be kept out for it on any open, relatively bare, unvegetated areas of damp,recently exposed mud on the margins of lakes, ponds, or even around ditches and puddles.
Reproduction
The tiny flowers are only 3-4 mm across and they are produced from July to September. E. hexandraflowers have three delicate petals, while those of its close relative E. hydropiper have four. Inboth species, the petals are usually pale pink, although occasionally they are white. The trimerousflowers (ie with parts in threes) are habitually self-fertilized and, indeed, if submerged they remainunopened and are cleistogamous (ie selfed while in bud). However, occasionally insect visitors mayachieve cross-pollination of aerially exposed flowers. Elatine flowers possess nectar glands,which suggests the flower family to which they belong was once terrestrial (Mitchell 1983). The fruit isa capsule with three (or rarely four) locules, each containing numerous minute seed. Hooker (1884)claimed the seed number generally lay between 24 and 36 and was a multiple of three (ie three loculeseach containing 8-12 seeds). A study by Salisbury (1967) examined the contents of 406 capsules and foundthe mean number present in each was 37.5 seeds. The number of seed per capsule varied between five and72. Salisbury also showed that loculi of the same fruit could have varying numbers of seed in them, andthat nutrition and the prevailing light climate probably were the major factors determining the numberof seed produced.
Seed production is so rapid that under favourable growing conditions two fruiting generations can oftenoccur in a single season. This creates an enormous dormant, overwintering seed population typical ofpioneer species of intermittently exposed mud (Salisbury 1967).
Buried seed survival
According to Mitchell (1983), a persistent, long-lived soil seed bank is formed, yet the survey carriedout in NW Europe does not include this species (Thompson et al. 1997). However, the behaviour ofthe species with its intermittent, almost explosive reappearance after a quasi-simultaneous germination,which in turn follows a drought period that provides a suitable open, muddy environment, stronglysupports the notion that long-term dormant seed survival exists in E. hexandra. Buried seedremains dormant until water levels drop at the right time of year and the light intensity is sufficientto trigger germination (Mitchell 1983).
Seed dispersal
Dispersal of the minute seed is readily imagined by flotation and by current movement in linked watersystems. Salisbury (1967) found that about 8% of freshly released seed in an experiment he conductedwere still afloat after 24 hours in standing water. In moving flood water, the tiny, lightweight seedwould probably remain floating longer than in still water and could certainly travel some distance.Since the seed coat possess mucilage and becomes very sticky when wet, the muddy feet and feathers ofwaterfowl are probably responsible for the spread of the species to new sites, such as those observed infreshly dug reservoirs or recently abandoned sand and gravel pits in S England (Salisbury 1967; Mitchell1983; Preston & Croft 1997).
Fermanagh occurrence
A number of detailed, systematic surveys made of aquatic habitats in Fermanagh in recent years have nowdiscovered four sites for this species in the VC. At the original station listed above the plant wasconfined to the summer exposed muddy shore of this large lake on the county boundary with Co Leitrim(H29). (NB Apparently there is no voucher.) The plant is so small and inconspicuous that at the time itwas believed it could easily enough have been overlooked at other suitable sites in Fermanagh, althougha great deal of time and effort had already been spent surveying aquatic habitats in the VC.
In 2006, a further survey of water quality at selected lakes in N Ireland was commissioned by theEnvironment & Heritage Service and this uncovered E. hexandra at three additional stations inNW Fermanagh, again near the county boundary but this time with East Donegal (H34): Lough Vearty, LoughRushen and Lough A Waddy. RHN refound the plant at Lough A Waddy in October 2010: it was picked upamongst lakeshore plant debris consisting mainly of Isoetes lacustris (Quillwort) plus otherisoetids and originally it was mistaken for a Callitriche (Water-starwort) species. Thisis a very difficult plant to identify by normal 'keying out' in a standard Flora, but Haslam etal. (1975) (British Water Plants) is recommended. The leaf venation is a gooddistinguishing character for separating vegetative material of Elatine from Callitriche(M. Jebb, pers. comm. 2010).
Irish occurrence
Elsewhere in N Ireland, since the early 19th century, E. hexandra has occurred rare and scatteredaround Cos Armagh, Down, Antrim and Londonderry (H37-H40), although in recent times it is principallyfound in a few lakes in Co Down (Flora of Lough Neagh, FNEI 3, NI Vascular PlantDatabase).
Considering how easily E. hexandra could be overlooked or mistaken for a Callitrichespecies, even with the modern identification guides available, it is all the more remarkable that H.C.Hart (1898) listed this minute aquatic species in his Flora of the Co Donegal in four of theeight subdivisions he made of the county (VCs H34, H35). One of the stations he listed is Bannus Lough,which is less than 500 m from the Fermanagh boundary and only 4 km from Lough A Waddy: Hart describedE. hexandra as being, "plentiful at Bannus Lough, 2½ miles [4 km] SW of Pettigo, at theroots of Equisetum limosum [= E. fluviatile] and Carex ampullacea [= C.rostrata] in mossy mud."
Elsewhere in Ireland, the New Atlas hectad map shows E. hexandra thinly and widelyscattered, mainly in coastal counties, and most frequent in the far south-west (Preston et al.2002).
British occurrence
The New Atlas hectad map shows that E. hexandra is a decidedly scarce species, thinly butwidely scattered across the whole latitudinal range of Britain from the Channel Isles to Shetland.Having said this, the distribution has a definite western preponderance, most especially displayed inScotland and in Wales (Preston et al. 2002).
Reasons for rarity
Knowing the huge potential seed population of E. hexandra from its biology, its considerabledispersal ability and prolonged dormancy in soil, the comparative rarity the species displays acrossBritain & Ireland cannot be entirely due to the plant's small scale, prostrate habit and consequentreduced competitive ability, plus its avoidance of hard, limestone waters and soils. Rather, theobserved scarcity must be attributed to the marked degree of intermittence of mud exposure in suitablesites, together with the rapidity with which such areas are colonised by taller, more vigorous plants ifthe exposure is prolonged (Salisbury 1967).
European occurrence
E. hexandra is very much confined to W & C Europe. On the W coast of Europe, itstretches from S Norway to Portugal, but is almost absent from Italy south of the Alps and entirelyabsent from the Balkans. It is also reported from the Azores and from Senegal, W Africa (Hultén &Fries 1986, Map 1347; Clapham et al. 1987; Preston & Croft 1997).
Names
The genus name 'Elatine' was an ancient Greek name used by Dioscorides for an unknown plant, but as hewas a medic employed in the Roman army, it must have had herbal properties. The name translates as'little fir tree' and is applied to the genus because of the appearance of one of the species unique inthe Family Elatinaceae, E. alsinastrum L. The latter is a continental European aquatic thatresembles a seedling conifer since it possesses an unbranched stem with whorled leaves (Gilbert-Carter1964; Mabberley 1987). The Latin specific epithet 'hexandra' means 'six stamened', which generallyapplies to this species (Gilbert-Carter 1964).
Threats
Cultural eutrophication, due for instance to runoff of agricultural fertilisers or slurry, causingnutrient enrichment that leads to massive algal blooms that can shade out and destroy suitable growingconditions for waterworts.
Introduction, neophyte, a very rare garden escape or discard.
8 July 2000; Northridge, R.H.; Riversdale Forest, Ballycassidy.
A popular, low-growing, evergreen, garden perennial subshrub native in Bulgaria and NW Turkey, theunderground rhizome of H. calycinum can spread rapidly through soil (Webb et al. 1996).The plant very easily outgrows its welcome therefore as a useful garden ground-cover and frequently endsup being uprooted and discarded. In the experience of the current author, it is shallow rooted and wasquite easily removed from a sandy soil in a Belfast garden when it became too invasive. Discarded plantsmay thus appear on waste ground or, as here in Fermanagh, where the solitary known record wasdiscovered, dumped in a secluded site in the coniferous plantation of Riversdale Forest.
Despite its low stature, in semi-shade H. calycinum is strongly competitive with native speciesand it is perfectly capable of forming dense clonal patches of long-persistence. It is known to havesurvived in some Irish sites for well over a century (Reynolds 2002).
H. calycinum tolerates a considerable range of soil conditions from dry to damp, moderately acidto neutral, but typically it prefers conditions mesic both in terms of nutrient- and base-status (Sinkeret al. 1985).
All plants of this species that grow in the wild in Britain & Ireland appear to derive from oneoriginal garden strain that was introduced to these islands in 1676 AD from a site near Istanbul by SirGeorge Wheeler (N.K.B. Robson, in: Green 1973). Fortunately, the flowers are self-incompatible to a highdegree and, therefore, Rose-of-Sharon very seldom sets seed in Britain & Ireland. The introductionof new garden forms of the species, however, may pose a threat, by overcoming this inherent barrier toseed production.
Plants are currently quite widely distributed throughout Britain and are particularly frequent in SEngland. Thankfully H. calycinum is very much more thinly recorded in Ireland and it seems to beabsent from much of the W, C and NW of the island. Despite its relatively large, showy flowers and anattractive sounding biblical name, Rose-of-Sharon remains a potentially invasive alien 'thug' (N.K.B.Robson, in: Preston et al. 2002).
The biblical 'Rose-of-Sharon' was most probably a bulbous rather than a woody plant: Narcissustazetta and Tulipa montana are two suitable candidates for this honour (Smit 1992).
Threats
A vigorous vegetative coloniser, it can form persistent clonal patches where it becomes established. Thepossibility of future seed production and rapid subsequent spread is quite worrying.
Native, frequent and widespread. Submediterranean-subatlantic, but widely naturalised, including in NewZealand.
1881; Stewart, S.A.; Cladagh River Glen NR (= the Marble Arch Glen).
Throughout the year.
Growth form and preferred habitats
A highly variable, perennial, lowland, stout, bushy subshrub 30-100 cm tall, with brownish-red erect orspreading shoots arising from a woody rootstock. Stems are ± round, but when young have two raised linesor ridges running lengthwise. Tutsan plants retain at least some of their paired, large, oval, obtuseleaves during our relatively mild oceanic winters and thus in W Ireland the species is a semi-evergreen,opposite-leaved plant.
Although there are more garden-worthy species and forms of the genus Hypericum available fromhorticultural suppliers, plants indistinguishable from the 'wild form' of H. androsaemum areregularly planted in gardens for their rather large yellow flowers with prominent yellow stamens in fouror five bundles and colourful, fleshy, berry-like fruit capsules. If, as very likely is the case,non-local forms 'escape' over the garden wall, the distribution of the native population inevitablybecomes obscured. The shrubby Hypericum species that occur in Britain & Ireland all have fouror five separate bundles of 10-25 stamens on long slender filaments that help make the flowersattractive to both gardeners and insect pollinators (Clapham et al. 1987).
Fortunately, the natural habitats of the species are fairly distinctive, being essentially characterisedby damp, semi-shaded conditions towards the more open areas or margins of woods and hedgerows,especially on base- or lime-rich soils. Another typical habitat of the species is sheltered areas inrocky ground or on cliffs, again particularly frequent in limestone districts. The shrub is completelyabsent from farmland, wetland and areas of strongly acidic peaty soils.
Variation
Leaves are sometimes tinged bronze with red pigment, in the same manner as the sepals generally aresemi-coloured. The leaf base sometimes clasps the stem (amplexicaul). Several named garden varietiesexist. Var. 'Albury Purple' has young parts suffused dull purple; var. 'Aureum' has leaves golden yellowlined; forma variegatum D. McClintock & C. Nels. has leaves variegated pink and white (Griffiths1994).
Flowering reproduction
Flowering takes place from June to August, the yellow flowers being solitary or in terminal,flat-topped clusters (umbellate cymes) of 2-11 blossoms. Individual flowers vary in size, usually 15-25mm in diameter, averaging around 20 mm. The green sepals are very unequal in width and are slightlylonger than the petals. They enlarge and become deflexed (bent back) when the plant fruits. In plantspossessing somewhat larger flowers than average, the sepals are sometimes tinged red by the sap thatgives the species its specific scientific name, 'androsaemum', meaning 'Man's Blood' (Robson 1973). Thered pigment is hypericin, and it is high concentrations of this compound that gives the black colour toglands on some other Hypericum species. The filaments of the yellow stamens of H.androsaemum ± equal the petal length. Petals and stamens are both deciduous, dropping off oncethe fruit begins to form, while the sepals and fruit remain attached to the plant all winter (Robson1990; Sell & Murrell 2018).
Although there appears to be little or no nectar present in flowers, abundant pollen attracts a widevariety of insect visitors including flies, butterflies, moths and bees (Fitter 1987). If insectpollination fails to occur the flowers self-fertilize. The fruit is a fleshy ovoid berry, either reddishbrown or bright red becoming shiny black as it matures. Each fruit contains numerous 1 mm long seeds,estimated at between 800-900 seeds per fruit (Lang 1987).
Fermanagh occurrence
H. androsaemum is frequent and widespread throughout Fermanagh, being often recorded in 204tetrads, 38.6% of those in the VC.
Seed dispersal
Although the black glossy fruits of Tutsan look as if they ought to attract birds, these visitorsare seldom, if ever, observed on the plant in the author’s experience, or in that of other naturalists.The English Hypericum specialist, Robson (1990), admits that he has never seen birds takeberries, and neither did Ridley (1930, p. 403), who discusses his observations at Kew. The 'berries'appear to often remain on the bush right through the winter and, to the human eye, they lookincreasingly unattractive as they shrivel with age. In the spring, they eventually split so that some ofthe seed may leak out, but this is hardly an effective dehiscence mechanism.
Circ*mstantial indirect evidence for presumed bird (or other animal) dispersal is implied by theobservation that H. androsaemum seedlings and plants frequently appear, even in gardens where noplants of it have been growing (Robson 1973). In New Zealand, this species has become a serious weedsubsequent to its introduction to the country around 1870. There, birds are presumed to take the berriesand transport the numerous contained seed, since the plant has frequently made the 'jump' from gardensinto wild wooded areas (Webb et al. 1988, p. 539).
Despite the lack of observations of birds taking the fruit in Britain & Ireland, there clearlyremains a strong possibility that this is happening, meaning that bird-sown plants of garden origin areescaping into the wild, at least in open habitats near habitation.
Locally in Fermanagh, the greatest probability of bird-sown seed would be around Enniskillen. There areno other large towns or villages in the county where seed dispersal of this nature is at all likely.
Fossil history
Fossil Hypericum seed can be identified to species level, but only one seed of H.androsaemum had so far been recorded in Ireland, at Gort, from sub-stage II of the Hoxnianinterglacial (a stage in Ireland renamed the Gortian), when Godwin (1975) was writing his major overviewof the topic. Nevertheless, this fossil is sufficient to comfirm Tutsan as a native plant at thatperiod. On its own, however, it does not allow us to say that it is indigenous in the currentFlandrian/Littletonian interglacial. Having said this, traditionally, at least, the species is regardedas native in Ireland, and while present throughout Britain apart from the far north, it is much morecommon (and more likely native), only in western and southern parts of the island.
British and Irish occurrence
The boundary between native and introduced Tutsan plants is not clear and we have to accept that in themore populated areas of Britain & Ireland, Europe and N Africa, H. androsaemum is widelynaturalised beyond its native range. As the overall distribution of this species is variously describedas Oceanic Southern (essentially a SW European group) (Matthews 1955), Atlantic-Mediterranean (Godwin1975), or Submediterranean-subatlantic (Preston & Hill 1997), clearly the mild aquatic climaticinfluences of the Atlantic and the Mediterranean are invoked and reflected in these geobotanicalgroupings.
The hectad distribution of H. androsaemum, as displayed in the New Atlas, is very much morewestern slanted in Britain than is the case in Ireland. This suggests a winter low-temperature growthlimitation exists, the shrub rather obviously avoiding the colder E coast of England and Scotland,except where the occurrence possibly (or probably) represents garden escapes. The distribution (bothsupposed native and presumed introduced), also peters out further north in W Scotland aroundUllapool (N.K.B. Robson, in: Preston et al. 2002).
European and world occurrence
H. androsaemum is native in W Europe from Belgium and Normandy southwards, becoming more local inSpain and Portugal. It stretches east through Italy, Corsica, Sardinia, Sicily, Yugoslavia and Bulgariato Turkey and Iran, the Caucasus and Asia Minor (Lebanon-Syria). It is also reported as being native inN Africa (Algeria, Morocco and Tunisia) (Tutin et al. 1968; Clapham et al. 1987;http://www.plantsoftheworldonline.org/taxon/urn:lsid:ipni.org:names:433187-1#distribution-map (websiteaccessed 8 February 2019).
H. androsaemum has been introduced to Maryland (USA) and to Chile. Tutsan was also introduced toNew Zealand (both N & S islands), the Antipodean Isles and Tasmania in the late 19th century as agarden ornamental due to its profusion of attractive yellow flowers. Once established in New Zealand, itproved very difficult to eradicate and it can become an invasive thicket-forming weed (Webb etal. 1988). In the last 20 years, it has spread into native scrub, poorer pastures and alongroadsides in several regions of New Zealand with high rainfall. It is most significantly affecting theRuapehu District and Bay of Plenty region in North Island, where it is a difficult and expensive alienthreat to the conservation of native vegetation including forest margins. Current research is lookingfor a suitable biological control mechanism for this weed(https://blog.invasive-species.org/2015/03/26/tackling-tutsan/ (website accessed 7 February 2019)).
Toxins
All members of the genus Hypericum contain a poisonous red, fluorescent glycoside pigment calledhypericin. It is a polyphenolic compound structurally similar to fa*gopyrin, the photosensitising agentfound in buckwheat (fa*gopyrum esculentum). Hypericin has been found to have antiviral andantidepressant properties. Animals eating plants containing hypericin may develop lesions on unpigmentedskin exposed to bright sunlight (ie photosensitisation occurs). This is most likely to occur on hairlessareas, such as eyelids, muzzles and udders. The skin reaction can take one to two weeks to develop insheep, cows, pigs and horses, and established lesions are slow to heal, perhaps taking months.
Lesions cause the animals to rub and scratch and open lesions can become scabbed and infected. Othersymptoms reported in horses include loss of appetite, debility, staggering and even coma. Post-mortemexamination of affected rabbits showed liver and kidney damage had occurred. Animals can recover, butonce affected by photosensitisation, the condition more quickly reappears after further consumption ofthe pigment (Cooper & Johnson 1998).
Other toxins present in Hypericum include a polyphenolic flavonoid, called hyperoside, plustannins and carotenes. However, it should be made clear that most cases of Hypericum poisoninginvolve H. perforatum (Perforate St John's-wort) and H. elodes (Marsh St John's-wort),rather than Tutsan (Lang 1987).
Uses
Medieval herbalists confused for some time H. androsaemum with the Agnuscastus (Chaste Tree) of Pliny and transferred the latter's supposed magic virtues to it,including the reputation of warding off evil spirits and of being an 'all heal', panacea for medicinalherbalists (Allen & Hatfield 2004). This undeserved reputation is reflected in the fact that in hercomprehensive book A Modern Herbal, Grieve (1931) ignores Tutsan altogether, although she doesgives a little space to uses for H. perforatum (Perforate St John's-wort). In other herbalsources, there is mention only of leaves being used to make a poultice for the prevention of 'marks' onthe body (bruises perhaps) (Allen & Hatfield 2004). Dried Tutsan leaves are slightly aromatic andthey were, or can be, used to perfume the pages of books, mainly as a luck token. The red sap and theberries can also provide a dye. Other magical folklore uses are mentioned below.
Names
There appears to be some confusion as to the origin and derivation of the genus name 'Hypericum'. TheClassical Greek form of the name is 'hypereikon' and it was used by Dioscorides (a medic in the Romanarmy) for an unknown plant. Differing spellings and pronunciations occur: eg Gilbert-Carter (1964) saysthat in England the 'i' is wrongly treated as short. Some derive the name from the Greek 'hyper', overor above, and 'ereike' a heath, possibly referring to the natural habitat of some species in the genus(Johnson & Smith 1946). Another suggestion is that the second element derives from 'eikon', meaninga picture, and thus 'Over or above a picture' might refer to the belief in the magic properties of theplant, which had the power to dispel evil spirits, and is why the Devil pierced the leaves with a needle(the translucent glands).
The flowers of some Hypericum species were placed above religious images or shrines to ward offevil at the ancient midsummer festival of Walpurgisnacht, which later became the feast of St John (24June), when they are in flower. Hence the name 'St John's-wort' (Stearn 1972; Gledhill 1985).
The Latin specific epithet 'androsaemum' is derived from the old Greek generic name of the plant'androsaimon', from 'aner', 'andros', man and 'haima', blood, thus translating as 'man's blood', areference to the red sap of the plant (Gilbert-Carter 1964; Stearn 1972).
The most frequently applied English common name, 'Tutsan', is a corruption of the French name, 'Toutesaine', meaning 'all wholesome' or 'all heal', a reference to the many supposed medicinal virtues of theplant. Tutsan is thus a plant of good reputation, but the species was ascribed its properties in error,having being mistaken by medieval herbalists for the plant that Pliny described as 'Agnus Castus' ,'Chaste Tree'. In reality, the latter is Vitex agnus-castus L., a tree-forming Mediterraneanmember of the Verbenaceae, the Vervain family, but it was not until the 17th century that herbalists inW Europe realised their earlier mistake in applying this name to H. androsaemum (Grigson 1987).
A list of 16 alternative local English common names from around Britain is supplied by Grigson (1987).These include variants of 'Tutsan', such as 'Tipsen', 'Tipsy', 'tit*um', 'Titzen', 'Touch-and-heal','Touch Leaf', 'Touchen Leaf' and 'Treacle Leaf'. Other names such as 'Amber', 'Sweet Amber' and 'SweetLeaf' probably refer to the fact that heavily crushed leaves emit a slight perfume from the containedoil glands, which is like ambergris. The odour, which is sharp, aromatic and rather reminiscent ofPhlox flowers in the sun, is retained and perhaps even enhanced when the leaves are dried. Driedleaves were placed inside books (including the Bible) for luck, and gave rise to names such as 'BibleFlower', 'Bible Leaf' and 'Book Leaf' (Grigson 1987).
Threats
None.
Introduction, neophyte, a very rare garden escape or discard.
28 July 2000; Northridge, R.H.; roadside W of Templerushin
Church, near Rushin Point, Upper Lough Macnean.
This semi-evergreen bushy garden shrub is a partially sterile hybrid, which is similar to H.androsaemum (Tutsan) in size and general appearance. There are up to six cultivars, some popularexamples like 'Summergold' and 'Elstead' in widespread garden use largely as decorative groundcover. Theplants carry 3 cm diameter yellow flowers and some forms have interesting leaf variegation, or berrycolour, or both.
Fermanagh occurrence
The first find of this hybrid was not made in Fermanagh until 2000 when RHN noted a large clump by theroadside near Rushin Point, on Upper Lough Macnean in the SW of the county. This was quickly followed bythree further records scattered across the shores of two other larger lakes in the VC, ie at twostations on Lough Melvin and one on the N shore of Lower Lough Erne. Details additional to the firstrecord above are: Gublusk Bay, Lower Lough Erne, 12 October 2002, I. McNeill; Rosskit 'Island' (actuallya peninsula), Lough Melvin, 14 June 2003, RHN & HJN; and Bilberry Island, Lough Melvin, 15 June2003, RHN & HJN.
Irish occurrence
Elsewhere in Ireland, this hybrid or its shrub group has been rarely recorded in a total of ten of the 40VCs, including Cos Tyrone, Down and Londonderry in NI (H36, H38 and H40). However, the Down record datesfrom 1860, and the recent records in Cos Tyrone and Londonderry were made by one field worker alsoactive in Fermanagh, namely Ian McNeill (Reynolds 2002). This strongly suggests that this taxon isregularly being over-looked.
Native, rare or very occasional, probably under-recorded. Eurosiberian southern-temperate, but widelynaturalised in both hemispheres.
1882; Stewart, S.A.; Carrick Td.
June to September.
Growth form and preferred habitats
This rhizomatous, much-branched, erect herbaceous perennial can reach 75 cm in height, although habitatfactors often restrict its growth and typical plants rarely achieve more than 30 cm. Some taprootspenetrate deeply in suitable soil, while other roots remain superficial. It overwinters as anunderground rhizome. It is only very occasionally recorded in Fermanagh and in other parts of NWIreland. Typically it is a herb of rather shallow, dry or well-drained, infertile, open or moderatelydisturbed, rocky calcareous grassland, wood margins, lakeshores and artificial habitats includingquarries. It prefers warm ground and therefore tends to avoid shade and wet or strongly acidic soils(lower than about pH 5.0) (Grime et al. 1988). These particular environmental requirementsnaturally restrict the occurrence of this species to some extent in the prevailing climate of W & NWIreland.
The presence of many translucent dots scattered over the leaf (best seen when the leaves are held upagainst the light) gives H. perforatum both its scientific name and its most frequent Englishcommon name. The glandular dots are oil sacs which extend almost the entire depth of the leaf blade fromupper to lower epidermis. When bruised the leaves release an aromatic scent from the oil glands. Therosin-like smell and the stickiness of the oil on the many-flowered, branched inflorescence, gives thespecies an alternative name in Yorkshire and the USA, 'Rosin Rose' (Salisbury 1964, p. 209).
Variation and apomictic reproduction
H. perforatum is a very variable species in terms of its physical form, size, vigour and degreeof 'weedy' behaviour. A number of varieties, subspecies or separate species have been recognised byEuropean taxonomists (N.K.B. Robson, in: Tutin et al. 1968). The subdivision appears to be basedmainly on leaf size and shape.
To a large extent, H. perforatum does not set seed in the normal sexual manner, ie meiosis(reduction division) to form haploid male and female gametes which fuse to form a diploid zygote embryothat develops into the next generation. Pollen formation is straightforward, the sex cell undergoingnormal meiosis (reduction division), but the resultant pollen can sometimes be up to 30% sterile. In theformation of the female ovules, 97% of the time meiosis fails to occur and the embryo-sac chromosomecomplement remains unreduced. The ovaries containing the unreduced ovules require pollination, but notfertilisation, in order to set seed. In this way, the normal reproductive cycle is short-circuited.
The fact that H. perforatum reproduction is not entirely apomictic means we should describe it a'facultative apomict' species (Crompton et al. 1988). The remaining 3% of ovules set seednormally after meiosis, pollination and fertilisation. Despite these reproductive abnormalities, flowersappear to form seed equally well either by cross-pollination or by selfing. This unusual pattern ofsexual reproduction strongly suggests that H. perforatum has originated as the result ofhybridisation, and one of the parent species very probably is the closely related H. maculatum subsp. maculatum Crantz (Imperforate St John's-wort) (Robson1990).
In their critical Flora, Sell & Murrell (2018) regard H. perforatum as a species aggregate andthey subdivide it into as many as four species. The three additional species are named: H.lineolatum Jord. (Narrow-leaved St John's-wort), H. densifolium P.D. Sell (Dense-leavedSt John's-wort) and H. microphyllum Jord. (Small-leaved St John's-wort). All four of these formsor species are apomictic.
Flowering reproduction
The yellow flowers, around 2 cm in diameter, are borne in many-flowered branched cymes from Juneto September. The flower contains many stamens in three bunches or fascicles and the superior ovary istopped by three long styles that spread outwards between the stamens. Although the rather showy flowersdo not contain any nectar, they attract many kinds of insect visitors that collect the abundant pollen.As the flower ages, the pollen-covered anthers bend inwards and make contact with the stigmas to achieveself-pollination, although as mentioned above, almost all reproduction in this species is apomictic andagamospermous, so that seed set is automatic after pollination, without any actual fertilisation takingplace (Proctor et al. 1996). However, since pollination is a necessary trigger for seeddevelopment in agamospermous species, this phenomenon is not easily detected. It is thus possible thatagamospermy may occur in many other species, and be much more widespread than we know about at present.
All the seeds produced by agamospermy will usually be genetically identical to the parent species (apartfrom rare mutations), so they are effectively a clone formed and dispersed by seeds (Proctor etal. 1996).
Seed production, dispersal and longevity
An average-sized plant produces about 360 seed capsules and has a seed output of c. 30,000 per year. Theseeds are the heaviest of any British terrestrial species, weighing 0.0001 gm (Salisbury 1942). The seedcapsule ripens and splits but there is no specialised seed dispersal mechanism. It is imagined that theseed, which has a somewhat gelatinous seed-coat, is merely blown by wind, carried on animal coats orflushed along by rainwater. The species can sometimes be found growing on walls at some height (Ridley1930, p. 29), and in areas like California where it has become a major, widespread weed, the initialinfestations appeared to often reflect animal movement (Crompton et al. 1988). Although some seedis only transient, other samples can survive and germinate after more than 45 years burial in soil(Thompson et al. 1997). Seed germinates readily in the spring after production.
Vegetative reproduction
H. perforatum plants also reproduce very freely by vegetative means. Even the finest of the moresuperficial horizontal roots can produce adventitious shoots. Animal browsing or any other check toaerial shoot growth stimulates profuse propagation by this means. This allows the species to extend inrelatively dense vegetation, whereas seedlings are restricted to more open, better illuminatedconditions (Salisbury 1942).
Weediness
In Britain & Ireland, H. perforatum essentially is a poorly competitive, but stress-tolerantspecies, rather than a successful weedy pioneer coloniser of disturbed, open habitats. Grime etal (1988) described it as being intermediate between two of their categories:competitive-ruderal and their so-called intermediate C-S-R strategist. In western N America and inAustralia, however, a taller and more vigorous form of the species has been introduced, or hasdeveloped, and there it has become a serious economic weed of rangelands, poorly managed pastures,roadsides, waste places, forest clearings and other non-arable sites (Crompton et al. 1988).
Toxicity
All members of the genus Hypericum contain a poisonous red, fluorescent glycoside pigment calledhypericin. It is associated with the visible black glands that occur on the petals, leaves and stems.Hypericin is a polyphenolic compound structurally similar to fa*gopyrin, the photosensitising agent foundin buckwheat (fa*gopyrum esculentum). Hypericin has been found to have antiviral andantidepressant properties. Animals eating plants containing hypericin may develop lesions on unpigmentedskin exposed to bright sunlight (ie photosensitisation occurs). This is most likely to occur on hairlessareas, such as eyelids, muzzles and udders. The skin reaction can take one to two weeks to develop insheep, cows, pigs and horses, and established lesions are slow to heal, perhaps taking months.
Lesions cause the animals to rub and scratch and open lesions can become scabbed and infected. Othersymptoms reported in horses include loss of appetite, diarrhoea, debility, staggering and even coma.Post-mortem examination of affected rabbits showed liver and kidney damage had occurred. Animals canrecover, but once affected by photosensitisation, the condition more quickly reappears after furtherconsumption of the pigment (Cooper & Johnson 1998).
Other toxins present in Hypericum include a polyphenolic flavonoid, called hyperoside, plustannins and carotenes. Given adequate forage, most livestock avoid H. perforatum, althoughsignificant grazing may occur when plants are young and succulent. Mild poisoning reduces livestockperformance, but the major significance of H. perforatum on N American rangelands, where it is anaggressive introduced alien weed, is that it reduces the carrying capacity of the land, rather thanbeing a dangerously poisonous plant (Crompton et al. 1988).
Fossil record
There is only a very meagre fossil record of three separate stages for H. perforatum listed byGodwin (1975); one (rather tentative) seed record from the Late Bronze Age, one from the Roman era andone from the much earlier than the present (Ipswichian) interglacial.
Fermanagh occurrence
There are records of H. perforatum in the Fermanagh Flora Database from a total of 13 tetrads(2.5%), nine of which have post-1975 records. As the distribution map indicates, it is very thinlyscattered around Lough Erne and on the western limestones.
The comparative rarity of recent H. perforatum records in Fermanagh suggests that it is overlookedto some extent or mistakenly identified as another Hypericum species. Five Fermanagh records arepre-1955 and the only records made by Meikle and his co-workers are from two quarries: Silverhill Quarryon the outskirts of Enniskillen (recorded 1953) and E.N. Carrothers's record of the same year fromClonmackan Quarry in the far SE of the county, near Clones town.
H. perforatum is distinguished from H. tetrapterum (Square-stalked St John's-wort) (thespecies in Ireland most like it) by its cylindrical as opposed to four-angled stem, which does, however,have two raised lines or ridges on opposite sides. In addition, Perforate St John's-wort has blackstreaks on some of the petals and leaves with intramarginal black or dark glands beneath. Thesefeatures, plus variable quantities of the translucent leaf oil glands, separate H. perforatumfrom all Britain & Ireland species except H. humifusum and forms of the rare H.linariifolium (Toadflax-leaved St John's-wort), which anyway does not occur in Ireland (Robson1990).
A considerable number of the very much more numerous H. tetrapterum records in Fermanagh are fromrather drier, shaded, more ruderal habitats than might normally be the case for this species in otherparts of these islands, eg on cliffs, in quarries and on roadsides. Some confusion may therefore existbetween these two species and conceivably this might also involve H. maculatum (Imperforate StJohn's-wort), which is equally seldom found in Fermanagh (see below).
Irish and British occurrence
In Ireland, apart from the western limestones of the Burren, Co Clare (H9), E Connemara (H16 or H17?) and the Ben Bulbin mountains in Co Sligo (H28) and CoLeitrim (H29), H. perforatum has a quite definite eastern predominance, reflecting lower rainfalland less acidic soils. In Britain, it is much more widespread in lowland southern England and Wales thanin Scotland, where it becomes rare N of the Solway Firth (Preston et al. 2002).
European and world occurrence
Abroad, a larger and very much more aggressively invasive form of this species is a major pasture weed intemperate areas of N America, Australia and New Zealand. Mowing and over-grazing reduced the very large,facultative apomictic seed production of the species, but promoted vegetative reproduction of freshshoots from shallow, spreading roots and the rhizome. Rather successful biological control, however, hasbeen achieved using chrysomelid beetles (Burdon & Marshall 1981; Crompton et al. 1988).
Uses
In herbal medicine, St John's Wort was used to treat all forms of pulmonary complaints,bladder troubles, dysentery, diarrhoea, worms, hysteria, nervous depression, bleeding and jaundice. Inchildren, a drink of tea made with the leaves was used before bed to treat or prevent bed-wetting(Grieve 1931).
The main use of St John's Wort was as a white magic token, active against the powers of evil, and linkedto St John the Baptist through his annual Saint's Day in the Christian calendar (24 June), when theplant was generally in flower. There are a number of folk traditions associated with the virtues of theplant, which involved lighting fires and smoking the leaves to further purify and strengthen theirmagical and medicinal powers. They were used to protect stables, cow-stalls, horses, animals or menagainst elves, devils, witchcraft, sickness and all evils. The plant is sometimes called an 'elf-chaser'or a 'devil-chaser', eg in France (chasse-diable). Undoubtedly some of this belief is a hangover frompre-Christian pagan times when similar ceremonies were invoked to protect the harvest, farm animals andpeople from storms, fires and devils.
The perforations observed in the leaves (the glandular dots), also became a 'signature' of wounds,reinforcing the signature of the red juice squeezed from the fresh stems and leaves, which is likened tothe blood of St John at his beheading (Grigson 1987).
Names
There appears to be some confusion as to the origin and derivation of the genus name 'Hypericum'. TheClassical Greek form of the name is 'hypereikon', and it was used by Dioscorides (a medic in the Romanarmy) for an unknown medicinal herb. Differing spellings and pronunciations occur: eg Gilbert-Carter(1964) says that in England the 'i' is wrongly treated as short. Some derive the name from the Greek'hyper', meaning 'over or above', and 'ereike', 'a heath', possibly referring to the natural habitat ofsome species in the genus (Johnson & Smith 1946).
Another suggestion is that the second element of the name derives from 'eikon', meaning 'a picture', andthus 'Hypericum' means 'over (or above) a picture', which might refer to the belief in the magicproperties of the plant, which was supposed to have power to dispel evil spirits, and is why the Devilpierced the leaves with a needle, ie a reference to the translucent glands which when the leaf is viewedagainst the light, make it look pierced with multiple pinholes. The flowers of some Hypericumspecies were therefore placed above religious images or shrines, to ward off evil at the ancientmidsummer festival of Walpurgisnacht, which later became the feast of St John (24 June), when severalspecies of the genus are in flower, and hence the origin of the name 'St John's-wort' (Stearn 1972;Gledhill 1985).
The Latin specific epithet 'perforatum' means 'pierced', or 'apparently pierced with small round holes'(Gledhill 1985), an obvious reference to the transparent oil glands in the leaves.
Seven English common names are listed by Grigson (1987), including two from North America where thespecies is a widespread and significant weed; 'Amber' and, as mentioned above, 'Rosin Rose'. The name'Amber' is said to derive from the scent given off by the dried leaves, which is reminiscent ofambergris (Grigson 1987). Two English names refer to the leaves being used to dress and heal wounds,'Balm of the Warrior's Wound' and 'Touch and Heal' (Grigson 1987). In Wales, the plant was called'Mary's Ladder' or 'Christ's Ladder', possibly a reference to the opposite leaves and its devil chasingproperties. Similar traditions and names occur in an Irish context, with the Virgin Mary, St Colum Cilleand St Columba linked to the plant. It was sometimes worn under the left armpit to ward off evil spiritsand death (Grigson 1987).
Threats
None.
Native, apparently very rare, probably somewhat overlooked. European boreo-temperate.
1900; Praeger, R.Ll.; Belcoo, Lough Macnean.
Identification difficulties and variation
In an article dealing purely with the genus Hypericum, Robson (1990) commented on theidentification of this species, writing, "It is safe to say that the variation in H.maculatum and H. perforatum (Square-stemmed St John's-wort) has caused more problems forthe British botanist than that of any other British or Irish species." Considering the context inwhich it was written, the current author believes that Robson made this statement with respect to anyother British or Irish Hypericum species, rather than from a more general botanical perspective.
The reasons for Robson making this claim involve the pattern of variation created by hybridisationbetween the two species (which are interfertile and backcross frequently) and by species breedingabnormalities that 97% of the time involve agamospermy (or pseudogamy) (ie seed formation requiringpollination but without fertilisation of unreduced ovules) (Robson 1990). Together these features haveresulted in an almost complete intergrading of these two St John's-wort species, which could make onewonder why these apomictic forms are regarded as separate at all. Nevertheless, taxonomists do describethem as if they were separate species and field botanists try to recognise them as best they can (Sell& Murrell 2018).
Growth form and preferred habitats: From the list of records in the Fermanagh Flora Database,H. maculatum is apparently a very rare species in the VC, having been recorded in only fivetetrads, mainly in the south of the county. However, on the evidence of our Irish field recording,Robert Northridge and the current author believe it quite possible that in Fermanagh, and probablyelsewhere in B & I, H. maculatum is also sometimes being mistaken for H. tetrapterum(Square-stalked St John's-wort), which in any event we suspect is locally over-recorded. Both these StJohn's-wort taxa possess a square, four-angled stem. However, the angles on the H. maculatum stemare not as prominent as those on H. tetrapterum, being unwinged. H. maculatum has longerpetals − which are at least twice as long as its sepals. H. maculatum also differs from H.perforatum in having distinctive, narrow, parallel-sided leaves which have few translucentglands or none (Webb et al. 1996).
Growing up to 60 cm tall, H. maculatum is a shortly rhizomatous perennial of damp grassland onbanks, in open woods and on roadsides − similar habitats in fact to H. perforatum (Perforate StJohn's-wort), although the latter tends towards more calcareous situations than H. maculatum everdoes.
In B & I, H. maculatum subsp. obtusiusculum is known to cropup in more ruderal habitats than the more restricted damp ground situations to which the diploid subsp.maculatum adheres, the former being additionally encountered in rough grassland, scrub, quarries,wasteground and wayside banks (N.K.B. Robson, in: Preston et al. 2002). These more ruderalsituations are just where we have on record a considerable number of Fermanagh finds for H.tetrapterum, which we believe may well prove to be errors.
Fermanagh occurrence
In Fermanagh, H. maculatum is very rare and has only been recorded in five tetrads, mainly in thesouth of the county. There is just one post-1947 record and thus Robert Northridge and the currentauthor believe it is being overlooked and under-recorded. Apart from the first find listed above, thedetails of the other records are: Crom Castle Estate, 1939, Praeger; Inver Lough, Tattygormican Td,1947, MCM & D; beside Old Ulster Canal, near Wattle Bridge, 1947, MCM & D; Colebrooke River,Killarbran Bridge area, 19 May 1990, I. McNeill. The habitats involved range from damp wood margins onlakeshores and riverbanks, to wayside hedge banks and ditches.
British and Irish occurrence
Throughout B & I, this species is unevenly but rather widely distributed, being most frequent inBritain in the W Midlands and C Wales (New Atlas). Two subspecies of different ploidy level (iechromosome number), are recognised in Britain, but only the auto-tetraploid, subsp. obtusiusculum(Tourlet) Hayek has ever been recorded in Ireland (Robson 1958b, 1990). The New Atlas hectad mapshows that this plant appears to be quite widespread in the southern half of Ireland, but in the northit is infrequent and much more scattered, except in ground on and around the shores of Lough Neagh.While H. maculatum subsp. obtusiusculum appears from the hectadmap particularly widespread in Co Armagh (H37), the Flora of the NE of Ireland, 3rd ed. accountcovering three of the other VCs around the lake (Cos Down, Antrim and Londonderry (H38-H40)), describesthe plant as, "rather rare and local … mainly about Lough Neagh and along the Lagan Valley, UpperBann Valley and the Newry Canal to Carlingford Lough" (ie including ground downstream, towards thecoast in both the N & SE of the lake) (Hackney et al. 1992). Taking account of this, thesubspecies is not so rarely found in N Ireland that one would expect just one post-1975 record of it inFermanagh.
Robert Northridge and the current author think it very possible that in Fermanagh H. maculatum issometimes being mistaken for H. tetrapterum (Square-stalked St John's-wort), which in any eventwe suspect is locally over-recorded. Further careful examination of this genus, and of H.tetrapterum in particular, is needed in our area.
Threats
None.
Native, common and widespread throughout the lowlands. European temperate, introduced in New Zealand.
1881-2; Barrington, R.M.; Devenish Island, Lower Lough Erne.
Throughout the year.
Growth form and preferred habitats
This is a hairless herb with stiffly erect, sharply defined square stems, the stem-lines being strong,forming definite wings. Stems are normally held erect from a decumbent base which produces slendercreeping stolons, although they can sometimes be entirely procumbent (Sell & Murrell 2018). H.tetrapterum can grow up to 100 cm tall, but more usually it is smaller, between 10 and 60 cmtall. Leaves are oval (ovate to orbicular) and possess numerous small translucent glands and a few blackones. The pale yellow flowers are numerous, borne in ± flat-topped cymes and are 9-13 mm in diameter.The sepals are very acute and the petals slightly longer than the stamens.
Throughout lowland B & I, this is a widespread and occasional to frequent or locally common plant,occurring in a wide range of damp to wet, reasonably fertile habitats. These include the sides ofditches, streams and other damp to wet eutrophic situations, in woods, marshes, lakeshore meadows andpastures.
Fermanagh occurrence
In Fermanagh, it is recorded in 265 tetrads, 50.2% of those in the VC. Although widespread and commonlyfound here, it is never abundant and more often than not it is only sparse, forming small, isolatedpatches. In addition to the typical damp ground habitats, a substantial proportion of the Fermanaghrecords of H. tetrapterum (approximately 50 of them, representing 11% of the species total) arefrom drier, more ruderal, wayside and waste ground habitats, including some from quarries, roadsides,cliffs, rocky ground and grassland in districts of predominantly shallow limestone soils.
Locally, H. tetrapterum is absent only from the most acid or regularly waterlogged soils. Thereare even a few records of it from acidic bogland. Several of these habitats lie beyond the normal rangeexpected for this species elsewhere in these islands. They are not readily explicable, except perhaps aserrors for other similar Hypericum species. In the absence of vouchers we cannot check theidentification of records from the exceptional habitats, but we feel that with just five records, H.maculatum subsp. obtusiusculum (Imperforate St John's-wort)appears decidedly under-recorded in Fermanagh (see the species account above).
Even if we discard around 50 of the Fermanagh H. tetrapterum records as possible errors, it stillremains a very frequent St John's-wort, second only to H. pulchrum (Slender St John's-wort) interms of frequency and distribution in the VC. This is a situation we couldconfidently expect, since there is no shortage of suitably damp ditches, stream-sides and marshygrasslands for it to colonise.
Reproduction
H. tetrapterum reproduces both vegetatively by stolons and sexually by insect pollinated flowers.The resultant fruit capsules of the sexual process release huge numbers of tiny, lightweight seed on thebreeze in autumn. Seed production is vast. Salisbury (1942) estimated a mean seed production of between23 and 32 thousand seeds per plant! Furthermore, estimates of seed longevity determined in NW Europerange from ephemeral to long-term persistent, ie from less than twelve months, to at least five years(Thompson et al. 1997).
Salisbury (1942, p. 108) concluded after reviewing the reproductive processes of ten members of thegenus, "The species of Hypericum ... support the view that reproductive capacity is apositive asset to the species, and that for species having no special means of vegetative spread thehigher the potential reproductive capacity, the commoner the species tends to be."
British and Irish occurrence
H. tetrapterum is widespread in B & I, but it becomes rarer or absent in northern Scotland,probably due to the mountainous ground and the huge extent of very wet, nutrient poor peatland. It isabsent altogether (as a native) north of Ross & Cromarty (ie all VCs from 88-112, apart from 94& 95). A comparative glance at the New Atlas maps of H. tetrapterum, H.pulchrum and H. perforatum (Perforate St John's-wort) indicates how very well distributedthese three St John's-wort species are in these islands. H. tetrapterum is second only to H.pulchrum in respect of dispersal (measured in number of map hectads with symbols: H.tetrapterum 2959; H. pulchrum 3323), although H. perforatum, possessing a weedyform and having the most marked ability of vegetative propagation of these three species, is by far themost widespread Hypericum worldwide.
European and world occurrence
H. tetrapterum is a member of the European temperate element and is restricted to W, C and SEurope and adjacent parts of Africa and Asia. Like so many members of the genus, it has been introducedto New Zealand, where it occurs on both islands (Hultén & Fries 1986, Map 1315; Webb et al.1988; Sell & Murrell 2018).
Threats
None.
Native, rare or very occasional. European temperate, introduced in S Africa and New Zealand.
1884; Barrington, R.M.; Gubbaroe Point, Lower Lough Erne.
April to November.
Growth form and preferred habitats
This short-lived perennial produces prostrate, non-rooting stems that are quite distinctive. However,trailing specimens of H. pulchrum (Slender St John's-wort) should not be mistaken for thisspecies. The best distinguishing feature between these two species is the unequal sepals of H.humifusum, two of which are shorter and narrower than the other three.
The most frequent substrate on which Trailing St John's-wort grows is bare sandy or gravelly ground. Thisis quite a good starting point for making a correct identification since H. humifusum isgenerally recognised as a pioneer of open or disturbed, light, dry or well-drained, slightly to somewhatacidic soils.
Thus it occupies a wide range of open habitats in B & I in situations where competition is virtuallyabsent. Suitable ground generally occurs in short turf with open, bare areas on moorland, grasslands,dunes, tracks, dry wayside banks, waste ground, sand- and gravel-pits, plus on the margins of woods orin clearings. Some English local Floras indicate it can also sometimes be found growing on walls and inlawns.
Fermanagh occurrence
H. humifusum is rare in Fermanagh and occurs sparingly, either as single individuals, or just afew plants at most of its 14 Fermanagh sites. It is thinly and widely scattered across 13 tetrads in theVC and only nine tetrads contain post-1975 records.
The Fermanagh soils it occupies provide dry, bare, sandy or well-drained peaty conditions and they occuron lakeshores, riverbanks, hedgerows, waste ground and in an old sand-pit near Pubble Bridge. These fitthe typical habitat range of this species in B & I quite well, except with respect to moors, woods,lawns and walls. In the case of the walls, we might substitute the cliffs at Carrickbeg, where it wasrecorded by Meikle and co-workers in 1947. However, on reflection, these cliffs are wooded at theirbase!
Apart from the first record above, the other Fermanagh station details are:
"Several localities about Boho", 1946, MCM & D; Carrickbeg Cliffs, E of Horse Rock, NW ofBoho, 1947, MCM & D; Drumderg Lough, SW of Teemore, 1950, MCM & D; Bank near Church with tower,1.5 km N of Boho, 1950, MCM & D; Pubble Bridge, Tempo River, 1 October 1988, RHN;Cradien/Ballydoolagh Tds, 3 km NE of Enniskilllen, 16 June 1989, RHN; Scarford Bridge, Colebrooke River,2 November 1989, RHN; Near children’s playground, Castle Archdale, 2 November 1990, RHN; Drumcreen,banks of Ballinamallard River, 16 April 1995, RHN & HJN; Coolbuck Td, near Lough Eyes, 2.5 km NE ofLisbellaw, 19 October 1997, RHN; Sand pit near Pubble Bridge, Tempo River, 20 August 1999, RSF &RHN; Dry bare shale bank above Sillees River near farm, E of Melly’s Rock, 14 July 2001, RHN; and LargyHill, 1 km S of Lack, 3 October 2002, I. McNeill.
Ecology
The typical lifespan of perennial H. humifusum in the temporary growing conditions associated withits pioneer status falls between two and four years. Its survival potential is enhanced by an adaptationenabling it to flower and fruit in its first year of growth and additionally the unopened flower budscan self-pollinate if weather conditions are unfavourable for normal open pollination by insect visitorswhen the flowers are ready to function (ie it displays facultative cleistogamy). However, the species isvery variable and seed germinating in early autumn can produce dwarf semi-erect individuals, which maybehave either as very ephemeral annuals, or as biennials if they manage to overwinter (Robson 1990).
British and Irish occurrence
H. humifusum is widespread but always local and more often scarce than abundant throughout most ofB & I, but is most frequent in the S & W and is absent from the extreme N of Scotland and fromthe N & W isles. There appear to be considerable losses in parts of B & I, which began around1950 and can probably therefore be attributed to changes in land-use including the intensification offarming. In parts of W Ireland, some apparent losses are more likely due to insufficient recording(N.K.B. Robson, in: Preston et al. 2002).
European occurrence
The wider distribution of H. humifusum is mainly Central European, but it extends into W Europeand NW Africa in Morocco, Madeira, the Canaries and the Azores (Robson 1990).
Threats
The closing of the sandpit at Pubble with subsequent loss of bare ground, plus the general tidying-up ofthe countryside, which could easily eliminate the few remaining stations.
Native, very common and widespread. Sub-oceanic temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form, identification and preferred habitats
This slender, erect, hairless perennial with smooth stems can grow up to 60 cm tall, but very often it ismuch smaller (c 15-25 cm). Leaves are cordate or truncate at the base and petals and sepals are fringedwith shortly stalked black glands and the backs of petals are sometimes red-tinged (Parnell & Curtis2012). Young or sterile plants are trickier to identify. In H. pulchrum, young plants (andsometimes mature ones also) have decumbent to prostrate stems with oblong leaves, making them rathersimilar to H. humifusum (Trailing St John's-wort). However, the leaves are more leathery than thelatter and they also lack the intramarginal black glands of H. humifusum, which should enableH. pulchrum to be distinguished. The absence of transparent pellucid glands from the basal partof the leaf nearest the stem is another helpful characteristic of H. pulchrum (Robson 1990).
In general, H. pulchrum occurs in habitats where a variety of stressful growing conditionsseverely limit biological competition from taller, more vigorous and more aggressive plant species. Ittolerates a wide range of dry, shallow, stony, infertile and often leached, sandy or peaty soils in along list of open, or semi-shaded habitats at almost all altitudes. Locally these include the marginsand open areas of woods and scrub, hedgerows, dry, rough grazing grassy heaths or moors, coarse grassybanks, roadsides, quarries and gravel-pits.
Fermanagh occurrence
H. pulchrum is the most common and widespread Hypericum species in Fermanagh by aconsiderable margin. It has been commonly recorded in 310 tetrads, 58.7% of those in the VC. Bycomparison its nearest rival, H. tetrapterum (Square-stalked St John's-wort), has two thirds therecord frequency of Slender St John's-wort and is distributed across 50.2% of the tetrads.
As the tetrad distribution map indicates, H. pulchrum occurs widely across the VC, but the dearthof records in lowland tetrads adjacent to the eastern shores of Lough Erne is rather surprising.
A possible limestone ecotype
While H. pulchrum more commonly grows on acid to neutral soils and elsewhere in B & I is oftenreckoned a definite calcifuge species (Robson 1990; Crawley 2005), in Fermanagh and other parts ofwestern Ireland, H. pulchrum occurs in some profusion on often quite bare limestone habitats.These include stabilised screes, rock ledges on cliffs and knolls and both on the surface and in thedeep crevices typical of limestone pavement.
On noticing this behaviour on the limestones in the Burren, Co Clare and in Connemara, Webb &Scannell (1983) (Flora of Connemara and the Burren) remarked on the possibility that a calcicoleedaphic ecotype may have differentiated, it becoming more tolerant of alkaline, base-rich soils than thenormal form of the species. If this is the case, then both ecotypes are present in Fermanagh, sinceH. pulchrum appears common on both acid soils and on alkaline, lime-rich ones. It thus appearsindifferent to soil base-status, a situation also described in the Shropshire region of England bySinker et al. (1985).
Other variation
H. pulchrum varies relatively little, but a dwarf, few-flowered, prostrate to procumbentform (f. procumbens (Rostrup) Beeby) is found rarely in exposed habitats at the NW extreme of thespecies range in the Faeröes, the Shetlands, Caithness, W. Ross and the Outer Hebrides in Scotland andon Clare Island and Achill Island off the W coast of Ireland (Robson 1990). This variant is said tobreed true, but a complete series of intermediates links it with the typical plant (N.K.B. Robson, in:Tutin et al. 1968).
Reproduction
In common with H. perforatum (Perforate St John's-wort), H. pulchrum can increase itspresence on a very local basis through vegetative reproduction by means of adventitious shoot budsproduced on its most superficial spreading lateral roots. The mean annual seed production of H.pulchrum is approximately 4 to 5 thousand, considerably less than the other two most widespreadSt John's-wort species in B & I, H. tetrapterum and H. perforatum (Salisbury 1942). Incommon with these latter species, however, buried seed of H. pulchrum survives for at least fiveyears in soil (Thompson et al. 1997).
British and Irish occurrence
Although it is often regarded as a 'locally common' species in most of B & I, on the evidenceprovided by the hectad maps in the New Atlas, H. pulchrum is the most widespreadHypericum species by quite a wide margin. The species map has record symbols in 3323 hectadsthroughout these islands, representing 85% of the total area. This is 364 hectads more than the nearestrival in the genus, H. tetrapterum (Preston et al. 2002). Having said this, H.pulchrum is absent or rare in the low-lying English Wash and in the limestone belt that lies tothe SW of it. In Ireland, it is similarly absent from around the Dublin conurbation (H21) and from thefar W of Co Mayo and W Galway (H16 & H27) where completely unsuitable wet, peaty soil conditions arevery common.
European occurrence
H. pulchrum is a suboceanic temperate species confined to NW Europe. It occurs in theFaeröes, coastal S Scandinavia and the region west and north of a line through Germany, Switzerland,France, Spain and C Portugal. There are also outlying stations further east from SW Poland to NW Italy(Hultén & Fries 1986, Map 1312; Robson 1990; Pignatti 1997, 1; Sell & Murrell 2018).
Introduced, deliberately planted, occasional, but very definitely deliberately ignored andunder-recorded.
10 July 1986; Austin, L.W. & McMullin, A.S.; Upper Lough Erne shore near Knockninny Hill.
May to November.
The New Flora of the BI (1997) notes that this large deciduous hybrid is one of the commonestplanted trees in Britain. The British tree expert Alan Mitchell (1996) describes it as, "anunfortunate and ill-favoured tree", since although it is a natural hybrid between two species ofconsiderable arboricultural distinction, it has inherited only the bad features from each parent. Themost obvious and very common defect is the presence of vigorous basal suckers and epicormic shootsemerging from burrs higher up the trunk. These disfiguring outgrowths are inherited from T.cordata (Small-leaved Lime), but on that tree species, they generally appear only on veryold specimens.
Common Lime is thinly and widely scattered in Fermanagh, mainly on large estates and in the grounds oflarger, older houses around both parts of Lough Erne, but it occasionally appears planted in roadsidehedgerows, along river banks and on or near lake shorelines. It is very seriously under-recorded asthere are only records of it from 16 widely scattered tetrads in the Fermanagh Flora Database, 3% of thesquares in the VC. The majority of recorders working in Fermanagh entirely ignore planted aliens.Neither Praeger nor Meikle and his co-workers recorded trees much at all and we ourselves have notbothered greatly keeping records of obviously planted specimens. Thus the real distribution of thispartially fertile hybrid tree and indeed all other alien tree species in the county, needs furtherconcerted investigation.
Half of the Tilia records were originally listed as T. cordata largely or entirely on thebasis of lower leaf and bud colour characters. Pigott (2003), however, points out that these are notsatisfactory features for distinguishing mature trees. Reliable identification requires material fromthe exposed (generally upper) leaf canopy, preferably also with supporting flowers or fruit. Since oldertrees are often very large, obtaining such specimens is difficult or totally impossible. In view of thisand of our experience of lime trees generally, we feel that none of the Fermanagh tree records is likelyto be true T. cordata, but rather they very probably all belong to one of the two widespreadclones of this hybrid. We have therefore judged it sensible to consolidate all the records into thistaxon.
Common Lime has a rather irregular crown when compared to either of the parent species and this, and theextensive suckering and burring, is often sufficient to identify the hybrid and possibly the individualcultivated clone of it, even from a distance and in silhouette (see the excellent comparative figures inPigott (1992)). Common Lime is widely planted across lowland areas of both B & I, although again,comparatively speaking, it is nothing like as frequent in the latter island (Preston et al.2002).
Threats
None.
Introduced, neophyte, a rare garden escape or 'wild-flower' seed plantation; probably only casual.European temperate, widely naturalised in N Europe, N America, Tasmania and New Zealand.
1950; MCM & D; Cornaleck, Upper Lough Erne.
July to November.
Growth form and preferred habitats
Musk-mallow is a medium tall (30-80 cm), herbaceous perennial bearing several erect, often purple-spottedstems from a thick, branching rootstock. The alternate leaves are rounded or kidney-shaped in outline,long stalked, 3-7-lobed and extremely variable, but often deeply cut into ± linear ultimate segments.Just below the flower calyx there are three small bracts, free for their entire length, which constitutean epicalyx, a useful identification feature for the genus.
M. moschata is a plant of dry or well-drained, sunny or lightly shaded, open or somewhat neglectedgrasslands, preferring, but not restricted to, light sandy loam soils of medium to high fertility. Itcan tolerate, or may actually require, moderate levels of disturbance, which helps to keep aggressivecompetition from taller and more vigorous species in check. Disturbance may take the form of lightgrazing or mowing on the margins of woodland and scrub, pastures, meadows, field margins, roadsideverges, hedge banks, river banks, lake shores and grassy waste ground (Sinker et al. 1985; J.H.S.Cox, in: Preston et al. 2002; Sell & Murrrell 2018).
Variation
M. moschata is very variable, especially with respect to leaves and theirdegree of dissection, degree of stem and leaf hairiness, types of hairs and shade of flower colour.Despite this, most Floras do not distinguish varieties. However, Sell & Murrell (2018) have recentlydescribed no less than four varieties, two of which occur in only a few scattered locations. Of theother two more widespread forms, var. tenuifolia Guss. is probably the native plant throughoutEngland, Wales and coastal Scotland, while var. undulata Sims was probably introduced in the last40 years, mainly as a component of wild-flower seed (Sell & Murrell 2018).
Flowering reproduction
M. moschata produces attractive, 3-6 cm diameter, rose-pink or white, musk-scented flowers inclusters at the tops of stems from June to August. Flowers attract bees and other insect pollinators,but they can also self-fertilise. As in other members of the genus, the fruit is a dry schizocarp, ie itconsists of a single ring of numerous (around 15) mericarp segments or nutlets, which split from oneanother when mature, each portion containing a single kidney-shaped seed (Stace 2010; Sell & Murrell2018). Seed can remain viable in the soil for many years (Thompson et al. 1997).
Seed dispersal
While in Fermanagh and most of Ireland, M. moschata is only a rare, casual plant in the wild, thereally interesting fact is that despite the lack of any obvious dispersal mechanism, the dry fruitmericarps which split off the disc-shaped schizocarp fruit, somehow manage to 'jump the garden wall'.The transport mechanism or vector enabling this is certainly not obvious. It is possible to imagine windand rain-wash might together move the relatively large fruit segment and its contained seed a metre orso from the parent plant, but it would require a very exposed plant and a real gale or flood to move thepropagule any greater distance right into the wild.
Considering seed dispersion in this genus, Ridley (1930, p. 361) relied mainly on cattle or otherungulates ingesting and transporting the fruit in their gut for his most probable mechanism. However,this solution would not apply in a garden setting. Ridley also mentioned work on the American Crow,Corvus brachyrhyncos, which is a major consumer of soft-fruit, yet which also eats many dryfruits and grains, including those of Malva species. Most seeds are probably destroyed bydigestion, but some might survive and appear in the bird's pellets.
Does anyone know of a bird taking an interest in Mallow fruits in B & I, thus serving as anoccasional seed vector? The only other vector the current author can suggest is human, possiblytransporting the fruit unknowingly in mud on boots, trouser bottoms, tyres, or amongst discarded gardenrubbish. I await with interest any helpful observations throwing light on the mystery of Malvadispersal.
Fermanagh occurrence
In the Revised Typescript Flora, Meikle et al. (1975) noted that Musk-mallow was notuncommon in Fermanagh gardens. Their solitary 1950 record at Cornaleck on the shore of Upper Lough Erne(not given in the 1951 Typescript Flora) was of a white-flowered garden form. Although itis a perennial, none of the plants at four of the five recent additional stations, all recorded by RHN,has persisted for long. They are also assumed to be garden escapes, although apparently M.moschata is now sometimes (regularly) associated with wild-flower seed mixtures (often ofimported origin), increasingly sown in both public amenity areas and in garden settings. We are not asyet aware of any such source of Musk-mallow in Fermanagh.
The remaining details of the five later Fermanagh stations are: roadside on the Enniskillen-Belfast roadnear Boyhill, 1 July 1997;RHN; waste ground, Enniskillen town, 22 November 1998; RHN; waste ground,Drumawill, Sligo Road, Enniskillen, 11 October 2003; RHN; near farm yard, Cam, 1.5 km W of CorrasloughPoint, Upper Lough Erne, 17 August 2004; RHN; and by wall on N side of Mill Lough near Ballanaleck, 13October 2010; RHN.
British occurrence
In England, Wales and S Scotland, M. moschata is a common and widespread perennial of lowlandgrassy places, generally regarded as native. In Scotland, north of a line between Girvan andBerwick-upon-Tweed, it is increasingly rare and is considered an escaped garden introduction (J.H.S.Cox, in: Preston et al. 2002).
Irish occurrence
In Ireland, by contrast, M. moschata is not usually a persistent plant. Nowadays it is recognisedas an alien introduction, generally of garden origin (Reynolds 2002). In the mid-west of the island,where it typically is a rare roadside occurrence, the authors of the Flora of Connemara and theBurren commented, "scarcely more than a casual; it is only in SE Ireland that this speciesis firmly established". (Webb & Scannell 1983).The longest known persistence of any Musk-mallowcolony in the wild in N Ireland is approximately 40 years, which happened along the banks of the CrumlinRiver near Lough Neagh, Co Antrim (H39) (Harron 1986; Hackney et al. 1992).
European and world occurrence
M. moschata originates from the Mediterranean region and probably parts of C Europe from Englandand Poland southwards (Tutin et al. 1968). It is widely cultivated in gardens and has spread wellbeyond its area of natural distribution to S Scandinavia, N Africa, N America, New Zealand and Tasmania(Hultén & Fries 1986, Map 1304).
Threats
None.
Introduction, archaeophyte, rare to occasional garden escape. Eurosiberian southern-temperate, but widelynaturalised in both hemispheres.
1884; Barrington, R.M.; roadside between Blaney and Poulaphouca.
June to October.
Growth form and preferred habitats
A colourful, if rather untidy, straggling, decumbent or nearly erect, 45-90(-150) cm tall, variablyhairy, introduced garden perennial or biennial, M. sylvestris has palmate stem leaves, each with5-7 deep lobes, indented about a third the way to their stalked base.
Like most mallow species in B & I, M. sylvestris tends to occupy fairly dry, even droughted,well-drained sites. It prefers moderately fertile or nutrient-enriched soils and sunny or only lightlyshaded situations on grassy waste ground, hedge banks, field margins and the like, often near houses orruins (J.H.S. Cox, in: Preston et al. 2002). It has a deep, thick, woody taproot which allows itto tolerate dry soils and drought remarkably well, considering the large amount of aerial tissue itusually supports.
Mallows are not very competitive and they all tolerate or appreciate some habitat disturbance, helping tokeep the ground open and limiting competition. Usually this involves grazing, trampling or mowing, butin this specific instance, on account of the proximity of many of our Fermanagh records to lakeshores,seasonal flooding may help create suitable conditions.
Flowering reproduction
Flowers are produced from June to September, either singly or in small clusters in the leaf axilsat the tip of the stem, forming an irregular raceme. Only one or two flowers are in bloom at any time.Individual flowers have five connivent sepals that fail to enlarge in fruit, so they never overlap. Thefive notched petals, 20-30 mm long, are 2-4 times the size of the sepals and are bright pinkish-purplewith darker stripes. The calyx is furnished with an epicalyx of three bracteoles beneath it. Pollinationis by bees and other insects searching for nectar; the fruit is a schizocarp composed of a single ringof wedge-shaped carpels around a central axis that eventually splits into separate mericarps or nutlets,each containing a single seed (Sell & Murrell 2018). Other than this, there is no special dispersalmechanism (see below).
Fruit dispersal
Although like all members of the family Malvaceae, M. sylvestris is not long-lived and it readilyreproduces, flowers and seeds freely. The fact that it is occasionally observed growing in wall creviceshigh above the ground proves that Common Mallow must have definite powers of dispersal to reach suchsites (Ridley 1930, p. 27). However, unless strong winds or birds transport the fruit nutlets, it isdifficult to imagine a reliable seed dispersal mechanism. Praeger (1913) experimentally showed thatseeds retain their viability in water for up to 36 hours, so they may also disperse along streams andaround sheltered lake shorelines by flotation (Ridley 1930, p. 201).
Variation
Although previously regarded as a single species (Stace 2010), or with a single variety with hairymericarps (var. lasiocarpa Druce) (Clapham et al. 1987), three varieties are nowrecognised by Sell & Murrell (2018). Var. sylvestris is either perennial or biennial and haserect stems and glabrous mericarps; var. socialis Griseb. is perennial and has spreading orprostrate stems, a densely hairy inflorescence and mericarps with simple eglandular hairs. Finally, var.incanescens Griseb. is an erect biennial with small leaves covered with stellate hairs beneath;it also has mericarps with simple eglandular hairs.
Fermanagh occurrence
The Revised Typescript Flora commented that Common Mallow was, "still cultivated in cottagegardens, and probably always is a recent escape elsewhere". In Fermanagh, M. sylvestris hasbeen recorded in a total of 18 tetrads (3.4%) and is occasional about Upper Lough Erne and rare andthinly spread elsewhere. Nine of the 17 post-1975 records were made on lakeshores (or possibly on nearbyaccess roads and fields), during the 1986-7 DOE survey of Upper Lough Erne. Two of the most interestingstations are the Green Lough limestone turloughs near Fardrum, where R.S. Weyl recorded it on 20 June1985, and Trien Mountain, a limestone hill above the Florencecourt estate, where A. Waterman discoveredit on 11 October 1989. Other Fermanagh sites are on roadsides and waste ground, generally nearhabitation.
Irish occurrence
M. sylvestris has long been considered a probable introduction in Ireland (Scannell & Synnott1987; Parnell & Curtis 2012), but in more recent years it has been recognised as a definite gardenintroduction (Preston et al. 2002; Stace 2010). In parts of the N & W of Ireland, CommonMallow shows a very definite coastal affinity, colonising and persisting in sandy and gravellysituations. On the shore of Co Down M. sylvestris is a regular member of the upper littoral plantcommunity (Hackney et al. 1992; NI Flora Website 2006).
British occurrence
While M. sylvestris is common and widespread in most of lowland England and Wales, theNew Atlas hectad map indicates it has a definite southern and eastern predominance. Previouslythe species was regarded as native in Britain (eg Stace 1997), but it is now recognised as an ancientgarden introduction (an archaeophyte) (Preston et al. 2002; Stace 2010). In Scotland, the speciesis much more thinly scattered, becoming confined to coastal areas, mainly in the east of the country asone travels northwards. It is absent from the Outer Hebrides, Orkney and Shetland (Preston et al.2002).
European and world occurrence
The original distribution of M. sylvestris was probably restricted to mainland Europe, N Africaand W Asia. However, it has long been cultivated as a garden subject and has been introduced to manycountries around the world, including N & S America, S Africa, E Asia, S Australia and New Zealand(Hultén & Fries 1986, Map 1302).
Threats
None.
Archaeophyte, casual, probably extinct. European temperate, widely naturalised beyond its native range inEurope and introduced in N America and Australia.
1900; West, W.; Levally House, 1 km SE of Roosky.
A habitually self-pollinating annual, Dwarf Mallow grows in the same sort of dry, sandy or gravelly,ecologically open situations which other Malva species typically frequent in B & I, either oninland waste ground or disturbed grassy roadsides, often near habitation, as well as in open, coastalsites.
It is an archaeophyte dating from Roman times in Britain and although it is fairly well established in afew places further south in Ireland, eg around Galway Bay and in the Dublin-Wicklow areas, it is amerely casual introduction in the rest of Ireland. Throughout Ireland, the majority of the widelyscattered stations are coastal, the plant often growing along the drift-line (Webb & Scannell 1983;New Atlas).
M. neglecta has not been seen in Fermanagh for over 50 years and we therefore regard it as locallyextinct. The only record in the Fermanagh Flora Database additional to the first listed above is: on arubbish tip, Enniskillen town, 1947, MCM & D.
In N Ireland, the only recent record of M. neglecta was made around 1999 by Ian McNeill, on wasteground on the coast at Portstewart, Co Londonderry (H40) (Reynolds 2002).
In Britain, Dwarf Mallow is widespread, frequent and established in S England, but elsewhere it again ismainly a casual species, decreasing northwards and becoming increasingly confined to the E coast,although the New Atlas map shows it stretches north beyond Inverness.
Native, frequent, widespread and locally common. Circumpolar boreo-temperate.
1836; Mackay, J.T.; bogland on Cuilcagh Mountain.
April to January.
Growth form and preferred habitats
A small, shallow-rooted, rosette-forming, bright red, short-lived, native perennial, D.rotundifolia has stalked leaves with orbicular laminae, 7 mm in diameter. Each leaf lamina bearsaround 200 stalked, sticky, red tentacles that attract and trap insects. The bright red colour ofDrosera leaves and stems is due to cyanidin glycoside, an anthocyanin pigment.
Round-leaved Sundew is a small plant of wet, unshaded or lightly shaded, relatively open, strongly acid,wet peatland. It is frequent and occasionally abundant at the edges of bog pools on damp to wet mosscarpets – often having to compete with Sphagnum moss; it also grows on damp, bare peat and on theseeping sides of drainage ditches cut in bogs dug for fuel turf. D. rotundifolia prefers full sunhabitats, but it can tolerate a degree of shade from Calluna vulgaris (Heather) and otherdwarf shrubs. It can survive overwinter completely shaded within Sphagnum cushions, after beingovertopped by autumn growth of the moss. Wide fluctuations in the water table, including immersion forup to two weeks are tolerated, but due to its very shallow roots, dry periods must be only of shortduration, if the plant is to survive. D. rotundifolia is replaced by D. anglica (GreatSundew) in the wettest areas of the bog, eg the bottoms of hollows and in intermittent or more permanentpools (Crowder et al. 1990).
Probably present on every patch of otherwise bare, acid peat bog and wet heath in the county, althoughsometimes it is rather scarce and needs to be actively searched out. With its tennis-racket shapedleaves this is the easiest sundew to identify and it is by far the most common of the threeDrosera species we have in B & I, around twice as widespread as D. anglica (= D.longifolia) (New Atlas).
Variation
No varieties are recognised in Britain, although three have been described from continental Europe(Crowder et al. 1990).
Insect capture, plant nutrition and performance
Insect capture by the abundant glistening sticky leaf glands is effective in stimulating plant growth,reproductive success and survival through supplying additional nitrogen and phosphorus, the usuallimiting nutrients on strongly acidic, oligotrophic bogland soils. However, plants without access toarthropods remain healthy and D. rotundifolia can take up inorganic nitrogen if it is available.
In Drosera, erect leaves have been found to be more efficient in capturing insects than horizontalleaves close to the soil surface. When an insect adheres to a leaf tentacle and struggles, thisstimulates the bending of neighbouring tentacles and the in-rolling of the leaf lamina itself, bothfurther entrapping the prey. These movements are believed to be responses to chemical rather thanmechanical stimuli and they are mediated by hormones. One to two weeks after prey has been caught, theleaf reopens and the blackened remains, mostly chitinous pieces of legs and wings, fall off or are blownaway (Crowder et al. 1990).
Observations of insect capture rate for D. rotundifolia in W Germany were higher than thosepublished for another insectivorous genus (Pinguicula (Butterworts)) and were correlated withtotal leaf number and leaf area. It was calculated that the plant obtained around 24-30% of its nitrogenfrom insect capture (Karlsson et al. 1987; Schulze & Schulze 1990). Despite the advantage oflarge plants in capturing prey, the seasonal development of plant size was not simply related to insectcapture but strongly influenced by leaf losses, flower-bud formation, flowering and seed set. All ofthese processes cost the plant photosynthetic and nutritive resources that take time to replenish beforerosette regrowth and size recovery can take place (Schulze & Schulze 1990).
Individual plants are smaller in exposed and upland sites and there must be a lower size limit at whichinsect capture rates are inadequate and plants become doomed through starvation. Size of plant isgenerally related to the local abundance of prey and plants on most Fermanagh bogs are small, rarelyover 6.0 cm tall, often with only 4-8 functional leaves, although exceptionally they can have around 20on well grown individuals (Crowder et al. 1990).
Experimental studies in S Germany showed that leaf abscission and replacement is affected by the rate ofinsect capture and subsequent plant growth rate. Thus leaf number remains fairly constant, while theamount of stored nitrogen in the hypocotyl (the storage region between root and shoot), increases withinsect supply. The experiments demonstrated that in Germany at least, insect capture is essential forgrowth of D. rotundifolia and that if small plants are continually deprived of insects, they willprobably die.
Despite these relations between growth and prey, reallocation of resources from old leaves appears to beas essential as insect capture for plant development. It is likely the case that as for otherinsectivorous species, the captured prey mainly provides the nitrogen and phosphorus requirements of theplant, while other elements, especially calcium, magnesium and potassium have to be obtained from thesoil (Schulze & Schulze 1990).
Reproduction
Rather tiny white flowers are produced in very small numbers on a leafless stem (ie a scape), from Juneto August, but they only open in really bright sunshine. Due to their closed nature, flowers arenormally self-fertilised. In cloudy W Ireland, flowers rarely fully develop or open, but insteadself-pollinate in bud (ie they are cleistogamous) (Garrard & Streeter 1983). Flowers that do manageto open, last only for a morning and are probably mainly selfed, although some degree ofwind-pollination is possible (Crowder et al. 1990). The proportion of plants flowering is veryvariable, both from place to place and from year to year. For instance, in one study the percentage ofplants flowering in Irish bogs varied from 35% near Ballymena, Co Antrim (H39) in 1965, to 73% atRossbeg (given as Rosbeg), Co Donegal (H35) in 1963 (Crowder et al. 1990). Another study of Irishplants from ten sites found there was a mean of 6.7 capsules per plant and capsules contained a mean of37 seeds (range 4-63) (Crowder et al. 1990).
Very little is known about seed dispersal, but the propagules are tiny and can float in water for months.Wind, water and animal feet (particularly those of birds), appear to be the most likely transportingagents.
In addition to seed, vegetative reproduction can take place by means of adventitious axillary buds thatform new rosettes beside and under the main leaf rosette of the plant. Decay of the stem axis separatesthese rosettes from one another, allowing them to grow on and establish distinct plants. Growth buds canalso develop on dying leaf blades, or on their petioles, forming penetrating roots as the parent leafbecomes moribund (Crowder et al. 1990).
In both D. rotundifolia and D. anglica new leaves grow in the centre of the rosette afterflowering, which makes old inflorescences look as if they arise laterally. This can result in possiblemisidentification if other characters are not checked (A. Culham, in: Rich & Jermy 1998).
Hybrid
A rare hybrid occurs with D. anglica (Great Sundew), named D. ×obovata Mert. & W.D.J. Koch (see account below).
Fermanagh occurrence
In Fermanagh, D. rotundifolia has been frequently recorded from 171 tetrads, 32.4% of those in theVC. It is very frequent and locally common on wet, acid bogland, including flushed areas of slightlyhigher pH, at all levels right up to Cuilcagh Gap near the summit of the highest mountain. Plants ofD. rotundifolia on most Fermanagh bogs are small, often with only four to eight functionalleaves, although exceptionally there can be around 20 leaves on well grown individuals.
Irish occurrence
Round-leaved Sundew is chiefly found in peaty uplands and on lowland raised bogs, so it is widelyscattered across the whole of Ireland, but more frequent in the N, W and C of the country (NewAtlas).
British occurrence
D. rotundifolia remains frequent in the N & W of Britain and formerly (pre-1970) it wasalso widespread in parts of C England (New Atlas). Losses in lowland S & E England continuedue to habitat destruction (F.J. Rumsey, in: Preston et al. 2002).
European and world occurrence
Round-leaved Sundew is widespread throughout Europe, including S Greenland, Iceland andScandinavia, but absent from most areas of the south continental mainland and all of the Mediterraneanislands except Corsica (Jalas et al. 1999, Map 2962). Outlying stations occur in the Lebanon andin the Caucasus. Elsewhere, it is widespread and circumpolar in boreal and temperate zones worldwide,extending across N Asia to Korea, Japan and New Guinea. In N America, it ranges from the subarctic toVancouver on the W Coast and Georgia in the east and thence to Alabama. It is absent from the RockyMountains (Hultén & Fries 1986, Map 999; Crowder et al. 1990).
Threats
While D. rotundifolia is tolerant of forms of disturbance such as grazing, occasional fires ortrampling, undoubtedly there have been population losses locally in Fermanagh and elsewhere in B &I, mainly due to drainage and peat-cutting. At higher altitudes, losses are the result of forestryplantation, over-grazing, peat burning and consequent bog surface erosion.
Native, extremely rare, but probably under-recorded. Eurosiberian boreo-temperate.
July 1949; MCM & D; bog at Meenatully Td, near Lough Vearty.
This hybrid has been recorded only once in Fermanagh, growing with both parents on a flat bog atMeenatully near Lough Vearty in July 1949 by Meikle and his friends (Carrothers et al. 1950). Theleaf-blades of this hybrid are more similar to D. intermedia than to either of the parentspecies, but it has a ± centrally arising inflorescence peduncle, which like that of the parent speciesis straight. Apart from this, the hybrid also differs in being completely sterile, having small, emptycapsules. Unfortunately in many sites the proportion of Drosera plants bearing flower scapes isoften quite low, making it very difficult or almost impossible to distinguish the hybrid. The hectad mapin the Hybrid Flora of the British Isles, shows the range of D. ×obovata covers most of the joint range of the parent species, with by far the greatest recordedfrequency of the hybrid in B & I being in NW Scotland (Stace et al. 2015). The same mapindicates that in Ireland, the hybrid has been recorded in a total of 27 hectads, 21 of which alsocontained both parents. Both parents plus D. intermedia are regularly, indeed rather frequentlyrecorded together on the same bog surface in Fermanagh. Obviously the hybrid should be actively lookedfor each time the parent species are found growing immediately adjacent.
The fact that all three Drosera species occupy very similar wet, acid bog situations, means thatthe shape and size of the leaves alone would not enable the hybrid to be discriminated from D.intermedia. Robert Northridge and the current author therefore have no doubt that D. × obovata is being regularly overlooked in Fermanagh andelsewhere. However, while we cannot tell the extent of such errors, we do know that in Britain thishybrid nowadays is usually regarded as relatively frequent, although it remains under-recorded (Staceet al. 2015).
In the Lough Neagh area, Harron (1986) mentioned just one very old (mid-Victorian) record at Sluggan Bognear Randalstown (given as 'Slogan'), undated but listed by David Moore in Cybele Hibernica −before the plant was fully recognised as being a hybrid (Moore & More 1866). The FNEI 3,however, lists three post-1969 sites for this hybrid in Co Antrim (H39) (Hackney et al. 1992).
In NE Scotland, Mary McCallum Webster's Flora of Moray, Nairn and East Inverness, has the authordescribe this hybrid as, "frequent where both parents occur". Closer examination of thisparticular Flora, however, indicates that a total of only ten sites are listed for the hybrid inthe large geographical area covered, three of which have 19th century dates only (McCallum Webster1978). Whenever D. × obovata is found, typically it occurs either asisolated individuals or in very small numbers.
Still we have to recognise that on account of the similarity of D. ×obovata to D. intermedia and also to small plants of D. anglica, this hybrid isvery easily over-looked.
Threats
Opening drains and cutting peat bogs.
Native, occasional and widely scattered. Circumpolar boreo-temperate.
1806; Scott, Prof. R.; Co Fermanagh.
June to October.
Growth form and preferred habitats
Mature plants of D. anglica are distinguished from D. intermedia (Oblong-leaved Sundew), bytheir proportionally narrower leaves which have a blade up to 5 cm long, the stiffer nature of the plantoverall, the presence of glandular hairs on the petioles and by the flowering stem being up to twice aslong as the leaves and arising centrally (though sometimes bent at the base and arising more at oneside). The leaves are much more erect or ascending than those of D. intermedia and they do notform a flattened, or even a definite rosette compared with those of D. rotundifolia (Round-leavedSundew), from which they also very obviously differ in shape and size.
Apart from inhabiting the very wettest conditions on Atlantic ombrogenous acid peat bog surfaces, whichoffer nutrient-starved growing conditions dependent upon dust brought down by rainfall (includingraised, valley and blanket bog types), D. anglica appears to avoid any degree of shade, whichalso helps to distinguish it from the ecologically more tolerant D. rotundifolia. It does nottolerate any degree of burial in peat, nor even partial overgrowth in Sphagnum hummocks.
Great Sundew also frequents somewhat more nutrient-rich sites than the other two Sundew species. Thisoccurs only in somewhat drier, climatically more continental parts of Britain − including conditions oflime- or base-rich fens. Although there is no scientific analysis available to prove it (see Crowderet al. 1990, p. 256), this appears probably more to do with water supply than nutrient levels,since what D. anglica really demands is a constantly moist substrate. It is often foundgrowing in standing water in bog pools and it tolerates being submerged. Flushing of bogs and fens withgroundwater springs supplies this irrigating moisture requirement, which would be particularlysignificant in the drier regions of B & I.
Webb & Scannell (1983) comment on the very remarkable and unique occurrence in an Irish context ofD. anglica growing in highly calcareous fens on the N & E shores of Gortachalla Lough, NNW ofGalway. This is replicated in some English base-rich fens and Webb recounts seeing the species growingon pure calcareous marl in Gotland, SE Sweden (Webb & Scannell 1983, p. 81).
In other parts of B & I, D. anglica may rarely be found growing on lakeshore sands or on rivergravel (Crowder et al. 1990).
Variation
No varieties are recognised in B & I, although in the Alps and in Arctic Europe small forms with justone or two flowers are sometimes referred to as var. pusilla Kihlm, or forma minorAbromeit (Crowder et al. 1990; Sell & Murrell 2018).
Fermanagh occurrence
There are records of D. anglica in the Fermanagh Flora Database from a total of 42 tetrads, 8% ofthose in the VC. It remains sufficiently frequent in Fermanagh to be described as occasional and it isquite widespread, mainly in the more upland W of the county. It is still present (or at least it wasonce recorded) in 29 post-1975 tetrads. However, the tetrad map suggests a definite decline of thisspecies during the last 40 years.
D. anglica has more limited ecological tolerances than D. rotundifolia (Round-leavedSundew) and in Fermanagh it grows in the wettest parts of bog pools and hummock surfaces. Ittends to be either submerged in the middle of intermittent pools, or occurs around the margins of morepermanent bog pools, or beside soakaways, the flushed seepage areas of valley bogs. It is also found indamp hollows on blanket bogs and more rarely and more sparsely in open areas on wet heaths.
Insect capture, plant nutrition and performance
The larger leaves of D. anglica can catch more insects simultaneously than D. rotundifoliacan manage, but while the leaf surface and its trapping hairs react more quickly, digestion is lesscomplete. Such differences in nutrition may eventually help explain the contrasting pH tolerances of thethree Sundew species, a topic which otherwise remains a mystery comparable with the differing ecologicalbehaviour of Schoenus nigricans (Black Bog-rush) in Irish bogs and British fens.
Deleterious effects of disturbance, drainage and drought
On account of its more strictly defined hydrological requirements and especially an absolute intoleranceof dry conditions, D. anglica is even more sensitive and vulnerable to drainage than the otherSundew species. The greatly disturbed habitat that is left after mechanical peat cutting using asub-surface boring auger, is extremely detrimental to the survival of this species: the pre-existingwater table is hugely disrupted and the shallow roots of D. anglica are completely unable tomaintain the plants in these conditions. Of the three Drosera species we are dealing with here,D. anglica is by far the most susceptible to drought, being confined to hollows that nevercompletely dry out on bogs and wetter heaths.
Reproduction
The floral biology of D. anglica is similar to that described for D. rotundifolia (see thataccount) and a detailed summary is provided by Crowder et al. (1990).
Seed longevity
Fortunately Drosera seeds appear to have a dormant survival period in the soil seed bank ofseveral years, eg, up to 5 years in both D. intermedia and D. rotundifolia. There does notyet seem to be any estimate of this for D. anglica in the literature and no records for anySundew from sites in B & I (Thompson et al. 1997).
Colonising ability
In common with D. rotundifolia, seedlings of D. anglica display colonising ability on bare,wet peat surfaces. Typically D. anglica roots in either wet peat or in Sphagnum moss andwhile the plant can tolerate flooding for periods of up to two months, it never forms dense mats ofplants under these conditions in the manner of D. intermedia (Oblong-leaved Sundew).Rather it tends to occur singly or in small groups.
British and Irish occurrence
The New Atlas map shows that D. anglica is increasingly confined to N and W parts ofScotland and Ireland, with scarce outliers in Cumbria and coastal parts of W Wales, southern England andEast Anglia. The species has suffered a major decline in both B & I, but it has been much moredrastic and widespread in England than in SE & E Ireland. This difference can undoubted beattributed to the particular sensitivity of D. anglica to operations involving drainage,peat-cutting and burning of bogs and wet heaths. Nowadays D. anglica, which is more confined tolowland situations than D. rotundifolia, is much more predominantly northern and western in itsdistribution on both islands than was previously the case (Preston et al. 2002).
European and world occurrence
D. anglica has a more restricted European range than D. rotundifolia, being absent fromIceland and much less widespread in France. It is rare in N Spain, ± absent from the Mediterranean basinand rapidly thins east of Switzerland (Jalas et al. 1999, Map 2963). Again, compared with D.rotundifolia, it is less extensive and more discontinuous in Asia although it does reach Japanand Kauai Island in the Hawaiian group (Crowder et al. 1990). D. anglica is alsocircumpolar and in N America is widespread in the more northern regions from coast to coast. However, itdoes not penetrate as far south on the continent as D. rotundifolia does (Hultén & Fries1986, Map 1000).
Threats
Drainage and cutting of peat bogs.
Native, occasional. Suboceanic temperate, but also present in N America.
1836; Mackay, J.T.; bogland on Cuilcagh Mountain.
May to October.
Identification and variation
Intermediate in size and leaf shape between D. anglica (Great Sundew) and D. rotundifolia(Round-leaved Sundew), the leaves of D. intermedia are held erect and have a wedge-shaped blade7-12 mm long and 4-10 mm wide. The leafless flowering stem (ie scape), arises laterally from below theterminal leaf 'rosette'. The scape is decumbent at its base and thus appears to curve up from the sideof the plant, whereas in D. anglica the scape originates from the centre of the loose rosette ofleaves.
Unfortunately, D. intermedia shows some variation in this often-used identification feature, soplants need to be examined very carefully. Significantly, the scape is usually much shorter in D.intermedia than in D. anglica, often scarcely overtopping the leaves − although this alsois not completely reliable, since it can rarely be up to three times as long as the leaves. Having saidthis, in general the scape is short in comparison with D. anglica, where it is usually two tofour times the leaf length (Crowder et al. 1990).
In both D. rotundifolia and D. anglica new leaves grow in the centre of the rosette afterflowering, which makes old inflorescences look as if they arise laterally. This can result in possiblemisidentification if other characters are not carefully checked (A. Culham, in: Rich & Jermy 1998).
On account of the variation involved, RHN and the current author fear that there may be some confusionbetween D. intermedia and D. anglica. Particularly with respect to leaf shape and size,potential errors also occur between D. intermedia and the hybrid between D. anglica andD. rotundifolia (= D. × obovata), which by its nature is oftenintermediate between its parent species. This problem is not unique to Fermanagh, however, and we do notbelieve that the potential for mistake greatly distorts the picture of taxa frequency and occurrencepainted here.
Growth and preferred habitats
Oblong-leaved Sundew usually forms rather dense mats of plants up to 100 cm in diameter on bare, wet,acid peat, typically on the bottom of shallow pools on both lowland raised and blanket bogs. Plants inthese mats are inevitably entangled, since vegetative propagation by means of axillary buds isundoubtedly involved in their production. Axillary bud formation and colony mats are probably morecommon in D. intermedia than in the other two sundews. The stem axis and leaf petioles growstrongly and, to a limited extent, this enables the plant to keep up with a rising water level orrapidly growing moss cushion. However, unlike D. rotundifolia, Oblong-leaved Sundew cannottolerate any degree of shading from shrubs (Crowder et al. 1990).
In terms of habitat preferences, D. intermedia more closely resembles D. rotundifolia thanD. anglica, ie in occupying shallow temporary pools in bog hollows and appearing around the edgesof deeper, more permanent pools and towards the base of mossy hummocks on valley bogs. Plants appearable to withstand flooding indefinitely and can therefore be considered as aquatics. More rarely D.intermedia appears in wet, flushed seepage areas, or beside paths on blanket bog slopes and onbare peat in disturbed scrapes on wet, peaty heathland. It is more of a lowland species than D.anglica and D. rotundifolia, reaching only 335 m above sea level in Co Donegal (vcunspecified) (Crowder et al. 1990).
D. intermedia appears to combine the ability of D. anglica to tolerate long periods ofimmersion, with the capacity of D. rotundifolia to survive on substrates that dry out for a timein summer, but D. intermedia does not quite match the competence of either in extreme situations.In the oceanic west of Ireland, however, rain is plentiful and well distributed in time, so that summerdroughts are rare and not usually very prolonged.
On most if not all of our lowland bogs, all three Drosera species occur together, often associatedwith Narthecium ossifragum (Bog Asphodel), Rhynchospora alba (White Beak-sedge),Potamogeton polygonifolius (Bog Pondweed) and either both common Cotton-grasses ((Eriophorumvagin*tum (Hare's-tail Cottongrass) and E. angustifolium (Common Cottongrass)), or justthe latter on its own.
Insect trapping
D. intermedia is an efficient insect trapper and Robert Northridge and I have seen patches of theplant in Fermanagh covered with the dead bodies of Coenagrion spp., damselflies and, on anotheroccasion, the bodies of Pieris spp., butterflies. Data from three field sites in Britain alsoshowed a correlation between the amount of insects captured and plant growth and reproductiveperformance. The percentage of plants that had captured prey on their leaves in these studies variedfrom 55%-96% (Crowder et al. 1990).
Reproduction
As in D. rotundifolia, flowers are bisexual and are usually self-fertilised. Some flowers do notopen (ie they are cleistogamous). However, the flowers are not as tightly closed as is the case in D.rotundifolia and all of them appear capable of opening when exposed to bright sunlight.Normally, however, cleistogamous flowers fail to open and they self-fertilise while in bud.
Seeds of D. intermedia are more ovoid than the other two British species and they are dark incolour and warted. The rough, warted nature of the seed coat makes them stick together, reducing theirability to disperse by wind. In water the seeds can float for several months and probably local watermovement represents the best chance there is for dispersal.
The proportion of plants flowering, capsule numbers and seed production per capsule are all highlyvariable from year to year and place to place making general statements pointless. The proportion ofseeds showing viable germination in experimental studies is often very low − in several studies it beingaround a mere 2%. When seeds are given a prolonged period of cold stratification of up to eight weeks,followed by a diurnal temperature regime and 14 hour daylength, however, germination rates were improvedup to 78-90% (Crowder et al. 1990).
Fermanagh occurrence
D. intermedia is more local and appears scarcer in Fermanagh than the other two native sundewspecies. Its frequency is merely occasional and it is mainly found on the Pettigo Plateau bogland andaround the Five Points area of lowland bogs W of Enniskillen. D. intermedia has been recorded ina total of 31 tetrads (5.9% of the total in the VC), but has not been seen in six of these during thepost-1975 period.
British and Irish occurrence and decline
John Harron in his 1986 Flora of Lough Neagh noted a similar decline of D. intermediapopulations around that large Ulster lake, where he described it as, "very local and becomingrare". This decline in frequency is reflected on the larger scale when one examines the B & Idistribution of D. intermedia displayed in the New Atlas hectad map, or indeed when theworld distribution of the three species native to these isles is considered (Crowder et al.1990).
D. intermedia is local and scattered throughout B & I and displays a strongly marked westerntendency of occurrence, more obvious than that shown by D. anglica and D. rotundifolia. Ithas suffered a long period of decline in both islands, dating from at least the end of the 19th centuryonwards. This is particularly obvious in lowland C & SE England and in C Ireland, where it hasbecome locally extinct in many previously occupied sites, but the phenomenon is much more widespreadthan this. Losses appear to follow drainage, peat cutting and afforestation operations, affecting inparticular the destruction of lowland heaths and bogs. The result of this has been to make the planteven more definitely western in occurrence in both B & I than previously was the case (F.J. Rumsey,in: Preston et al. 2002).
Sinker et al. (1985) suggested that D. intermedia is the Sundew in these isles leasttolerant of competition, but it is uncertain on what scientific basis this verdict is given. It maysimply be a subjective comment based on field observation, such as I have made above. On the other hand,an experimental culture study made in a bog lake in Ontario by Wilson & Keddy (1986) examined thecompetitive ability of D. intermedia when it was paired in turn with six non-insectivorousspecies in containers (no species names given when reported in Crowder et al. 1990); it was foundto be a poor competitor, unable to tolerate much natural stress or disturbance in the particularconditions involved.
European and world occurrence
Beyond B & I, D. intermedia has a much more restricted occurrence than either D.rotundifolia or D. anglica. It is present in NW and C Europe, but is absent from NScandinavia and from most of the Mediterranean area, except a small area in Tuscan Italy (Jalas etal. 1999, Map 2964). It is also rare in E Europe, although it has been recorded from the easternend of the Black Sea (Hultén & Fries 1986; Crowder et al. 1990). Other sources also mentionTurkey (Sell & Murrell 2018), although there is not unanimity about the more eastern occurrences inthe published maps and other literature. The overall phytogeographic distribution is described asbelonging to the Suboceanic Temperate element (Preston & Hill 1997).
D. intermedia is also amphi-Atlantic, being widespread in eastern N America. However, there areonly a few isolated records from the N and NW of that continental landmass, although to the south it isrecorded down into Florida and on the island of Cuba (Hultén & Fries 1986, Map 1001).
Threats
As with all Sundew species, drainage, cutting of peat bogs and afforestation are the major problemsrestricting available habitats.
Introduction, archaeophyte, a rare garden escape. European temperate, but also widely naturalisedincluding in Japan, N & S America and New Zealand.
1885; Hart, H.C.; Waterfoot, where the River Letter enters Lower Lough Erne.
April to July.
Growth form and preferred habitats
V. odorata is a patch-forming, perennial with long, creeping stolons and flowers usually (but notalways) sweetly perfumed. It grows in a range of habitats from damp, shady banks under trees, or inplantations, to more open conditions in hedgerows, scrub margins, on roadside banks and along laneways,usually near habitation from which it often arises as a garden escape. It very rarely occurs inso-called, native woodland. Sweet Violet is generally considered to favour fertile, base-rich,calcareous, alkaline to neutral, moist soils, but it seems to have quite wide tolerances of both soilconditions and levels of light and shade (Porter & Foley 2017).
V. odorata is the only scented violet species in the British Isles − all the other violets inthese islands are of the so-called 'Dog-' or 'Horse-' variety, ie less highly regarded, wild, scentlesstypes (Genders 1971; Grigson 1987).
Variation
Six named varieties based on leaf size and flower colour are listed by Porter & Foley (2017)and Sell & Murrell (2018).
Flowering reproduction
V. odorata is the earliest of the Viola species in B & I to come into flower, flowersoften appearing as early as February. The flowers range in colour, but usually are dark violet, orwhite, or something in between. The spur is stout, pale lilac, violet or purplish-green, sometimeshooked upwards and it is not notched. Flowering can continue into late April and sometimes there isanother flush of flowers in the autumn. The sweet fragrance attracts bees which collect nectar from thespur and pick up pollen as they do so (Proctor & Yeo 1973). The fruit capsule is rounded or slightlythree-sided and is densely covered with short, patent (ie erect), hairs.
The flowers of V. odorata are hom*ogamous and compatible with their own pollen. However,self-pollination and self-fertilisation is prevented by the irregular, two-lipped, zygomorphic structureof the flower. Nevertheless, in late spring and summer Sweet Violet produces small flowers that neveropen, the styles and stamens of which remain short and poorly developed. These flowers self-pollinatewithin the unopened bud, ie they are cleistogamous, and they set additional seed.
Seed dispersal
The globular capsule lies on the ground and splits open, but does not explode to release its seeds.Rather, the valves of the split capsule dry and shrink, spilling out the seeds under or near the parentplant.
V. odorata seeds bear a conspicuous 'elaiosome' or nutrient oil body that attracts ants (Ridley1930). Studies show that seed carried to ant nests has a greater probability of seedling emergence thanseeds placed randomly on or slightly below the soil surface. The advantage of ant dispersal is therelocation of the seed to a 'safe site', ie one with favourable soil moisture and nutrient conditions,enabling better seedling establishment than otherwise. Micro-environment conditions at the scale of theseed can vary dramatically over distances as small as 10 cm. Movement by ants reduces predation bybutterfly larvae, small mammals and birds, all of which are distance-related, rather thandensity-dependent, mortality factors (Beattie & Lyons 1975; Culver & Beattie 1978, 1980; Howe& Smallwood 1982).
Fermanagh occurrence
In Fermanagh, V. odorata has been recorded in a total of 16 tetrads (3% of those in the VC),scattered throughout the lowlands, usually near habitation. The largest and most established stand inFermanagh is at Old Crom Castle, where Sweet Violet covers a considerable area under lime trees. Nine ofthe stations in the VC are pre-1955 and only at Crom does the species appear long persistent, havingsurvived there for at least 50 years.
British status and occurrence
The native range of V. odorata in B & I is clouded by a very long history of introduction andcultivation. Both Genders (1971) and Grigson (1987) state that in ancient times it was in commercialcultivation for the sweetening properties of its perfume, and by the Middle Ages it was in use forculinary, medicinal, cosmetic and perfumery purposes (Grieve 1931).
Irish status and occurrence
In Ireland, it is traditionally regarded as possibly native in some areas of the south – but perhaps alsoin the Midlands, as suggested in the BSBI Atlas (Perring & Walters 1976). This position isnot followed in the New Atlas, where the supposed native Irish occurrence is more confined to theSE quarter of the island (Preston et al. 2002). There may also be something of a decline withinthe Irish native area, since many of the records are pre-1970 (28% of them by RSF's calculation). In theN & W of Ireland, V. odorata has long been acknowledged as a garden escape (CybeleHibernica 1866; An Irish Flora 1996). The Irish Census Catalogue, with its,"not always native" contribution, is uncommitted on the matter (Scannell & Synnott 1987).
With its creeping habit, colonising ability and sometimes long persistence, V. odorata might bestbe considered in Ireland, "doubtfully native, but thoroughly established" (Bunker 1950). TheCat Alien Pl Ir (Reynolds 2002, p. 323) simply reports that V. odorata is,"considered an introduction or garden escape in many places".
Cultivation, uses and status of Viola odorata: The status of any Violaspecies is always likely to be decided on circ*mstantial evidence since the flowers produce littlepollen and thus are very unlikely to appear in the fossil record (Godwin 1975; Webb 1985). The statusquestion is made even more ambiguous when, as in the case of V. odorata, there is a very longhistory of cultivation and human use. Although it is sensible to be wary when identifying ancient plantreferences with modern species (Raven 2000), it seems probable that V. odorata was a plant knownand referred to by the ancient Greek pastoral poet Theocritus (c 300-c 260 BC) (Lindsell 1937). BothGenders (1971) and Grigson (1987) state that in ancient times V. odorata was in commercialcultivation for the sweetening properties of its perfume.
By the Middle Ages, Sweet Violet was in use for culinary, medicinal, cosmetic and perfumery purposes(Grieve 1931), and it was held in great affection by both Gerard (1597) and Shakespeare, who made asmany as 20 references to it in his plays and poems (Ryden 1977). Medicinal uses of the plant were many(Grieve 1931), most significant perhaps being as a laxative mild enough for use with infants. Thespecies also has a historical use as a bronchial and cough remedy. Like its relative, Violatricolor (Wild Pansy), it was also used to treat mild skin conditions. The fresh leaves of V.odorata have been used to relieve the pain of cancer, especially in the oesophagus where otherpain relievers have failed. A syrup made of the flowers has been used to treat eye inflammation,sleeplessness, jaundice, coughs, bronchial ailments and throat pain (Grieve 1931; Vickery 1995).
European and world occurrence
It is clear from the above that the native range of V. odorata is clouded by its long history ofcultivation and introduction, yet Hultén & Fries (1986) remark that it, "originates mainly fromS and W Europe". The latter authors indicate that they can only tentatively delineate its originaldistribution − using a dotted line on their map to mark the supposed northern limit of the native rangeof the species (Hultén & Fries 1986, Map 1318). Their map shows the species as being native inEngland and Wales, S of a line between Hull and Liverpool, plus France and all territory S of the Alps,including the Balkan peninsula and eastwards to the shores of the Caspian Sea. Isolated stations arealso shown around the E and S shores of the Mediterranean in Asia Minor and N Africa. There is somedispute as to the status of V. odorata in Denmark and S Sweden, but it is definitely regarded asa naturalised introduction in N & C Europe, Japan, N & S America, the Philippines and NewZealand (Hultén & Fries 1986, p. 1092).
Names
The name 'Viola' is the Latin name used for various sweet-scented flowers including stocks andwallflowers, and is derived from the Greek 'ion', which in an earlier form had an initial lettercorresponding to 'v' or 'w', the digamma, which was later lost (Stearn 1992). Even without being alinguist, it is clear that the identification of ancient flowers by translation alone is an extremelyrisky undertaking. The Latin specific epithet 'odorata' means 'sweet-smelling' or 'fragrant'(Gilbert-Carter 1964). The English common name 'Violet' is of 14th century origin, derived from the OldFrench 'violette', a diminutive of 'viole', from the Latin name 'Viola', which as already mentioned,appears to have originally been given to Matthiola incana, now called Hoary Stock − the parentspecies of the garden Stocks, which have an aromatic spicy perfume − as well as to Viola odorata,Sweet Violet (Genders 1971; Grigson 1974).
Threats
None.
Native, common, widespread and locally abundant. European temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year, but mainly April to October.
Growth form and preferred habitats
This rosette-forming, wintergreen perennial is very variable in form, but when in flower is usuallyrecognised by its broad, overlapping violet-coloured, upper petals decorated with much-branched,spreading, purple veins on the broad, lowest petal. The flower also has lanceolate sepals, 1.5 mm ormore and enlarging when in fruit, and a stout, pale, yellowish-green spur that is conspicuously notchedat the rear. In comparison with V. reichenbachiana (Early Dog-violet), with which it is mostlikely to be confused, the leaf stipules offer a useful distinction: in V. riviniana they arenarrow (but broader than those of V. reichenbachiana), and they are fringed with coarse fimbriae(marginal teeth) that are distinctly shorter than those of Early Dog-violet.
V. riviniana tolerates a very wide range of soil type and nutrient status, though it is mostfrequent and abundant on moist, semi-shaded, infertile, unproductive habitats, especially in lowlanddeciduous woods and margins of scrub. V. riviniana is also occasionally found in disturbedhabitats on roadside banks and verges, beside paths, on waste ground and in gardens. It is also frequenton calcareous screes and on coastal sand hills. It is really only absent from situations of extremewetness, dryness, disruption, or acidity below about pH 4.0. In pasture situations, it is little grazed(Grime et al. 1988).
The establishment strategy of V. riviniana in England was described by Grime et al. (1988)as intermediate between a stress-tolerator and a more general C-S-R (a midway balance of Competitor,Stress-tolerator and Ruderal species). The exceptionally wide habitat range it displays in Fermanaghhelps to confirm this view of its general, intermediate ecology and competitive, colonising andsurviving ability.
Under deciduous woodland canopy, V. riviniana is not sufficiently vigorous to compete successfullywith dense patch-forming vernal species such as Hyacinthoides non-scripta (Bluebell),Ranunculus ficaria (Lesser Celandine) and Anemone nemorosa (Wood Anemone). It canhowever become plentiful and conspicuous on more shallow, rather drier soils on banks or near rocks, andalso in ground where relatively large amounts of tree litter accumulates, a situation that excludes mostother herbaceous species.
An experimental study in England of the effect of leaf litter on garden transplants of five ground floraspecies including V. riviniana, found that it was slow to emerge from the covering layer, butshoots did eventually emerge from all of the transplants, even in plots where the largest experimentalamounts of litter persisted. Thanks in part to the orientation and storage reserves of their short,stout, erect stock, the violet plants were able to respond to varying litter depths by elongation of theleaf petioles, and there was no significant effect of the litter depth or persistence on plant yieldcompared to exposed control plants (Sydes & Grime 1981).
In woodland, on heavy, often base-rich soils, V. riviniana regularly occurs along with V.reichenbachiana, although the latter is somewhat more shade-tolerant, flowers a few weeksearlier and produces fewer seeds (Grime et al. 1988). The two species occasionally hybridise (seebelow).
Variation
V. riviniana is not just the most common and widespread British dog-violet by a long way; it isalso the most variable and 'plastic' in its response to environmental conditions (Valentine 1949). Forexample, some plants can form adventitious buds and shoots on the roots whilst others do not. Thisallows plants with the necessary gene(s) to reproduce vegetatively, forming clonal patches.
Although two forms of the plant have been described within this species in British Floras for over 70years (Valentine 1941, 1950), and they have been variously rated as subspecies, ecotypes or varieties(Clapham et al. 1962, 1987; Stace 1991, 1997), I will not follow Prof. Valentine into thelabyrinthian obscurities of g-Ecospecies, a-Ecospecies, or cytotypes on this one! (Valentine 1950). Hissplit of subsp. minor and subsp. riviniana has not had very much attention in an Irishcontext − not even getting a mention in our main field Flora, An Irish Flora (8th ed., Parnell& Curtis 2012). Indeed, the general consensus throughout B & I now appears to be thatValentine's subsp. minor or var. minor, which is dwarfed to about half size in bothflower and all of its vegetative characters (Porter & Foley 2018), is merely a phenotypic reactionto exposed, dry or coastal ecological situations and, at most, it is a habitat induced ecotype (Stace1991, 2019). Porter & Foley (2018) mention and illustrate three colour forms or cultivars that arein garden cultivation: pure white, purple and pink.
Flowering and pollination
Like several other Viola species in these islands, V. riviniana plants often flower twicein a season. In spring, flowers are abundantly produced from mid-March to May, and although they arefunctionally self-compatible, pollination is only achieved in these flowers after insect visitation. Ageneral lack of seed set by the spring flowers is probably due to a scarcity of insect visitors early inthe year, even though the flowers do provide copious nectar, which is reserved in the spur specificallyfor longer-tongued insects (Beattie 1969a). It is interesting to note that the flowers do not appear topossess any scent to attract visitors (Press et al. 1994).
A great deal of extremely detailed research by Beattie into the pollination of V. riviniana andits near relatives (Beattie 1969a & b, 1971, 1972) has shown that the mechanism of flowering in thespring follows two distinct phases with respect to pollen presentation: an initial very precisepollination syndrome involving small amounts of pollen and "brief and irregular visits" by afew specialised long- or medium-tongued insects, chiefly hover-flies (Diptera: Syrphidae), bee-flies(Diptera: Bombyliidae) and bees (Hymenoptera: Apoidea) (Beattie 1969b). This is followed after four tosix days by a second phase, which may last between two and nine days depending upon weather conditions,when the pollen is released from the anthers onto the surfaces of the petals. During this latter stagethe pollination mechanism becomes completely unspecialized, and many more insect species (up to 40 ormore) may become involved. During the second stage, selfing or chance crossing are much more readilyachieved than before, although selfing is probably still predominant. In reality, however, althoughout-crossing can and does operate in the normal 'open pollinated', chasmogamous spring flowers of thisspecies, viable seed production in this type of flower is frequently almost negligible (Beattie 1969a).
Late season flowering
In late-summer and autumn, the same or different plants often develop a second flowering period of mainlycleistogamous flowers which are automatically self-pollinated while still in the bud. Since theseflowers do not require insect visitors to function, they are often very inconspicuous and may remaintucked away at the base of the plant, more or less concealed by the leaves of the characteristicnon-flowering central rosette. Thus in any season the majority of violet seed is produced by selfing andprobably most of it is produced by the obligately inbreeding cleistogamous flowers (Valentine 1941,1962; Beattie 1969a, 1971).
Fruiting and primary seed dispersal
After fertilization of the ovules, the ripening fruit capsule of both types of flower (ie chasmogamousand cleistogamous ones) generally become elevated above the leaves by reorientation and elongation ofthe pedicel or flower stalk. Seed release is often described as 'explosive', but it is more the casethat they skite out of their already opened capsule by a sudden release of lateral pressure, in a mannerrather reminiscent of playing 'tiddlewinks' but with spheres involved instead of flat plastic counters.Perhaps it is better described as being like a marble being squeezed between the thumb and finger. Toexplain this matter, the Viola fruit is a three valved capsule which splits along very obvioussutures or lines of weakness, and then opens out to form boat-shaped valves containing a total of up toabout 20 smooth, ovoid, hard-coated, pale yellow seeds (Butcher 1961). The three somewhat elastic valvesdry and shrink shortly after they open and, as they do so, they exert lateral pressure on the containedseeds, so that one by one, or in small groups of individuals, they are forcefully ejected. The firstseeds expelled are usually from the central row in each valve and, not being under direct pressure fromthe valve walls, they often fall closer to the parent plant than those which remain to be expelled bydirect pressure (Leavitt 1902, quoted in Beattie & Lyons 1975). Violet seeds may fly distances up to2 m or so, depending upon whether or not the flight path is clear of leaves or other obstructions.
Secondary seed dispersal by ants
The seed coat of many Viola species has an attached oil-rich food body or elaiosome which may beof various origins; for example, often it is a portion of the funicle or stalk of the ovule, butsometimes it is an outgrowth from the seed coat. Irrespective of its origin, the elaiosome alwayscontains a high percentage of lipids, sometimes with starch and proteins also present. The size of theelaiosome on offer is relevant to its dispersal potential: larger ones being more attractive to antsthan small ones, but often in violets the difference is not significant (Beattie & Lyons 1975; Mark& Olesen 1996). Viola seeds may thus undergo secondary dispersal, being located andtransported by the ants, in many cases all the way back to their nest (Culver & Beattie 1980). Inother situations, they are transported a minimal distance, but even if they are simply scarified and thefood body removed, this enhances subsequent germination (Culver & Beattie 1980).
Almost irrespective of the distance the seeds are physically moved, this 'myrmecochory' (from the Greek,'myrmeco' = ant and 'khorein' = to spread around), confers several benefits on species which undergo theprocess; these involve not just additional dispersal from the parent plant, but avoidance ofpost-dispersal seed predators (which frequently include mice, voles and a considerable variety ofinvertebrates) and competitors, as well as scarification assisting germination. If removed to the insectnest or to ant refuse heaps near the nest, then additional moisture plus nutrients such as nitrogen andphosphate may become more available to the germinating seed (Culver & Beattie 1980; Beattie 1983).In practice this process is unlikely to transport the seeds very far in terms of distance, ie often onlya few cm, but there remains the potential for a few seeds to travel a moderately long-distance (perhaps50-100 m) to the ant nest.
Seed germination and buried seed survival
V. riviniana seed has a pronounced chilling requirement for germination so that seedlings onlyappear in spring after winter frosts are past (Thompson & Grime 1979). The majority of studiessuggest V. riviniana seed is transient, surviving overwinter and germinating more or lessimmediately under a wide variety of environmental conditions once the chilling requirement has been met.However, a handful of reports indicate longer persistence may occur, ie more than one year and, in twoinstances, for more than five years (Thompson et al. 1997).
Fermanagh occurrence
V. riviniana is by far the commonest violet in Fermanagh. It has been recorded in 448 tetrads,over 84.9% of those in the VC, growing in a very wide variety of habitats – chiefly (but notexclusively) shaded or semi-shaded. This includes almost all kinds of woody vegetation, plus establishedgrassland at all altitudes, rocky slopes, cliffs and screes.
British and Irish occurrence
The New Altas shows the species is almost omnipresent throughout B & I, apparently absent onlyfrom the prairie-like arable agricultural landscape areas of South Lincolnshire, West Norfolk andCambridgeshire (VCs 53, 28 & 29) (Preston et al. 2002).
European and world occurrence
Beyond the shores of B & I, V. riviniana is distributed widely in W, N, S. and C. Europeincluding Iceland and Madeira. In the Mediterranean, it stretches to S Greece, Crete and to coastal andmountainous parts of N Africa, but it is absent in the eastern Balkans, Cyprus and Asia Minor, where itis replaced by the closely related V. sieheana Beck. (Hultén & Fries 1986, Map 1326).
Names
The specific epithet 'riviniana' is the Latinised form of the name August Quirinus Rivinus (1652-1723), aProfessor of Botany at Leipzig, in whose honour the plant is called (Gledhill 1985). The English commonname 'Dog Violet' and variants of it, all refer to the fact that such species are scentless non-gardenplants, and as such are regarded as inferior. The name was apparently first given by John Gerard, whosimply translated the botanical name 'Viola canina'. Thus 'Dog Violet' is really a good exampleof a book name, ie one invented by a book writer, rather than originating from common parlance or folkusage (Grigson 1987).
Threats
None.
Native, very rare, but possibly over-looked.
1957; MCM & D; in woods at Drumcose Td, Ely Lodge, Lower Lough Erne.
Although this is the most widely recorded Viola hybrid in England, it is notoriously difficult toidentify due to the similarity of the parent species. It can combine floral characters closer to V.riviniana (Common Dog-violet) with broad petals and larger sepal appendages, but with a dark,more delicate spur of the much rarer V. richenbachina (Early Dog-violet), or else it mayhave narrow petals and shorter sepal appendages, bringing it closer to V. reichenbachiana. It canalso combine the darker spur colour of V. richenbachiana with the notched and furrowed spurtypical of V. riviniana (Stace et al. 2015).
This hybrid is very rare in Fermanagh, there being just three records made by Meikle and co-workers in1957. However, it is easily over-looked and could perhaps be quite frequent since the parent speciesfrequently occur together in Fermanagh woods and on shady banks, especially over calcareous substrates.While V. reichenbachiana flowers up to three weeks earlier in the spring than V.riviniana, the flowering periods nevertheless do overlap considerably.
Stace (New Flora of the BI) notes that the hybrid is intermediate between the parent species inthe length of sepal, but it has a dark spur and is highly sterile (but not completely so). D.H.Valentine (in: Stace 1975, p. 157) reckoned that the hybrid is widely distributed in England, but theNew Atlas map for it shows that V. × bavarica is very muchrarer than one might imagine considering how frequently the parent species grow together. It is possiblethat the hybrid is being regularly overlooked, or that it is being mistaken for V.reichenbachiana, which it more closely resembles.
The New Atlas hectad map and that in the Hybrid Flora (Stace et al. 2015) bothindicate a thin and very uneven smattering of records of this hybrid over England and Wales south of aline between Teeside and Lancaster, with an almost total absence in the industrial midlands. In Ireland,there are a total of just nine hectads plotted in Cos Limerick (H8), Kildare (H19), Antrim (H39) andLondonderry (H40). The records in the latter two VCs are from around Lough Neagh, date from 1946-7 andwere discovered by the original three Fermanagh Typescript Flora recorders before R.C. Davidsonjoined them, R.D. Meikle, E.N. Carrothers & J.McK. Moon (MC & M) (Flora of Lough Neagh;FNEI 3). In addition to the first Fermanagh record listed above, there are two others also madeby Meikle and co-workers during July 1957. The sites are: Castle Coole NT estate, Enniskillen; Lisgoole,on the shore of Upper Lough Erne, which lies on the SE outskirts of Enniskillen town.
The fact that there are no recent records for this violet hybrid does not mean that it has locallydisappeared. It only signifies that during the last 60 years no one has been confident enough todiscriminate it and claim its presence. Several observers, including ourselves, have reported findingviolets intermediate between V. riviniana and V. reichenbachiana, both in Fermanagh and inthe glens of Co Antrim, but only MCM & D have definitely recorded them as this hybrid(Beesley 2006).
Native, frequent but probably still under-recorded. European temperate.
1913; Carrothers, N.; the Enniskillen-Belfast Road, near Lisbellaw.
March to June and August to October.
Growth form and preferred habitats
This perennial dog-violet flowers early in the year – in late February, about a fortnight before theother, much more common violet, V. riviniana (Common Dog-violet). Like the latter, some V.reichenbachiana plants produce a second flush of entirely self-pollinated (ie cleistogamous)flowers late in the summer. The flowers are usually pale violet and have very narrow upper petals thatdo not overlap and are sharply bent backwards (reflexed). The lowest petal is marked with unbranched orlittle-branched dark veins and the attached deep purple, or deep lilac spur is slender and darker incolour than the rest of the flower. These are the most distinctive identification features of thespecies when in flower. When not in flower, the plant can be recognised by its narrow leaf stipules,which have marginal fimbriae considerably longer than those of V. riviniana (Porter & Foley2017).
V. reichenbachiana is a plant of lowland deciduous woodlands, hedgerows and wood edges. It is alsomuch more lime-tolerant than V. riviniana. Like the latter, it normally competes best under shadeconditions. However, Early Dog-violet also occurs more rarely in open, well-lit situations in limestonegrasslands and even on calcareous screes, provided there are other environmental factors that limit thevigour of its ecological rivals, eg a shallow, free-draining substrate, or low or unbalanced soilnutrient levels.
Variation
In the past, V. reichenbachiana was combined with V. riviniana in V. silvatica Fr.,but cytological studies have shown that in fact V. reichenbachiana is a diploid with 2n=20chromosomes, whereas V. riviniana has various chromosome numbers, ie 2n=35, 40, 45, 46, 47 (butmost often 2n=40). Plants of V. riviniana with more than 40 chromosomes in their nuclei areaneuploid forms which have five, six or seven small supernumerary or B-chromosomes present (Gadella1963).
Sell & Murrell (2018) mention an occasional form of the species with purple blotches on the lowerpetal, named var. punctata (Rouy & Foucaud) P.D. Sell, and Porter & Foley (2017) detail arare, almost pure white flowered form, sometimes called var. leucantha Beck.
Flowering reproduction
Flowers are without scent and seed production from the spring anthesis is low. The closed flowers, thatare sometimes produced later in the year, self-pollinate in bud (ie they are cleistogamous) and theyproduce more seed than the open pollinated ones (Sell & Murrell 2019).
Seed dispersal
The seed of V. reichenbachiana bears a small elaiosome food appendage and therefore presumably it issecondarily dispersed by ants to some extent. (For more detail of this mechanism, see the speciesaccounts of V. riviniana and V. odorata (Sweet Violet)).
Fermanagh occurrence
Our experience of Early Dog-violet in Fermanagh suggests that it is much commoner than most authoritieswould have us believe. V. reichenbachiana has been frequently recorded in 130 Fermanagh tetrads,24.6% of those in the VC. We reckon that despite our efforts it remains under-recorded due to its earlyflowering and probably it is even more widespread in lowland Fermanagh and especially so on limestonesoils.
The current survey shows it is a very characteristic plant of hazel woods in springtime, where it flowersfrom March to May, with a notable peak in April. However, it is identifiable and is recorded in muchsmaller numbers right through the summer months, even into October. Typical habitats are in deciduouswoods and shady hedgebanks along roads, rivers and by lakeshores.
British and Irish occurrence
V. reichenbachiana is common in suitable habitats across much of both England and Ireland althoughit thins markedly northwards and is rare in Scotland and absent from most of W Wales and SW England(New Atlas; Porter & Foley 2017).
European occurrence
V. reichenbachiana is restricted to Europe and adjacent parts of Africa (including the CanaryIsles) and W Asia (Hultén & Fries 1986, Map 1324). It is widespread but uncommon in suitablehabitats in W & C Europe from S Sweden southwards to Greece and eastwards from Spain to the Caucasus(Sell & Murrell 2018).
Name
The specific epithet is the Latinised name of the Dresden botanist and taxonomist, H.G. LudwigReichenbach (1793-1879), in whose honour the plant is named.
Threats
None.
Native, rare, possibly under-recorded. Eurosiberian boreo-temperate.
1884; Barrington, R.M.; Devenish Island, Lower Lough Erne.
June to August.
Growth form and preferred habitats
V. canina is a ± hairless perennial with ovate-lanceolate, ± triangular-shaped, dark green leaves.The shiny leaves are strongly reticulate below. The flowers, produced from April to June, are a fineclear blue or bluish-grey colour, without a dark purple zone outside the whitish base of the lowestpetal. They are furnished with a bright yellow, or more rarely greenish-yellow spur, usually deeplynotched, but occasionally not. The vegetative plant has no stolons and no central leaf rosette, allleaves being borne on the flowering stems (Porter & Foley 2017; Sell & Murrell 2018). Greencleistogamous flowers or flower buds are produced in summer.
As its English common name suggests, Heath Dog-violet is chiefly found in unshaded acidic habitats and,apparently nowadays, it is mainly a plant of coastal distribution on more open areas of older, fixed,grey dunes and dune heaths across both B & I. However, the species does also occur scattered andmuch more rarely in open, vegetation gaps in inland situations including woods, stony heaths andgrasslands over dry, shallow or sandy soils and in open, stony ground, or in crevices and ledges on rockoutcrops. It can also occur on damp to moist, peaty ground near high water mark on stony riverbanks andon shingle around lakeshores. The waterside types of inland habitat are especially well represented inScotland and Ireland, and the soils in them may be much more calcareous or base-rich than is the normfor this species elsewhere (Corner 1989; New Atlas).
In most or all of its sites, coastal and inland, the associated vegetation needs to be kept fairly openby periodic grazing to permit V. canina to thrive and flower. In its lake shore and riverbankhabitats, competing plants are additionally limited by being subjected to occasional flooding afterheavy or prolonged rainfall (Corner 1989; M.J.Y. Foley & M.S. Porter, in: Preston et al.2002).
Variation
Two subspecies are recognised in Britain at least, if not confirmed for Ireland. Subsp. canina isthe widespread form in B & I, while subsp. montana appears to be confined in these islands toa couple of fens in Cambridgeshire. The latter form has somewhat larger flowers than the more commontype (15-22 mm long, as opposed to 7-18 mm in subsp. canina) and the stipules of the middleleaves of subsp. montana plants are less than half the length of the petiole (Sell & Murrell2018).
Fermanagh occurrence
In Fermanagh, V. canina has been recorded from 14 tetrads (2.7%), but only seven of them havepost-1975 dates. This suggests that Heath Dog-violet, which was always a very local species here, hasdeclined during the last half century. The downward trend also appears elsewhere in B & I during thesame period, and losses of this widespread but rather thinly scattered violet appear particularlyobvious at its previous inland sites across these islands (M.Y.J. Foley & M.S. Porter, in: Prestonet al. 2002). Nevertheless, RHN and I believe V. canina is probably under-recorded inFermanagh, at least to some extent, perhaps on occasions being mistaken for V. riviniana (CommonDog-violet) (Rich & Woodruff 1992). We suggest that modern botanists, including ourselves, have notsufficiently searched likely inland habitats for this violet, whose flowers are a fine clear blue with awhitish or pale yellow spur.
Irish turlough occurrences
One of the most typical habitats of V. canina in Ireland is in an open, well illuminated zone onthe upper shore levels of turloughs or vanishing lakes. These grassy or rocky hollows in limestone areasare intermittently flooded after heavy rainfall and they drain vertically into cave systems (Praeger1932). Almost a third of the Fermanagh records of V. canina (five of the 17) are from threeturlough sites in the Ely Lodge area, NW of Enniskillen. Unfortunately, in the last ten or 15 years,despite their listing as ASSI conservation sites, these limestone hollows have become overgrown bytaller,
herbs and dense, turf-forming, strongly competitive graminoid species, so that active survival of a shadeintolerant violet like V. canina has become improbable or impossible. However, V. caninaseed can persist in the soil for many years (Thompson et al. 1997) and the plant populations maytherefore recover at a later date if the ground is cleared sufficiently and the vegetation reopened.More active management is required to conserve the floral biodiversity of the unique turlough habitat.
Ecological overlap and hybrids
V. canina typically occupies an open, well illuminated zone on turlough shores about a metre or soabove the much rarer and more ecologically restricted V. persicifolia (Fen Violet), or inIreland, 'Turlough Violet', which, as the latter name indicates, is another characteristic species ofthis unique habitat. In the intermediate zone where these two violets meet, hybrids are sometimes foundin some abundance (Praeger 1932; Webb & Scannell 1983). Hybrids are also quite commonly found whenV. canina and V. riviniana meet in grasslands or heaths.
Other local sites: Elsewhere in Fermanagh, apart from the turloughs, V. canina has beenrecorded, although very rarely, on the shingle or rocky shores of most of the larger lowland lakes.Praeger's 1900 record from the village of Garrison (no habitat details given) is possibly from the rockyor stony riverbank. However, even this record (mention of which appears only in his monumental book,familiarly referred to by Irish field botanists by the initials ITB (Praeger 1901c)), might welloriginate from the nearby shore of Lough Melvin, since Praeger rowed to the Trolliuseuropaeus (Globeflower) site on the shore of the lough from Garrison on 27 June 1900, after anovernight stay in the village (Praeger 1901a).
Fossil record
Although the genus Viola is not well represented in the fossil pollen record for obvious reasons,its seeds are commonly met in such deposits, although distinguishing the individual species is anothermatter. Godwin (1975) lists records for V. canina from glacial deposits at Kirkmichael, the Isleof Man dated to 10,000 BP (zone I/II transition and zone II) and there are tentative earlier recordsfrom the Lea Valley arctic beds in Essex, plus finds from Bronze and Iron age sites in Britain. Thesmall, scattered and, now with the prospect of global warming, increasingly vulnerable populations ofthis violet are thus very probably relicts of long past, possibly early post-glacial, more widespreadopen habitats with unleached, nutrient-rich soil conditions (Pigott & Walters 1954). Thesepopulations have managed to survive locally in pockets of specialized, relatively uncompetitive growingconditions, despite massive long-term changes in climate, vegetation and soils (Corner 1989).
Clearly this is a species worthy of conservation. The current population status and behaviour, withrespect to land use in its inland sites in our area, is worthy of further more intensive investigation,as indeed is also the case elsewhere in these islands, at least in inland sites.
European occurrence
The wider distribution of V. canina subsp. canina extends overmuch of boreal and temperate Europe stretching from Iceland and N Fennoscandia eastwards to middlelatitudes in Siberia and southwards into the northern half of Italy, Corsica, Sardinia and extendingeastwards at these latitudes to the shores of the Caspian Sea (Hultén & Fries 1986, Map 1327).
Threats
The Fermanagh turloughs are now ASSIs, as is much of the shore of Upper Lough Erne, but in recent yearsdisputes with landowners and lack of adequate grazing regimes have resulted in several of theseimportant sites becoming rather overgrown with taller vegetation. I am seriously concerned that rare orrelatively scarce species like V. canina and V. persicifolia will be, or have alreadybeen, ousted by more vigorous competitors.
Native, very rare or locally extinct.
1919; Praeger, R.Ll.; The Green Loughs, near Fardrum.
June.
This hybrid is closer to V. canina (Heath Dog-violet) in flowers and to V. persicifolia(Fen Violet) in habit, and it is sterile and very vigorous. The hybrid is said to be characterised byoften having shrivelled brown corollas on both normal open, sexual flowers and closed, cleistogamousones and it also displays a decurved spur of intermediate length (Stace et al. 2015). The leavesare longer and the stipules are larger than those of the more common parent (V. canina). Being ahybrid between two such rare parents, unsurprisingly it is also very rare and seldom found, there beingonly a few known sites in B & I.
Fermanagh occurrence
There are only four rather old local records from three sites in Fermanagh. The parent species overlap oncalcareous lakeshores and turloughs in the VC and locally these conditions occur around Lough Erne andat Fardrum, near the Ely Lodge estate to the NW of Enniskillen.
Apart from the first record above, the remaining details are: Green Lough turloughs, near Fardrum,refound here on 20 June 1985 by R.S. Weyl; Corrard Peninsula and Coolbeg Td, both latter sites being onthe seasonally flooded shores of Upper Lough Erne, where the hybrid was found by Meikle and co-workers,1946-53 (no specific years listed).
Although there are records of this rare hybrid violet from three Fermanagh tetrads, it has only beenrecorded once in the last half century and therefore it may be locally extinct. Despite theirconservation status as ASSI sites, the turloughs in the Fardrum area are currently very overgrown withtall invasive vegetation of sedges, grasses and herbs. The previously open area of the turlough basin isfast being reduced by scrub colonising from the marginal hedgerows. The re-excavation of these turloughswill be essential if the rare, low-growing violets, the hybrid and its parents, are ever to beseen here again.
Irish occurrence
In addition to the three Fermanagh sites, elsewhere in Ireland V. ×ritschliana has in the past been recorded in only five hectads scattered around turloughs andlimestone lakes in Connemara and the Burren (H9 & H14) (Webb & Scannell 1983; Stace etal. 2015). The map in Porter & Foley (2017) indicates that the hybrid was recorded in fourhectads in Connemara and the Burren post-1987, all other records being previous to this date.
British occurrence
In England, V. × ritschliana has been rarely recorded at a very fewsites in Oxfordshire, Cambridgeshire and Huntingdonshire (VCs 23, 29 & 31).
European occurrence
On the continent, this hybrid is said to be widespread in C & N Europe as far north as southernSweden (Porter & Foley 2017).
Threats
The turlough habitats are protected by ASSI status, as is much of the shoreline of Upper Lough Erne.Regular monitoring and active management to open up the vegetation and reduce encroachment by moreaggressive species is absolutely essential to maintain biodiversity.
Native, very rare, declining and possibly locally extinct. Eurosiberian temperate.
June 1919; Steele, Rev. W.B.; Fardrum Green Loughs, NW of Enniskillen.
May to July.
Growth form and preferred habitats
This small, creeping, perennial is one of the rarest and most vulnerable species in Fermanagh. It is aplant of seasonally wet, base-rich soils in fens and unimproved water meadows on lake shores overlyingfen peat. It avoids permanently wet or waterlogged ground. It is a Red Data Book species in bothIreland and England (Irish Red Data Book; Wiggington 1999). It is so very rarely found that wefear it is teetering on the verge of local extinction, at least at its oldest station in Fermanagh – theFardrum turloughs or 'Green Lakes'. As far as we are aware, V. persicifolia has not been seen atany of our local turloughs since 1992. On a visit in 2004, albeit at the end of August, one of theFardrum turloughs visited was completely covered in tall, rank grassy vegetation, which leaves nophysical or ecological space for a weakly competitive, low-growing herb like V. persicifolia.
Fermanagh occurrence
There are a total of 15 records for the species in the Fermanagh Flora Database, nine of which datebetween 1919 and 1957. The species tetrad map shows that apart from their date range (six are pre-1975and four of more recent date), the record distribution falls into two groups. These comprise theoriginal stations at the Fardrum turloughs, near Lower Lough Erne, and secondly, and considerably morewidespread, at nine sites on rocky limestone grasslands around the middle section of Upper Lough Erne,an area subject to seasonal flooding. The site names and record dates around Upper Lough Erne are asfollows: Inish Rath Island, 1953; Gubdarragh on the southern extremity of the Corrard peninsula, 1953& 1954; promontory ENE of Tiraroe Jetty, 1953; Coolbeg Td, 1954; Corraslough Point, 19 July 1957;Inishroosk, 21 June 1985; Inishbeg shore, Corrard peninsula, 27 July 1986; Aghinish Island, 16 June1987; Trannish Island, 16 June 1987. Most of the 1950s records were made by MCM & D (RevisedTypescript Flora) and the more recent records by members of the EHS Habitat Survey Team.
At the series of turloughs in the townland of Fardrum, Steele (1919) described the plant as occurring,"in great profusion around the edges of the little loughs", but on a revisit 28 years later in1947 he found it, "very sparingly". Richard Weyl rediscovered the species at the Fardrum sitein June 1985 and RHN recorded it again there on 30 May 1992.
English occurrence
Populations of Fen Violet in Cambridgeshire consistently observed for many years have shown that seed ofthe species is long persistent. The plant reappeared at sites following vegetation clearance andsoil disturbance after periods of five decades of apparent absence (Rowell et al.1982; Rowell 1983). Studies in England also indicate a strong natural cyclical tendency in terms ofviolet population numbers (Croft 2000).
In England, V. persicifolia was previously known from more than 20 sites in fens and rivervalleys, mostly in SE England, but nowadays it seems to have been lost from Norfolk (VCs 27, 28),Suffolk (VCs 25, 26), Yorkshire (VCs 61-65), Lincolnshire (VCs 53, 54) and Nottinghamshire (VCs 56). Bythe early 1990s its only surviving sites appeared to be in Huntingdonshire at Woodwalten Fen (VC 31) andin Cambridgeshire at Wicken Fen (VC 29), but since 1997 it has revived at another site, Otmoor inOxfordshire (VC 23), where after decades of apparent absence, it is now the strongest English population(Porter & Foley 2017).
European and world occurrence
In Europe, Fen Violet is widely distributed in cool temperate regions from southern Scandinavia to NSpain and eastwards to W Russia. It is absent from the hotter and drier regions including theMediterranean basin and the south-east. Its distribution in Asia is local and is only poorly known as itbecomes confused with V. pumila Chaix (Hultén & Fries 1986, Map 1329).
Threats
Excessive nutrient enrichment and/or inappropriate drainage or disturbance (either too little or toomuch) may destroy suitable habitat or encourage aggressive competitors.
Native, common and locally abundant. European boreo-temperate, but also present in eastern N America.
1806; Scott, Prof. R.; Co Fermanagh.
March to November.
Growth form and preferred habitats
V. palustris is the only violet in B & I that produces long, slender, creeping rhizomes thatbear rather large, broad, almost circular, cordate or reniform, hairless stem leaves in rosettes of 1-4on very long petioles (stalks). The leaves start pale green and darken as they age and they continuouslyenlarge throughout the growing season, from around 2.0-4.0 cm across during spring flowering up to 7 cmor more later in the summer. They are hairless (glabrous) and the leaf margin bears very shallow, bluntteeth. The creeping stems also bear flowers on individual stalks that are pale blue, pale lilac orwhite, the lowest petal being streaked with purple veins (Porter & Foley 2017).
V. palustris occurs over a wide altitudinal range in wet, acidic soils in bogs, damp heaths,marshes and swampy woodland including alder and scrubby willow and birch fen-carr, especially besidelittle runnels and where the typical peaty or clayey substrate is flushed by groundwater springs. In wetwoods and scrub it thrives in partial shade. Marsh Violet is often found growing through spongy cushionsor carpets of Sphagnum moss and on account of its creeping mode of growth and single blossom perflower stalk, it cannot, or only very rarely, flowers in profusion.
In coastal regions of B & I it can sometimes be found in non-calcareous dune slacks. However, thepreferred wet, acid habitat conditions are most frequently met in northern and western parts of theseislands (New Atlas). Previously, V. palustris was also a common component of species-richfen meadows, but it has declined in this particular habitat during the last 50 years, particularly in S& E England and also in C Ireland where it was never very common. The decline in the species followsmajor changes in land use, including long-term peat cutting for fuel on bogland, plus drainage ofmarshes and wet fields and the ploughing and reseeding of meadows as part of agriculturalintensification on lower slopes and hollows taking place over the last 50 years or so.
Studies using American species have shown that predation (ie herbivory and seed destruction) andinsufficient soil moisture, tend to be the most frequent causes of death in violet populations. However,long persistence in the soil seed bank is a characteristic feature of this genus and is a verysuccessful adaptation to uncertain environments, permitting re-emergence of the species if and when moresuitable growing conditions return (Cook 1979, 1980, p. 126).
Fermanagh occurrence
Locally, V. palustris is common and has been recorded scattered across 152 Fermanagh tetrads,28.8% of the VC total. It is widespread in peaty areas throughout Fermanagh, but is especially commonaround the southern half of Upper Lough Erne and on the slopes and summit plateau of Cuilcagh mountain.
Flowering reproduction
The small spring flowers, which appear from April through to June or July, open in the normal manner forcross-pollination by insect visitors, although they produce no scent to attract them (Hutchinson 1972).Like other species in subgenus Viola, V. palustris also produces small, closed, bud-like,self-pollinated cleistogamous flowers later in summer and early autumn. Both flower types shed seeds,but unlike most members of the genus they either do not bear elaiosome food bodies (or only very smallones), since ants do not frequent boggy ground (Clapham et al. 1962; Jonsell et al. 2010).
V. palustris has also become regionally endangered in some other European countries. Recentexperiments in an abandoned fen meadow in NW Germany showed that V. palustris recruitment andgrowth in damp fen-grassland conditions is promoted by a regular disturbance regime, ie by moderatedegrees of grazing or cutting. Sinker et al. (1985) had also noted that the species is fairlytolerant of grazing and trampling in the Shropshire region (VC 40). This form of management preventssuccessional changes which lead to an increase in standing crop (ie the development of rank, dense,taller vegetation) and the development of a deeper litter layer clothing the soil surface. Both thesefactors can oust V. palustris through suppression of seedling recruitment and excessivecompetition for light and other survival essentials (Jensen & Meyer 2001).
Vegetative reproduction
It should be remembered that many perennial violets including V. palustris also reproduceeffectively by vegetative growth and spread of above-ground stolons, or of the subterranean rhizome.This provides perennial violets with a pattern of growth and increase which Bell & Tomlinson (1980)recognised as "opportunistic". Horizontal stem growth is indeterminate in terms of stolonnumbers, distance (ie stolon length) and direction.
European occurrence
Beyond our shores, elsewhere in Europe, V. palustris is widespread in northern temperate latitudesfrom S Greenland to Scandinavia and it stretches southwards to Spain and Portugal, N Italy and Bosnia,plus the mountains of Morocco and the Azores (Hultén & Fries 1986, Map 1333).
World occurrence
It is present in N Africa and eastern N America, although it is still unclear whether or not the plantpresent in the latter is V. palustris (Hultén 1958, Map 103), or the closely related (and alsoAmphi-Atlantic) V. epipsilla Ledeb., of which there are two subspecies (see Hultén 1958, Map 104and Hultén & Fries 1986, Map 1334 and p. 1093). If we combine the ranges of these two species toform V. palustris sens. lat., the combination would then be almost circumpolar (for a discussionof the problem, see Hultén 1958, p. 122).
Name
The Latin specific epithet 'palustris' means 'growing in swampy places' (Gilbert-Carter 1964).
Threats
Habitat changes favouring secondary succession, leading to competitive exclusion or burial by plantlitter or overgrowth.
Native, rare and probably declining. European temperate, but widely naturalised including in N Americaand New Zealand.
1881-2; Barrington, R.M.; Gubbaroe Point, Lower Lough Erne.
April to October.
Growth form and preferred habitats
This attractive little annual violet is a casual ruderal species of disturbed, sparsely vegetated, dry orwell-drained, light, sandy acid or neutral ground in pastures, banks, quarries and stony or sandylakeshores. Unlike the closely related and rather similar V. arvensis (Field Pansy), it avoidscalcareous soils. Although very variable, V. tricolor is recognised by its deeply lobed stipules,the mid-lobe narrowly elliptic to oblanceolate, ± flat-faced flowers (several produced per stem betweenApril and September), with individual petals not more than 12 mm long. The typical blossom is a mix ofviolet, blue, cream and yellow colours, some yellow usually being present. Plants have erect, oftenbranched stems from a short (or sometimes absent) rhizome and the habit ranges from lax and sprawling tocompact (Parnell & Curtis 2012; Porter & Foley 2017).
Elsewhere in B & I, Wild Pansy is regarded primarily as a native herb of dunes, acid heaths andhillsides, all presumably well-drained, mainly open, grassland habitats.
Variation
The species aggregate V. tricolor is very variable and includes perennial forms. Petal size andcolour vary most conspicuously with development and with season. Petals are small, relatively wide andwhitish or pale coloured at the beginning of the flowering season, but they almost double in size,elongate and become increasingly intense in colour over several days until the flower is fullydeveloped. Later in the season, individual plants frequently produce smaller, narrower and lesscolourful flowers than those of early summer. As the species name suggests, flowers are often two orthree coloured from a shade palette ranging from violet and sky-blue to cream and yellow. Althoughentirely violet flowers are also quite frequent, pure creamy-white or completely pale yellow ones aremuch rarer. Studies show there is not any obvious geographical or ecological pattern to the flowervariation (Jonsell et al. 2010).
Variation in plant habit is more significant than in the flower in this case. Coastal plants and thosefrom sandy soils are usually perennial, rather than the annuals found elsewhere. When sand blows overthe lower stems of the perennials, they root and produce much-branched subterranean or procumbent growththat can eventually develop large clonal colonies. This form of the plant, which maintains its featuresin cultivation, is widespread around the coasts of Scotland, Wales & Ireland, but much moreoccasional and scattered along English coasts. It is described by Jonsell et al. (2010) andPorter & Foley (2017) as a sandy ecotype, although previously it was very often recognised as subsp.curtisii. Indeed, examination of the BSBI Big Database (accessed 2 January 2020), shows itremains a subspecies in that forum. What some regarded as 'true' subsp. curtsii (E. Forst.) Symeis now considered very much rarer and it may indeed be endemic to its original English station, BrauntonBurrows in N. Devon (VC 4), and to some sandy heaths in Breckland, East Anglia (VC 26 & 28) (Porter& Foley 2017). The variation pattern here and elsewhere over the whole species range needs furtherstudy to clarify the taxonomy of these forms.
Hybrids
Three hybrids involving V. tricolor are known, one of which is extremely rare in Britainand has not been recorded in Ireland at all (V. lutea Huds. × V. tricolor L.). Morecommonly, V. tricolor hybridises with V. arvensis Murray to form V. × contempta Jord., but while this is widely recorded, if scatteredacross much of Britain, it has only been found once in midland Ireland. The third hybrid is theextremely variable Garden Pansy (V. × wittrockiana Gams ex Kappert),which is the product of three violet species, V. lutea, V. tricolor and V. altaicaKer Gawl. This has over 400 named garden varieties and is very frequent and widely recorded in Britain,but much less so in Ireland, rare in the north of the island and never recorded in Fermanagh (Staceet al. 2015).
Status in Britain
V. tricolor is regarded as indigenous in Britain on the basis of the fossil evidence of a longpre-agricultural presence (Godwin 1975, p. 138). Changed land use, intensive agriculture from the 1950sonwards, herbicides and, locally in Fermanagh, a major decline in arable cultivation, have produced awidespread decline of V. tricolor during the 40 years between the two BSBI atlases (Walters &Perring 1962; Preston et al. 2002). This decline is most marked in SE England but has happenedthroughout B & I. As a result, V. tricolor has nowadays been pushed into an increasinglyrestricted, occasional, ruderal role, as an early colonist of open vegetation associated with disturbedsituations in artificial man-made habitats. These include cultivated ground, thin patchy garden lawns,waste places and other temporary, dry, disturbed growing conditions near habitation (M.S. Porter &M.J.Y. Foley, in: Preston et al. 2002).
In Ireland, V. tricolor subsp. tricolor has always been chiefly aplant of northern regions, associated with arable cultivation (An Irish Flora 1996). It remainswidespread but rare and declining in this role, principally found nowadays around the Lough Neagh basin(M.S. Porter & M.J.Y. Foley, in: Preston et al. 2002; Day & Hackney 2004). Like otherViola species, it seeds prolifically and it can survive dormant in the soil seed bank forprolonged periods, perhaps for many decades. Thus it cannot be written off as locally extinct until avery long time after its last sighting (Thompson et al. 1997).
Fermanagh occurrence
While V. tricolor subsp. tricolor has been recorded in 17 Fermanagh tetrads, 3.2% of thosein the VC, the only place in the county where we can rely on regularly finding it is in the disused sandpit at Pubble near Tempo. Twelve of the 27 records in the Fermanagh Flora Database are pre-1975 and,apart from the Pubble sand pit, since 1975 it has only been seen at ten other disturbed or urban sitesthinly scattered around the VC. It is, however, still perfectly capable of reappearing from the soilseed bank, whenever and wherever growing conditions prove favourable.
European and world occurrence
In continental Europe, V. tricolor s.l. occurs from Iceland and Scandinavia southwards tothe northern shores of the Mediterranean and eastwards through Turkey to the Ural and Caucasusmountains. In Scandinavia, V. tricolor is regarded as native or an archaeophyte in the south andwest, but in the north and east it is considered a neophyte and often is merely casual in its occurrence(Jonsell et al. 2010). It also occurs in the mountains of NW Africa and has spread withcultivation and settlement to become naturalised in parts of SW Asia (including India and thePhilippines), N America (mainly in eastern states) and in New Zealand (Hultén & Fries 1986, Map1337).
Threats
Changing land use and agricultural intensification.
Native, very rare and almost certainly extinct, but also possibly a mis-identification. Suboceanictemperate.
1884; Barrington, R.M.; Tully South Td, SW of Lisnaskea, Upper Lough Erne shore.
There is only one 1884 record for this mainly coastal subspecies and it may have been amisidentification, although Barrington says of his record, "This species [sic] has been named byProfessor Babington.", proving that he had a verified voucher specimen. We do not know what becameof this specimen, however.
Circ*mstantial support for this inland record of a normally maritime species comes from several othersimilar occurrences in Fermanagh, including Carex distans (Distant Sedge), Plantagomaritima (Sea Plantain), Asplenium marinum (Sea Spleenwort) and Bolboschoenusmaritimus (Sea Club-rush).
Additional support comes from the fact that V. tricolor subsp.curtisii also occurs around Lough Neagh with an assemblage of otherwise almost exclusivelymaritime species (FNEI 2; Flora of Lough Neagh). Nevertheless, and despite this, in theRevised Typescript Flora, R.D. Meikle wrote of it, "... should probably be referred to V.tricolor, true V. curtisii has not been seen in Co Fermanagh".
Native, extremely rare and almost certainly locally extinct.
1949; MCM & D; sandy fields below Gortaree, Slieve Rushen.
There is just the above, solitary, old Fermanagh record for this partially fertile violet hybrid. V.tricolor (Wild Pansy) occurs mainly on acidic and V. arvensis (Field Pansy) mainly onbasic soils, so that they very rarely meet. In England, the hybrid generally occurs as one would expectalong with its parents, but occasionally in the absence of one or both. Although intermediate incharacter, the hybrid is not easily distinguished from some variants of V. arvensis. The plantscan really only be separated by their degree of fertility. The hybrid can often occur as a single plantamongst a population of V. arvensis in arable fields and on other cultivated land and wasteground (D.H. Valentine, in: Stace (ed.) 1975; Stace et al. 2015). However, the parent species arenow even more rare and of casual occurrence in inland Ireland than they were before, so that theirhybrid is extremely unlikely to be refound in landlocked Fermanagh.
Although Stace (ed.) (1975) lists both Fermanagh and Down (H33 & H38), as Irish VCs with records ofthis rare hybrid, only the Fermanagh record is known in detail (see FNEI 3, p. 146) and it aloneis plotted in the New Atlas. It is not even mentioned in the Vascular Plant Register forCo Down (Day & Hackney 2004).
Introduced, archaeophyte, a very rare casual of arable cultivation. Eurosiberian temperate, but verywidely naturalised in both hemispheres.
1900; Praeger, R.Ll.; Co Fermanagh.
August to September.
Growth form and preferred habitats
A winter or summer annual weed with a slender taproot, V. arvensis differs from V. tricolor(Wild Pansy) in its smaller flowers (6-15 mm across), which have petals that are very slightly shorterthan the sepals and are mostly creamy-white with a yellow eye, variably streaked with purple or orange.The habit of V. arvensis is sprawling, but shrubby and, in comparison with V. tricolor, itis more dense and more leafy in the upper part of the plant, having shorter internodes. The blossom iscup-shaped, not flat-faced as in V. tricolor and the small, white to pale cream, sometimesflushed with blue flowers have sepals that are longer than the petals (Webb et al. 1996; Porter& Foley 2017).
However, despite its rarity in our survey area, V. arvensis seed is capable of surviving in soilfor up to 400 years (Cook 1980, p. 114), so that this little weed could still be found as a casual onany piece of open, cultivated or frequently disturbed waste ground, well-drained, light, sandy orgravelly ground, including in gardens, parks, sand-pits and quarries and the plant should be activelylooked for in these situations in Fermanagh. In the disturbed habitats to which it appears restricted,V. arvensis shows a definite preference for base- and nutrient-rich calcareous or neutralconditions in comparison with less fertile, moderately acid soils. This explains why it is only everreally frequent and abundant on farmland.
Fermanagh occurrence
With a total of just six Fermanagh records, V. arvensis was never more than an occasional speciesand is now very rare in the VC. The Fermanagh Flora Database has only three post-1975 records that arewidely spaced in both space and date, each observation representing just a handful of individual plants.
Additional to the first record given above, the other five Fermanagh records in the Fermanagh FloraDatabase are: sandy fields below Gortaree, Slieve Rushin, 1949, MCM & D; fields above Corragh Lough,S of Lisnaskea, 1951, MCM & D; Kilmore South, Derrychaan shore, Upper Lough Erne, 6 August 1986,S.J. Leach & A.S. McMullin; by border bridge, Muckle Rocks, near County Bridge, 13 August 1995, RHN;waste ground N of road near Rossharbour, Bigwood Td, N shore of Lower Lough Erne, 29 September 2001,RHN.
British and Irish status
Previous Floras of B & I regarded V. arvensis as indigenous (An Irish Flora; NewFlora of the BI), but the New Atlas is the first document which recognises Field Pansy asan 'archaeophyte', an ancient (pre-1500 AD) accidentally introduced weed of cultivation (M.J.Y. Foley& M.S. Porter, in: Preston et al. 2002). The argument for reclassifying 157 previous nativespecies as probable archaeophytes was subsequently published (Preston et al. 2004). In view ofits very strong links with agriculture and with human disturbed sites, the correct status of V.arvensis is, very probably, as an archaeophyte and this now appears to be generally accepted(Porter & Foley 2017).
Irish occurrence
In Ireland, V. arvensis is largely a plant of the eastern half of the island, although Hackneyet al. (1992) found it occurred rather frequently as a weed of lowland arable fields and wasteground in the NE counties of Down, Antrim and Londonderry (H38, H39 & H40) (FNEI 3). On theother hand, around the Lough Neagh lowlands in the SE centre of these three VCs, Harron (1986) (Floraof Lough Neagh) found there were, "few records … widespread but sparinglydistributed".
The New Atlas map shows the species scattered throughout most of Ireland, but the spread of dateclasses clearly indicates a widespread decline to local extinction in most of W & C Ireland, a factwhich emphasises this violet's current easterly Irish distribution.
British occurrence
Field Pansy is very much more common and widespread in Britain than ever was the case in Ireland. V.arvensis is present throughout most of England, Wales, the Isle of Man and the Channel Isles onsuitable soils on lowland and at moderate altitude. It is also frequent in E Scotland as far north asthe Moray Firth. It is much less frequent in the west and far north of Scotland and on the offshoreScottish islands, where suitable soils and growing conditions are rare or absent due to the acidity ofthe rock structure and the widespread development of peat in the strongly oceanic climate of the region.
European and world occurrence
Examining the world map of the species and its frequency, this variable pansy certainly must haveoriginated in open habitats in SE Europe and adjacent parts of Asia. It has been widely spread withcultivation beyond this, for instance to N Scandinavia, where it is regarded as a casual archaeophyte(Jonsell et al. 2010) and to N & S Africa, N America (widespread, mainly present in easternand midland states), Iceland, Greenland, New Zealand and Tasmania (Hultén & Fries 1986, Map 1338).This spread of the species by agricultural man strongly supports the case for archaeophyte status in NWEurope, including in B & I.
Threats
None.
Introduced, neophyte, deliberately planted, rare, but under-recorded.
21 August 1986; McMullin, A.S., Corbett, P. & Phillips, J.C.L.; Inisherk Island, Crom Castle Estate.
August to October.
The leaves of White Poplar are very decorative, presenting strongly contrasting upper and lower surfaces− dark glossy green on the upper side and startlingly white-tomentose below. The tree or large shrub ismade all the more attractive since like the native Aspen (Populus tremula), the leaves veryreadily flutter in the breeze. This appearance sufficiently recommends the species for planting, and itis popular in parks and for amenity planting around public buildings and on roadside embankments, inboth urban and more rural settings. Less frequently, it is seen on the boundaries of larger privategardens, most probably on account of the deserved notoriety all poplars have for suckering abundantlyand for producing roots that seek out and block drains and sewer pipes. It grows best in damp to wetsituations, in heavy soil unlikely to dry out in summer, or in ground near water.
The New Atlas hectad map indicates that P. alba is very much more frequently planted andwidespread in lowland Britain than is the case in Ireland. Its Irish use is only occasional andscattered, and in Fermanagh P. alba appears only rarely, around large demesnes (eg on the CromCastle Estate), or in fairly recent landscape plantings in the grounds of public buildings. Examples ofthe latter occur at the Lakeland Forum and at Chanterhill, both in Enniskillen town, and again inamenity planting around Rosslea village. There are just seven records in six tetrads included in theFermanagh Flora Database and they all date between 1986 and 1990.
We are certain P. alba is being simply ignored by recorders on account of the species always beingdeliberately planted and of little or no interest to the naturalist since it fails to naturalise in thewild. Virtually all the White Poplar trees in B & I are female, the catkins appearing in Februarylong before the leaves emerge. Thus the trees fail to set seed.
The species is regarded in B & I as a fairly early tree introduction of unknown date, but most likelypost-1500. Very possibly it arrived with us from Holland (ie in status, it is a 'neophyte'). P.alba is widely distributed in temperate C & SE Europe and eastwards into C Asia. However,even within this range, in many areas it is an obvious planted introduction, so that the naturaldistribution of the species has become obscured (Meikle 1984; Brendell 1990).
Threats
None.
Deliberately planted introduction, very rare, possibly under-recorded.
15 August 1985; Leach, S.J., McMullin, A.S. & Wolfe-Murphy, S.A.; shore of Knockballymore Lough.
May to August.
This very vigorous hybrid, which originated in the continental Asian or European range of P. alba,is capable of producing viable seed which theoretically could allow mature specimens to successfullyback-cross with either parent species. There is little or no evidence that this is happening in B &I, however, or else it is an extremely rare event. Even the spontaneous hybridisation of the parentspecies appears very rare and thus the trees we see are deliberately planted nursery stock. As might beexpected, the foliage is intermediate between that of the parents; the leaves of long shoots (ie ofrapid growth) are large and lobed, but not deeply so and they are covered on the undersurface withgreyish down which readily rubs off. Leaves on short shoots are orbicular and quickly become almostglabrous.
The hybrid, like its parents, readily suckers and vegetative reproduction therefore appears to be itsdominant mode of increase. The vast majority of trees in B & I are male, females being regarded asvery rare (Meikle 1984).
Grey Poplar was introduced to B & I sometime around 1700 AD. Originally it was regarded as a timbertree on account of its rapid growth rate (Mitchell 1996, p. 278). However, foresters soon found that thehybrid proved difficult to propagate from cuttings, which is their normal method of stock production,and although it can grow on poor, sandy soils, it really requires better, more fertile,nutrient-rich, lowland soils than those usually available for timber plantations. Thus P. × canescens quickly fell out of favour for forestry purposes andnowadays it is planted purely for decorative amenity purposes, or in situations where rapid screening isrequired and the suckering drawback does not matter.
Fermanagh occurrence
There are only three sites in Fermanagh where this hybrid poplar has been recorded: two isolated stationsin the southern lowlands, plus one on a landed demesne. Apart from the first record listed above theother details are: Aghavea Td, 2 km E of Maguiresbridge, 30 May 1990, RHN; Templehill Wood, Belleisleestate, 27 June 1992, RHN. In common with other deliberately planted woody species, it is very probablyseriously under-recorded in our survey.
British and Irish occurrence
The New Atlas hectad map shows that elsewhere in Ireland this hybrid is only occasional, beingrather thinly and widely scattered across the island. In Britain, it is common mainly in SE England,with occasional occurrences in the remainder of England and Wales, becoming increasingly scarce andcoastal northwards to Inverness. The largest tree of this hybrid in B & I (ie 'the Champion tree'),is on the Birr Castle estate, Co Offaly; its height in 1999 measured 42 m (Johnson 2003).
Threats
None.
Native, occasional, but also deliberately planted. Eurasian boreo-temperate.
1882; Barrington, R.M.; Lower Lough Erne.
April to February.
The fluttering leaves of this potentially large species moving in the slightest breeze make it such anattractive decorative feature that Aspen is often chosen as a specimen tree in demesnes, parks andlarger gardens (thus avoiding trouble in urban areas with fluffy windblown seed). It is one of the lasttrees to break its buds in the spring, usually in late April after the oak (Hatfield 1957). It growsbest when positioned near open water features, but must be sited well away from piped drains, which theroots have a tendency to invade and block. Roots and suckers can also affect building foundations bycausing subsidence on predominantly clay soils. These organs can travel up to 30 metres, although 90% ofdamage is caused by trees growing within 20 m (Thomas 2000).
P. tremula is usually found in seasonally wet or damp, low-lying ground near lakes and rivers,occasionally in or on the margins of old or long-established woods or bogs, or in hedgerows where it maysucker along and further spread by self-seeding. Seed is liberated from the capsules of all fertilePoplars before the leaves are fully open, and it must settle on moist earth at once or it quicklyperishes. Thus the species does not grow naturally in dry, infertile, stony or sandy soils, or underwaterlogged conditions, or in shade (Hatfield 1957). Although this tree is indigenous it is oftenobviously planted in parks, gardens and along roadsides. On account of the unknown extent of pastplanting regimes P. tremula is definitely native only on the more remote parts of rockylakeshores, uninhabited, small, lake islands, in gullies in mountain glens and on cliff faces.
Fermanagh occurrence
Around Fermanagh, Aspen is a frequent and widespread tree, or else a large, suckering shrub, having beenrecorded in 118 tetrads, 22.4% of those in the VC. P. tremula is the only native poplar speciesin Fermanagh.
Biology and ecology
Like its close relatives the willows, P. tremula is dioecious, the catkins on both male and femaletrees being pendulous. Likewise, it is not a long-lived species, perhaps surviving 50 years. Also, itreadily forms hybrids with introduced members of the genus (Brendell 1990).
P. tremula is not regarded as being very competitive, especially with regard to light and shade.It is capable of rapid growth given full illumination, but
in deep shade it even fails to sucker let alone flower and fruit (Rackham 1980).
As a native species it primarily behaves as a pioneer colonist of seasonally wet or damp, open ground,which frequently involves unstabilised soil in upland situations, eg among boulders on scree, or asisolated individuals in cliff crevices and on ledges (Brendell 1990; Jonsell et al. 2000).Although Poplars have plumed seed that can travel long distances by wind, most P. tremulaexpansion occurs by vigorous suckering to form often dense, shrubby thickets (Milner 1992; Thomas 2000).On the other hand, seed and seedling mortality, even on bare soils, is described as very high (Jonsellet al. 2000). Some of these seedling losses will be due to an unfavourable physical environment,but predation is also highly significant (Milner 1992). In Fermanagh, apart from occasional treesgrowing on scarps, there really is no definite current evidence of this colonising ability and P.tremula has not been recorded in the typical habitats available for invasion, eg in oldquarries, sand-pits, and damp disturbed waste ground.
British and Irish occurrence
The New Atlas hectad map shows Aspen is extremely widespread throughout Britain, being morefrequent in the N & W. It is recorded even on the exposed conditions of the Scottish Isles,including the Outer Hebrides (where it is confined to cliffs) and on Shetland (Pankhurst & Mullin1991; Preston et al. 2002). However, in Ireland the distribution is very much patchier. It isonly really consistently recorded in the north of Ireland, and there principally in VCs W Donegal,Fermanagh, Tyrone and Londonderry (H35, H33, H36 & H40).
European and world occurrence
The world distribution of P. tremula is the widest of any Poplar, stretching from Iceland throughEurope (except the S Iberian peninsula) to Greece and the Caucasus and across temperate Asia as far asChina and Japan (Jalas & Suominen 1976, Map 264). It reaches from montane woodlands in theMediterranean to the limit of tree growth in the Arctic tundra (Hatfield 1957; Meikle 1984). In Asia, itis largely represented by subsp. davidiana (Dode) Hultén, although the two forms (subsp.tremula & subsp. davidiana) overlap (Hultén & Fries 1986, Map 617; Jonsell etal.2000).
Threats
None.
Probably planted, rare, but at least some identifications need verification. Circumpolar boreo-temperate.
1912; Druce, Dr G.C.; Co Fermanagh.
June to October.
Growth form and preferred habitats
Black Poplar is an uncommon, large, rough-barked, dioecious, deciduous tree of lowland hedges on floodplains along rivers and around lakes. Previously, Black Poplar was considered indigenous only in SEngland and E Wales, although even here the tree has also been regularly and widely planted as anamenity or specimen tree in urban locations, since it is very tolerant of industrial pollution(Milne-Redhead 1990; Cottrell et al. 1997). The previous regular plantation of Black Poplar,which was highly prized for its timber, appears to have been superseded in B & I by the introductionof hybrid trees as early as the end of the 18th century (Hobson 1993).
Recently, Black Poplar was recognised as a native tree in Britain under imminent threat of extinction dueto the old age of many specimens, the general lack of new plantations for very many years and the factthat planting female trees had been avoided due to the local mess made by the dispersal of their fluffyseed. The realisation of the tree's rarity then stimulated surveys which have now been carried out inboth Britain (Milne-Redhead 1990) and in parts of the RoI by Hobson (1991, 1993). The results of thesurvey in Britain recognised that the majority of trees were derived from planted cuttings, frequentlyresulting in local clones of a single sex. Very few locations are known in Britain where the two sexesco-exist and therefore the vast majority of reproduction is vegetative, either by root suckers or byfragmentation and re-rooting. In the very rare locations where viable seed is formed, seedlingsurvival requires a prolonged period of bare wet mud substrate, habitat conditions which are rarely metnowadays in the intensively managed lowland wet landscape of Britain. For these reasons, seedlings andsaplings are almost unknown in the British survey (see Milne-Redhead 1990, p. 3).
Fermanagh occurrence
Ten of the 13 Fermanagh records in eleven tetrads that exist for Black Poplar in the Fermanagh FloraDatabase are from the 1986-7 DOENI Upper Lough Erne survey, but probably all of the local recordsof this taxon require verification. The Fermanagh trees are principally located on or near the CromCastle estate at the southern end of Upper Lough Erne and we are fairly confident that if they arecorrectly identified in the first instance, they would almost certainly be of planted origin in such asetting.
Irish occurrence and status
Hobson's survey in southern Ireland found that the tree was exclusively associated with river valleys andflood-plains, often in sites remote from farms and other habitation. The tree was widespread in thesesituations, which were not as expected, concentrated in 'the English Pale' around Dublin and theadjacent eastern counties. On this basis, Hobson (1991) suggested that Black Poplar might possibly be anative Irish tree. If this is so, it is interesting that young specimens not more than ten years oldwere also located during the survey. These probably indicate that some Irish farmers continue to plantlocal Black Poplars instead of using imported hybrid stock. The useful timber the tree produces helpsexplain its plantation in these cases (Cottrell et al. 1997). Hobson (1993) found a quiteextensive Irish population with a well-balanced age structure, which again suggests ecological stabilityhere. Furthermore, around Lough Ree and Lough Allen he discovered it is still naturally regenerating inthe wash zone of winter storms.
Threats
None.
Introduced, neophyte, deliberately planted, but very much over-looked and seriously under-recorded.
1989; Northridge, R.H.; Lakeland Forum, Enniskillen.
RHN and the current author tend not to bother recording obviously planted cultivars in urban settings,but there is a row of this distinctive, narrowly columnar fastigiate tree by the lake in the centre ofEnniskillen town that even we could not completely ignore. Shamefully we admit that they are the onlyrecord of this tree listed in the Fermanagh Flora Database.
Cultivars of fastigiate, upright growth forms, like the Lombardy-poplar, being of garden origin arealways single sex clones (Lombardy-poplars are entirely male). Female trees of similar shape in B &I are most likely hybrids with other forms of poplar, or else they represent another fastigiate cultivarof Black Poplar from the E Mediterranean, P. nigra var. afghanica Aitch. & Hemsl.(Meikle 1984). Being a strict, columnar growth form, Lombardy-poplars typically do not sucker andtherefore they fail to reproduce or naturalise themselves in the wild.
Again, as with most other members of the genus, this is not a long-lived tree. It is abundantly clearfrom the scanty records displayed over the whole of Ireland in the relevant New Atlas hectad map(which is a composite map aggregating records of all the fastigiate cultivars of Populus nigra),that other recorders on the island generally ignore the presence of these conspicuous landscaping trees.
Anyone wishing to learn more about the five fastigiate forms of Black Poplar is advised to start byconsulting the late Alan Mitchell's excellent account (Mitchell 1996, pp. 281-5).
Populus nigra × P. deltoides (P. × canadensis Moench), Hybrid Black-poplar
Introduced, neophyte, deliberately planted, rare but very probably under-recorded.
1912; Druce, Dr G.C.; Co Fermanagh.
June to October.
Meikle (1984) comments that named cultivar clones of this variable, fast-growing hybrid (all of which aresingle sex, propagated from cuttings and do not sucker) are the most commonly planted poplars in B &I. The history of this hybrid and some of its most frequently recorded clones is provided in summary byMitchell (1996) and Stace et al. (2015). Originally this hybrid was favoured over the nativeBlack Poplar (P. nigra subsp. betulifolia) because of its fastergrowth and a long trunk clear of burrs, giving good timber lengths. While previously clones of thishybrid were frequently planted, they eventually fell out of favour in forestry circles due to theirsusceptibility to both rust fungi and canker, plus the advent of even faster growing hybrid cultivarsbred in Belgium that are better suited to our climate and soils (Stace et al. 2015). Beingcrosses between poplars from two continents, the maintenance of rapid growth rates in Euro-Americanpoplars depends upon a combination of hot summers and moist soil. These conditions are not often foundin these islands except in low-lying parts of S England, a fact reflected in the New Atlas map(Brendell 1990; Mitchell 1996). These hybrid cultivar clones are perfectly hardy, however, and they willgrow, albeit more slowly, as far north as Inverness and even on Orkney. These hybrid clones have beenmainly planted as amenity trees in parkland and along linear habitats such as roadsides, railways,river-, stream- and canal-banks and in hedges. They have also been planted for screens, shelterbelts andas timber trees in plantations (Stace et al. 2015).
In Fermanagh, we have just eleven records for this hybrid in ten tetrads. Apart from Druce's originalrecord above, for which there is no site whatsoever, all the remainder are either from the islands orthe shores of Lower Lough Erne (LLE). Ignoring Druce's inadequate record, four of the others were madeby Matthew Tickner (MT) surveying the lake islands for the RSPB, while the remainder were made by RHNand RSF. We believe that this distinctive tree is greatly under-recorded, and further work is requiredto determine this question. The record details are: Crannoges Island, LLE, 1989, MT; Hare Island, LLE,1989, MT; Inishmakill Island, LLE, 1989, MT; Lough Nawalsky, NW of Lough Scolban, 27 July 1990, RHN;Drummony Bridge, Bannagh River, 9 July 1991, RHN; Keenaghan Lough, Tievalough Td, 7 July 1991, RHN &RSF; Horse Island, Killadeas Bay, LLE, 1992, MT; disused quarry, Roosky, 17 September 1994, RHN;roadside, Mullaghmore Bridge, 19 October 1996, RHN; Old Castle Archdale estate, August 2001, RHN.
The New Atlas hectad map shows P. × canadensis is widespread,frequent and scattered throughout Ireland, but with a high proportion of older pre-1970 recordsunconfirmed.
Threats
None.
Introduced, neophyte, deliberately planted, a rare garden cultivar.
1947; MCM & D; roadside by cut-over bog, SW of Lisnaskea.
Again only a female clone is known of this taxon in B & I, a species or hybrid of obscure anddisputed origin. Stace et al. (2015) believe it is P. deltoides Marshall × P.balsamifera L. Whatever its parentage, it is supposed to have been introduced to these islandsfrom America about 1773 (Sell & Murrell 2018). P. × jackii (=P. candicans or P. × candicans) (Balm-of-Gilead) has a strongbalsamic smell similar to P. trichocarpa Torr. & Gray ex Hook. (Western Balsam-poplar), butit tends to produce suckers prolifically. Partly for this reason, but also because it suffers extremelybadly from a bacterial canker, this plant is no longer popular in gardens (Meikle 1984).
A huge-leaved, white, cream, pale green and pink variegated form of the plant, usually called P. × candicans 'Aurora' in the horticultural trade, is vastlyover-popular, however, and is very common in gardens throughout N Ireland. It too is extremelysusceptible to canker, and therefore does not survive very long (More & White 2003).
The three widely separated Fermanagh records of P. candicans in the Fermanagh Flora Database werefound planted in damp ground on roadsides near habitation. The details of the other two records are:Cargin Lough, 20 August 1986, L.W. Austin & A.S. McMullin; farm east of Melly's Rocks, near LoughNavar Forest Park, 14 July 2001, RHN.
Threats
None.
Native, common and widespread. Eurosiberian boreo-temperate, but widely planted and naturalised beyondits native range, including in eastern N America.
1836; Mackay, J.T.; Florencecourt.
April to November.
Growth form and biology
The oval, glossy, bright or dark green, leathery leaves of this species, its young branchesreddish-brown, glossy and shining as if varnished and highly polished are very distinctive features, andonce identified are easily recognised in future. This large shrub or smallish dioecious tree deservesits English common name 'Bay Willow', since the resemblance to Laurus nobilis, the culinary BayTree, is real, even down to the fact that its twigs and crushed leaves are fragrant, although not spicylike the true culinary Bay. In more northerly regions (eg Scandinavia), the leaves make a suitablekitchen substitute for the less hardy Mediterranean leaf (Newsholme 1992).
Catkins are produced along with the leaves around late May or June, which is considerably later thanother willows. Both male and female catkins produce nectar and attract bees and other insects. Malecatkins also produce copious amounts of conspicuous yellow pollen which the insects transfer to thefemale catkins. Some wind pollination may also occur. Seeds are shed in late autumn or winter, and theygerminate in the spring after a period of overwinter dormancy (Jonsell et al. 2000).
Fermanagh occurrence and preferred habitats
Bay Willow is common and widespread in Fermanagh, being recorded in 192 tetrads, 36.4% of those in theVC. It is, in fact, the fourth most frequently found willow in the county after the three nativesallows, in their order of commonness, S. cinerea subsp.oleifolia (Rusty Willow), S. aurita (Eared Willow) and S. caprea (GoatWillow). However, it is not quite so widespread in the area as the introduced S. viminalis(Osier).
Bay Willow chiefly grows in a wide variety of damp to wet ground habitats, and it seems fairlyindifferent to soil reaction. It is perhaps slightly better grown on calcareous soils in the area,although it is definitely not more frequent on them. While Bay Willow can and does occur on acidic mosspeat, eg on the margins of raised bogs or on rather drier cut-over bog surfaces, in such situations itis often very obviously associated with flushing conditions associated with springs and surfacedrainage. Movement of water at the roots would certainly alleviate the otherwise very poor mineralnutrition in bogland sites. If anything, S. pentandra is rather more frequent on higher groundthan the other willows in the county.
Apart from mainly occupying ground near water like other willows do, S. pentandra is also found indrier situations along roadsides, and also quite frequently in neglected ground, eg in at least sixdisused quarries and sand-pits in Fermanagh. It also invades waste ground in several Fermanagh villagesand towns, as well as in or near a few rural churchyards.
Irish occurrence
The New Atlas hectad map shows that in Ireland S. pentandra occurs much more frequently inthe northern half of the island (especially throughout NI), becoming increasingly scarce and moreprobably introduced southwards to S Kerry (H1).
British occurrence
In Britain, S. pentandra is most frequent and is regarded as indigenous in middle latitudes, fromStaffordshire north to the Great Glen in Scotland. Beyond this section of the country to both north andsouth, it gradually becomes scarce, coastal and more obviously introduced (Preston et al. 2002).The absence of the species from much of N Scotland is very puzzling, especially seeing that it is sovery widespread in Scandinavia, where it can reach above the timberline as a shrub (Jalas & Suominen1976, Map 201; Jonsell et al. 2000, Map p. 123).
Since male plants bear large numbers of very beautiful, cylindrical, golden-yellow catkins in late May,they induce people to plant it purely for this ornamental feature. Despite its decorative use, thereappear to be no named cultivars (Griffiths 1994). As a result it is difficult or impossible to discernthe true native distribution of the species on these islands.
European and world occurrence
S. pentandra s.s. originally occurred in Europe and W Asia. Today, it is widespread in N & CEurope and western parts of Central Asia, although absent from the Iberian peninsula and almost all ofthe Mediterranean basin (Meikle 1984; Hultén & Fries 1986, Map 589). It is also introduced ineastern N America.
Hybrids
These occur with S. alba and S. fragilis and a triple hybrid involving all three species.All of the hybrids are of garden origin, although those with S. alba and the triple hybrid canalso occur spontaneously (Meikle 1984).
Threats
None.
Introduced, archaeophyte, deliberately planted, occasional. Eurosiberian temperate but widely naturalisedin N Europe, N India, N America and New Zealand.
1939; Praeger, R.Ll.; Enniskillen Town.
May to November.
Taxonomic difficulties
In the Revised Typescript Flora, Meikle et al. (1975) state that typical S. fragilisL. (sec. Sm.), that is the Smith 'type', var. fragilis, had not been seen by Meikle in Fermanagh.In his Willows and Poplars Handbook, where he enlarges upon the situation regarding this complexgroup of large, tree-willows in a B & I context, Meikle affirms that in Ireland S. fragilisvar. fragilis is generally replaced by var. russelliana, exclusively female and thecommonest and most widespread form of the four variants (var. fragilis (dioecious), var.furcata (male), var. russelliana (female) and var. decipiens (male)) of thisspecies in these islands (Meikle 1984, p. 28; Meikle 1992). The three single-sex variants, in Britain atleast, are individually so uniform that each appears to be represented by a single clone. Thus from thisargument, almost certainly the records in the Fermanagh Flora Database for S. fragilis refer toeither var. russelliana (most probable), or just possibly to the other form that Meikle reckonsoccurs in Ireland, var. decipiens (Hoffm.) W.D.J. Koch. The latter is part of the taxonrecognised by continental botanists as the 'type' for the species S. fragilis L.
Meikle (1992) cogently discussed this problem, listing the points where his opinion at that time differedfrom that of the continental taxonomists. In a recent Scandinavian critical Flora (Jonsell et al.2000), where the editors take a narrow Linnaean view of S. fragilis, it appears that three ofMeikle's four variants of S. fragilis (all except var. decipiens), fit into what theyrefer to as 'S. × rubens Schrank', albeit along with several other named forms andhybrids. S. fragilis var. decipiens W.D.J. Koch is now referred to by Stace (2010, 2019)as S. euxina I.V. Belyaeva, and the three remaining 'fragilis' variants, that previouslywere referred to as S. × rubens Schrank, have been transferred to S. × fragilis L.
This continental reworking might prove sensible here, since it greatly simplifies life and makes thethree single-sex Meikle S. fragilis variants redundant (which we believe are of horticulturalorigin anyway). This approach is encouraged by the fact that nowhere in B & I does S.fragilis s.s. nor any of the Meikle species variants occupy a naturally occurring plantcommunity. Rather, they are always, or almost always, found in obviously planted situations. Someproportion of the less obviously planted tree clusters might possibly be fertile hybrids with S.alba (White Willow), which can back-cross with S. fragilis and create additionalidentification problems.
The critical Flora of Great Britain and Ireland, 1 (Sell & Murrell 2018) takes theEuropean approach to S. fragilis (or S. × fragilis, S.× euxina), but lists numerous more splits and subdivisions. Three ofMeikle's four subspecies are listed as nothovars, and two new ones are added, nothovar.basfordiana (Scaling ex Salter) P.D. Sell, which is a hybrid between S. alba var.vitellina and S. euxina. Within this latter nothovar, two formas are named, formabasfordiana and forma sanguinea (Meikle) P.D. Sell. The fifth nothovar is rubens(Schrank) P.D. Sell, a hybrid between S. alba var. alba and euxina. Meikle's var.decipiens (Hoffm.) Koch becomes a separate species, S. euxina I.V. Belyaeva.
Growth form and preferred habitats
S. fragilis var. fragilis is a sturdy, round-headed tree with wide-spreading branches thatcan grow up to 15 m tall. Young twigs are brittle at their point of attachment to the older branch. Seebelow for the comparative growth form of the var. russelliana in a separate account.
All variants of S. fragilis tend to occur in linear collections or plantations by water, or indamp hedgerows near ditches, typically, and rather obviously, in low-lying situations. Occasionallytrees appear singly or in small groups in wet hollows in fields, where they may have self-sown, orrather since the most likely form of the tree we have in Fermanagh (Meikle's var. russelliana) isentirely female and does not set seed, where wind or animal broken and transported branches may haveself-rooted. Where clusters of the tree occur we presume the original planted individual has fallen overand layered itself, or else it has formed new plants through branches breaking off at joints andre-rooting, something they readily do. On the other hand, it is possible that some proportion of theseless obviously planted tree clusters may be hybrids with S. alba (White Willow).
The fragile nature of S. fragilis, ie the ease with which young side branches snap off at thejoint with the stem, is a much over-rated species distinguishing character (Howitt & Howitt 1990).This feature should not be relied upon to distinguish the tree, since it also applies to the other verycommon large tree willow, S. alba. The difference between these two taxa in terms of brittlebranches is simply a matter of degree.
Fermanagh occurrence
As the tetrad map shows, S. fragilis (ie probably largely, or almost entirely, Meikle's var.russelliana), has been recorded in 72 tetrads in Fermanagh, 13.6% of the total number in the VC.It is widely scattered throughout the county along riverbanks, lakeshores and roadways, but with adefinite concentration towards the SE of the area around the Upper Lough Erne basin.
Irish occurrence
Because of the confusion of S. fragilis var. fragilis with var. russelliana and var.decipiens and other forms and hybrids (especially that with S. alba), it is almostimpossible at present to map with any exactitude the distribution of S. fragilis var.fragilis in B & I (Meikle 1984). However, for what it is worth, the New Atlas hectadmap for S. fragilis in Ireland shows that it is widespread and scattered throughout, with themost consistent presence being in NI. In the light of what Meikle (1984) explains, very probably thismap really shows the occurrence of var. russelliana, rather than that of var. fragilis.
British occurrence
The same New Atlas map (with the same disclaimer applied), displays how very widespread the treeis in lowland Britain, the distribution however petering out in N & W Scotland.
Threats
None.
Archaeophyte, deliberately planted, rare, but possibly occasional, probably over-looked andunder-recorded.
1949; MCM & D; Ports Lough, Derrymacrow Td, N of Crom.
Inadequate data.
In the Revised Typescript Flora, Meikle et al. (1975) described this particularvariety of S. fragilis as being frequent in the SE quarter of Fermanagh, ie south of Upper LoughErne. However, only five of the total of ten records for this variety of the species made by Meikle andco-workers, could even remotely fit into this sector of the county as they subdivided it. Apart fromtheir ten records of var. decipiens dating from 1949-53, there is just one additional record madein 1993 by Ian McNeill in the NE of the county and two made by RHN and HJN on Inishmore Island, UpperLough Erne. Thus var. decipiens has been only rarely recorded in 13 tetrads and with just threerecords in the last 50 years, it is very probably being over-looked to some extent by local recorders.
In addition to the first record given above, the remaining record details are: between Rosscor and CastleCaldwell, 1950; Carnmore Lough, 1950; Lough Tawy, NW of Rosslea, 1950; Galloon Td, Upper Lough Erne,1951; bogs W of Clontymullan Td, near Arney, 1952; Arney River at Clontymullan Td, 1952; CorrardPeninsula, shore of Upper Lough Erne, 1953; Killyrover Td, SW of Maguiresbridge, 1953; roadside by LoughMelvin near Garrison, 1953; W of Imeroo, by the Tempo River, 7 August 1993, I. McNeill; shore of TullyTd, NE Inishmore Island, Upper Lough Erne, 21 May 1994, RHN & HJN; Drumhirk Td, Inishmore, 21 May1994, RHN & HJN.
S. fragilis var. decipiens forms a twiggy bush, usually only 5-7 m high. Meikle (1984)describes the year-old twigs as being pale ochre-coloured and distinctly lustrous, quite unlike anyother fragilis segregate. The twigs are brittle as in other variants of S. fragilis.Fresh, unripened shoots are often stained crimson. The leaves are shorter and broader than those of var.russelliana or var. fragilis, rarely exceeding 9 cm in length.
On account of the few Fermanagh records, S. fragilis var. decipiens appears very rare andthinly and widely scattered in wet or damp ground along riverbanks, lakeshores and roadsides. As withvar. russelliana, most of these trees will have been deliberately planted, but single or remoteclusters of trees may have established themselves, most probably through vegetative reproductioninvolving transported broken branchlets rooting and establishing themselves.
Introduction, archaeophyte, deliberately planted, very probably under-recorded and possibly occasional.
30 August 1967; Parker, R.E.; roadside near Lisnaskea.
There is only the solitary record listed above for this hybrid in the Fermanagh Flora Database. A voucherexists in BEL, but the identification needs to be expertly confirmed. It is likely that some ofthe plants that are listed as S. fragilis, but are growing in unlikely planting sites, may reallybe this hybrid, rather than the usual form of S. fragilis in our area, ie var.russelliana. The entirely female var. russelliana crosses with pollen from S. albato form this intermediate fertile hybrid. Back-crossing can occur with both parents, creating a hybridswarm of intermediates. Fortunately this development has never been suggested in our area, although morework is definitely required on the variation within Fermanagh tree willows.
Introduction, archaeophyte, deliberately planted, occasional. Eurosiberian southern-temperate, but widelynaturalised.
1900; Praeger, R.Ll.; Co Fermanagh.
January to November.
Growth form and preferred habitats
A tall, graceful tree, 10-25 m tall of varying crown shape which forms a distinctive and well establishedfeature of wet to damp, lowland, field, riverbank and roadside hedges around Upper Lough Erne and theLisnaskea area in particular. S. alba has a more upright crown and is clothed with moreattractive, silvery-white-coated foliage than S. fragilis (Crack Willow), yet it appears slightlyless frequently planted than the latter in Fermanagh. Long regarded as an alien introduction in Ireland,S. alba is always planted, or originates from planted material which may have reproduced andspread to some extent along or away from original, deliberately chosen, waterside sites.
Catkins appear with the leaves in late April or early May and after insect pollination the capsules ripenin July or early August (Meikle 1984). The only critical requirements for willow establishment are dampor wet ground for seed germination and seedling growth, and full sunlight. All willows require fullsunlight and they will not grow well without it, indeed, any shaded part of a plant soon dies.Otherwise, most Salix species are environmentally relatively undemanding, although S.pentandra (Bay Willow) and S. fragilis (Crack-willow) and their various forms do notperform well on calcareous soils and tend to avoid them. When planting willows, it should always beremembered that all of them age quickly and tend to become unsafe. An old willow wood always containsleaning and broken specimens, reducing the ornamental value of the stand (White 1992).
Apart from its ornamental value, S. alba was previously valued for basketry and undoubtedly one ormore cultivars of it will have been planted locally for this purpose. Until basket-making declinedsometime in the 1930s or earlier, some plantings of S. alba varieties would have been coppiced,or less likely in the area, pollarded at intervals to produce canes. There does not seem to be anysurviving evidence of this practice locally in Fermanagh, as far as we are aware, but while willows areextremely rapid in growth they are not long-lived trees (White 1992), and we therefore feel that thisabsence is not significant.
Fermanagh occurrence
Despite being of more attractive appearance, S. alba appears slightly less frequently planted thanS. fragliis (Crack-willow) in Fermanagh. The greatest difference in their occurrence is thatWhite Willow is very much more localised than Crack-willow, being almost entirely concentrated aroundUpper Lough Erne. S. alba is recorded from a total of 57 tetrads, 10.8% of those in Fermanagh,while the comparable figure for S. fragilis is 13.6%, although the latter is much more widelyscattered. Unfortunately we do not know to what extent S. alba is fully naturalised and capableof seeding itself in Fermanagh, but it does so rarely elsewhere in B & I. More local study isrequired.
Meikle (1984) describes three varieties that are widely planted in B & I, but we have no details ofthese in our survey.
Irish occurrence
The New Atlas map shows S. alba is widely distributed throughout Ireland, although thereare a large number of hectads plotted with only pre-1970 records, especially in counties on or close tothe western seaboard. These areas have a predominance of strongly acid, peaty soils of poor nutrientstatus and in coastal areas there is excessive wind exposure. Even if it were planted in the west, boththese factors would limit or exclude the growth of this normally large tree species, which naturallydemands reasonable substrate fertility and stability.
British occurrence
In Britain, until very recently, White Willow was another of those species, which despite a long historyof widespread plantation managed to retain native status, perhaps more from tradition than through anyrational scientific analysis (Stace 1997). After an examination of the status question, it is nowrecognised as an archaeophyte throughout B & I (Preston et al. 2002; Preston et al.2004). S. alba is widespread and locally common in suitable lowland waterside habitats inBritain, stretching across the whole latitudinal range. It does, however, become more scarce as onetravels N & W, and it appears to be declining in N & W Scotland in the same manner as occurs inW Ireland (G. Hutchinson, in: Preston et al. 2002).
European and world occurrence
Although this species is often planted, making it difficult to be certain where it is native, S.alba appears widespread in Europe, extending across middle latitudes from southern Denmarkeastwards to C Asia and south to the Mediterranean region (Meikle 1984; Jalas & Suominen 1976, Map203).
Threats
Many of the larger trees are over-mature and will soon become subject to windfall or simply die, sincethis is not a long-lived species.
Introduced, archaeophyte, deliberately planted, very rare.
1836; Mackay, J.T.; Florencecourt.
September.
Growth form and preferred habitats
This small, bushy tree or fairly robust, spreading shrub which grows in wet to damp, lowland situationsand can reach 10 m in height, remains the preferred willow species cultivated for basketry canes.Polymorphic and genetically variable, many named varieties exist and numerous continental clones wereimported into B & I over many years. Typical plants bear lanceolate to elliptic, pointed leaves morethan three times as long as wide, with their margins finely toothed. Leaf stipules are large, broad andpersistent, and the mature bark is smooth, peeling off in large irregular flakes like the London Plane(Platanus × hispanica) (Meikle 1984; Parnell & Curtis 2012). Theshrub or small tree is rare and grows in tickets and hedges in wet ground.
Commercial basket cane production
The major period of commercial willow cane production was during the Napoleonic wars and for a hundredyears afterwards. Peak activity was around 1900, after which production declined rapidly as importsbegan from E Europe and the Argentine (Meikle 1984; Newsholme 1992). Irish commercial cane productionwas concentrated at the SE end of Lough Neagh in NI and further south along the Rivers Shannon nearLimerick, and the Suir and the Blackwater in SE Munster (Hogan 2001). A recent account of Irishbasket-making suggests S. triandra only began to be widely grown in Ireland around 1900, althoughvarieties called 'Mauls' and 'Spaniards' were being grown S of Lough Neagh some years earlier (Hogan2001).
Apart from intensive osier beds, however, S. triandra and varieties of the other two major canespecies, S. viminalis (Osier) and S. purpurea (Purple Willow) would have been planted fortheir own and local use by many farmers with suitably damp ground. Lakeland Fermanagh gave virtuallyunrivalled opportunity for growing osiers in these small-scale 'sally gardens' and we can safely assumefrom folk memories and literature accounts that this also happened in many other parts of Ireland.
Fermanagh occurrence
In the Fermanagh Flora Database, there are just six records for this introduced and previously regularlyplanted willow species, all but the first and last of them made by Meikle and his co-workers. Thehabitats include riverbanks, roadsides and a gravel-pit in the south of the VC, the latter at leastindicating some degree of spread and naturalisation. Apart from the first record listed above the otherrecord details are as follows: road between Newtownbutler and Crom Estate, 1945; Gortaree gravel pits,Slieve Rushen, 1949; roadside at Galloon Td, Upper Lough Erne, 1950; around Newtownbutler 1951;Derryvore Td, NW shore, Upper Lough Erne (which is just across from Crom Castle), 11 September 1986, A.Waterman.
In the Revised Typescript Flora, Meikle et al. (1975) commented that S.triandra is frequent about the Newtownbutler area, which suggests either that modern botanistshave been overlooking it, or that the species may have greatly contracted since the 1950s. The NewAtlas hectad map gives evidence of a similar decline throughout B & I. Locally, the factthat these relicts of cultivation remained confined to the near vicinity of two large demesnes,Florencecourt and Crom, is significant, providing circ*mstantial evidence of where the species wasoriginally planted and indicating its rather limited ability to spread. Even more interesting is theability of the species to survive untended for perhaps 50 years or more.
S. triandra requires more fertile soil than the other commonly grown basketry willows and it isalso reputed to be more prone to rusts and insect damage, a fact which could account for its apparenteventual demise in most of Fermanagh.
Status in Britain and Ireland
Until very recently, S. triandra was regarded as native in British Floras, at least in C & SEngland, but as a probable alien introduction in other parts of Britain and in Ireland (Scannell &Synnott 1987; Stace 1997). Without giving any explanation, Scannell & Synnott (1987) consideredS. triandra might possibly be native in unspecified counties in SE Ireland, where indeed it has astronger presence. This could merely reflect, however, the extent of its previous cultivation along theriver basins of the Suir and the Blackwater, which were major areas of intensive osier productionthroughout the 19th and early 20th centuries.
The New Atlas editors first recognised that S. triandra is an ancient introduction, anarchaeophyte, and they mapped it as such throughout B & I.
Irish and British occurrence
In Ireland, S. triandra is rather thinly scattered throughout with a slight eastern and southernpredominance and with a considerable number of older pre-1970 records displayed. In Britain, thedistribution is much more decidedly south-eastern, although again the species is widely distributedreaching both Inverness and the Outer Hebrides. Over 26% of the hexads plotted for B & I were basedon pre-1970 records and a further 14% were pre-1986. Taken together, these facts demonstrate thewidespread decline of S. triandra, a species that without a major revival of basketry andreplanting is certain to continue towards local extinction (Preston et al. 2002).
European and world occurrence
S. triandra is widespread throughout most of Europe, the distribution thinning out north of 60°latitude and southwards into the Iberian peninsula, Italy and Greece. From Europe, it spreads eastwardsthrough Turkey and Iran into C Asia (Meikle 1984; Jalas & Suominen 1976, Map 204).
Threats
The current lack of planting fresh canes and the observed decline of existing populations will inevitablylead to local extinction.
Both native and deliberately planted, occasional. Eurosiberian temperate, but widely planted and probablyalso naturalised.
1836; Mackay, J.T.; Florencecourt.
May to November.
Growth form and preferred habitats
The linear-oblong leaves, and the associated buds and branches are very unusual and distinctive for awillow in being borne opposite or sub-opposite, rather than in the alternate and spiral arrangement moretypical of the genus. Although the specific epithet is 'purpurea' and the English common name atranslation, 'Purple Willow', in reality the twigs are usually yellow or grey, only sometimes tingedwith purple or red.
In the Typescript Flora and Revised Typescript Flora, Meikle et al. (1957, 1975)considered this variable shrubby species to be frequent in marshy ground and waterside banks inFermanagh. They commented that it was often planted along with other suitable, but more decidedly alienSalix species, such as S. viminalis (Osier) and S. triandra (Almond Willow), in wetparts of lowland farms (ie 'sally gardens'). Stems were harvested both as basket canes, or withies andfor thicker, lightweight timbers, which were put to a great variety of uses described in farming folkliterature (Evans 1942, 1957). Uses included live material for rapid screening and as shelter forlivestock, or in some species for animal fodder. S. purpurea is an exception in this last case,being much too sharp-tasting, even for rabbits and hares (Newsholme 1992). S. purpurea produces awoodier, tougher rod than S. viminalis and thinner rods of it are also more pliable than thelatter when fresh or partially dried (Hogan 2001).
Fermanagh occurrences
Meikle and his co-workers (1975) noted that in Fermanagh S. purpurea also grew spontaneously andapparently indigenously on wet hillsides, occasionally remote from habitation, eg in hedgerows, besidestreams and in damp hollows in the uplands. In the immediate post-war period, when they were recording,S. purpurea was particularly frequent around the Carnmore area, between Lisnaskea and Rosslea inthe E of Fermanagh. Our longer term survey has recorded S. purpurea in a total of 63 tetrads,representing 11.9% of those squares in the VC. As the tetrad map shows this shrub is very widelyscattered throughout, but with a slight concentration of records in the more actively farmed lowland SEof the county. The fact that there are 19 tetrads with only pre-1976 records suggests there has been aloss of suitable habitat, or competitive ousting has taken place.
British and Irish occurrence
The New Atlas hectad map shows that S. purpurea is fairly common and widespread throughoutB & I, although the distribution really is rather patchy. Furthermore, since this species was muchplanted in the past we really cannot discriminate native occurrences.
In Ireland, the hectad map suggests it is much more prevalent in NI and in the Midlands, ie acrossthe country from E Mayo to Kildare (H26-H19). The Cen Cat Fl Ir 2 suggests the status ofthis species is, "possibly introduced", with the additional debatable comment considering it,"probably native in the Centre". Further S & W in Ireland, S. purpurea is eithervery much more scattered or virtually absent.
Typical wet ground habitats in Britain include woodland margins, shingle beside rivers and streams and inmarshes and fens, including osier beds (G. Hutchinson, in: Preston et al. 2002). The distributionin Britain is curiously concentrated along the spine or middle regions, becoming less prevalent to botheast and west of the central line as far north as the Scottish Highlands. North of Glasgow and Edinburghit becomes much more scarce and coastal, although reaching one isolated hexad in Shetland, adistribution suggesting a temperature limitation is in force.
The map of the European distribution suggests that in B & I, S. purpurea is only indigenous inN England, S Scotland and NW and C Ireland (Jalas & Suominen 1976, Map 255). Meikle (1984) suggestsS. purpurea has a much greater claim to be indigenous than either S. viminalis or S.triandra, and the current author would not argue with that expert assessment. With regard tonative occurrence, it is significant that occasionally S. purpurea grows in damp hillside andstreamside sites quite remote from both habitation and cultivation, where it presumably must haveself-sown (Meikle 1984).
From the published hectad maps, there appears to have been little or no change in the S. purpureadistribution during the 40 years between the two B & I flora atlases (Perring & Walters 1976;Preston et al. 2002) − a rather surprising finding in view of the considerable loss of wetland habitatsdue to drainage and herbicide use associated with the intensification of farming during this period.
European and world occurrence
The suggestion of a temperature limit to distribution is confirmed by an examination of the Europeanrange of the shrub which thins markedly beyond 55°N; it scarcely passes anywhere beyond 60°N, and isplotted as 'introduced' in most of such areas. It is therefore more or less absent from Scandinavia, andit is also so from much of C Europe. Several closely related species occur in Asia (Meikle 1984; Jalas& Suominen 1976, Map 255).
Threats
None.
Introduced, archaeophyte, rare, but very possibly under-recorded and perhaps occasional. Eurosiberiansouthern-temperate, but also naturalised.
1949; MCM & D; roadside near Drumrain [sic, ie Drumram] Lough, SW of Donagh Crossroads.
According to Meikle (1984), this is one of the more common Salix hybrids "found here andthere throughout the British Isles". One might add, 'indeed across almost all of Europe'. The mainreason for this is that it was once a popular cane-providing shrub with basket makers, grown inosier-beds in a wide variety of clonally reproduced cultivars. Despite this fact it is said to varywithin a narrow compass and appears intermediate between its parent species.
All four records of the hybrid in the Fermanagh Flora Database are pre-1975 and were made by Meiklehimself and his co-workers. Thus we, and all the other field workers in Fermanagh since 1953, appear tobe completely overlooking this hybrid. Apart from the first record above, the remaining details are:Drumkeenagh, Black River, ie at NW end of Upper Lough Macnean, 1950; streamside, upper part ofDonaghmore Glen, S of Cooneen, 1950; Many Burns River, N of Many Burns Bridge, 1953.
The few habitats we have evidence of are not unusually associated with damp or wet ground on roadsides,near lakeshores and by river and stream banks. However, this suggests to us rare spontaneous crossesarise in open, more or less competition-free ground, with or without the parent species, probably innaturalised thickets rather than in previously cultivated ground. Having said that, all but three typesof spontaneous willow hybrids are occasional to rare or very rare, and S. × rubra is merely one of the occasional ones (R.D. Meikle, in: Stace1975, p. 305). The possibility of some of these stations representing the margins of old osierplantations cannot be definitely decided upon at this remove, but no comment claiming this is made ineither version of the MCM & D Typescript Flora (1957, 1975).
For comparison, the FNEI 3 gives brief details of a total of 14 stations in the three counties itcovers, spanning the whole period from John Templeton (active 1793-1825) until 1990. The very thinlyscattered recorded occurrence displayed for this hybrid in the New Atlas hectad map (in Irelandlargely confined to northern counties), suggests to the current author that either Meikle over-estimatedthe extent of this hybrid, or else it is frequently overlooked in both B & I.
Introduced, deliberately planted, very rare.
1948; MCM & D; Lough Aguse, Tullygerravra Td.
May to August.
Meikle (1984) states that until 1959 he had only seen female plants of this erect vigorous hybrid andthat males are very rare indeed. The true nature of this hybrid is still a bit of a puzzle, but itappears to be the product of three parents: the native S. cinerea var. oleifolia(probably?) (Grey Willow) crossed with two alien, archaeophyte basket osiers, S. purpurea (PurpleWillow) and S. viminalis (Osier). The hybrid is thus a relict of basket-cane cultivation and mostlikely arose from a cross between S. × rubra (Green-leaved Willow)and S. cinerea (Stace et al. 2015). It first came to scientific notice in the very earlyyears of the 19th century, although probably it was familiar to willow growers well before 1804 when SirJames Smith described and named it after Rev. Joseph Forby who had brought it to his attention.According to Smith it was locally known as, 'Fine basket osier', but when it was subsequently introducedto osier-beds on the Thames it proved too coarse for fine basket-work (Bean 1980). Really fine basketsare capable of holding water like a bucket.
The current author has no idea when this hybrid was introduced to Fermanagh, but the fact that it is alsofound in three other VCs in NI and that the New Atlas map plots it in a total of 29 hectads andStace et al. (2015) likewise in 34 hectads in this corner of Ireland, shows that it is by nomeans isolated. This hybrid is particularly well represented in Co Antrim (H39), around Ballymena andClough Mills (Hackney et al. 1992).
Fermanagh occurrence
In W Fermanagh, S. × forbyana has persisted at several sites on wetground along the banks of the Roogagh River for over 50 years. There are just seven records in theFermanagh Flora Database spread across five tetrads scattered across the VC, east and west of LoughErne. Three records were made by Meikle and his co-workers between 1948-50, and four by RHN in the early1990s and 2006. In addition to the first record given above, the remaining details are: lane nearLisnaskea Railway Station (railway now disused), 1950; frequent along the course of the Roogagh Riverand connecting streams, 1950; Tullyedderdamore Td, Roogagh River, 23 May 1992, RHN; left bank of RoogaghRiver, 100 m below Roogagh Bridge, 20 June 1993, RHN; forest track, S of Lough Namanfin, 28 August 1993,RHN; Lough Skale, Clogtogle Td, 5 June 2006, RHN & HJN.
Introduction, archaeophyte, deliberately planted, common.
Eurasian temperate, also widely naturalised.
1900; Praeger, R.Ll.; Co Fermanagh.
April to December.
Growth form, status and preferred habitats
The long, narrow, untoothed leaves, with their narrowly inrolled margins, covered underneath withadpressed, silky, silver hairs, make this a very distinctive tall shrub. S. viminalis is one ofthe most common willow species of damp ground throughout lowland B & I.
Variation within the species is not great, but hybrids are freely formed with numerous other sallows andosiers and several of these crosses have been rarely recorded in Fermanagh. Fortunately the species iseasily distinguished from the hybrids by its long leaf with straight, silky hairs on the under-surface.
In Ireland, Osier, or Common Osier, has long been regarded as a much-planted introduction for basketcanes, eg Mackay (1836) and Cybele Hibernica 1866 (p. 269), the latter commenting,"scarcely wild". This has probably also been the supposition in Scotland (New Flora of theBI). Very recently, S. viminalis has also been recognised as a naturalised ancientintroduction (ie an archaeophyte), throughout the whole of these islands (New Atlas;Preston et al. 2004).
S. viminalis is a common and widespread relict of cultivation and a naturalised escape throughoutIreland, found in a wide range of wet to damp ground situations usually near water. In our damp Atlanticclimate, the species behaves as a pioneer colonist, self-seeding, naturalising and increasingvegetatively once it becomes established in well-lit spots in hedgerows, along roadsides, in ditches andon any damp waste ground, eg in disused quarries and gravel pits.
Fermanagh occurrence
In Fermanagh, S. viminalis has been recorded in 228 tetrads (43.2%), making it the fourth mostcommon and widespread willow in the VC. It is tolerant of a wide range of moderately acidic to neutral,calcareous soils of variable nutrient status, but it prefers moderately fertile, sunny, sheltered sites.While it is frequently found on the somewhat higher ground of the Western Plateau, like other willows itdoes not venture into shade anywhere, nor on to exposed high ground or very acidic, ombrogenouspeatland. Full light is critical for sustained growth of any kind of willow, tree or shrub, species orhybrid, and any shaded part of a plant soon dies away (White 1992).
Fermanagh never seems to have had the large-scale, well-developed cane willow industry one might expectin a county geographically so well suited to it. The main area of intensive willow cane and commercialbasket production in NI of the 19th and early 20th centuries was in the SE corner of Lough Neagh nearLurgan. The other major Irish willow plantations were along the River Shannon in Co Limerick and in thefertile river basins of the Suir, the Blackwater and their tributaries in E Munster (Hogan 2001). Anample water supply is essential for successful seedling development of willows, but established matureplants are much less demanding in this respect, although wet ground also facilitates or enablesvegetative reproduction, involving layering, suckering and rooting of detached twigs.
European origin and modern uses
S. viminalis is probably of Russian or N Eurasian origin and Meikle (1984) suggests it may havespread westwards with early man, since its long, straight, strong and very flexible young shoots are, orrather, were, very widely planted and harvested, chiefly for basket-making. Often Osier stools did notyield ideal basket canes however, since the very rapid growth rate tended to make them too brittle.Nowadays, named clonal forms and hybrid S. viminalis varieties are widely planted along roadsidesand around buildings for amenity landscape or screening purposes. The material used by landscapecontractors is invariably imported from European horticultural sources and inevitably this willintroduce additional alien genetic variation, some of which might prove invasive or introduce associatedpests or diseases (White 1992). Cultivars of S. viminalis are also being used for experimentalshort-rotation biomass projects in research areas including alternative energy sources, or processed ascattle fodder.
Threats
None.
Introduction, deliberately planted, very rare but probably under-recorded.
1947; MCM & D; laneside hedge, Lowery Bay, Lower Lough Erne.
There is only one record of this tall erect shrub or small tree in Fermanagh, made by Meikle andco-workers almost 60 years ago. The current young twigs and one-year-old shoots of this hybrid aredensely covered with greyish, silky, velutinous hairs and leaves are narrowly lanceolate orlinear-lanceolate, dull green and sparsely pubescent on the adaxial side (ie the upper surface, towardsthe apex). The leaf under-surface is densely coated with white or ashy hairs. All plants in B & Iare female and they possibly belong to one clone (Stace et al. 2015).
The plant is a basket-cane hybrid between alien and native species, of perhaps still slightly dubiousparentage (R.D. Meikle, in: Stace 1975; Stace et al. 2015). It was widely planted and stilloccurs in these islands, but only rarely and thinly scattered in damp lowland hedgerows and scrub acrossNE Ireland, although in a wider variety of habitats in parts of N England and S Scotland (Stace etal. 2015). There are records in FNEI 3 from a total of 22 stations in the three VCs(H38-H40).
S. × stipularis usually occurs in deliberately planted areas of thecountry, and very probably it remains under-recorded, being similar in appearance to S. viminalis(Osier). It is distinguished by its broad persistent stipules, densely grey-pubescent(velutinous) leaf under-surfaces and its relatively large villose female catkins (R.D. Meikle, in: Stace1975; Stace et al. 2015; Stace 2019).
Introduction, archaeophyte, deliberately planted and possibly also rarely occurring spontaneously; rarebut probably under-recorded.
1949; MCM & D; laneway by Lough Melvin near Garrison.
June to November.
This erect hybrid shrub or small tree is very variable and therefore difficult to distinguish from S.× holosericea Willd. (Silky-leaved Osier), the cross between S.viminalis (Osier) and S. cinerea subsp. oleifolia (RustyWillow). This means it is very probably under-recorded in areas of B & I where basket caneswere previously cultivated (Meikle 1984). The previous confusion in the application of the name S.× smithiana was resolved by Larsson (1995). In our Fermanagh FloraDatabase, we have a total of 14 records of S. × smithiana in separatetetrads, but in comparison only seven records of S. × holosericea.
S. × smithiana is thinly and widely scattered across damp lowlandareas of Fermanagh, growing in roadside and laneway hedgerows, stream-sides, rock outcrops andlakeshores. As also noted in the account of S. × holosericea, thesehybrids probably arise rarely and spontaneously whenever the parent species happen to occur together.Like S. viminalis itself, however, they may often be relicts or escapes from previously plantedimports in cane cultivation plots. One of the Fermanagh S. ×smithiana sites, south of the road at Kilturk, contained a collection of around a dozen largeshrubs of this hybrid, which suggested that this patch of ground might originally have been one of thesmall, local, cane plantations referred to as a 'sally garden'.
The FNEI 3 regarded S. × smithiana as "widespread",and the editors commented that, after S. viminalis, this hybrid is (or was) the commonest basketwillow in the region of that Flora. Their text follows this remark by listing just one station in CoDown (H38), three in Co Antrim (H39), but as many as 15 from Co Londonderry (H40).
The New Atlas map and that in Stace et al. (2015) indicate that S. × smithiana is very widely, but rather patchily, distributedthroughout B & I. It occurs spontaneously with or without both or one of its parent species,appearing in hedgerows, thickets and waste ground. It also is found as relicts of cultivation, survivingin or near more or less obvious osier plantations, their remnants or, increasingly nowadays, theirmodern reconstructions. Previously, S. × smithiana was valued forcoarse heavy-duty basketry, eg of the type used for turf (peat fuel) creels, thatch ties and manurecarriers. Lately, interest in this hybrid has reawakened and it is now considered suitable forshort-rotation biomass production (G. Hutchinson, in: Preston et al. 2002).
Threats
None.
Introduction, neophyte(?), deliberately planted, apparently very rare but probably under-recorded.
1833; Mackay, J.T.; Florencecourt estate.
In the Revised Typescript Flora, R.D. Meikle gave his opinion that Mackay'sidentification might possibly be correct, but he also made it clear that the specimens had not beenlocated and expertly examined. Mackay (1836) listed it as S. acuminata Sm. (Long-leaved Osier)and stated that, "it had become a tree 20 feet high, although growing in an elevatedposition". It is still possible that a herbarium specimen may have survived, most likely in eitherTCD or DBN. The Revised Typescript Flora suggests that Mackay collected theplant from hedgerows and considered it either planted or an escape, but as far as we are aware this ismerely surmise.
There are no subsequent records for Fermanagh of this tall, erect hybrid shrub or tree, the putative (butuncertain), parent species of which are the common shrubby sallows, S. caprea (Goat Willow),S. cinerea subsp. oleifolia (Rusty Willow) and S.viminalis (Osier), ie two native and one alien species. The hybrid exists as a purely femaleclone, probably a single genome, of necessity reproducing vegetatively. Stace et al. (2015)discuss the possible parentage of this hybrid in detail, without coming to a definitive conclusion.S. × calodendron is always imported and planted, rather than arising spontaneouslywhere the parent species meet. The reasons for planting in the past are also uncertain, but it was mostlikely admired by some for its handsome, crowded, upwardly curved female catkins (Bean 1980).
While the specific epithet, calodendron, translates as, 'beautiful tree', Howitt & Howitt(1990) described it as the only ugly willow they know. These workers found that a small patch of S.× calodendron was grown in every English willow holt they hadstudied in the Trent valley, although they could not really fathom the reason why. One suggestion theymade was that the timber of this hybrid is very easily longitudinally split, or it does so naturallyonce the branches reach a diameter of around 10 cm. It seems very likely that basket makers had aspecific use for this split timber and accordingly they valued and cultivated this hybrid.
Although the date, place of origin and reason for introduction are all unknown and rather mysterious, theNew Atlas editors still consider S. × calodendron as being aneophyte, ie an introduction of recent, or at least very probable post-1500 AD date. Their hectad mapfor Ireland shows the plant recorded almost exclusively in coastal areas, the major area on the islandbeing the NE counties of Down and Antrim (H38 & H39). Mackay's old Fermanagh record represents aunique inland Irish station for this hybrid. In S England, there are numerous scattered examples ofinland sites for the plant, again always planted.
In recent years, S. × calodendron plantation has been renewed for useas a biomass crop, managed on a short-rotation coppice for fuel (Stace et al. 2015).
Threats
None.
Both spontaneous and an introduced and deliberately planted archaeophyte, rare, but very probablyover-looked and under-recorded.
1948; MCM & D; Lough Aguse, Tullygerravra Td, 6 km SE of Garrison.
July and October.
This is regarded by Meikle (1984) as one of the more common Salix hybrids in B & I, but as healso points out, it is an easy one to confuse with S. caprea × S.viminalis (S. × sericans) (Broad-leaved Osier). There areonly eight records of S. × holosericea in the Fermanagh FloraDatabase and all but two date from the 1948-53 period and were made by Meikle and his co-workers. Apartfrom the first record given above, the remaining details are: near Kesh, 1950; Roogagh River nearTullynanny Lough, 1950; Ross Lough near Carr Bridge, 1950; Mullyduff Lake, E of Newtownbutler, 1951;roadside between Ballindarragh and Lough Digh, 1953; Goladoo disused quarry, 20 October 1996, RHN, det.P. Hackney, Herb. RHN; upstream of bridge, near Rosscor viaduct, 2 November 2010, Herb.RHN.
Thus in Fermanagh, S. × holosericea is extremely thinly and widelyscattered in damp or wet habitats on lakeshores, river banks and roadsides.
In the Flora of Lough Neagh, this willow hybrid is regarded as locally frequent in hedgebanks,ditches, streams and lakeshore thickets and it is mapped in 17 tetrads around the lakeshore. The FNEI3 account also describes the shrub as "frequent", although the number of sites listedfor the three component VCs (including some very old 19th century records) amounts to a total of just50.
S. viminalis (Osier) was introduced in ancient times and widely and commonly cultivated throughoutB & I for basket cane production. Both the above mentioned osier hybrids involving this introductioncan arise spontaneously through its ready interbreeding with large and widespread native populations ofS. cinerea subsp. oleifolia (Rusty Willow) in this instance, andsimilarly with S. caprea (Goat Willow) to create S. × sericans(Broad-leaved Osier). Clones of both these hybrids were also commercially propagated and around acentury ago were widely used for basket canes. Very probably some of the hybrids found nowadays are theoutcome of rarely occurring indigenous local crossings, but certainly the majority of them are relictsof cultivation.
The map of S. × holosericea in the New Atlas shows a largemajority of its Irish records are confined to NI. However, as the brief species account accompanying themap suggests, this hybrid is very probably under-recorded throughout B & I (G. Hutchinson, in:Preston et al. 2002). In the current author's view, the same can be said of all willow hybridsand, indeed, of some willow species too. In recent years there has been renewed interest in this hybridfor biomass production.
Introduced, archaeophyte, deliberately planted, but may also occur spontaneously, very rare either way.
July 1947; MCM & D; laneway hedgerow near Tattycam Lough, in the SE of the VC.
This is another shrubby, sallow-osier hybrid, this time combining the distinctive leaf characters of twowillow species of very different appearance. Meikle (1984) considers S. ×fruticosa to be a purely native and spontaneous hybrid in Ireland, the result of crossingbetween introduced clones of the basket-cane S. viminalis (Osier) and the native S. aurita(Eared Willow). Crossing would not be easily achieved, however, since the parent species have verydifferent flowering periods. Meikle (1984) commented that this is an uncommon but widely scatteredhybrid in B & I, perhaps more frequent than might be expected in view of the significant differencebetween the normal flowering periods of the two parents (S. aurita – March and April, S.viminalis – April and May), giving them just one month of overlap. As is the case with severalwillow hybrids, sometimes progeny of one sex only are produced: in this instance only female plantsoccur in B & I.
Fermanagh occurrence
Until recently, we had only one record for this plant in the Fermanagh Flora Database, made by Meikle andhis co-workers during their regular July visit, on this occasion as long ago as 1947. Then RHN and HJNmade two additional records as follows: roadside hedge S of Clonelty, 15 September 2010; and hedgebeside stream opposite old house Tullygarry, near Meenatully, 4 October 2010. In both these cases, thehybrid was growing with both parents and was obviously intermediate. Voucher specimens were collected.
British and Irish occurrence
There are a total of 74 hectads with records plotted in the New Atlas map which shows S. × fruticosa very widely but also very thinly scattered across Britain,with a just discernable western tendency in the albeit rare presence of this hybrid. In Ireland, the 30plotted hectads form three discrete clusters dispersed across the island. The most prominent of thesegroups lies within NI, although here it is chiefly in Tyrone (H36). There are two discrete hectadclusters in the RoI, in the Midlands (Longford-Roscommon (H24 & H25) and possibly also on theborders of the adjoining VCs), and further SW in Limerick (H8).
The widely scattered distribution of this uncommon hybrid throughout these islands which Meikle (1984)predicted in his Willows and Poplars Handbook, does exist, but it remains a rather rare plant.Willows are not long-lived plants and although hybrids possess extra vigour in comparison with species,even these will tend to die out through shading by taller trees and shrubs, unless light levels aremaintained by active management of the hedges and thickets it frequents.
The present day B & I distribution of S. × fruticosa probably owesmore to recent and continuing experimental plantation of selected clones for biomass production, as itdoes to the appearance of naturally occurring hybrids, or to the long-term survival from old basket-caneplantations. Undoubtedly S. × fruticosa is just as under-recorded asall other willow hybrids, which always needs to be borne in mind when making any such comment.
Threats
None.
Native, rare, but under-recorded and probably scattered alongside both parents.
1989; Tickner, M.; thickets on Stony Islands, Lower Lough Erne.
May to July.
Fermanagh occurrence
Nine of the twelve records for this hybrid in the Fermanagh Flora Database are credited to MatthewTickner and were made by him on the rocky islands in Lower Lough Erne in the summer of 1989. In hisWillows and Poplars Handbook, Meikle (1984) concluded that this hybrid is very common,that pure S. caprea (Goat Willow) is much less common than generally supposed, and that indisturbed habitats (eg in felled woodland), this hybrid with S. cinerea subsp. oleifolia (Rusty Willow), plus the latter species itself,often replace the pioneering colonist, S. caprea.
British and Irish occurrence
In the three NE counties of Ireland, Hackney reckoned that S. ×reichardtii is more frequent than pure S. caprea in some districts, eg in E Down (FNEI3). A very similar report was given in the recent Flora of Berkshire (Crawley 2005), butother recent local Floras from throughout B & I have generally tended to play down the presence andsignificance of this hybrid (eg Swan 1993; Woods 1993; Trueman et al. 1995; Brewis et al.1996; Flora of Co Dublin; Beckett et al. 1999). In the Shropshire region, on the otherhand, Sinker et al. (1985) considered that this hybrid is less common than is often imagined,although they acknowledge that hybrid swarms do occur, "where the parent species occur together inopen communities with plenty of space for seedling establishment".
In Cumbria, Halliday (1997) reported 14 records of S. × reichardtiifrom several localities in the NE of the region, including around Penrith. It was particularly frequenton disturbed ground and was recorded chiefly by one particular field worker. Apart from this, Halliday'scomment was, "otherwise apparently rather scarce". Nevertheless he reckoned that the positionin Cumbria was a distortion of the true picture, blaming the laziness of most recorders who on findingboth parent species in a tetrad, "not unnaturally tend to pass over inconvenientintermediates".
Exactly the same can be said of the Fermanagh experience and the patchy distribution of this hybridplotted in the New Atlas hectad map indicates that this situation is commonplace and that ourrecording is undoubtedly deficient. We believe that Meikle (1984) was very accurate in his assessment ofthis hybrid and its parents. The problem is that the hybrid can be hard to distinguish from each of itsparents and, where the two species occur together (or previously did so), they hybridize and backcrossto produce an unbroken series of polymorphic, intermediate hybrid forms that intergrade between theparents (Meikle 1984; Sell & Murrell 2019). There is particular difficulty in separating the hybridfrom S. caprea, a fact which undoubtedly inhibits recording. This can readily be seen in the mapprovided by Stace et al. (2015), where clustering of records denotes areas of the country visitedby confident willow recorders.
Threats
None.
Native, common. Eurosiberian boreo-temperate, introduced rarely in eastern N America.
1881; Stewart, S.A.; Co Fermanagh.
This is a common enough shrub of wet ground, lake shores, thickets and the margins of woods. There are 48records from 21 tetrads simply recorded as S. cinerea without reference to the subspecies. Thegreat majority of these records were made by the EHS Habitat Survey team around Upper Lough Erne and onlake shores in the SE of the county. Detailed comments are made below under the two subspecies of S.cinerea.
Possibly introduced, and either extremely rare, or more likely, absent and mis-identified. Eurosiberianboreo-temperate, introduced rarely in eastern N America.
21 June 1985; EHS Habitat Survey Team; marsh to east of Inishroosk Td, Upper Lough Erne.
June to September.
British and Irish occurrence
This subspecies, although it is the typical, 'nominate' form of the species and is common and locallyabundant over most of continental Europe, is decidedly uncommon in the British Isles. Here it is largelyreplaced by the widespread Atlantic subsp. oleifolia (Rusty Willow) (Meikle 1992). By contrast,in B & I subsp. cinerea is only widely distributed in the base-rich fenlands of S and SEEngland. However, the New Atlas map indicates that it has recently been recorded from other morescattered areas of the British Isles, including from Orkney and Shetland. In Ireland, this increaseddistribution includes a few specimens from Co Down (H38), which were found by John Harron shortly afterMeikle published his excellent aid to identification, the BSBI Willows and Poplars Handbook in1984. The identity of the Co Down specimens, however, was considered uncertain in the subsequentlypublished FNEI 3.
Status in Ireland
Although subsp. cinerea may turn out to be more common than the few records in Co Down and thesolitary Fermanagh find claimed by Daniel Kelly in 1992 (voucher in TCD) would lead us tobelieve, at the moment the current author is not convinced that this subspecies occurs in Irelandat all. Robert Northridge and the current author are not surprised that the editors of the NewAtlas ignored the supposed 26 Fermanagh records of subsp. cinerea from 23 tetradsthat were made between 1985 and 1996 by the EHS Habitat Survey Team. We very seriously doubt theiraccuracy ourselves. The map does however show three rather lonely older date class symbols in Co Down.In the brief text accompanying the New Atlas map, G. Hutchinson comments that on the western edgeof its range, subsp. cinerea becomes more difficult to distinguish from subsp. oleifolia,a fact probably related to hybridization between these two forms.
The characteristic morphological differences between the two subspecies, eg in the size and persistenceof leaf stipules, plus the breadth of their ecological ranges, are not clearly demonstrated in Irishspecimens of putative S. cinerea subsp. cinerea. We suspect thatthere really is only the one very variable Atlantic subspecies of S. cinerea present in Fermanaghand throughout Ireland (ie subsp. oleifolia), plus the several hybrids and their parentalbackcrosses which this form of S. cinerea makes with four other local shrubby sallows. Wetherefore regard all the Fermanagh records for S. cinerea subsp. cinerea as mis-identifications.
Native, common, widespread and locally abundant. Suboceanic temperate.
1882; Stewart, S.A.; Co Fermanagh (as S. cinerea L.).
Throughout the year.
Growth form, preferred habitats
Familiarly and locally very well known as 'Sally' or 'Sallies' in Fermanagh, S. cinerea subsp. oleifolia almost always appears as a multi-branched shrub,rather than as a tree with a solitary basal trunk. The tree form is more likely found in betterilluminated parts in the shrub layer of relatively undisturbed woodland, which, on account of prevailinglocal grazing practices, is an uncommon habitat in Fermanagh. Until one is practiced at willowidentification and familiar with this plant, the leaves of subsp. oleifolia are so extremelyvariable that fully mature summer ones are recommended for its proper, secure identification: the springand early summer leaves often appear quite different from the mature leaf, not only in shape and size,but also in the details of the leaf margin. This great variability is reflected in the complicatedhistory, numerous names and frequent changes of status which this taxon has undergone, all clearlydescribed in Meikle's excellent 1984, BSBI Handbook of Willows and Poplars.
Subspecies oleifolia is an extremely variable plant, but generally it, − and unfortunately fromthe identification point of view, the range of hybrids it forms with other willows − is readilyrecognised by the presence on the underleaf of sparse rusty-brown hairs on the veins. However, whitehairs may also be present. The very much rarer subsp. cinerea, which is either very rareor maybe entirely absent in Ireland (see above account), is distinguished by the lack of these brownhairs and by the possession of twigs covered with very short, velvet-like, grey hairs, which remaindensely pubescent for over a year (ie they are very persistently pubescent) (Meikle 1984).
S. cinerea subsp. oleifolia shrubs occupy a large number ofhabitats, tolerating damp to occasionally flooded, but not permanently waterlogged, lightly shaded oropen sites on moderately acid to base-rich or calcareous soils, but in the latter case, the soils arealways moisture-retentive. Habitats range from relatively undisturbed semi-natural, swampy to marshyfen-carr, to drier scrub and wood margins and meadows on moderately acid soils, to more disturbed, bare,wet mineral soils, together with artificial, open wayside and waste ground situations, eg hedgerows andold quarries.
Reproduction
In more open, moderately disturbed situations such as waste ground and quarries where bare soil surfacesbecome available, high seed production and efficient plumed wind-dispersal enable this subspecies torapidly invade as a pioneer colonist. Once established, the remarkable vegetative reproductive abilitycharacteristic of all willows enables it to spread and form dense thickets by layering and by there-rooting of any detached parts.
Fermanagh occurrence
This essentially Atlantic subspecies of S. cinerea is by quite a long margin the most common,widespread, locally abundant and sometimes dominant type of shrub willow in Fermanagh. It occurs almosteverywhere in the VC except on very strongly acidic, truly aquatic or excessively dry, heavily disturbedor very exposed ground. It is the fourth most widespread woody plant in Fermanagh behind Hawthorn(Crataegus monogyna), Ash (Fraxinus excelsior) and Alder (Alnus glutinosa). Subsp.oleifolia is represented in 478 Fermanagh tetrads, almost 90.5% of those in the VC and so itcould almost be described as 'tetrad ubiquitous'!
British and Irish occurrence
At the hectad level of discrimination plotted in the New Atlas, the Fermanagh ubiquity is mirroredjust about everywhere in B & I except Norfolk and adjacent East Anglian VCs, where subsp.oleifolia is more or less replaced by subsp. cinerea (Willows and Poplars Handbook;New Atlas).
Hybridization
In wet or periodically waterlogged conditions around Fermanagh lakeshores, subsp. oleifolia veryoften grows alongside the two other common and most ecologically undemanding sallows, S. aurita(Eared Willow) and S. caprea (Goat Willow) and they may regularly form hybrids. We are aware thatwe and other local recorders tend to habitually overlook these shrubby willow hybrids and we acknowledgethat they are under-recorded here, as they also are elsewhere in B & I.
Threats
None.
Native, rare but very possibly under-recorded, perhaps occasional.
2 August 1967; Parker, R.E.; bogland E of Boa Island, Lower Lough Erne, BEL.
June to August.
Fermanagh occurrence
This hybrid was described by Meikle (1984) as very common, found everywhere the parent species occur. Ifthis really is the case, it is distinctly odd that it was not noted in Fermanagh by anyone (includingMeikle himself) before 1967. Up until 2010, the cut-off date for records in The Flora of CountyFermanagh that was published in late 2012, there were only eleven records of this hybrid in theFermanagh Flora Database. These few records were thinly and widely scattered across nine tetrads in theVC, which suggested that we were not recognising and recording this hybrid, or else that it was notanything like as common in Fermanagh as appeared to be the case further east in Cos Down and Londonderry(H38 & H40). The few records in Fermanagh at the time of the Flora publication (Forbes &Northridge 2012), came from a surprisingly wide range of habitats where the parent species overlap,including wood margins, thickets and hedges on lakeshores, by rivers, streams, in quarries or onroadsides. This rather strongly suggested that the hybrid was more frequent than had so far beenrecorded in the VC and this has proven to be the case since the online BSBI Database (accessed February2020) now lists a total of 41 records, although a few of them appear to be duplicates and others requirefurther validation before they can be fully accepted.
Genetic introgression following frequent back-crossing may have established a continuous series ofintermediates forms between the parent species, leading non-specialist Salix recorders, includingourselves, to widen in our minds the variation acceptable within these two species. Thus we may be'shoe-horning' shrubs into one or other of the species, rather than recording them as putative hybridforms.
Hybrid occurrence elsewhere in Britain and Ireland
Having acknowledged this quite definite possibility, the evidence provided by the New Atlas map ofthis hybrid suggested that Meikle (1984) might have overstated the likelihood of finding this taxon inFermanagh. The recorded hectad distribution of S. × multinervisappears more scattered than that of either of its parent species and, while the New Atlas editorsregard the map as 'incomplete', our slight Fermanagh record in 2012 did not appear very differentfrom, nor totally inadequate in comparison with many other VCs in B & I. The considerable increasein Fermanagh records in subsequent years, plus an addition of earlier records by experienced recordersnow recognised as belonging here, means Meikle has been proven correct in his prediction.
The experience of Scandinavian botanists is similar and relevant to the above position: Jonsell etal. (2000, p. 181) concluded that, "most specimens identified as this hybrid belong to oneof the parents". On the other hand, Howitt & Howitt (1990) claim that in Nottinghamshire (VC56) and in some other lowland areas of Britain, including East Anglia, S. aurita has,"hybridised itself out, or has been lost through drainage". It has to be said, however, thatthe New Atlas maps of S. aurita and of this hybrid indicate that even to begin with S.aurita was never very strongly present in the lowland areas of the Midlands and SE England(Preston et al. 2002).
The updated hectad map of B & I provided by Stace et al. (2015) confirms that S. × multinervis is more common and widespread throughout both islandsthan previously was thought. These authors consider it unlikely that hybridization directly causes anydecrease in S. aurita, but rather that replacement of the latter is more probably due todecreasing fitness of the species to cope with the current rapidly changing environment (Stace etal. 2015).
A further examination of the records listed in the FNEI 3 indicated that only a very few recordersin the three Irish VCs covered by this Flora were able or willing to recognise this hybrid. Fordiscovering this and other willow hybrids in NI, John Harron deserves particular mention (Hackney etal. 1992). The suspicion that this hybrid is being regularly overlooked or mistaken for itsparents by many recorders in Ireland and only recognised by a few, is strongly reinforced by the factthat Daniel Kelly (1985) published records of S. × multinervis in sixother Irish VCs apart from Fermanagh (ie in N Kerry (H2), Mid Cork (H4), N Tipperary (H10), SE Galway(H15), Offaly (H18) and Leitrim (H29)). The habitats he lists ranged from damp mixed woodland marginsand hedgerows to the cut-away margin of a bog (Kelly 1985).
Additional evidence supporting Meikle's contention comes from Green (2008) who found this hybrid was thesecond most common willow after S. cinerea subsp. oleifolia in CoWaterford (H6). He found it occurred from sea level to high in the mountains, and only rarely appearedalong with both parents. It is clear from this, and from the wide range of habitats in whichS. × multinervis has been recorded, including along rides in coniferplantations, banks of rivers and streams, in disused quarries and railways and on rock faces in mountaincoums (Green 2008), that R.D. Meikle, who verified some of the Irish vouchers, is absolutely correct inbelieving this hybrid to be of frequent occurrence in Ireland.
Threats
None.
Probably introduced, very rare, but probably also over-looked.
22 September 1990; Riley, Dr D.H.; Edenmore, 1 km NW of Tempo village.
Fermanagh occurrence
This rare hybrid willow has been found only once in the VC – in a roadside hedge near a stream justoutside Tempo village which drains Derrin Lough. The specimen's identity has been confirmed by R.D.Meikle. The erect shrub was found by Dave Riley, BSBI VC Recorder for Co Londonderry (H40), who hasspent more time studying willow hybrids than most other local recorders. He also found a single stationfor this hybrid in adjacent Co Tyrone in 1990, at Maghera Lough, near Castlederg (Harron 1992). S.× puberula was previously recorded along with both its parentspecies by an upland stream in Co Antrim (H39) by A.W. Stelfox in 1931 and, more recently (1984-9), byJohn Harron (the other current NI willow expert), in Cos Antrim and Londonderry (H39 and H40).Consequently the FNEI 3 lists a total of eight stations for this particular shrub hybrid, all butone in Co Antrim.
British and Irish occurrence of S. myrsinifolia: In B& I, S. myrsinifolia Salisb. (Dark-leaved Willow), a Boreal-montane species, is mostfrequently recorded in northern regions and is very rare and scattered in Britain south of a linebetween Fleetwood and Scarborough. The hybrid is completely absent south of this same line. The mappedrecords of S. myrsinifolia and this hybrid, produced most recently by Stace et al. (2015),suggest that S. × puberula is at least occasional within the rangeof S. myrsinifolia. The species itself almost always occurs in very small numbers (or as solitarybushes), growing in open, unshaded banks, usually rooted in damp but free-draining, peat-rich, stony orgravelly soils on river banks and lakeshores (R.D. Meikle, in: Stewart et al. 1994). Generally,but not always, S. myrsinifolia grows sufficiently near lowland streams that its roots areunlikely to dry out. While rare and rather widely scattered, it is regarded as native in the N & NEof Ireland. Elsewhere in Ireland, S. myrsinifolia is even more rare and it is assumed to beeither naturalised or perhaps deliberately planted, eg when in or near hedgerows, as it appears inFermanagh. S. myrsinifolia readily hybridizes with accompanying but definitely indigenoussallows, S. cinerea subsp. oleifolia (Common Sallow), S.aurita (Eared Willow) and S. caprea (Goat Willow). Rather often only these hybridbushes can be found.
Harron (1992) reckoned that S. myrsinifolia is being hybridised out of existence in Co Antrim(H39) at least. It has already died out along several glen-side streams where it previously grew and,based on morphological characteristics, in 1990 only the last remnant genetic traces of S.myrsinifolia remained in some of these particular sites. The most thriving surviving station inNI for S. myrsinifolia and its hybrids appears to be in Co Londonderry (H40), along the banks ofthe River Roe between Dungiven and Corrick Bridge above Carn and further up along some feeder streams(Harron 1992).
Native, common throughout on acid soils, locally abundant. European boreo-temperate.
1881; Stewart, S.A.; Co Fermanagh.
January to November.
Growth form, appearance and preferred habitats
One of the more distinctive shrub willow species of Fermanagh and indeed so in a B & I context, thesmall, broad, wrinkled, dark green leaves, softly pubescent below, with prominent impressed, net-likenervation and with persistent, ear-like stipules, make this a very readily identifiable much-branchedshrub. Howitt & Howitt (1990) warn that the auricles on their own are not a reliable guide.Much better is the always twisted tip of the leaf, a character which is also seen, however, in thehybrid S. aurita makes with S. cinerea subsp. oleifolia(Common Sallow, Rusty Sallow or Sally) (ie S. × multinervis).
The preferred habitats include around lake shores, the margins of upland woods and scrub, onmoors, heaths, bogs, cliffs, stream-sides, limestone pavement, quarries and roadsides of the WesternPlateau. It is found in both limestone and peaty, acid districts, but much more frequently in thelatter. At higher altitudes and in more exposed sites it often becomes considerably dwarfed and care isthen needed to distinguish it from S. repens (Creeping Willow).
Fermanagh occurrence
S. aurita ranks as the second most frequently recorded and widespread sallow in Fermanagh, beingfound in 308 tetrads, 58.3% of those in the VC. It is widely distributed throughout the county, but isvery much more frequently recorded around the shores of Upper Lough Erne and is common across much ofthe Western Plateau.
The Upper Lough Erne and Western Plateau prevalence of S. aurita is very likely in part anartefact reflecting the pattern of intensive surveys made by government field workers prior toconserving sites of scientific interest in these two areas of the county. However, in our view it cannotentirely be attributed to this factor, since throughout its widespread B & I range S. auritais known to be one of the shrubby willows that generally performs best on upland, acidic, sandy topeaty, unproductive, infertile soils. It can become locally dominant on wet heaths and on the driermargins of fens and peat bogs, including cut-over examples (Meikle 1984; Sommerville 1992).
Hybridisation is the only thing that occasionally blurs the identification of this species, but even whenthe species do overlap and interbreeding and introgression occurs with the considerably more commonsally, S. cinerea subsp. oleifolia, both parent species aregenerally very much more abundant than their intermediate hybrid, so they can be readily distinguishedfrom it.
British and Irish occurrence
S. aurita is widespread and frequent to abundant in suitable acidic soils throughout B & I,including the northern and western Scottish Isles (New Atlas). It is least common in the morepopulous and most intensively farmed areas of SE England, where habitat loss has been ongoing for almosta century.
Threats
None.
Native, very rare.
1946-54; MCM & D; on the shore of Lough Melvin, near Garrison.
Growth form, appearance and preferred habitats
The conspicuously ear-shaped stipules and the entire or sub-entire (ie scarcely toothed) leaf margins,the sometimes obliquely twisted leaf tip, the leaves at first coated on the upper surface with silky,white, adpressed hairs and the small size of the sprawling plant are all important recognition features.The habitat is acidic or slightly base-enriched heaths and moors, or in limestone grassland, occurringwhere the parent species overlap, although as with other willow hybrids, more often where one parent ismore common than the other, rather than where they are equally represented (Willows and PoplarsHandbook).
Fermanagh occurrence
There is only one record for this low, sprawling hybrid in our Fermanagh Flora Database made by Meikleand his co-workers on Lough Melvin near Garrison. It is undated but must fall sometime in the immediatepost-war period of this group's regular summer visits. In his BSBI Willows and Poplars Handbook,Meikle (1984) regarded this hybrid as being frequent in B & I, so perhaps it is being regularlyoverlooked.
Irish occurrence
The FNEI 3, published eight years after the Willows and Poplars Handbook, lists ten recordsof this hybrid from Co Antrim (H39), all of which were made by one recorder, John Harron, during the1970-88 period. The FNEI 3 account describes S. × ambigua asbeing, "frequent and widespread on the Antrim Hills". However, the Flora of Co Dublinmakes no such claims regarding this subshrub, merely mentioning that a previous record (no details) wasnot refound on Mountpelier in District 7 of the Flora. The Irish representation of this hybrid is verypoor, with just six hectads in Co Antrim (the Harron records mentioned above), and solitary hectads infour other Irish VCs, Co Kildare (H19), E Mayo (H26), Co Fermanagh (H33) and W Donegal (H35).
British occurrence
Despite Meikle's assertion that this hybrid is of frequent occurrence, the New Atlas hectad mapdemonstrates only a thinly scattered record of S. × ambiguaexists across northern parts of B & I, with a quite marked Scottish predominance, especially onthe N & W Isles. The hectad map in the Hybrid Flora of the British Isles shows a considerableincrease in records right across both B & I, but the pattern of scattered sites is similar, althoughwith more hybrid presence in the S of England. It is described by these authors as, "one of thecommoner willow hybrids in northern Britain, particularly near the west coast where the parents oftengrow in close proximity in exposed coastal heathland and moorland" (Stace et al. 2015).
Clearly, a greater and much more focussed recording effort is required to ascertain the true extent ofthis hybrid, but Howitt & Howitt (1990) point out the significant and relevant fact that S.repens (Creeping Willow) flowers later than S. aurita (Eared Willow) (ie usually aboutthe end of April), which obviously time-limits the opportunity for cross-fertilisation. The northerndistribution of the records in B & I may reflect a modification of the timing of flowering of thesewillow species, perhaps significantly increasing the period of overlap and enabling a greater frequencyof crossing.
Native, but a mis-identification. Eurosiberian boreo-temperate.
1990; Montgomery, J. & Foster, S.; Castle Caldwell FNR, swamp and carr.
This is regarded by us as a clear case of mis-identification. The EHS Habitat Survey recorders whobelieved they found this very rare willow wrote the species name on their record card rather than tickeda box. However, they did not appear to realise that S. phylicifolia is such a rare species, thatany new record requires a voucher, and after expert verification it would then deserve separatepublication in the Irish Naturalists' Journal.
Irish specimens of this species were previously referred to as S. hibernica Rech. f., but thecurrent view of most botanists in B & I familiar with the group is that S. hibernica is notsufficiently distinct and that it can be accommodated within S. phylicifolia. However we name it,this shrub is known in Ireland from only two damp limestone mountain cliffs on the Ben Bulbin range inCos Sligo and Leitrim (H28 & H29). The willow populations there are chiefly on cliff 'slips', whichoriginally formed part of the face of the main cliffs but which have slid downwards sometime in thecurrent post-glacial period. These slips now appear either as cliff stacks, separate but close to themain cliffs, or else they lie slumped at the base of the original cliff. These cliffs and slips carry arich assemblage of relict alpine plants. It is thought by some botanists at least, that thealpines and this willow survived here in situ on surfaces that were high enough to protrude abovethe glacial ice during the previous cold period (Stelfox 1965; Synnott 1983). Personally, the currentauthor finds this hard to credit, although there is evidence that suggests one or more nearby refugiamay have existed (see Vegetation History in The Flora of County Fermanagh) (Forbes &Northridge 2012).
The habitat of the questionable Fermanagh record at Castle Caldwell was listed as swampy fen-carrwoodland, which in view of the above is completely inappropriate for S. phylicifolia. Othermistaken 19th century records have previously been claimed for VCs W Mayo (H27), W Donegal (H35), Antrim(H39) and Londonderry (H40). The surviving vouchers for the latter are of, "remarkably poorquality" (Stelfox 1965), but have been re-determined as shiny-leaved forms of S. cinerea subsp. oleifolia (Common Sallow) (given away by the presence of afew characteristic brown hairs on the leaf under-surface), or rarely as S. myrsinifolia Salisb.(= S. nigricans Sm.) (Dark-leaved Willow), or possibly its numerous hybrids (Synnott 1983).
On these grounds, we must discard the Castle Caldwell finding of S. phylicifolia. Unfortunately,there is no voucher specimen so we cannot clarify how or why the error was made, but most probably theplant was a form of the extremely variable sally, S. cinerea subsp.oleifolia.
In Britain, S. phylicifolia is a very much more widespread species, but even here it ispredominantly a Scottish plant of waterside habitats on moist rocky ground. It is also found in Englandas far south as Lancashire and Yorkshire. It occurs at altitudes from near sea-level to 670 m in theHighlands (Meikle 1984). S. phylicifolia is widespread in Scandinavia and Iceland, where itappears indifferent to soil reaction. It is the dominant Salix species in middle and north borealriverbank communities, being frequent in abandoned damp pastures and meadows, plus along roadsides(Jonsell et al. 2000).
Native, occasional, widely scattered, but especially frequent around and to the W of Lough Erne.Eurosiberian boreo-temperate.
1884; Barrington, R.M.; Owl Island, Lower Lough Erne.
May to November.
Growth form and preferred habitats
This attractive, polymorphic, creeping, sprawling, or low but still erect or ascending, shrubby willow islocal, occasional and widely scattered in a range of acidic heath and sandy habitats in B & I, butrare or absent elsewhere. The prostrate form of the plant (var. repens, or var.ericetorum) with leaves that soon become nearly hairless (glabrous) is typically found on acidheaths, bog margins, cliffs, upland stream-sides and on stabilised scree, rocky slopes and stony orsandy, generally acidic lough shores and in nearby pastures. The more ascending growth form, var.fusca, although very much less obviously differentiated than the procumbent form, is generally aplant of lowland fen margins and stony lakeshores and can grow up to 1.5 m or exceptionally 2.0 m inheight, bearing densely covered, silky-haired leaves. It has only ever been recorded by John Harron ontwo occasions at one site, Carrigullian Lough, Co Down (H38) (Meikle 1984; BSBI Database accessedFebruary 2020).
Fermanagh occurrence
S. repens is recorded in 46 Fermanagh tetrads, 8.7% of those in the VC. However, although weregard it as occasional, it only occurs in any great quantity in two sites: around the damp, heathyshores of Lough Glencreawan, above the Cliffs of Magho (also known as 'Poulaphouca') in the Lough NavarForest Park and secondly in the hills above Florencecourt. Elsewhere, the prostrate growth form ismainly, but quite sparingly found on acid heaths, bog margins, cliff ledges, upland stream-sides androcky slopes or on stony or sandy, acid lough shores. As our tetrad map demonstrates, S. repensis widely, but rather thinly scattered across the VC, but is mainly met around both lakes of Lough Erne,plus on the shores of Lough Melvin and the shallow, heathy peatlands overlying limestone aboveFlorencecourt and the blanket bog slopes of Cuilcagh mountain. More widely it is less frequently foundon heaths, bogs, moorland tracks and cliffs on the Western Plateau.
Variation
Taxonomically, the distinction, status and correct naming of the variant forms are complicated andunclear (Meikle 1984, p. 144). In the Flora of Connemara and the Burren, for instance, Webb &Scannell (1983) found that they could not reliably correlate the distinguishing characters and separateany different forms at all. Locally in Fermanagh, however, one rather different habitat of the sprawlingplant (whether or not we distinguish it with a name) is in and out of the gryke (or grike) fissures inlimestone pavement on the summits of Knockmore and Knockninny hills.
S. repens is a very variable species with respect to small-scale environmental differences (Fowleret al. 1983). The Fermanagh forms of the plant are very different from the maritime var.argentia, whose leaves are covered with silky hairs and which is quite common on the not verydistant W Donegal coast.
In exposed upland situations, the Fermanagh plants can easily look like dwarfed specimens of S.aurita (Eared Willow), a species that may well also be present. The leaves need to be examinedcarefully to determine whether they are rugose, as is the case in S. aurita.
There are ten Fermanagh tetrads where S. repens has not been recorded in the post-1975 period,which suggests there has been some loss of suitable habitat.
Threats
None.
Native, very rare. European arctic-montane, also in N America.
1950; MCM & D; Cuilcagh summit ridge.
July to October.
Growth form and preferred habitats
The smallest willow species of B & I, S. herbacea is a dioecious, prostrate, mat-formingshrublet with an extensive branching underground rhizome. The dwarfism is very striking, aerial branchesof the shrublet rising only 5-10 cm above the surface of the ground. Woody aerial shoots can bedistinguished from the generally (but not always), subterranean rhizome, by their being segmented bybud-scale scars (Meikle 1984; Beerling 1998).
Being very low-growing, S. herbacea frequents disturbed, unstable or physically severe habitatsthat prevent or limit overgrowth by taller, more competitive species. Thus it is characteristic ofsummit plateau and other very wind-exposed, high-altitude situations, generally involving some degree ofsnow cover, frost solifluction, unstable scree slopes, rock crevices or regular sheep and goat grazing,allowing the development and maintenance of permanently open, sparsely vegetated, montane-heathcommunities.
In NW Ireland, protective snow cover is seldom present for long in winter due to the mild, wet oceanicclimate, and consequently S. herbacea occurs only very sparingly and rarely in shallow, rocky orstony, strongly acid, nutrient-poor, leached podsol soils, in fully exposed, sheep-grazed vegetation.The openness of the vegetation and the growth-limiting severity of the physio-chemical environment areessential to the montane survival of S. herbacea which, on account of its small size, isbiologically a very poor competitor (Beerling 1998). Plants confined to north-facing rock crevices areconsiderably more sheltered than those on the typical mountain summits and slopes and thanks to regular,high amounts of rainfall and their shaded situations, they are never going to suffer dehydration.
Fermanagh occurrence
S. herbacea is decidedly rare in Fermanagh, having been recorded in two tetrads only, both onCuilcagh mountain. It occurs at several spots on the summit ridge and also slightly lower at TiltinbaneTd on the northern flank of the mountain, where it grows with lichens amid sparse cover of othersubshrubs such as Erica cinerea (Bell Heather), Vaccinium myrtillus (Bilberry) andEmpetrum nigrum (Crowberry) in open, prostrate, sometimes moss-dominated montane heathvegetation.
The Robert Northridge and the current author have together seen the plant at Cuilcagh Gap, where it growson the north side of two deep clefts or gullies in the rocks of the summit ridge itself. In one ofthese, there are six flattened mats of this tiny, dwarf shrublet scattered in moist crevices across onlyabout 7 m of cliff. The plant appears able to spread in these crevices by means of its unusually woodyrhizomatous growth, but, unfortunately, we have no idea as to whether or not it manages to set seedhere, or if so, with what measure of success it establishes new plants.
Practically every aerial shoot the shrublet produces, however short it is, ends in a small, 3-9 flowered,either male or female catkin. These are sheltered and warmed between two subtending, shining, leatheryleaves. An exhaustive Biological Flora account of S. herbacea in B & I produced byBeerling (1998) suggests that seed reproduction is rare and existing clonal populations are purelymaintained by vegetative spread.
As has been found in other clonal species, both herbaceous and woody, individuals may be thousands ofyears old, older indeed than any of the most often quoted patriarch trees. These plants may befunctionally immortal (Korner 1999, p. 289). However, just how comprehensive reliance on vegetativereproduction is throughout the very wide range of this arctic-alpine species, and whether anypopulations in B & I are exceptions to this general picture, are unanswered questions.
Irish occurrence
The New Atlas map shows that these Cuilcagh stations are the most inland extant sites of DwarfWillow anywhere in Ireland, all other sites for the species being much closer to the coast. It is an oddfact of geography that the highest mountains of Ireland all lie on or close to the coast and it isessentially on the highest ground and on N-facing cliffs, sometimes at rather lower altitudes, that thistypical arctic-alpine species survives (The Botanist in Ireland). The New Atlas map forIreland displays post-1987 date-class records in 33 hexads, while both it and the previous BSBIAtlas (Walters & Perring 1962) display a total of 35 Irish hexads which had pre-1930 (andtherefore pre-1970) records. Clearly the only comforting thing about such statistics is that the lossesthat occurred prior to 1930 have not continued at the same pace, or at least not at the hectad scale. Along list of willow hybrids involve S. herbacea (Stace 1975; Beerling 1998), so that fertility isnot in question.
Fossil history
Fossil remains of S. herbacea have been found locally in Fermanagh in a full-glacial freshwaterdeposit of Middle Midlandian age, radio-carbon dated to 30,500 BP, discovered at Derryvree, nearMaguiresbridge (Colhoun et al. 1972). The flora and fauna of this deposit indicated that opentundra vegetation and a periglacial climate prevailed at the time when it was laid down. S.herbacea is thus a glacial survivor which probably spread to Fermanagh in the very earlypost-glacial period, perhaps as much as 12,000 years BP, and through changing climate and land use ithas clung on ever since. Current rapid climatic warming poses the greatest threat to its continuingsurvival, since the degree of change may well release potential competitors from their presentconstraints.
Conservation
The duty of conservation stewardship requires that the remaining small Fermanagh populations should bestudied to determine not only their extent, but to determine the ability of the species to reproduce andmaintain itself. It would also be interesting to know even an approximate estimate for individuallongevity and have some measure of rate of population turnover. Studies of this nature are alreadyunderway in Scotland and the Environment and Heritage Service in Northern Ireland should considerfollowing suit.
Threats
Dwarf Willow is protected by its small size and the remoteness and physical position of its mountainstations. Occasional grazing by sheep keeps the high montane vegetation open and some populations areprotected from erosion by shelter in rock clefts near the summit ridge. The real threat is from globalclimate warming stimulating increased competition from more vigorous perennials.
Introduction, neophyte, a very rare casual.
European southern-temperate, widely naturalised beyond its native range.
1951; MCM & D; Newtownbutler football ground car park.
May to July.
Growth form, origin and history of introduction
S. orientale is an alien neophyte annual that was first recorded as introduced to Britain in theearly 18th century as a contaminant of imported grain, very probably including hen food. It is a nativeof N Africa, Macaronesia, the Near East to W Himalaya and S Europe from Portugal to Bulgaria and Turkey(Garrard & Streeter 1983; Jalas & Suominen 1994, Map 2123). It has been widely introducedelsewhere in both hemispheres, including to B & I and on the coast of Norway to 68N. It has beenrepeatedly reintroduced to B & I in more recent times along with wild bird food seed and also as asheep wool adventive (Salisbury 1964, p. 141; Clement & Foster 1994). S. orientale was notrecorded in Ireland until the very end of the 19th century (Cat Alien Pl Ir).
Irish occurrence and preferred habitats
Although it can commonly become naturalised in parts of SE England, in Ireland, Eastern Rocket isregarded as a casual spring or winter annual which in NI only rarely becomes more or less naturalised indocks, roadsides, railways, waste ground and other disturbed, open, well-illuminated, moderately dry tomesic soil habitats, including on and around walls (FNEI 3; Urban Flora of Belfast; Hillet al. 1999). On account of this strong habitat association with commerce, transport and largeurban areas in general, S. orientale is more frequently recorded in eastern Ireland, although itreally is only regarded as a fairly common casual. Reynolds (2002) lists it as having been recorded atleast once from 25 of the 40 Irish VCs.
A very much better estimate of the species Irish occurrence is demonstrated in the New Atlas,where the hectad map displays records of S. orientale very thinly scattered on this side of theIrish Sea. Of the approximately 83 hectads of any age-class plotted for Ireland, 35 are from east coastor nearby districts.
Fermanagh occurrence
S. orientale first appeared in Fermanagh in 1951 in a football ground car park close to a railwayyard. The railway is long gone, having closed in autumn 1957. The second of our four VC recordswas made by RHN exactly 40 years later, also on waste ground in Enniskillen town. This very clearlyindicates just how rare a casual it is in our area. The two remaining records were also made by RHN atOld Crom Castle in 1995 and 1999, when the plant was found growing quite abundantly on a wall at onespot and subsequently on disturbed ground just north of the castle.
British occurrence
Eastern Rocket is very much more common in Britain than in Ireland, at least in the area S of a linebetween Liverpool and Hull, and there is a similar widespread representation around the large Scottishconurbation of Glasgow and Edinburgh (Preston et al. 2002). S. orientale appears to havefirst come into prominence during the 1940s Blitz in London, where and when, along with many otherruderal species, it rapidly colonised vacant bomb sites (Salisbury 1964).
In Britain, and possibly also in the Dublin and Belfast urban areas, S. orientale appears to havespread considerably during the last 50 years. In Britain at least, its status has graduated to itbecoming 'a persistent casual' (Rich 1991; D.A. Pearman, in: Preston et al. 2002).
Biology
The main period of germination is in autumn, the young plant overwintering as a small leaf rosette with aslender tap-root (Clapham et al. 1962). Flowers are produced over a long season, from May toSeptember or even longer in warmer, milder parts of these islands (Rich 1991; An Irish Flora1996). However, apart from this observation and mention by Rich (1991) that the flowers areself-incompatible (no reference quoted), the current author has been unable to locate any furtherinformation regarding the reproductive biology or population behaviour of the species. S.orientale is resistant to some herbicides. Clearly little or nothing is known of the seedbiology, since the species does not appear in either the meta-survey of soil seed bank data for NWEurope (Thompson et al. 1997), nor in the germination trials of Grime et al. (1981). Itthus appears that Eastern Rocket is a subject where school project observations in areas of B & Iwhere the plant is more common, might add worthwhile data to the basic biology of the species.
Threats
None, since it is a very rare casual.
Introduction, archaeophyte, occasional and thinly scattered. European southern-temperate, but so widelynaturalised it is now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
April to December.
Growth form and preferred habitats
The species is not very variable except in size and degree of hairiness, but it can appear either as asmall, erect, simple stemmed rosette-forming annual with a slender taproot, or more commonly as a tall(60-90 cm), straggly and unevenly, stiffly branched biennial, which overwinters as a much larger basalrosette of deeply lobed leaves. The more hairy examples of the plant have as Grieve (1931, p. 570)points out, "a peculiar aptitude for collecting and retaining dust". The tiny lemon-yellowflowers overtopping the very distinctive, small, appressed fruits, and the straggling, tangled habit ofthe plant together readily allow definite identification of the species (Rich 1991).
In either growth form, Hedge Mustard is almost always found near habitation in both urban and ruralsituations. It grows on regularly disturbed, dry, open, sunny sites, over more or less fertile,moderately acid to calcareous or base-rich soils (Sinker et al. 1985; D.A. Pearman, in: Prestonet al. 2002). Hedge Mustard is essentially a lowland species, almost always confined below 305 m(= 1,000 ft). Its stations are almost always closely associated with man's activities and the vegetationand soil disturbance that result (Salisbury 1964).
Fermanagh occurrence
Although Hedge Mustard has been recorded in 43 Fermanagh tetrads, 8.1% of the total in the VC, it is onlyoccasional and sporadic and it never appears in any great quantity. Although most often found nearhabitation, it can also occur in more natural or semi-natural habitats along river and canal banks.However, even in these situations, in Fermanagh, it is generally found near bridges or along paths – iebeside another man-made feature! S. officinale has been recorded, however, in a definitelysemi-natural setting on the Knockmore limestone cliffs, near the main woodland block on the slopes. Wehave almost no arable agriculture in Fermanagh, but before the advent of herbicides in other parts ofthe B & I Hedge Mustard was closely associated with such growing conditions.
Status
Although long regarded a native plant (eg Clapham et al. 1962; New Flora of the BI), thisruderal is now belatedly recognised as an archaeophyte (ie an ancient, accidental introduction), theonly fossil record available to Godwin (1975, p. 136) being a seed excavated from a Roman well!
Flowering reproduction
Like many other weedy members of the Brassicaceae (the Cabbage family), reproduction is entirely by seed,flowering taking place from April to September, but with the main flush from mid-June to late July. Theself-compatible flowers automatically self-pollinate, but are said to be occasionally visited by flies,hoverflies and small bees (Clapham et al. 1962; Proctor & Yeo 1973, p. 84), so there remainsat least some possibility of crossing, even if it is only extremely rarely achieved.
Seed production
The very numerous pods or siliquae borne on a plant each contain an average of 15, small, light,wind-dispersed seeds and a large plant can produce more than 9,500 potential offspring (Ridley 1930;Salisbury 1964, p. 275).
Seed viability and longevity
Reports on the duration of seed longevity vary from transient to long-term persistent (that is, over fiveyears), a not unusual picture of the available data obtained from the survey carried out on the topic inNW Europe by Thompson et al. (1997). However, experimental studies by Roberts & Boddrell(1983), listed in the mentioned survey, did show that in replicate experiments begun over foursuccessive years, between 2.8% and 6.8% of seed sown remained viable and dormant after burial for fiveyears (an average of 4.2%). This study also showed that most seedlings emerged in the field during theperiod March-September, with peak emergence often in July or August, some of the flushes followingcultivation. Roberts & Boddrell (1983) pointed out that this pattern of predominantly summeremergence contrasted with other published data by Chancellor (1979), who found greatest emergence duringMarch-June, with practically no seedlings appearing later. Montegut (1975) found that S.officinale germination occurred mainly in autumn and late winter.
These differing results highlighted the surprising fact that relatively little detailed study hadpreviously been made of the basic biology and ecology of this very common weed, but during the last 20years S. officinale has become, like Arabidopsis thaliana (Thale Cress), an importantexperimental organism and many research studies have been published on the complex physiologicalmechanisms and interacting factors controlling germination, dormancy and secondary dormancy in thisspecies.
British and Irish occurrence
As an ancient introduction, Hedge Mustard has become very widespread in the British Isles (being found in99% of VCs). However, the distribution thins decidedly, or is completely absent from, large areas of Cand NW Scotland. It also becomes much less frequent in western parts of Ireland at all latitudes on theisland (Preston et al. 2002). The reasons for this restricted distribution in B & I are notobvious since the range of the species in Europe extends very widely from the Mediterranean to around62°N in Norway, and indeed as an introduction in the latter, it extends to almost 65°N.
European occurrence
S. officinale is believed to be native in Europe, N Africa and the Near East (Rich 1991), butbeing so closely associated with man, it really appears difficult to trace its true centre of origin. Inthe past, S. officinale occurred in Iceland as an introduction, but did not become established.It is commonly found and distributed well into continental E Europe and Russian parts of Asia (Jalas& Suominen 1994, Map 2129). In a scattered manner, it has been introduced and become naturalisedboth eastwards and southwards in many parts of the world, including China, Japan and W & E regionsof N America. It is also naturalised in the southern hemisphere in extensive areas of S America, C &S Africa, Australia, Tasmania and New Zealand. It has been transported to very remote oceanic islandsand become established in Hawaii and in the Falkland Islands (Hultén 1971, Map 253).
Uses
As the species name indicates, S. officinale has a history of use in herbal medicine, having thereputation of being an infallible remedy for the loss of voice. A strong infusion made from the wholeplant used to be taken in the 18th century for all diseases of the throat, so that the French referredto it as the 'Singer's Plant' (Grieve 1931). The leaves have also been used as a green vegetable (Mabey1996), although the plant has also been implicated in contact-dermatitis.
In recent years, forms of Hedge Mustard in some parts of the world have developed resistance to certaintypes of herbicide used to control broadleaved weeds and currently this is a subject of researchinterest. It is still possible to control it with hormone based herbicides provided they are applied atthe seedling stage.
Names
The generic name 'Sisymbrium' is derived from the Greek 'sisymbrion' and was an ancient name referring toWatercress and possibly other species of the Brassicaceae, together with species of aromatic herbs ofthe Lamiaceae considered sacred to Venus (Gilbert-Carter 1964; Hyam & Pankhurst 1995).
The Latin specific epithet 'officinale' indicates that the plant was once listed as medicinal and sold byapothecaries in their 'shops' ('officina' in the singular) (Gilbert-Carter 1964).
The species has numerous English common names, several of which are far more interesting than the currentboring 'Hedge Mustard'. They include 'Hedgeweed', or 'Bank Cress', from the frequently encountered hedgebank habitat of the plant. Prior (1879) includes 'Crambling Rocket' which he explains as being like,"a spurious Crambe or mustard, (as vetchling is a spurious vetch", plus the fact thatthe plant has, "leaves of rocket". Britten & Holland (1886) point out that one of theplant's synonyms was Eruca peregrina, so that there seems little doubt that 'crambling' is to beunderstood as a corruption of 'scrambling', or wandering 'rocket', and the name was also sometimesapplied to quite a different plant, Reseda lutea (Wild Mignonette). Another interesting namelisted by the latter authors is 'Lucifer Matches', which originated in Worcestershire, from the effectof the small clusters of pale yellow flowers resembling the sulphur used in the manufacture of matches(Melderis & Bangerter 1955). When the plant was introduced to Tasmania, the characteristic stiff,tangled side branches gave rise to the rather apt local name 'Wiry Jack' (Salisbury 1964).
Threats
None.
Native, locally frequent. European temperate, but widely naturalised.
1872; Stewart, S.A.; Enniskillen Town.
Throughout the year.
Growth form and preferred habitats
This tall winter annual, but more generally rosette-forming monocarpic, tap-rooted biennial, oroccasionally short-lived perennial can reach a height of 120 cm in woods or along ± open, butsemi-shaded, occasionally disturbed, linear habitats, often forming dense stands. It is often mostabundant on damp, shaded soils although it can cope with a very wide range of soil moisture content(Cavers et al. 1979). The white racemes of flowers are extremely conspicuous in April, May andearly June. Garlic Mustard has variable, round or heart-shaped, mainly stalked, bluntly-toothed leavesup the stem, which smell strongly of garlic when crushed.
A. petiolata appears most abundantly on roadsides, hedgerows and woodland margins, where the morefertile the soil, the denser and taller the stand it develops. It is interesting that A.petiolata can also tolerate very much drier and less fertile conditions in sand or gravelquarries and on scree slopes. The inherent disturbance of these types of unstable sites is clearlysignificant, and presumably it must assist the species survival by limiting the growth of potentiallyvigorous competitors.
Moderate disturbance is experienced by A. petiolata when roadsides are occasionally mown, andalong stream and river banks where the ground is kept sufficiently open by seasonal flooding. Thespecies is probably completely excluded only from the most acidic or the very wettest soils, or fromgreatly disturbed situations of ploughed, closely grazed or heavily trampled agricultural ground (Caverset al. 1979; D.A. Pearman, in: Preston et al. 2002).
Flowering reproduction
The biennial form of the plant typically spends its first year of growth slowly developing a basal leafrosette. However this is followed in the very early spring of the second year by a period of rapidflowering shoot elongation (ie bolting), of up to 1.9 cm/day. It has been suggested that the presence ofglucosinolates and a range of similar compounds in the tissues of A. petiolata may permit thespecies to grow vigorously at near freezing temperatures in late autumn, and again in early spring(Cavers et al. 1979).
In the spring of the second year of growth, plants initiate inflorescence production, and they flowerfrom early April, reaching a peak of flowering around mid-May. Flowers remain open for two, or veryrarely three days, but they produce nectar and are visited by insects mainly in day one. The species iscapable of both cross- and self-pollination, and pollen release and stigma receptivity occur before theflower bud opens. Thus, while flowers are visited by a range of midges, short-tongued bees and flies forfood and pollination, autogamy might appear to be the most likely breeding system (Anderson etal. 1996). Self-pollination can occur as early as mid-afternoon on day one. Older flowers havelittle or no nectar present, which deters insect visitors (Cruden et al. 1996). However,self-pollination is not automatic, and indeed there is evidence that selfing is biologically delayed,presumably to favour out-crossing. A high degree of out-crossing would enable rapid adaptation of thepopulation to local growing conditions (Clapham et al. 1962; Cavers et al. 1979; Crudenet al. 1996).
Mature plant size and seed production varies enormously within each type of habitat the species occupies.The long slender fruit capsules (siliques), ripen and split to release seed from the middle of Julyonwards to around early October. All the flowering plants die off in the autumn, and dormant seedoverwinters to germinate in early spring (usually concentrated in April).
Seed production
Mature plant size and annual seed production varies enormously within each type of habitat the speciesoccupies, but it is considerable when, as they often do, plants grow in dense stands. Measured seedproduction ranges between 15,000 in woodland to 107,000 seeds/sq m in more open conditions (Cavers etal. 1979).
Seed dispersal
There is no specialised seed dispersal mechanism. Wind dispersal is very ineffective and the seed doesnot float well either. However, seeds stick together when damp and they readily adhere to small crumbsof soil. Thus it is very probable that the major means of seed dispersal involves epizoochory (ieexternal adherence on passing animals), including man and his machines (Cavers et al. 1979).
The high rates of seed set and production typical of the species often enable A. petiolatapopulations to colonise fresh sites in suitable vegetation around existing individuals. In several partsof N America it has become a major alien invader, ousting native species from what appears to berelatively undisturbed woodlands (Cavers et al. 1979; Anderson et al. 1996). On the basisof similar evidence in England, Grime et al. (1988) classified its reproductive strategy as a'competitive-ruderal' species.
Seed dormancy and germination
Dormancy is controlled by physiological features of the embryo. Cold stratification at temperaturesbetween 1° - 6 °C for a period between 60 - 97 days is required to break dormancy (Lhotská 1975). Mostseeds germinate in the spring after they are produced following exposure to winter conditions, but someviable seeds are retained in the seedbank and germinate over a period of 3 to 4 years (Cavers etal. 1979; Baskin & Baskin 1992). In several studies the range of seedlings survivingto flowering maturity varied from 2 - 7.5 %, the great majority of plants dying off at the seedlingstage during dry periods in May (Cavers et al. 1979; Anderson et al. 1996).
Vegetative reproduction
Salisbury (1964) mentions that A. petiolata, like several other English hedgerow plantsincluding Hypericum perforatum and Pimpinella major, is capable of a certain degreeof vegetative reproduction. Should the flowering stem become damaged early on in the second season ofbolting growth mentioned above, the plant may form secondary shoots from adventitious buds on the rootsto replace the injured one. The frequency of this occurrence is probably not very great, and itssignificance should not be overestimated, indeed, from the evidence reported by Cavers et al.(1979) in Canada, it may not happen at all in some populations.
Fermanagh occurrence
In Fermanagh, A. petiolata is locally frequent and has been recorded in 93 tetrads, representing17.6% of the area. Garlic Mustard is most commonly found scattered throughout the VC in lowland, shaded,somewhat disturbed ground involving moderate to high exposures of bare soil. It grows best in fertile,moist situations. The fact that ten scattered tetrads have pre-1976 records only is not regarded assignificant in view of the biology and ecology of the species.
British and Irish occurrence
In Ireland, A. petiolata is much more thinly scattered, eastern and central in its distribution incomparison with the much more widespread occurrence in Britain (Perring & Walters 1976; Prestonet al. 2002). Allen (1984) described the plant as calcicole in Ireland, but although it is morewidespread on the limestone districts in the south of Co Dublin (Colgan 1904), little else seems tosupport this notion, and the current author prefers to consider it a species of fertile, moderatelybase-rich soils.
The New Atlas hectad map shows that in Britain A. petiolata is much less frequent or absentfrom the more strongly acid conditions prevalent in the N & W of Scotland, and on wet, infertile,heavily leached high ground throughout.
European and world occurrence
A. petiolata is native, common and widespread throughout Europe and eastwards to Asia Minor andthe Himalaya. It is also considered native along the N coast of Africa. The European distributionextends in an increasingly scattered manner northwards up the west coast of Norway to around 68°N, andit thins markedly southwards in Greece. Garlic Mustard is absent from Crete, the Balearic Isles andSardinia, although it is present in Sicily (Jalas & Suominen 1994, Map 2139). It also extends(native or not), to India and Sri Lanka, and is a definite introduction in NE parts of Canada, theUnited States and Australiasia (Cavers et al. 1979; Anderson et al. 1996; Rich 1991).A. petiolata is considered a rather invasive alien weed in the eastern states of N America. Itwas also introduced to one site in New Zealand at the end of the 19th century, but it only survivedthere for about ten years (Webb et al. 1988).
Uses
Like other wild crucifers, Garlic Mustard has been used as a condiment, boiled and eaten with mutton, asa salad, or to make various sauces, so that one of the fifteen alternative English names listed byGrigson is 'Sauce Alone', apparently originating in Somerset (Grieve 1931; Grigson 1987). The leaves andtop of the plant just before flowering have a higher value of vitamin C, on a weight basis, thanoranges, and the leaves at all times of year contain more vitamin A than spinach (which has the highestlevel of all the widely marketed garden vegetables) (Zennie & Ogzewalla 1977). The leaves arealso reported to contain unusually high concentrations of N, P, Ca, Fe and Na (Survey of InorganicElements in Foliage: J.P. Grime & S.R. Band unpublished - quoted in Grime et al.1988, p. 70).
In herbal medicine, A. petiolata leaves were used internally as a sudorific (ie to promotesweating), and as a deobstruent to relieve obstructions of the intestinal tract. It was also appliedexternally as an antiseptic treatment for gangrene and ulcers. Leaf juices taken alone, or boiled into asyrup with honey, were also used to treat dropsy (Grieve 1931).
The plant is the specialist food plant of the caterpillars of both the Orange-tip and Green-veined whitebutterflies (Garrard & Streeter 1983).
Names
The genus name 'Alliaria' was given to the plant by the herbal writer Fuchs, referring to thegarlic-like smell of the crushed plant, and is derived from 'Allium' or 'Alium', theclassical name for Wild garlic (Gilbert-Carter 1964). The Latin specific epithet 'petiolata' translatesas 'furnished with a leaf-stalk', and usually means the plant involved possesses a particularly longpetiole (Stearn 1992).
The English Common name 'Garlic Mustard' is given on account of the fact that its rather variable leavesdo smell rather strongly of garlic when they are crushed, as indeed does any part of the plant. There isno odour unless the plant is crushed. Another well known name is 'Jack-by-the-hedge', and there areeight similar references to the hedgerow habitat in the names listed from around B & I by Grigson(1987). One of the more interesting names given by Britten and Holland (1886) is 'Swarms' fromYorkshire, possibly referring to the dense, tall stands the plant regularly produces on fertile ground.
Threats
None.
Native, occasional and uncommon, but very possibly a recent arrival and still spreading. Eurosiberiantemperate, but so widely naturalised it has become circumpolar.
1976; Dawson, Miss N.; Kesh village.
March to December.
Growth form and preferred habitats
A small, slender, variable, rather straggly grey-green leaved, winter- or summer-annual weed, or veryoccasionally a biennial (Salisbury 1964), with unexciting tiny white flowers in a raceme up to 30 cmtall, Thale Cress sporadically colonises bare ground in numerous types of open, disturbed, usuallyunshaded, acid, lowland habitats where there is little or no plant competition. It most frequentlyoccurs on dry or well-drained, sunny, light sandy, or sometimes stony soils in wayside situations.Populations are seldom large. It thrives in well-drained soils, but is prone to phosphorus deficiencies.Poor nutrition frequently causes reduced height, induces early flowering and causes low set set (Wilsonet al. 1991). Salisbury (1964) reckoned that ripe seed could be produced within four weeks ofgermination. Typical habitats include on rocks, walls and ledges, in river shingle, in pavements, onwaste ground, along the edges of paths and roadsides, or in garden plots and pots.
Flowering reproduction and use in genetic and physiological research
A. thaliana is a small, self-compatible crucifer with a relatively simple genome comprising 2n=10chromosomes. In suitable growing conditions, it displays a rapid life cycle that makes it an ideallaboratory organism and it has been very extensively used for plant molecular genetics and physiologicalstudies. Some races of the plant can flower within two or three weeks of germination and set seed aftera further two weeks, so generations can be rapidly advanced. While individual plants will set seed byself-pollination, seed can also be obtained by experimental cross-pollination by hand. Because a singleplant can produce thousands of seeds in less than eight weeks it is very easy to create segregatingpopulations and to amplify mutant seed stocks in experimental studies.
A. thaliana has the smallest plant genome known to science it being only five times that of theyeast genome. A majority of the 76 mapped mutations confer readily apparent phenotypes such as loss ofsurface wax, leaf morphology or colour, siliqua morphology, growth habit, seed colour or photo-periodicresponse (Estelle & Somerville 1986). Projects are in progress to isolate genes of potential use toplant breeders and include genes for disease resistance and male sterility (which ensures seed occurs bycross- rather than self-fertilisation). It is hoped that these genes may be used to induce similartraits in plants of commercial value (Wilson et al. 1991).
In the wild, a proportion of flowers are visited by short-tongued insects including hover-flies, midgesand bees and cross pollination and fertilisation achieved. However, insect visits probably are rare,being limited by the presence of only rudimentary floral nectaries (Wilson et al. 1991). Anynectar that is produced is confined to the first day of opening (Cruden et al. 1996). Either way,seed production is usually described as 'copious', each fruit siliqua (or silique) normally releasing30-60 seeds. A study by Salisbury (1964) in England found a mean seed number of 33 seeds per cylindricalseed pod. A mature plant under favourable growing conditions can produce around 200 siliquae and thusgive rise to several thousand small seeds, each less than 1 mm long (Lawrence 1976).
In common with many other members of the cabbage family, the A. thaliana seed coat possesses amucilaginous layer, so that it sticks to surfaces when wetted, and can thus achieve some measure ofsecondary dispersal in mud or on tyres or other passing objects.
Seed dormancy, germination and longevity
Physiological strains of A. thaliana occur that display differing photo-periodic responsesdetermining the timing of germination. Seed from wild populations of A. thaliana tends to remaindormant, failing to germinate without a prior cold period of overwintering stratification, thus behavingas spring annuals. However, some plants behave as winter annuals, germinating in the autumn after aperiod of warm weather, overwintering as leaf rosettes and then flowering and seeding in the earlymonths of the following year. This growth pattern avoids possible drought periods which the fibrous rootsystem of the plant probably would not survive. Other physiological forms of the plant do not requirecold treatment to germinate or induce flowering, and they behave as summer annuals. The differing formsmean the species can produce seed almost all year round, although Thale Cress remains most common in thespring. Most of the plants used in laboratory research studies of molecular biology are summer annuals(Wilson et al. 1991).
A persistent seed bank is also formed in cultivated soil, an unknown proportion of buried seed survivingfor up to five years and serving as a buffer against local extinction (Lawrence 1976; Roberts &Boddrell 1983; Grime et al. 1988; Thompson et al. 1997). In terms of its ecologicalstrategy (a measure of its competitive ability), A. thaliana is classed as a stress-tolerantruderal by Grime et al. (1988).
Fermanagh occurrence
It is hard to believe that this currently widespread weed was not recorded anywhere in Fermanagh until1976. Currently we have records from 62 tetrads, over 11.7% of the squares in the VC. It is thinlyspread in suitably open, disturbed habitats throughout the lowlands, but with a noticeable linearroadside pattern. Prior to 1976, A. thaliana was known to occur elsewhere in NI on the exposed,sloping, rocky ground of the basalt scarps in Cos Antrim (H39) and Londonderry (H40), and in other openhabitats providing sunny, dry soils.
The previous lack of Fermanagh records might possibly have been a consequence of its relatively earlyspring flowering maximum – ie generally before any visiting field botanists became active in the county.The majority of A. thaliana plants (certainly those in the more nearly natural habitats whichmight possibly be distinct ecotypes), follow a winter annual reproductive strategy with a rapidflowering and fruiting cycle from April to June (Grime et al. 1988; Rich 1991). In addition,Thale Cress frequently occurs as isolated individuals or in small quantity and only rarely does itbecome abundant, even in disturbed fertile garden soils.
Taken together, this behaviour might allow the species to be overlooked. However, the spreading, elongatedead seed heads are very distinctive and long persistent, making it rather difficult to imagine that ifthe species were previously present, no one in Fermanagh noticed it about the garden, or in otheropen, urban areas, including roadsides and waste ground earlier than 1976. Indeed, like other small,annual cruciferous weeds, when it appears in the most disturbed habitats the plant is free of day-lengthand seed vernalisation constraints, and it can germinate and flower virtually all year round (Grimeet al. 1988).
These arguments lead Robert Northridge and the current author (RSF) to conclude that the apparent recentarrival and spread of Thale Cress must be real, and it could possibly be associated with the riseof Garden Centres around the county in the 1970s and 1980s. A. thaliana is strictly anon-competitive pioneer therophyte and, like Cardamine hirsuta (Hairy Bitter-cress), it is verymuch associated nowadays with garden and nursery cultivation, especially on light, sandy or stony, oftencalcareous and generally shallow, dry soils (Sinker et al. 1985; Rich 1991).
British and Irish occurrence
A. thaliana is common throughout most of Britain, thinning northwards into Scotland, butabsent from a few areas on the E coast of England. In Ireland, it is much more frequent in the easternhalf, becoming scarce or absent towards the W of the island (New Atlas). It is probably stillspreading in Ireland, possibly with the assistance of the horticultural trade.
European and world occurrence
In common with numerous other members of the ruderal weed community, A. thaliana is thought tohave most probably originated in temperate Eurosiberia where it is very widespread. A.thaliana is regarded as native throughout most of Europe and the Mediterranean basin (includingmost islands), the Macronesian isles, N Africa and the Far East, Asia and E Africa (Rich 1991; Jalas& Suomnen 1994, Map 2141). Thale Cress has spread along with man and his agricultural systems andbecome naturalised and Circumpolar in its distribution. It has been introduced in N America, C & SAfrica, S Australia and New Zealand (Hultén & Fries 1986, Map 909; Webb et al. 1988). Rathersurprisingly, in the NE United States and adjacent Canada it is regarded as a pernicious, aggressive,invasive weed of moist woodlands (Cruden et al. 1996).
Names
The genus name 'Arabidopsis' is Greek meaning 'like or resembling Arabis' (Gilbert-Carter1964). The Latin specific epithet 'thaliana' may either be derived as a memorial to the GreekMuse Thalia, who personified 'luxuriant, blooming' features (Melderis & Bangerter 1955), orDr Johannus Thal (or Thalius), a sixteenth century German physician whom Linnaeus commemorated in thegiven name since he had published a catalogue of the plants of the Hartz mountains (Gilbert-Carter 1964;Gledhill 1985; Prior 1879). Druce (1932) gives the first record of the species (which he callsSisymbrium Thalianum (L.) Gay), as 'Johns. Merc., 59, 1634', which refers to Thomas Johnson(1595/1597-1644), and his account of one of his botanical journeys published in 'Mercuriusbotanicus' in 1634, the year after his famous greatly improved edition of 'Gerard'sHerball' (Henrey 1975, Volume 1, p. 94).
English Common names for the plant include 'Thale Cress', which is a mere book-name translation byLightfoot of the Latin epithet (Britten & Holland 1886), or 'Common Wall Cress', 'Mouse-ear Cress'(Wilson et al. 1991), or just plain 'Wall Cress', although this name is more usually given tospecies of Arabis, another genus within which this species has occasionally resided in the past.
Threats
None to it, since it appears to be on the increase.
Introduction, archaeophyte, a very rare casual. A widely naturalised species with a circumpolarboreo-temperate distribution.
1899; West, W.; Florencecourt Railway Station (now defunct and gone).
Growth form and preferred habitats
Treacle-mustard is a spring or autumn annual of open, well-illuminated conditions. It prefers light orsandy, medium dry to moist soils, but avoids more acidic ground (Fitter 1987). It can be frequent inarable fields, but is commonly found on lowland waste ground, including on walls, docks, roadsides,along railways and in gardens, often present in only small numbers (Rich 1991).
Flowering reproduction
Flowering occurs from June to September, or even for longer in milder areas. While the yellow flowers doproduce nectar and insects visit them, essentially the species is automatically self-pollinated (Claphamet al. 1962; Garrard & Streeter 1983; Webb et al. 1996). Well-grown plants can eachproduce around 15,000 seeds, but in N America where this weed is common, the average productivity isgreater at around 25,000 seeds per plant. The seeds are very bitter tasting, yet horses and cattle eatthe plant and its fruit, since seed has been recovered from their droppings, indicating some degree ofsecondary dispersal takes place (Salisbury 1964, pp. 66 & 102).
Seed survival and germination
Buried seed of Treacle-mustard can survive for up to six years in soil (Salisbury 1964, p. 328). Roberts& Boddrell (1983) found that about 3% of seed were dormant and viable after five years burial.Almost all seedlings emerge during spring and summer with a peak usually occurring in March or April.Lauer (1953) recorded spring and summer emergence in Germany, and included E. cherianthoides inthe group of species requiring relatively high temperatures for germination. Treacle-mustard issusceptible to modern herbicides and consequently it probably has declined in B & I during the past50 years as agriculture has intensified (Rich 1991).
Fermanagh occurrence
There are only three records of this weed in the Fermanagh Flora Database, all from the mid-south of theVC. In addition to the first record listed above the details are: Gortaree gravel pits, Slieve Rushen,1948, MCM & D; disturbed ground at Farnaght, SE of Tamlaght, 21 August 1989, RHN.
Irish occurrence
In N Ireland, Treacle-mustard is an exceedingly rare casual weed of disturbed lowland arable or wasteground. The New Atlas hectad map for Ireland shows it much more frequently represented S of aline between Dundalk and Galway and, especially so, in more inland Midland counties and around portdocklands. Nevertheless, Reynolds in Cat Alien Pl Ir describes its Irish occurrence as being,"a fairly widespread but uncommon casual or arable weed ... nowhere thoroughly established".
British occurrence and status
E. cheiranthoides is frequent in SE England, occasional in the Midlands, SW England and in coastalparts of Wales. Elsewhere in Britain it is rare and casual − although it has been recorded in ascattered manner from the whole range of latitude (Preston et al. 2002).
The native range of E. cheiranthoides worldwide has been a matter of speculation and doubt forsome time. It had long been given 'native' or 'doubtfully native' status in B & I, this at leastapplying to its stronghold in SE England (Jalas & Suominen 1994, Map 2244). With reservations, itwas still accepted as native by Clement & Foster (1994) in their book Alien Plants of the BritishIsles. After recent reassessment it is now considered an ancient introduction (ie anarchaeophyte), fossil evidence proving its presence in both Bronze Age and Roman settlements (D.A.Pearman, in: Preston et al. 2002).
In common with Sisymbrium orientale (Eastern Rocket) it is regularly (but probably as seedtechnology improves) increasingly rarely reintroduced as a contaminant of grain, animal feed and pastureseed mixtures, particularly those involving Clover from N America (Rich 1991; D.A. Pearman, in: Prestonet al. 2002; Cat Alien Pl Ir).
European and world occurrence
Beyond our shores, E. cheiranthoides is regarded as 'probably native' in E Europe, Siberia andwestern N America, but it has become so widely naturalised that it has colonised W Europe from 68Nsouthwards to N Africa and has developed a circumpolar boreo-temperate distribution (D.A. Pearman, in:Preston et al. 2002).
Names
The genus name 'Erysimum' is a classical one, given to some plant, possibly of the same family, byTheophrastus, the so-called 'Father of Botany'. It is also possible to derive the name from the Greek'eryo' or 'eruo', meaning 'to draw up' or 'to draw out', the name then supposedly alluding to theblister raising properties of some members of the genus (Johnson & Smith 1946; Gilbert-Carter 1964;Hyam & Pankhurst 1995). The specific epithet 'cheiranthoides', means 'like or resemblingCheiranthus', that is, 'Wallflower-like' (Hyam & Pankhurst 1995).
The English common name 'Treacle-mustard' from the 19th century onwards has been applied to this speciesbut formerly it was a name given by Turner (as 'triacle mustard'), to either Lepidium campestre(Field Pepper-wort) (Britten & Holland 1886), or more likely, Thlaspi arvense (FieldPenny-cress) (Stearn 1965; Grigson 1974). 'Thiacle' came from the Greek 'theriake', meaning 'antidoteagainst a poisonous bite', so that the name 'Treacle-mustard' meant, 'mustard for a treacle or remedy'(Grigson 1974). According to Prior (1879), 'Treacle-mustard' or 'Treacle wormseed' was one of 73ingredients used in making 'Venice treacle', a famous vermifuge and antidote to all animal poisons,which was in great vogue during the Middle Ages. Britten & Holland (1886) list another name,'Tarrify' from the Cambridgeshire Fens, given, they supposed (on unfathomable grounds), "because itterrifies the farmer or the weeder".
Uses
In a recent dictionary of plant names, Watts (2000) comments that E. cheiranthoides is certainly avermifuge (ie a wormer), "though it can be quite a dangerous one". AS WITH ALL SUCH HERBALMEDICINES, EXTREME CAUTION AND USE ONLY UNDER PROPERLY QUALIFIED SUPERVISION IS ESSENTIAL.
Threats
None.
Introduction, archaeophyte, a very rare garden escape or discard.
1980; Northridge, R.H.; on an old wall in the W of Enniskillen town.
Growth form, preferred habitats and likely Centre of Origin
Nowadays, this is a rather unfashionable garden perennial, but it is a colourful, traditional subjectnevertheless and has been in continuous horticultural use in B & I since at least the 13th century(Harvey 1990). The Wallflower's garden popularity may possibly extend right back to the Roman period inBritain, since the species, which is considered a native of Greece and the E Mediterranean is veryprobably of hybrid origin (Snogerup 1967). Garden varieties are very variable in flower colour and size,but naturalised plants usually lie within the yellow, red or orange spectrum (Rich 1991). The plantreadily naturalises itself on walls, cliffs and rocks when introduced to fresh territories throughout C,W & S Europe.
British and Irish occurrence
Naturalisation of E. cheiri commonly occurs in the southern two thirds of Britain and inmost of Ireland, but the species becomes rare and decidedly coastal the further north one travels, whichsuggests it is near its lower temperature limit (Garrard & Streeter 1983; Clement & Foster 1994;Preston et al. 2002). Having said that, the New Atlas hectad map indicates a presence justN of Inverness (VC 106)!
Reynolds (Cat Alien Pl Ir) lists E. cheiri as being recorded at least once (but sometimesindeed just that), from 33 of the 40 Irish VCs, which does not include Fermanagh (H33).
Fermanagh occurrence
As far as the current author can tell, E. cheiri only very rarely escapes from gardens in thecounty. One of the only two records in the Fermanagh Flora Database (both discovered by RHN) was indeedgrowing in the lime mortar of an old wall in Enniskillen. The second record was found in Newtownbutlervillage in 1988. In the FNEI 2, Praeger commented that in Cos Down (H38) and Antrim (H39),Wallflower showed a distinct preference for ancient walls, eg those in castles and tower houses.This reflects the affinity the species has for calcareous substrates (Rich 1991), the preference beingmet by the crumbling lime mortar of these types of old walls.
The paucity of E. cheiri records in Fermanagh is probably misleading, but the extent of anyunder-recording due to prejudice against introductions is unknown. We believe that earlier recorders(including many who were employed on conservation surveys from the 1960s onwards), considered gardenescapes and other aliens as transient, casual organisms of no conservation significance and thereforeunworthy of note. If this was their thinking they would be doubly wrong: E. cheiri is a perennialwith a woody stem base which makes it not only tolerant of dry soils of very indifferent fertility, butalso allows it, once established, to be remarkably persistent − always provided the most essentialgrowth requirements of the species are met − a warm sunny spot with little competition.
European and world occurrence
Having been in cultivation for many centuries, the precise origin of this species is unknown, but veryprobably E. cheiri is of hybrid origin, linked to several Aegean species. E. cheiri ismapped as an archaeophyte introduction across parts of W Europe − mainly in France, C Italy, Britain& Ireland. The distribution becomes increasingly scattered southwards towards the Mediterraneanbasin and it is absent from Denmark, Scandinavia and all of N & E Europe (Jalas & Suominen 1994,Map 2172).
Threats
None.
Introduction, neophyte, an occasional, widely scattered garden escape.
1899; West, W.; Derrygonnelly village.
April to November.
Growth form and preferred habitats
This attractive, large and distinctive perennial or occasional biennial with its often 100 cm tall, erectleafy stems and night-perfumed white, pinkish-lilac or purple flowers is a common garden escape.
It colonises open, disturbed habitats on waste ground, in old quarries, woods, hedgerows, roadside vergesand damp streamsides and riverbanks. In B & I, it is never found far from habitation (Rich 1991).
Fermanagh and Irish occurrence
Dame's-violet is widely scattered in Fermanagh. It has been recorded in 31 tetrads, 5.9% of those in theVC. Robert Northridge and the current author consider it uncommon and occasional in suitable open,disturbed ground near gardens, past or present. While catalogued as having occurred at least once in 36of the 40 Irish VCs, it is regarded in Ireland as widespread throughout – but rarely well-established inthe wild (Cat Alien Pl Ir).
Until recent decades all such garden escapes, and indeed most other definite aliens, have tended to beunder-recorded in many parts of B & I due to a widespread and lingering prejudice against them feltby earlier generations of botanists (eg Praeger 1901). For this reason we believe that Dame's Violet is(or was) probably somewhat more frequent and widespread in Fermanagh than the survey data and the tetradmap suggests.
British occurrence and status
A native of S Europe and W Asia, H. matronalis has a very long history of decorative gardencultivation and has been introduced for this reason to many parts of the globe. It has been grown ingardens in B & I "from earliest times" according to Genders (1971), and documentary proofshows this means at least since 1375 (Harvey 1981). Despite its early garden introduction, Dame's-violetwas not recorded from the wild in Britain until 1805, making it a neophyte here. The New Atlashectad map indicates H. matronalis is a very frequent and widespread garden escape in Britain,although more commonly recorded from some areas than others. Northwards it becomes less prevalent, orindeed absent in parts of C & NW Scotland. While it is often well-established, this only happens inrelatively open, preferably damp or shaded sites, where competition remains slight (D.A. Pearman, in:Preston et al. 2002).
Flowering reproduction
The numerous flowers produced from mid-May to August possess a powerful scent, especially at night. Theperfume is almost absent during the day. It smells rather similar to the Sweet Violet (Violaodorata) and hence the English common name 'Dame's-violet'. However, it also carries spicyundertones of cinnamon, since cinnamic alcohol is present. The perfume and the generally light flowercolour attracts night-flying Lepidoptera (moths), though the flowers are also capable ofself-pollination, the anthers of older, unvisited flowers bending inwards to cover the stigma withpollen (Genders 1971). The current author has found little else definite about the floral or populationbiology of the species in the literature.
The spreading, long, thin, torulose (twisted) fruits are indented or constricted like a row of beads and,when ripe, they split to release the seeds (Rich 1991). The seeds are light and are readily dispersed bywind. Like the Wallflower (Erysimum chieri) they are sufficiently numerous to be able regularlyto jump the garden wall (Ridley 1930; Salisbury 1942). Germination is immediate and there does notappear to be any data on a buried soil seed bank (Thompson et al. 1997).
European and world occurrence
Variation in H. matronalis is such that its taxonomic status is often described as 'unclear' andJalas & Suominen (1994) refer to it as a species group, ie consisting of several related species andsubspecies whose relationships require further study. Of course this makes mapping the supposedly nativeoccurrence of the plant difficult, to say the least, yet their map for H. matronalis in the broadsense indicates a distribution stretching from N Spain, S France, Italy and the Balkan states to the Ncoast of the Black Sea and further NE into continental Russia (Jalas & Suominen 1994, Map 2252).This map also indicates just how widespread the naturalised garden plant is in the rest of Europe, andthe species (or species group) is also described as widely naturalised in N America and Australasia(Rich 1991).
Uses
Grieve (1931) appears to confuse H. matronalis with Eruca sativa (Garden Rocket or SaladRocket), and apart from her comment on the perfume (that the plant represents deceit in the ancientlanguage of flowers, since it gives out a lovely perfume in the evening, but in the daytime has none),she should be ignored on the edible or medicinal uses of this species.
Names
The genus name 'Hesperis' is an ancient name given by Theophrastus to a plant flowering in the evening,being derived from the Greek 'hespera', meaning 'the evening' (Gilbert-Carter 1964; Stearn 1992). TheLatin specific epithet 'matronalis', means 'of or belonging to a married woman' ('matrona')(Gilbert-Carter 1964). The dame in question almost certainly refers to Saint Mary, the Madonna.
The English common name 'Dame's Violet' was first used by Lyte (1578) and is a straight translation ofthe botanists' Latin 'viola flos matronalis', which Grigson (1974) points out was in turn a translationof a Greek name in Dioscorides herbal for some such (ie unknown) plant. The original Dioscoridean nameindicated a plant used by the herbalist to treat diseases of women. Another suggestion of the origin isthrough an error associated with a similar name 'Damask Violet', for Viola damascena fromDamascus in Syria. This plant was named in French 'Violette de Damas' and misunderstood as 'Violette desdames' (Prior 1879).
Sixteen English common names are listed by Britten & Holland (1886), including 'Damewort' and 'Dame'sGilliflower' (Britten & Holland 1886). Surely the most ridiculous name ever given to any plant mustbe 'Close Sciences', perpetrated on this plant by Gerard (1633, p. 463). Prior (1879) gives us an ideaof how it may have developed from another name for H. matronalis, the single form of the flowerbeing called 'Single Sciney', and the double variety 'Close Sciney' in Parkinson's (1640) TheatrumBotanicum (p. 628). 'Sciney' probably arose from the one time specific name 'Damascena',understood as 'Dame's Scena', and then corrupted to 'Sciney'. Gerard excelled himself and made the namefor the double form into his 'Close Sciences'!
Threats
None.
Native, frequent and locally abundant. Eurosiberian temperate, but widely naturalised including inFennoscandia, N America and C Africa.
1881-2; Barrington, R.M.; Derryargon Td, NW of Enniskillen.
March to December.
Growth form and preferred habitats
B. vulgaris is a widespread, sometimes abundant biennial (infrequently a winter annual), or ashort-lived perennial weed that avoids competition by frequenting disturbed habitats. The very dark,shining basal rosette leaves, shallowly lobed stem leaves and deep to bright yellow flowers with aslender style make Winter-cress a distinctive wintergreen rosette-forming crucifer, readily enoughdistinguished from B. intermedia (Medium-flowered Winter-cress), which is the onlyother member of the genus widely found in N Ireland (Rich 1987a; An Irish Flora 1996; NewAtlas).
Typically, it grows in disturbed ruderal sites such as roadsides, waste ground, farmyards, aroundquarries and on damp, winter-wet soils in field margins, ditches and beside woodland paths, or othershaded or semi-shaded waysides or waterside habitats. The plant develops a stout taproot which oftenexceeds 50 cm in depth, and it also produces an extensive branched fibrous secondary root system thatallows it to readily cope with dry soil environments.
Winter-cress has no specific substrate requirements and will grow on a very wide range of soil texturesand levels of fertility. It is absent mainly from the more extreme acidic soils and also from bothvigorous closed-turf grassland (where it cannot compete effectively) and full-canopy woodland (whereautumn leaf litter cover encourages slugs which devour its young seedlings). Essentially it is anopportunistic coloniser of open habitats and individuals of B. vulgaris grow most rapidly, livelongest and reproduce most prolifically on recently disturbed, moist, fertile soil (MacDonald& Cavers 1991).
As with poppies (Papaver spp.), a widespread persistent seed bank exists allowing B.vulgaris to often appear in a recurrent manner, sometimes in great abundance on freshlydisturbed ground, eg along road widening schemes and by recently cleaned or canalised waterways.
Weedy properties
The weediness of B. vulgaris is based on a combination of three factors: (a) the ability toincrease numbers rapidly through high rates of seed production (ie there is an estimated seed productionof between 40,000-116,000 seeds per plant in a hayfield situation, or 88,000 per plant in a tilled,weeded site (Kott 1963 (in Russian), quoted in MacDonald & Cavers 1991); (b) inedibility ofthe rachis of the flowering stem at maturity, due to the presence of toxins; and (c) long-term seeddormancy - surviving over five years burial in soil (Roberts 1986), and according to MacDonald (1977),it can survive for at least 10-20 years, thus permitting intermittent recruitment to the population whenconditions are favourable for germination and establishment.
Temperature as a limiting factor
The geographical range of Winter-cress is limited to areas where its obligatory requirement for seedvernalisation is met. This requires exposure to temperatures of 5°C or less for several weeks to breakseed dormancy, which itself is imposed in the autumn after seed production by either drought and/or cooltemperatures, ie below 10°C. This cool temperature requirement for successful overwintering seedsurvival sets a limit to the southern spread of the species, so that recurrent populations are bestmaintained in areas with periods of low rainfall in late summer and cool temperatures from late summerto early spring.
Rosettes of B. vulgaris will grow at or below temperatures of 5°C, but bolting of the floweringstem only takes place after ambient temperatures rise above 10°C for more than one week (MacDonald &Cavers 1974). The northern limit to its range is about 60°N (Jalas & Suominen 1994), and thisis probably related to the length of the reproductive season permitting seed-set after this temperaturehas been reached (MacDonald & Cavers 1991). The New Atlas hectad map makes it quiteobvious that Co Fermanagh is close to the climatic NW limit of the species' overall European range.
Fermanagh occurrence
In Fermanagh, B. vulgaris is a widespread and locally abundant weed species. It has been recordedin 134 lowland tetrads, 25.4% of those in the VC.
Fossil record and status
B. vulgaris is regarded as native in both B & I, fossils from previous interglacial periods(the Hoxnian and the Ipswichian), lending weight to the case for the current interglacial, although theonly record listed by Godwin (1975) came in fact from a Roman settlement, and could therefore easily beargued to indicate an ancient introduced species (ie an archaeophyte).
British and Irish occurrence
B. vulgaris is the commonest of the four species of the genus found in B & I, with only theintroduced B. intermedia (Medium-flowered Winter-cress) rivalling it in distribution andabundance, and even then it does so only patchily and apparently very much more frequently in N Irelandthan in the RoI (New Atlas). Although certainly widespread in the both B & I, B.vulgaris becomes rarer or absent in westerly regions of both islands throughout the wholelatitudinal range. In Scotland, it becomes more coastal the further north one goes and eventually itpeters out towards the NW, at least as a presumed native species (Preston et al. 2002).
European and world occurrence
Beyond B & I, the species, which is probably of Mediterranean origin, is considered native acrossmost of Europe south of latitude 60°N, extending east to the Himalaya. B. vulgaris is absent fromthe Balearic Isles, rare in Sardinia and absent also from Crete. It is a widespread introductionthroughout Scandinavia and to a much lesser extent in Iceland (Jalas & Suominen 1994, Map2304). Elsewhere, it is an introduction in N Africa, N America (where it is classified as a noxiousweed) and Australasia (Rich 1991).
Growth and flowering reproduction
In B & I, most germination and vegetative growth of B. vulgaris occurs early in the spring,but seedlings emerge throughout the year from seeds brought to the surface by disturbance. In milder,sheltered areas of these islands, growth of the rosette is almost continuous (Baskin & Baskin1989). When germination occurs in favourable autumn conditions, the plants produced will be eitherwinter annuals or perennials, while spring germinating plants tend to be biennial or perennial. Veryshortly after spring temperatures reach 10°C, the rosette, provided it is large enough and hasaccumulated sufficient stored photosynthetic resources, will bolt and produce the flowering stem.
The flowers are primarily out-crossed, possess nectar and attract Hymenoptera (bees and butterflies) plusbeetles and flies. Probing by these insects for flower foods causes both self- and cross-pollination. Indull, cool weather when insect visitors are rare, the two shorter stamens of the six in each flower,bend towards the stigma and effect self-fertilisation (Proctor & Yeo 1973; MacDonald& Cavers 1991).
Seed production and dispersal
The seeds are small and light and the number per fruit (ie siliqua) ranges from 3 to 21 with a mean of 13(Salisbury 1942). As previously mentioned, given suitable ecological conditions, the overall seed outputper unit area can be enormous, but the plant has no specialized dispersal mechanism and the majority ofseeds are simply dropped from the septum within a metre of the parent plant. The actual primarydispersal distance of seed depends on plant height, surrounding vegetation, temperature, humidity andwind speed (MacDonald & Cavers 1991). There may be, however, some very occasional secondarydispersal, since the seed coat is covered with an adhesive mucilage that, when moistened, may eitherinhibit dispersal by sticking the seed to soil particles, or promote transport byattachment to the surfaces of passing animals.
Ridley (1930) quotes records of B. vulgaris growing as an epiphyte along with 75 other species inthe soil accumulated on the tops of pollarded willows in Cambridgeshire (VC 29), which one presumed musthave been blown on high, or have been carried aloft by birds. Winter-cress seeds are said to be bittertasting, yet they have been recorded in the droppings of numerous animals including horses, cattle, pigsand rabbits. The seed remains viable after passage through the gut of these animals, providing secondarydispersal by internal zoochory (ie endozoochory) (Salisbury 1964). They can also survive continuousimmersion in water (and presumably also in flooded soil) for up to eight weeks (MacDonald & Cavers1991).
Vegetative reproduction
Occasionally, and particularly after physical damage to the inflorescence, B. vulgaris produceswhat have become known as 'cauline rosettes' on the bolted stem of the inflorescence late in the seasonafter flowering has been completed. This can lead under certain circ*mstances to asexual reproductionand dispersal (MacDonald & Cavers 1974; Rich 1984). Cauline rosettes are similar to smallbasal rosettes but are produced in the axils of stem leaves and, if they come into contact with theground, they can form adventitious roots and may become independent, thus establishing new clonalplants. If these rosettes succeed in overwintering, they will flower the following year. The fact thatprior damage to the plant appears to be involved probably indicates that cauline rosette formation istriggered by a physiological or hormonal imbalance, but it is rather surprising that these asexualorgans have only been reported in this one crucifer species (Rich 1984).
Variation
B. vulgaris is a very variable species, and in terms of both size and robustness it reflectshabitat conditions so strongly that Jackson (1916) recognised four varieties within it. Rich (1987a)discussed these forms fully, but reckoned that the subdivisions require more study over the wholegeographical range of the species and, until this is completed, he felt it better regarded as a singlehighly polymorphic species. Rich (1991) mentions that the species is even more variable in the rest ofEurope than in Britain, a not unexpected fact considering the greater range of environmental conditionsavailable there. However, in the absence of a more convincing native fossil record, this lends furthercredence to the possibility that in common with the three other Barbarea species found in B &I, B. vulgaris might well be an ancient introduction. Double-flowered and variegated garden formsof B. vulgaris have also been described.
Names and uses
The genus name 'Barbarea' is derived from the fact that the plant was once dedicated to SaintBarbara (Herba Sanctae Barbarae), the patron saint of artillerymen and miners, and protectress inthunderstorms (Gilbert-Carter 1964; Stearn 1992). The reason for the dedication is probably because theplant is wintergreen and has generally grown to a substantial size around the Saint's feast day (Dec. 4- old style), when people collected rosettes to eat as a source of fresh green vegetable, a custom thatdates back to the 14th century. A 100 g serving of the fresh green leaf provides 130 mg of vitamin C(more than twice the 60 mg recommended daily human requirement) (Zennie & Ogzewalla 1977; MacDonald& Cavers 1991). The plants do however produce a number of mustard oils, including sinigrinwhich gives it a rather tangy taste, and too large an intake would prove toxic, although there are noreports of this actually happening in B & I (Cooper & Johnson 1998). The Latin specific epithet'vulgaris' meaning common, is quite appropriate in this instance.
Several of the English Common names include 'Cress', a general name applied to many members of theCabbage family, and known from all the Germanic languages from the earliest times, also adopted into theRomanic and other dialects, but of quite unknown origin (Prior 1879). The names include 'Wintercress',of obvious relevance, 'Land Cress', as opposed to 'Watercress', 'French Cress' 'Yellow Cress' and 'St.Barbara's Cress'.
Several other names include 'Rocket' (e.g., 'Yellow Rocket', 'Winter Rocket' and 'Wound Rocket' ), whichis derived from a supposed similarity to the hot-tasting plant Eruca sativa, Rocket, which wascalled in Italien 'ruca', diminutive 'ruchetta', and in French 'roquette', and thus very readily became'Rocket' in English (Grigson 1974). The name 'Wound Rocket' was given by Turner in the Names ofHerbes (1548), since he felt it had leaves like Rocket and was good for a wound remedy, but thename was never adopted and the current author can find no reference giving the plant any medicinal use(Grigson 1987). One of the most unusual names from southern England is 'Cassabully', for which thecurrent author can offer no explanation, except that 'caisse' is French for a purse or money-box, likethe Latin 'capsa' (Britten & Holland 1886).
Threats
None.
Introduction, neophyte, a widely naturalised weed, but rare and only casual here.
1939; Praeger, R.Ll.; around Enniskillen.
April to November.
Growth form and preferred habitats
Although this mainly but not exclusively biennial, naturalised alien species can generally bedistinguished from B. vulgaris (Winter-cress) by its more dissected upper stem leaves (examinethe top ten stem leaves and bracts) and, when in fruit, the shorter, stouter floral style (ie 0.6-1.7 mmlong (average around 1 mm) in B. intermedia, rather than 1.7-4.0 mm long (average around 2.5 mm)in B. vulgaris). Also, B. intermedia starts flowering earlier (in March), than B.vulgaris (April onwards) (Rich 1991). These two weed species occupy the same sorts of disturbedhabitats and they are sometimes rather difficult to separate in reality. For this reason, RobertNorthridge and the current author both feel we cannot rule out the possibility of confusion in some ofour own records of these two species, and that we need to bear this in mind when commenting on its localoccurrence.
Fermanagh occurrence
The Fermanagh Flora Database contains B. intermedia records from twelve tetrads (2.3%), ten ofwhich have post-1975 dates. This represents one-tenth the recorded presence of B. vulgaris. Themain habitats of this winter-cress are disturbed ground on roadsides and waste ground, but they includea garden and a lakeshore too. As the tetrad distribution map indicates, this species is rare and casualin Fermanagh and appears very thinly scattered around the lowlands, with no obvious factor determiningits distribution.
Rich (1987a) pointed out that nowadays in B & I, B. intermedia tends to colonise open habitatsin disturbed ground on roadsides and building sites, whereas previously it was most frequentlyencountered as an arable weed. It is rarely if ever abundant, tending to occur as isolated individuals.Being both uncompetitive and ruderal, in Fermanagh at least, B. intermedia appears almost alwaysas a casual species, cropping up in small numbers and soon disappearing.
Irish occurrence
B. intermedia is frequent to very frequent and widespread in N Ireland, especially invice-counties Tyrone (H36), Armagh (H37), Down (H38) and Antrim (H39). It is particularly frequentaround Belfast. The species is recorded to a much lesser extent in Fermanagh (H33) and Londonderry (H40)(Flora of Lough Neagh; FNEI 3; Urban Flora of Belfast). Around Lough Neagh, JohnHarron occasionally found it colonising lakeshore gravel and riverbanks, mostly in well-drainedconditions. Apart from one 1957 record at or near Lough Melvin, none of the Fermanagh Database recordsmatch this type of habitat, where instead one might well expect to encounter B. vulgaris(Winter-cress).
B. intermedia was first found in the wild in the British Isles near Ballymena, Co Antrim (H39)around 1836. It might be more accurate to say it was first distinguished from B. vulgaris thereat that time. It most likely arrived as a crop seed contaminant and became established in NEIreland around the end of the 19th century (Cat Alien Pl Ir).
In the second edition of the Flora of NE Ireland, Praeger described B. intermedia occurringon waste ground and the margins of fields as, "a very local plant in Ireland, occupying a wedge ofcountry extending from Galway to Tyrone and Down, and frequent only in the NE, where it is widespreadbut never abundant." (Praeger & Megaw 1938).
Perusal of the New Atlas map suggests that B. intermedia is particularly sparse throughoutthe RoI when compared to N Ireland to such an extent that David Pearman suggests it may be seriouslyunder-recorded there (D.A. Pearman, in: Preston et al. 2002). The species has been recorded atleast once from 28 of the 40 Irish VCs, and a hectad map of the BSBI database (accessed 1 April 2020)now shows it better represented in the RoI than in the New Atlas, especially in the SE countiesof Waterford (H6) and Wexford (H12) which are expertly surveyed by Paul Green (Green 2008).
British occurrence
In Britain, B. intermedia is a frequent and widespread casual throughout the whole range oflatitude, although it remains somewhat patchy in its distribution and becomes decidedly coastal in the N& W of Scotland (New Atlas; BSBI database accessed 1 April 2020). There has been animpressive increase in our knowledge of this weedy crucifer in recent decades. The index of changebetween the 1962 and 2002 Flora atlases (Perring & Walters 1962; New Atlas) has beencalculated as + 1.92, indicating a very nearly doubling in records over that 40 year period. In theiraccount of alien plants in B & I, Clement & Foster (1994) wrote of it being, "Locallyestablished and persistent on arable and waste land in widely scattered localities throughout theBritish Isles; increasing."
Comparison of the hectad maps of the New Atlas survey (2002) and the current BSBI database 2020survey shows there has been an increase in the cover of B. intermedia in both B & I, butespecially so in S & W England. This suggests that this species is increasingly frequent and morewidespread yet, as indicated above with respect to the Fermanagh experience, it is possible that it maysometimes still be overlooked or confused with the generally much more common and widespread B.vulgaris. Thus, perhaps what the hectad maps are really displaying is better fieldidentification than was the case in previous surveys. Probably both these suggestions are true.
European and world occurrence
As with other species that have a history as arable weeds, it is impossible to be certain of the nativerange of B. intermedia. The species is considered 'probably native' in S & C Europe from NPortugal to S Denmark and S Germany and also in N and E Africa (Rich 1991). It is introduced in NDenmark, S Sweden and Switzerland and has a few additional outliers in C Europe (Jalas & Suominen1994, Map 2307). It is also said to be introduced in Australasia (Rich 1991).
Threats
None.
Native, common and locally abundant.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Taxonomy and recent changes
Formerly all white-flowered water-cresses were regarded as belonging to a single species, Nasturtiumofficinale R. Br. s.l., and Druce (1932), in his Comital Flora of the British Isles,recorded it as occurring in every VC in B & I. This entity was subsequently split in the mid-1940sinto two species and their intermediate sterile hybrid (Howard & Manton 1946). The three formssubsequently changed genus from Nasturtium to Rorippa, section Cardaminum, so thatfor a time we recognised them as R. nasturtium-aquaticum, R. microphyllum (Narrow-fruitedWater-cress) and R. × sterilis (Hybrid Water-cress) (Rich 1991; Stace1997). Recent analysis of chloroplast DNA and nuclear encoded ITS sequences has confirmed, however, theseparation of Nasturtium from Rorippa Scop. (Franzke et al. 1998; Bleeker etal. 1999), so we have seen a return to Nasturtium nomenclature for these three taxa(Stace 2010, 2019; Sell & Murrell 2014).
In terms of their vegetative form, specimens of the two white-flowered water-cress species and theirhybrid are very variable with respect to their growing conditions, ie they are phenotypically veryplastic with respect to their environment (Rich 1991). This means that when we are making observationsin the field we are forced to accept that we cannot distinguish non-flowering, vegetative water-cressmaterial of N. microphyllum from either N. officinale, or their hybrid, and we are obligedtherefore to record such indeterminate plants simply as N. officinale s.l.
Following what has now become the preferred botanical practice, we will continue to keep records of thespecies aggregate separate from N. officinale (= R. nasturtium-aquaticum s.s.).
Fermanagh occurrence
The situation in Fermanagh is that what we now call N. officinale s.l. has been commonly recordedin a total of 249 tetrads, 47.8% of those in the VC. It is very widespread throughout, being present inall forms of marshy ground, including wet meadows, by lakes, streams, springs and ditches.
A very interesting Biological Flora account of Nasturtium officinale R. Br., by Howard& Lyon (1952a), describes the ecology and biological behaviour of the plant in detail, but since theinformation given really applies to the species in the strict sense, it will be dealt with by usunder that species account.
European and world occurrence
N. officinale s.l. is also common and widespread throughout W & S warm temperate Europe, NAfrica and eastwards to C Asia, but due to the change in taxonomy of the species aggregate, plus thefact that water-cress has been cultivated for at least 2,000 years, it is not really possible to delimitit* native occurrence (Hultén 1971, Map 195). The plant in this broad taxonomic sense has also beenintroduced to N & S America, S Africa, the Far East and to Australasia (Rich 1991).
Uses
Watercress has been eaten either raw as a rather spicy, peppery salad, or boiled as a green vegetable, bypeople of all ranks for many centuries. Following another of the periodic slumps in agriculturalfortunes, it has been commercially farmed in the midlands and the south of England since about 1808, andfound a niche in suitable places with a supply of clean, flowing, lime-rich water (Grigson 1987; Thrisk1997, pp. 204 & 295 ). The fact that cultivation has really only been successful along those streamsthat draw their water from deep in the English chalk may be due in part to the limey composition, purityand alkalinity, but it is also true that in winter these deep-welling waters tend to be warmer thantheir surroundings, giving the aquatic plant a more favourable growing environment than streams fed bylocal rainfall or snow melt (Edlin 1951).
Watercress farmers used to grow both N. officinale, 'Green Cress' and N. xsterilis 'Brown Cress' for salad, and actually prefer the latter, but much earlier than itscultivation as a salad crop Watercress had a deserved reputation among herbalists for the prevention ofscurvy, few plants being richer in vitamin C. Watercress was traditionally picked wild from the edges ofstreams and ditches, where it can grow in thick drifts. It is, or rather used to be, sufficientlyimportant and valued a plant for numerous English settlements to be called after it, e.g., namesbeginning with 'Kes', such as 'Kersey' in Suffolk, derived from 'Cress island', and 'Kershope' inCumbria, derived from 'Cress valley' (Mabey 1996).
Perhaps one of the reasons for the longstanding popularity of watercress salad in B & I is suggestedby Glenny (1897), who noted that compared with Continental Europe and the Far East, where (at leastuntil fairly recently), people traditionally ate far more salad herbs than we do, and they dress thebowl with "Lucca oil, vinegar, mustard and salt, or most of these, properly incorporated with theraw herbs", whereas in these islands, Watercress is "at least amongst the lower and middleclasses, preferably eaten in a simple and primitive way", ie alone, unadorned, and often not evenmixed with other greens as, "it loses its individuality when partaken with any other herb"(Glenny 1897, p. 608). Mabey (1972, p. 75-77) and (1996, p. 147-148) recounts interesting first handdescriptions of the Watercress trade in Victorian London. He also gives sensible warning advice on thechoice of plant material to avoid the dangers of liver-fluke, which due to the presence of a snail as asecondary host of the parasite on the leaves, often infests Watercress plants anywhere near sheep orcattle pastures.
In more recent times Watercress has several times gone in and out of fashion, its culinary use sometimesreduced to just a few sprigs garnishing steak, until it is promoted by a celebrity chef and suddenly itsuse revives. Until the mid-1950's hybrid 'Brown-' or 'Winter-watercress', R. x sterilis was thepreferred plant on acount of its somewhat greater frost-resistance and year-round leaf production, butcontinual reliance on vegetative propagation led to susceptibility to disease, and a mosaic virus andtwo fungal pathogens causing 'Crook root' and 'Cabbage black ring spot' quickly led to its abandonment.Farmers turned instead to N. officinale, 'Green watercress', which is fertile and whosepropagation from seed avoids virus infection. Selective breeding of N. officinale sens.str. since then has been directed towards disease resistance, prolonging the vegetative stage by delayedflowering, and developing a growth form which facilitates mechanical harvesting (Bleasdale 1964; Crisp1976).
At the same time there has been a reduction in the area of commercial growing in England, withBritish-based companies developing watercress cultivation abroad in both continental Europe and in NAmerica in order to maintain supplies in wintertime. There is also evidence on the World Wide Web thatcommercial Watercress cultivation, sometimes involving hydroponic, is currently being explored andencouraged by government agencies in New Zealand, and there are now at least 24 franchised growers therespread over both islands. In North America too, Watercress cultivation is carried on in five easternstates on a large scale.
Names
The English name 'Cress' generally refers to this plant, ie 'Watercress', or 'Garden Cress' (Lepidiumsativum). The name derives from the Old English (OE) or Anglo-Saxon 'caerse', 'cerse', or'cresse', with similar cognate names in other Germanic languages, all based on an Indo-european wordmeaning to nibble or to eat. In OE, N. officinale sens lat. or the species aggregate wasdistinguished as 'ea-cerse', meaning 'stream-cress', or 'wielle-cerse', 'spring-' or 'brook cress'(Grigson 1974).
Grigson's famous 'Englishman's Flora' lists eleven English Common names chiefly based on the OEname, although he also includes the Yorkshire 'Tang-tongues', which Britten & Holland give as'Teng-tongues', referring to 'Water-cresses as being pungent to the taste' (Grigson 1987; Britten &Holland 1886). Another English Common name Grigson lists, which originated in Gloucestershire, is'Carpenter's Chips', a clear reference to the chip baskets (ie baskets of unbarked (or stripped)willow), in which the Watercress crop was often shipped to market (Edlin 1951, p. 59).
Threats
None.
Native, probably frequent. Eurosiberian southern-temperate, but widely naturalised and now circumpolar.
1947; Moon, J.McK. & Carrothers, E.N.; Castle Coole.
May to November.
Growth form, identification and preferred habitats
This white-flowered, creeping, emergent or floating perennial water-cress species s.s. can only bedistinguished from the species in the broad sense (s.l. or agg.), when the plant has well-formed fruitsand seeds. The seed pods of N. officinale s.s. are short and round, and the brown seedsthemselves are in two rows per locule (Howard & Lyon 1952a). While these are good, reliableidentification characters, Rich (1987b, 1991) warns that confusion can still occur with N.microphyllum (Narrow-fruited Water-cress), as its seeds, although typically forming a singlerow, may also occur in two rows, but they only do so at the base of the fruit pod.The most reliable distinguishing characteristic of this section of the genus is the sculpturing of themature seed coat, and since a microscope is required to examine the seed, this is not a field character.
N. officinale is typically found in clean, flowing, often shallow waters, and experiencing fairlywell-illuminated growing conditions. These conditions are met in lakes, streams, springs and ditches,and on marshy ground and flushes beside such waters, including seasonally wet grazing meadows and quarrydrains.
Associated species include the very much more commonly recorded and ecologically similar Apiumnodiflorum (Fool's-water-cress), Myosotis scorpioides (Water Forget-me-not), Veronicabeccabunga (Brooklime), Mentha aquatica (Water Mint) and Filipendula ulmaria(Meadowsweet).
When considering altitude as a delimiting ecological factor, there is a tendency for N. officinaleto occupy more or less exclusively lowland sites, being replaced at higher altitude by either theclosely related N. microphyllum, or by their hybrid, N. ×sterilis (Howard & Lyon 1952; Grime et al. 1988; Preston & Croft 1997).
Variation
N. officinale s.s. is vegetatively extremely plastic in response to varying environmental growingconditions. Such variation extends from the overall scale of the plant (ranging from dwarf to giant(6-200 cm)), leaf shape, size and colour, growth-form, life-form which can be emergent or floating, andphenology being perennial or annual (the latter generally appearing in drier sites) (Rich 1991).
In the past, a number of varieties were distinguished, some of which had a genetic basis, others merelygrowth forms, so that confusion reigned (see Howard & Lyon 1952, p. 228). Both Manton (1935) andHoward & Manton (1946) recognised that Nasturtium officinale s.s., is a fertile tetraploidspecies (2n=32).
Flowering reproduction
Flowering and fruiting occurs freely under both dry and moist growing conditions, stretching from the endof May through to September, or even November in a mild year. However, the reproductive process isrestricted or prevented in very wet or totally submerged situations. The terminal racemes of small whiteflowers are self-compatible and seed set is normally abundant. No doubt at least some seeds result fromcrossing achieved with the assistance of insects visitors, since numerous species of flies and a fewbees and beetles are attracted by a plentiful supply of nectar and pollen (Howard & Lyon 1952). Seedproduction is very variable, but normally it is abundant. Dispersal is usually achieved by floatation inwater, effected by an air bubble formed around the seed coat. Seed may also be dispersed over longdistances and between water bodies embedded in mud, which is then transported by animals including birdsand man. There is no dormancy requirement for germination, but buried seed can remain viable in the soilfor between one and five years (Howard & Lyon 1952; Thompson et al. 1997).
Disturbance and vegetative reproduction
Water-cress plants are frequently grazed by a wide range of invertebrates, and in some of its seasonallydrier, grassier sites, plants are also very obviously subject to grazing and trampling by cattle whichmust limit vegetative growth and frequently prevent flowering and fruiting. However, all water-cresses(ie both species plus their hybrid) have great powers of vegetative reproduction and patch formationthrough their stems creeping and rooting profusely at their nodes. As with associated plants of similarvegetative properties, detached portions of plant occasionally disperse by floatation, particularlyafter spates or floods, compensating to some degree for the loss of seed production under suchcirc*mstances (Ridley 1930, pp. 182 & 546).
Fermanagh occurrence
In Fermanagh, N. officinale s.s. is slightly more frequently recorded than N. microphyllum,ie appearing in 61 tetrads (11.6%), compared with 44 (8.3%) for the latter, but this is hardlystatistically significant. N. officinale s.s. is thinly but widely scattered throughout the VC,but is a little more frequently found on the Western Plateau.
Irish occurrence
The New Atlas map shows N. officinale s.s. as scattered but locally frequent in Ireland,having a minimum of one hectad with recent records in every Irish VC except NE Galway(H17). At the same time, several Midland and western Irish VCs appear to have little more than thislevel of presence recorded. It is possible that this taxon's apparent preference for, or in some partsof England, restriction to, eutrophic or mesotrophic calcareous waters and soils, and its almost totalavoidance of peat (both acidic and alkaline) may, at least in part, explain this distribution pattern.Airy Shaw (1948) first observed this ecological tendency distinguishing N. officinale s.s. fromN. microphyllum, but he cautioned that, "it can in no way be expected to hold good over thewhole country".
British occurrence
In Britain, the New Atlas map shows N. officinale s.s. as being common and widespread inlowland areas, apparently becoming more coastal in occurrence further north, particularly in NE Englandand E Scotland. This distribution might well reflect the species' known sensitivity to frost, but sinceit is recognised that there is an unknown degree of under-recording of this and the related taxa, thereis a danger, however slight, that it could simply represent the extent of the local recording effort.
It must also be remembered when attempting to interpret the mapped distribution that water-cress has beencultivated for thousands of years (see also our account of the species aggregate for historical uses),the plants commercially involved being this species, and to an even greater extent, the hybrid, N.× sterilis, and the major region of large-scale cultivation in thelast 150 years has been in the S and E of England (Howard & Lyon 1952; Preston & Croft 1997). Wemight therefore expect a greater possibility of human dissemination of these plants in and around theareas of cultivation and their markets.
European and world occurrence
The European distribution illustrated in Jalas & Suominen (1994, map 2327), shows this specieswidespread and native in S & W Europe, southern parts of C Europe, but absent from Norway, Swedenand Finland except as an introduction in the extreme south of Sweden. In contrast to the situation inBritain, N. officinale s.s. is the more generally cultivated water-cress in European countriesand, in particular, this form of the plant has been widely introduced, cultivated and naturalisedelsewhere in the world, including in N & S America, many parts of Africa, the Far East andAustralasia (Hultén 1971, Map 195 and p. 204). The species was first recorded in New Zealand, forexample, in 1852 and it is now common in streams, ditches and drains throughout (Webb et al.1988).
Names
The name 'Nasturtium', or rather the Latin 'Nasturcium', was given by the Classical Roman author, Pliny,to a plant with a distinctly pungent flavour or odour. It is derived from two Latin words 'nasus' noseand ' torquere' meaning 'to twist' or 'torture' (Gilbert-Carter 1964; Grigson 1974), descriptive of thereaction the original plant invoked, and one that some palates might still present to the peppery biteof water-cress. The name 'Nasturtium' became linked with the S American genus Tropaeolum, aflower in a completely different family, simply on account of the pungent smell and flavour of itsleaves. The first Tropaeolum species was introduced to Europe through Spain from Peru in the late16th century and it was originally given the name Nasturtium indicum, Indian Cress (Gerard 1633,p. 251-2; Grigson 1974).
Threats
None apart from the possibility of excessive pollution or eutrophication.
HAS CHANGED NAME SINCE THE FLORA WAS PUBLISHED
Nasturtium microphyllum (Boenn) Rchb. (= Rorippa microphylla(Boenn.) Hyl. ex Á. Löve & D. Löve), Narrow-fruited Water-cress
Native, occasional. Native world distribution uncertain, but probably so in Europe and in C Asia: verywidely introduced and naturalised beyond this range.
1860; Smith, T.O.; Ardunshin.
June to November.
Growth form, identification and preferred habitats
Like the two other forms of water-cress, N. microphyllum is a patch-forming perennial foundfloating in clean running, productive rivers, streams, springs, ditches and in lakes and ponds subjectto fluctuating levels, plus as an emergent on marshy ground beside such waters. Unlike the other forms,it is not commercially cultivated and has not been spread in that manner.
Vegetatively, the two water-cress species (N. officinale s.s. and N. microphyllum) are toosimilar and too variable to distinguish from one another. Although this plant bears the specific epithet'microphyllum' (Greek, meaning 'small-leaved'), the name is a very misleading one, since apart perhapsfrom the terminal leaflet, its leaves and leaflets are not significantly smaller or narrower than thoseof its close relative N. officinale s.s. In reality, leaf shape and size are absolutely valuelessfor distinguishing the two species and their hybrid. In comparison with N. officinale s.s.,however, the seed pods of N. microphyllum are long, thin and curved and they contain a single rowof seeds − except sometimes near the base of the fruit valve. The seedcoat also has a much finer grainedtexture or network of reticulations than N. officinale s.s., having 12-20 depressions or cellalveoli across their greatest width (Rich 1991, pp. 152-3; Stace 1997, photomicrograph Fig. 254, 11).
N. microphyllum occupies very similar moderately nutrient-rich, productive, somewhat disturbed,shallow flowing water or waterside marsh habitats to those of N. officinale s.s. (Water-cress),but while the two may co-exist in more definitely calcareous situations, this species has a rather widerecological range, being tolerant of moderately acidic and less-calcareous environments than the latter.All forms of water-cress are absent from unproductive, strongly acidic, very shaded or highly disturbedor exposed habitats (Grime et al. 1988). N. microphyllum is a little more shade tolerantand is slightly the more frost resistant of the two species, and therefore can occasionally be found athigher altitudes − ie the current altitude record in B & I stands at an exceptional 550 m near MoorHouse in Westmorland & Durham (VCs 66 & 69) (Preston & Croft 1997; Preston et al.2002).
Seed dispersal and vegetative reproduction
It should be borne in mind, that since it is not cultivated, N. microphyllum is less likely to beintroduced or spread by man than the two cultivated forms, N. officinale s.s. and the hybrid,N. × sterilis (Howard & Lyon 1952). On the other hand, since allforms of water-cress grow on muddy stream banks, seed of the two species may be relatively easilytransported over long distances embedded in mud adhering to birds' or other animals' feet, or on theirexternal surfaces.
Broken stem portions of all three forms of water-cress, including the sterile hybrid, have an amazingability to form adventitious roots within a matter of hours, even on portions of the inflorescence,allowing the plants to reproduce very readily.
Fermanagh occurrence
As is the case in nearly every part of B & I, Robert Northridge and the current author are sure thatN. microphyllum is under-recorded in Fermanagh. It really should be more frequent than theoccasional, limited records in the Fermanagh database spread across 44 tetrads (8.3%) suggest. As thetetrad distribution map indicates, it is thinly and widely scattered throughout the VC. At the sametime, it is certain that it is nothing like as common as N. officinale s.l. (Water-cress), a notterribly surprising conclusion, since this entity represents the vegetative forms of three closelyrelated taxa, which unfortunately cannot be properly distinguished without ripe fruits.
Unusual ecological behaviour of N. microphyllum in New Zealand: While thevast majority of N. microphyllum populations are recorded in or beside shallow water less than 75cm deep, in New Zealand Michaelis (1976) found the species growing in and around very clear coldsprings, not only as an emergent and as a mat-forming, floating plant near the shoreline but, mostunusually, a somewhat dwarfed growth form was also growing completely submerged and rooted in gravel.This had translucent, dark green, rolled up leaflets, and occurred at depths ranging down to 6.5 m. Atdepths below 100 cm it was, of course, non-flowering. It also showed little seasonal change in growthform, other than a slight elongation of the internodes in summer (Michaelis 1976).
A brown winter pigment form
Plants of N. microphyllum have a strong tendency to turn a deep purple brown anthocyanin colourduring the colder winter months, a feature which when combined with lesser vegetative vigour than'Green' Water-cress, N. officinale s.s., or the even more hardy 'Brown' or 'Winter' Water-cress(the hybrid N. × sterilis), has excluded its use in commercialcultivation (Howard & Lyon 1952, p. 242).
Genetic makeup and species origin
N. microphyllum was originally thought to be an autotetraploid, ie the result of spontaneouschromosome doubling within a species, N. officinale being put forward as the putative ancestor(Manton 1935). However, N. microphyllum is now recognised to be an allopolyploid octoploidspecies (2n=64), which has arisen through hybridization between N. officinale s.s. and an unknownspecies. The unknown parent was presumed to be a member of the closely related genera Rorippa orCardamine (Howard & Manton 1946). On account of the seeds being arranged in the pod in asingle row and of its morphological resemblance to N. microphyllum, Oefelein (1958) suggestedCardamine amara (Large Bitter-cress) might be the unknown parent. A recent large-scale isozymeanalysis of Nasturtium and Rorippa species has proven that the unknown parent is aRorippa species. R. sylvestris (Creeping Yellow-cress), R. amphibia (GreatYellow-cress) and R. palustris (Marsh Yellow-cress) were considered likely candidates, and all ofthem carried the marker alleles of N. microphyllum, while Cardamine amara did not and isdefinitely not involved in the parentage (Bleeker et al. 1999).
Being octoploid, and thus having paired chromosomes, N. microphyllum is perfectly fertile; likeN. officinale s.s., it is self-compatible and sets copious seed. Also, like N. officinales.s., it has a very long flowering and fruiting season, but it comes into flower about two weeks laterthan the former. The larger white flowers of N. microphyllum are probably even more attractive toinsect pollinators than those of N. officinale s.s. The first flowers of water-cress plants thatopen set only a few seed and, especially in N. microphyllum, plants set very little seed if theyare crowded or growing in less than favourable conditions (Green 1962). Several properties, includingvegetative reproductive ability, seed dispersal, germination and longevity, all appear almost identicalin the two water-cress species (Howard & Lyon 1952).
British and Irish occurrence
While there certainly still is an unquantified degree of under-recording of the two white-floweredwater-cresses and their hybrid, both species are undoubtedly common and widespread throughout B & I(Preston et al. 2002). In Britain, N. microphyllum is the less frequent and abundantspecies in the southern half of the country, but on account of its wider ecological range, it probablyis the more common and abundant of the two further north, and especially so in E Scotland.
In Ireland, N. microphyllum appears to be slightly less common overall than N. officinales.s., but it is probably the more frequent of the two species in the central region of the island and onhigher ground throughout. In these two situations, winters are noticeably colder, and peat soils andtheir acidic influence on drainage water are very common and widespread (Preston et al. 2002).
European and world occurrence
Beyond B & I, N. microphyllum is very probably under-recorded in Europe,where it is mainly noted in the west from N France, Belgium, the Netherlands, Denmark, NW Germany,Switzerland and N Italy and very thinly scattered further south and east (Jalas & Suominen 1994, Map2328). The native range is said to extend to C Asia and E Africa and it has been introduced to S Africa,N America and Australasia (Rich 1991).
Threats
As we have an imperfect record of the presence of Narrow-fruited Water-cress throughout B & I, wecannot readily ascertain what population changes are due to the widespread environmental pressures onwetlands, such as local drainage operations and any form of pollution (including nutrient enrichment).However, we can predict that either or both these factors would quickly result in serious losses ofWater-cress through suitable habitat destruction and increased competition from taller wetland species(Grime et al. 1988).
Rorippa islandica s.s. (Oeder ex Gunnerus) Borbás, Northern Yellow-cress
Native, very rare. Eurosiberian boreal-montane.
28 September 1995; McNeill, I.; on dumped material, roadside waste ground, Crocksoult Td, SE of GlenLodge.
June to September.
Taxonomy and growth form
R. islandica s.s. and R. palustris (Marsh Yellow-cress) are two morphologically andecologically very similar pale yellow flowered taxa which originally were combined, and from the pointof view of the field botanist and the ecologist, they very conveniently remained so until the late1960s. At that date, the Scandinavian taxonomist Bengt Jonsell revised the NW European species withinthe genus, and he split the previous R. islandica s.l. group into two species of different ploidylevel. The tetraploid form, now recognised as R. palustris, has become recognised as by far thecommoner and more widespread taxon in comparison with the less weedy diploid plant, R. islandicas.s. (Jonsell 1968). Plants of both species are habitually inbreeding and generally perform as annualspecies, although they can sometimes also behave as biennials, or even as short-lived perennials.Jonsell (1968) found that individuals could survive in cultivated plots for up to five years.
Identification
R. islandica s.s. is quite difficult to distinguish from R. palustris and, rather rarely,the two taxa can be found growing together (Goodwillie 1995; Chater & Rich 1995). Microscopicexamination of the coat of ripe seed is used to confirm their identification, the seed coat of R.islandica s.s. being more finely colliculate, ie having smaller indentations than those of R.palustris, and thus forming a finer, less-distinct, net-like pattern (Jonsell 1968; Rich 1991).Those familiar with the plant find that it can be distinguished from R. palustris by its fruitcharacters, even at a distance of several metres. Fruits of R. islandica s.s. are sometimessecund and swept downwards (ie the whole fruiting inflorescence appears one-sided), and since the stalksof the fruits are relatively short, the arrangement looks rather dense. In contrast, the fruits of R.palustris are arranged evenly around the stem and they are not so densely crowded. The fruitshapes are also distinctly different to the practised eye or hand, those of R. islandica s.s.having a more rectangular, square-ended appearance in section – as can be ascertained by trying to rollthem between finger and thumb (Rich 1991; Chater & Rich 1995).
Irish occurrence and preferred habitats
R. islandica s.s. is regarded as being fairly rare in B & I, although not quite as rare as waspreviously supposed (Rich 1999; New Atlas; BSBI Database accessed 21 June 2020). Surveywork associated with the BSBI Atlas 2000 project found that since its recognition as a separate species,R. islandica s.s. has been quite frequently discovered in Ireland, having previously beenoverlooked, a situation undoubtedly also the case elsewhere in these islands.
The habitat of the first Irish station in Connemara which Jonsell himself discovered in 1964 (or ratherrediscovered, since herbarium specimens in BM and DBN indicated that R.S. Shuttleworth ofBerne had collected the plant in the same area as early as 1831 and 1832 (Scannell 1973)), was bare(probably muddy) ground near the Atlantic coast at Renvyle, W Galway (H16). Here, the plant grew as asmall, prostrate or decumbent annual in association with Potentilla anserina (Silverweed),Cardamine pratensis (Cuckooflower), Ranunculus repens (Creeping Buttercup) andNasturtium officinale s.s. (Water-cress) (Scannell 1973).
In 1971, Maura Scannell herself found the second Irish station for the plant in a somewhat more inlandsite, where it occupied bare ground towards the centre of a dried out turlough (ie a vanishing lake − agrazed grassy hollow which regularly floods during wet periods), in the Burren limestone region of CoClare (H9) (Scannell 1973). A total of 15 additional turlough records soon cropped up in W Ireland, allof similar habitat, ie seasonally inundated wet meadows, in cattle- or sheep-grazed, trampled and oftenmanure-enriched, muddy ground.
By 1995 the Irish distribution of R. islandica s.s. included a further four west coast Irish VCs(SE Galway (H15), NE Galway (H17), Roscommon (H25) and E Mayo (H26)), plus another in Co Kilkenny (H11)further to the SE of the island (Goodwillie 1995). Over the space of several visits to the Kilkennystation, Goodwillie noticed small specimens which he reckoned probably germinated in mid-August,reaching the fruiting stage at the end of October. This observation provided evidence of phenologicalplasticity, which fitted this annual species well for existence in ground subject to intermittentflooding and the other forms of habitat disturbance already mentioned. Most Irish records of R.islandica s.s. have been associated with seasonally or periodically inundated, cattle- orsheep-grazed, trampled and often manure-enriched, sometimes muddy pastures around lakes and turloughs inlimestone areas like the Burren, Co Clare (Scannell 1973; Goodwillie 1995).
Recently, this taxon has begun to turn up at numerous locations in the north of Ireland, thanks almostexclusively to the plant hunting efforts of Ian & David McNeill together with John Harron and JohnFaulkner (McNeill & Hackney 1996; McNeill 1998). The habitats of these widely scattered NI sitesinclude some very heavily disturbed amenity areas in public parks, including well-trodden paths andverges, cracks in concrete caravan park hard standings, beside a sheep dipping enclosure and in dumpedmaterial on waste ground.
Fermanagh occurrence
The first Fermanagh county record listed above is supported by a verified voucher specimen. Three earlierrecords by an EHS Habitat Survey Team that locate the species in muddy situations at Inishroosk, UpperLough Erne in both a marsh and on the lake shoreline on 21 June 1985, and subsequently from TrannishIsland, Upper Lough Erne on 16 June 1987 might well have been genuine discoveries. However, withoutherbarium vouchers they were not fully acceptable first county records, and originally Robert Northridgeand the current author as BSBI Recorders for Co Fermanagh, noted them with caution. However anothersurvey of selected sites in Upper Lough Erne commissioned by NIEA and carried out in 2006 by ENSIS(English Nature Site Information System) recorded R. islandica at four sites, including TrannishIsland, 2 July 2006, which of course supports the 1987 record at that site made by the EHS HabitatSurvey Team. The details of three other ENSIS sites are: Belleisle ASSI, 5 July 2006; Castle Lough, 22July 2006; and Derrykerrib Lough, 22 July 2006.
The outcome of this is that we now have a total of ten records from five tetrads as shown in the Floraof Co Fermanagh map: the two McNeill records to the north of the VC are quite remote from theremainder around Upper Lough Erne. Examination of the BSBI database shows just one additional site atPortora Boathouse made by RHN and Tim Rich in 2013.
British occurrence
The New Atlas hectad map showed R. islandica s.s. as rare, scattered and decidedly northernand western in its distribution in B & I. In the last 20 years, it has been much more widelyrecorded throughout the whole latitudinal range of Britain from Plymouth to Orkney though it remainsmost concentrated in the same northern and western range although thinly scattered elsewhere. In 1991,R. islandica s.s. was recorded growing in rock crevices beside a river in SW Wales and,subsequently, over the next few years nine additional stations were found in the same Welsh region,occupying a surprisingly wide range of habitats. These included damp depressions along a river floodplain, around muddy pools in either nutrient-poor or rich conditions, ditches and hedgerows and, muchmore unexpectedly, on somewhat drier but still disturbed conditions on waste ground and rubbish tips.The only obvious shared factor in these situations is their ecologically open, fairly damp character,conditions maintained by periodic disturbance which most often involves flooding, grazing and trampling,but which occasionally arises as a consequence of human activities as varied as dumping, trampling andfishing (Chater & Rich 1995; Rich 1999). The Welsh representation is still very noticeable and evenmore widespread today (BSBI Database Map accessed 21 April 2020).
European and world occurrence
Beyond B & I, R. islandica s.s. is similarly confined to the NW extremities of Europe(including Nordland fylke in N Norway, Iceland and S & W Greenland). However, it does also crop upas very disjunct populations around alpine lakes in the mountain ranges of S Europe, stretching W to Efrom the Cantabrian Mountains to the Northern Pindhos (Jonsell 1968, Fig. 7; Jalas & Suominen 1994,Map 2323).
Jonsell also recorded an area of R. islandica occurrence in Russian Siberia, but since hedistinguished and named this particular geographical form subsp. dogadovae (Tzvelev) Jonsell, itmust not be quite the same plant that is met in B & I, which is referable to subsp. islandica(Jonsell 1968).
Seed dispersal
Jonsell (1968) remarked that R. islandica s.s. is especially common in Iceland and is particularlyabundant around Lake Myvatn, a noted wildfowl site. This fact, together with the then known distributionof the plant, led him to suggest that geese might well be transporting agents for seed of the speciesduring their annual migration, either internally in their gut, or else embedded in mud on their feet.Unlike many other Brassicaceae, the seed coat epidermis of R. islandica, and of R.palustris for that matter, does not produce sticky mucilage when wetted, but rather the cellsare hollow, which may help confer the prolonged buoyancy they demonstrate in water (Jonsell 1968; Chater& Rich 1995).
The probable means of seed transport in this species, both local and long-distance (eg water flow, mudand birds) are entirely conjectural at present. In the nature of stochastic properties, these thingsalways are. However, the relevant arguments have been discussed very clearly by Chater and Rich (1995).
Native, common. Circumpolar boreo-temperate.
1806; Scott, Prof. R.; Co Fermanagh.
May to November.
Identification
The short fruits (only up to 9 mm long) and small flowers with tiny yellow petals, which are as long as,or just shorter than the sepals, clearly distinguish the R. palustris from all other species inthe genus except the very much rarer R. islandica sens. str. Previously these two morphologicallyvery similar taxa were combined in the R. islandica s. lat. group until Jonsell (1968) split themon grounds of differing ploidy level; the tetraploid taxon now recognised as R. palustris becameby far the commoner and more widespread construct in comparison with the diploid plant, R.islandica sens.str. In B & I the form present is subsp. palustris (Sell & Murrell2014).
Discrimination of R. palustris from R. islandica sens. str. is difficult, and while plantswith an upright habit, sepals longer than 1.6 mm, and with fruits less than twice as long as theirpedicels and arranged all round the stem (rather than held to one side, ie secund), are most probablyR. palustris, the only absolutely certain method of distinguishing the two species involveslaboratory study: either chromosome counting, or a microscopic examination (x 25-50) of the seed surfaceand its comparison with verified reference material (see Rich 1991, p. 48-49; Chater & Rich 1995).
Growth form and preferred habitats
This distinctive summer annual sometimes biennial or, occasionally, a short-lived monocarpic perennial isa pioneer coloniser of open, unshaded, marshy, or damp, muddy bare ground, exposed in summer around lakeand pond shores and river banks. Various floras consulted differ considerably with respect to thelife-form and perennating ability of the species: e.g., Grime et al. (1988) describe it inEngland as a 'summer-annual, semi-rosette therophyte', whilst in New Zealand, Webb et al. (1988)feature it as a 'perennial, rarely annual, taprooted herb.' Rich (1991) regards it as 'annual (rarelyperennial)', and Clapham et al. (1968) describe it (albeit as part of R. islandica s.lat), as 'an annual or biennial herb with a pale slender tap-root'. There clearly is a little projectneeded to clarify the definitive position on this matter, or to measure the range of variation itencompasses.
Marsh Yellow-cress spreads from the natural habitats listed above, to other more disturbed sitesassociated with man, eg beside lakeshore jetties, ditches and canal banks, although again, generally onground that is seasonally flooded (Chater & Rich 1995). In drier wayside areas, and in other siteswhere the mud hardens early in the growing season, the species shows considerable phenotypic plasticity.Plants in these drier conditions are often greatly dwarfed, flowering sooner, but then only managing toproduce comparatively few seeds (Grime et al. 1988).
Marsh Yellow-cress is often found in association with other typical pioneer colonists of bare mudhabitats, eg Agrostis stolonifera (Creeping Bent), Alopecurus geniculatus (Marsh Foxtail),Chenopodium rubrum (Red Goosefoot), Littorella uniflora (Shoreweed) and Ranuculusscleratus (Celery-leaved Buttercup). A moderate degree of trampling, grazing, or otherdisturbance may be required to keep growing conditions sufficiently open for plants of low competitiveability like R. palustris to thrive (Grime et al. 1988; Rodwell et al. 2000, p.428).
Fermanagh occurrence
In Fermanagh, R. palustris has been quite commonly recorded in 107 tetrads, 20.3% of those in theVC. It is particularly common in muddy, marshy, often seasonally or periodically flooded, mesotrophic toeutrophic sites around both Upper and Lower Lough Erne, although as the tetrad map shows, it does occurin a few other similar sites scattered across the VC.
Isolated plants or small populations also occur uncommonly or occasionally as a weed of winter-wet,moderately fertile ground along other artificial, man-made habitats more remote from water bodies,including old disused railway lines, on waste ground, roadsides and even sometimes in damp areas ingardens.
Irish occurrence
In Ireland, R. palustris is recorded as common and widespread particularly in NI, but also in Midand East Cork (H4 & H5). It appears to be very scarce or absent down most of the W coast wherestrongly acidic peat soils predominate, and likewise in the drier, warmer, more agriculturally favouredSE of the island. This rather peculiar distribution pattern fails to match any of the 23 categories ofvascular plant diversity in the flora of B & I examined by Arnold et al. (2002) in Chapter 5and mapped in Chapter 6 of the New Atlas. It somewhat resembles the map for the number of hybridsrecorded in each hectad in Chapter 5. It is possible that the Irish distribution of this species may atthe present time simply reflect local recorder effort and field skills.
British occurrence
In Britain, R. palustris is common and widespread in most lowland damp mud in waterside areas,becoming rare or absent only in N Scotland, although it is also uncommon in Wales and SW England. Onaccount of the identification difficulty with respect to R. islandica s. str., R.palustris might eventually prove to be somewhat under-recorded (Garrard & Streeter 1983;Rich 1991; Preston et al. 2002).
Flowering reproduction
R. palustris flowers from May to October and, while it attracts small bees and flies, generallyit is automatically self-pollinated so that even isolated plants achieve a good fruit set (Clapham etal. 1962; Rich 1991). The average sized plant bears around 200 fruits each of which containsaround 65 seeds, making a total seed production of 13,000 (Salisbury 1942, pp. 202-3).
Seed dispersal, longevity and germination
Seed release is passive after the pod splits, and the propagules are buoyant and remain afloat in waterfor up to twelve days. Like many other members of the Brassicaceae, the seedcoat may become gelatinousand sticky when wet, or the seed adheres externally in mud to waterfowl and other waterside animals.This feature enables at least the very occasional long distance jump-dispersal event to take placebetween different water systems, and also the colonisation of isolated ponds (Ridley 1930, pp. 201-2& 545).
Seed persists in mud for up to six years (Roberts 1986; Thompson et al. 1997). Germination ispromoted by light and fluctuating temperatures and it has twin peaks in March and September. Inwinter-flooded water margin situations, only the former is significant (Grime et al. 1981, 1988;Roberts 1986).
Vegetative reproduction
Effectively reproduction is entirely by seed, but detached portions of vegetative shoot or dislodgedwhole plants can rapidly re-root in wet mud, thus enabling broken or uprooted plants to re-anchor aftertransport by flood water, so that a modicum of vegetative reproduction and dispersal may occasionallysupplement the sexual process. This could therefore permit a very limited amount of clonal developmentof the species to arise (Grime et al. 1988).
Hybridization
Being a comparatively short-lived plant, even when behaving as a perennial, and highly autogamous in itsreproductive strategy, R. palustris is not much involved in hybridization (Jonsell 1975).However, a very rare sterile triploid hybrid is known to occur with R. amphibia on the banks ofthe Thames around Kew, but it has never been reported anywhere in Ireland (Rich 1991).
European and world occurrence
The distribution of R. palustris beyond B & I is widespread throughout Europe except in theMediterranean basin where it becomes very much scarcer and scattered (Jalas & Suominen 1994, Map2324). Elsewhere, the species is more or less cosmopolitan, being considered native in most of theworld, including New Zealand where it was first collected by Banks and Solander in 1769 or 1770(Garnock-Jones 1978), but other populations are also probably introduced there (Webb et al.1988). The same or a similar situation may well apply in S America and in other parts of Australasia(Rich 1991).
Names
The genus name 'Rorippa' originates as a Latinised form of an old Saxon (East German) vernacular orcommon name 'Rorippen', mentioned by Euricius Cordus (Stearn 1992). The Latin specific epithet'palustris' is derived from 'palus' meaning a bog or swamp and thus indicates a plant that lives inswampy places (Gilbert-Carter 1964). The modern English common name 'Yellow Marsh-cress' is a book namethat tells us just a little about the flower colour and habitat.
Threats
None.
Native, common but rather local. European temperate, but also widely naturalised.
1806; Scott, Prof. R.; Lough Erne.
April to November.
Growth form and preferred habitats
R. sylvestris is a yellow-flowered perennial species, 15-60 cm tall, that generally develops thickwhite tap-roots and storage roots from a basal 'crown' immediately below the rosette leaves. Sometimesit also produces additional very slender, horizontal, creeping roots which are branched off from thevertical tap-root and help the plant to spread and form clonal patches. Typically, plants of R.sylvestris also have rather weak stems, which appear to 'lodge' or become decumbent under theweight of the inflorescence, particularly when growing in loose, disturbed or moist, muddy soils. Thesemore or less prostrate stems generally bear a sparse smattering of short adventitious roots, even on theflowering portion of the stem but, nevertheless, they could hardly be accurately described as'rhizomatous', as they sometimes are (eg Webb et al. 1988; Rich 1991).
This often sprawling perennial occupies very much the same range of damp ground, waterside open habitatsas R. palustris (Marsh Yellow-cress), but it is sometimes found growing amongst fairly dense,obviously shading, shoreline vegetation. However, it does also appear as a pioneer weedy colonist ofdrier, more open ground in villages, harbours and along waysides. All Rorippa species tendto disappear when the open vegetation they originally colonised becomes closed, since the competitionthey subsequently experience becomes too strong for them to survive (Jonsell 1968). The need for moreopen conditions explains why Rorippa species appear to favour ± disturbed situations, and thusthey are often very closely associated with human activity, even sometimes to the extent of theirbecoming completely dependent weeds. This colonist behaviour limits both the type of cultivated ordisturbed habitats they occupy, and their means of seed dispersal. Creeping Yellow-cress regularlyoccurs along with R. palustris, but generally it occupies somewhat drier, somewhat less open bareground than the latter typically frequents (Jonsell 1968).
Identification
As with R. palustris, the flowers of R. sylvestris are small and yellow, yet with petalsabout twice as long as the sepals. The fruits are longer than those of other Rorippa species thathave regularly divided (ie pinnate) leaves, and taken together these features enable R.sylvestris to be readily distinguished from other yellow crucifers. Considerable morphologicalvariation within the species has frequently been observed and led to the description of infraspecifictaxa, none of which, however, are based on NW European material
Jonsell (1968). Much of the variation described, particularly in C Europe, is obviously due tohybridization, above all with R. amphibia and R. austriaca (Jonsell 1968).
Variation
Jonsell (1968) found there is considerable variation in leaf shape, plant habit and fruit shape,associated with polyploidy within the species (tetraploid, hexaploid and hybrid pentaploid forms occur).The pentaploid forms are particularly vigorous, aggressive weeds, but they are very rare (Rich 1991).From his study of pollen types and variation in vegetative characters, Jonsell (1968) also showed thatboth tetraploids and hexaploids are composed of numerous biotypes. Oddly, perhaps, while the tetraploidis the most common form in Europe (68% tetraploid versus 20% hexaploid), the hexaploid is most common inN America (65% hexaploid versus 29% tetraploid). Pentaploid hybrids are much more common in the UnitedStates (21%) than in either Canada (10%) or Europe (3%), yet even so, the usual situation of abortingsiliques in nature indicates that at most sites there is just a single chromosome race propagatingvegetatively (Mulligan & Munro 1984).
Fermanagh occurrence
This is another Rorippa species that is very much concentrated in the Lough Erne basin and aroundthe Upper Lough in particular. It is most frequently found in wet or damp ground by lakeshores, oftennear points of shoreline access such as quays, jetties and bridges or as a weed of drier ground invillages and along waysides. R. sylvestris is common in these types of habitat and has beenrecorded in 78 tetrads, 14.8% of those in the VC.
Flowering reproduction
Unlike R. palustris, R. sylvestris is known to be self-incompatible, but it is variably so(Jonsell 1968). Three chromosome races in R. sylvestris have been reported in Europe, but beyondScandinavia where hexaploids are the norm and pentaploids very rare, tetraploid forms dominate. MostR. sylvestris clones have reduced pollen fertility, but this is more the case in hexaploid thanin tetraploid clones, and only among the latter are there clones of good quality pollen. Jonsellregarded this tendency to be expected in plants derived by a process approaching autopolyploidy, whichwould be valid for the hexaploids.
Curran (1984) reported that most studies of R. sylvestris reproductive ability indicate itachieves only a small proportion of its seed production potential. It appears to be the case thatpopulations or colonies very often consist of one genotype only, and thus seed set tends to be severelylimited in such circ*mstances (Webb et al. 1988; Rich 1991). The small Irish population Curranstudied illustrates this effect very well, with less than 10% of fruits setting any seed, and many ofthe fertile fruits having only small seed numbers (ie between one and eight seeds present in them)(Curran 1984).
The sexual reproductive ability of the populations around both parts of Lough Erne has not been examinedin this respect, but the large number of records and the concentration of them particularly around theUpper Lough, suggests that reproduction, by vegetative means or seed or both, is perfectly healthy, andis more than sufficient to support the species.
British and Irish occurrence
The New Atlas hectad map clearly shows that R. sylvestris is less frequent than R.palustris throughout these islands, yet it is widespread in lowland Britain, becoming rarerfurther north in Scotland and the Western Isles. In Ireland, it is very much more scarce and scatteredthan R. palustris. The distribution of R. sylvestris in semi-natural habitats is probablystable, but it has become more widespread as a weed in B & I since the 1950s.
The index of species change in the 40 years between the two BSBI Atlases is + 0.73, a figure whichreflects the fact that R. sylvestris is now recorded in many more hectad squares than was thecase in the earlier BSBI Atlas (Walters & Perring 1962).
European and world occurrence
Beyond B & I, R. sylvestris is regarded as native and widespread in temperate latitudes of W& C Europe, but it becomes scarce or absent in the Mediterranean basin, and it is only of introducedstatus in most of Scandinavia, having originally arrived as ship ballast in many harbours (Jonsell 1968;Jalas & Suominen 1994, Map 2319). Rich (1991) lists R. sylvestris as native also in W Asiaand N Africa and it is widely introduced elsewhere in the world, including in NE USA where it was firstrecorded in 1818, SE Canada in 1894 (Mulligan & Munro 1984), and New Zealand in 1952 (Webb etal. 1988). R. sylvestris rarely produces seed in N America, and it appears therefore tospread there along streams, rivers, ditches, railroads and highways almost entirely by vegetativereproduction, assisted by transport of garden and nursery stock (Mulligan & Munro 1984).
Names
The Latin specific epithet 'sylvestris' in botanical Latin always simply means 'wild', rather thansuggesting any woodland connotation (silva = woodland) (Gilbert-Carter 1964). The plant has no medicinalor other uses and therefore has not attracted any folk names.
Threats
None.
Native, rare, but very probably under-recorded.
1945; MCM & D; Corraslough Point, Upper Lough Erne.
This was declared the most common hybrid Rorippa in B & I by Stace et al. (2015). Ithas petals much longer than the sepals (so differs from R. palustris (Marsh Yellow-cress)),pinnate leaves (so not R. amphibia (Great Yellow-cress)) and non-clasping leaf bases and arelatively long terminal leaf lobe (so not R. sylvestris (Creeping Yellow-cress)). The slenderfruits stand out from the inflorescence stalk at almost 90o (patent) and are shorter (4mm)than the pedicels that bear them (7 mm) (Crawley 2005).
Just seven records of this hybrid from separate tetrads exist in the Fermanagh Flora Database, but asboth parent species are common in the VC, very probably it is simply overlooked and under-recorded. TheNew Atlas map displays records from five Fermanagh hectads in three date classes, but it shouldactually display one more. The plant is found on damp to wet lakeshores and so far it has only beenfound scattered around Lough Erne and the nearby Derryhowlaght Lough.
When, in the 1960s, Jonsell looked at Rorippa forms in B & I during the research for his 1968taxonomic revision of the genus (from which, it is important to remember, he omitted sectionCardaminum, ie the white-flowered water-cresses), he found that records in these islands of theR. sylvestris × amphibia hybrid were sparse, both in herbaria and inthe literature (Jonsell 1968). However, his own field study and examination of herbaria specimens inboth B & I showed that this hybrid, intermediate in form between its parent species, and like bothits parents, variably fertile, was not rare in the areas he studied. It had merely beenpreviously overlooked (see also C.A. Stace, in: Hybridization, p. 152).
Jonsell found both tetraploid and pentaploid forms of this hybrid in English rivers, clearly indicatingthat both tetraploid and hexaploid cytotypes of R. sylvestris take part in hybridisation withtetraploid R. amphibia. Both of these cytotypes are partially fertile, producing some good seed,but are not fully fertile (Jonsell 1968). Backcrossing to both parents, particularly with R.amphibia, has been detected in several places (Jonsell 1975). In the Thames valley, forinstance, Jonsell found that, "from a number of these rivers repeated collections of intermediatehybrid specimens exist". Elsewhere in S England, eg in the River Severn from Shrewsbury down toTewkesbury, he found evidence of extensive hybridisation, leading to distinct signs of introgression inR. sylvestris, "so that pure specimens of the latter [ie R. sylvestris] were in somecases not found at all" (Jonsell 1968, pp. 131-2).
In Ireland, R. × anceps was first reported from Co Leitrim (H29) byPraeger in 1909. Subsequently, it was discovered in Cos Waterford (H6), Clare (H9), Roscommon (H25) andFermanagh (H33) (Cen Cat Fl Ir 2). Three voucher specimens of this hybrid discovered by R.D.Meikle exist at K (Nos 116 & 2123 plus one other), collected in 1945 and determined byJonsell (Jonsell 1968, pp. 132-3). On the basis of his examination of Irish herbarium material, Jonsellquestioned whether the curious serrated leaf-shape typical of Irish R. amphibia plants is not anintrogressed character from R. sylvestris as a result of their frequent hybridisation andconsequent genetic mingling (Jonsell 1968, p. 133, and see his Fig. 15, H & I).
The details of the other six records are: Gubbaroe Point, Lower Lough Erne, 5 July 1945, MCM & D,K; marshy shore, River Erne near Enniskillen, 13 July 1945, MCM & D, K; Duross Point,Lower Lough Erne, 16 August 1946, R. Mackechnie, det. T.C.G. Rich, RNG; jetty at Tully Bay, LowerLough Erne, 22 July 1984, M.J.P. Scannell, det. T.C.G. Rich, DBN; Derryhowlaght Lough, UpperLough Erne, 1988, NI Lakes Survey, det. T.C.G. Rich, BEL; Geaglum Jetty, Upper Lough Erne, 28August 1988, C.D. Preston.
In Britain, R. × anceps is most frequently recorded on the banks ofthe larger rivers and by drains in wetlands that flood in winter. There are also records from the sidesof reservoirs and ponds, water meadows and occasionally from rubbish dumps (Stace et al. 2015).In Europe, R. × anceps is widespread in C & E Europe and local inthe W and S. It is, however, absent from many islands and from most of the Mediterranean region (Jalas& Suominen 1994, not mapped).
Threats
None.
Native, common. Eurasian temperate, but widely naturalised.
1806; Scott, Prof. R.; Upper Lough Erne.
May to October.
Growth form and preferred habitats
Individuals of this rather large, stout but straggling, clump-forming, stoloniferous perennial show adegree of variation in leaf shape that approaches heterophylly, but without there being sharp limitsbetween the leaf types. Early in the growing season, the plant has submerged pinnatifid or deeplypinnately-cut basal leaves, which tend not to persist, but its aerial leaves are simple, quite broad andeither shallowly lobed or distinctly serrately toothed. In Irish material of this species, the leafmargins have many small, deeply incised but irregular teeth (see Jonsell 1968, p. 105, Figs 15, H &I). The curiously dissected leaf-shape of Irish specimens of R. amphibia might represent acharacter introgressed from R. sylvestris (Creeping Yellow-cress) through hybridisation, plusseveral (perhaps numerous) generations of subsequent back-crossing.
The leaves of Great Yellow-cress are also of a distinctive bright green or yellowish colour and, whenflowering, the 3–6 mm yellow petals are about twice the length of the sepals (Clapham et al.1962; Stace 1997).
R. amphibia prefers fertile growing conditions and, as its botanical name suggests, it toleratesfluctuating water levels, often growing on wet ground that only dries out at the surface for shortperiods in summer, if at all. It frequents tall, emergent swamp and fen margin vegetation on shelteredlakeshores and riverbanks. Along slow moving rivers and streams, it grows where the emergent watersidevegetation provides sufficient shelter to allow at least temporary anchorage and maintenance ofshallow-rooted species like R. amphibia (Haslam 1978; Holmes 1983).
The species also copes well with semi-shaded conditions and thus is frequently associated with tall-herbfringing vegetation in eutrophic and generally calcareous waters. It also occurs on muddy clay or siltedsoils of similar chemistry, which again are always subject to at least winter-flooding conditions(Preston & Croft 1997).
Fermanagh occurrence
In Fermanagh, R. amphibia has been commonly recorded from 104 tetrads, 19.7% of those in the VC.It is locally abundant amongst tall, emergent swamp and fen vegetation on sheltered lakeshore andriverbank sites in the county. The fluctuating levels and decidedly eutrophic, calcium-rich waters andmuddy, clayey, sandy or silted shingle shores of the more sheltered, shallow inlets of Upper Lough Erne,provide eminently suitable growing conditions for R. amphibia, as they also do for several otherRorippa species.
However, R. amphibia also occurs in a number of other situations scattered throughout the VC,including the lake shore and margins of wider ditches around Lower Lough Macnean. It also occurs in thesomewhat taller, wetter, 'floodgrass' vegetation around the turloughs (ie vanishing limestone lakes) atRoosky and near Fardrum, plus beside calcareous marl lakes and ponds along the River Finn (known as theFinn Floods). Other populations occurs along the lower stretches of the Silles and Swanlinbar riverswhere the water flow is slow and the substrate is sufficiently deep and stable to support emergentvegetation amongst generally taller, more firmly anchored aquatic plants such as Sparganiumerectum (Branched Bur-reed), Schoenoplectus lacustris (Common Club-rush) or Glyceriamaxima (Reed Sweet-grass).
Flowering reproduction
R. amphibia flowers in its second season of growth, from June to September, and both vegetativegrowth and fruiting reaching their maximum development in late summer. Like R. sylvestris, theflowers of R. amphibia show a high degree of self-incompatibility. Seed set is, therefore,variable; it is often good, but sometimes rather poor despite the conspicuous flowers, the clumpedgrowth of the plant and the consequent plentiful pollination by small bees and flies of various sorts(Jonsell 1968; Rich 1991).
Vegetative reproduction
In addition to seed production, or to compensate if the sexual process should fail, the plant alsoregularly reproduces vegetatively by means of axillary rosettes, which are formed later in the season onlateral stolons at the base of the aerial stems. These vegetative plantlets either enable the formationof clonal clumps, or else they assist seed dispersal in transporting this semi-aquatic species furtherafield, through their breaking off from the parent plant and subsequent floatation (Preston & Croft1997; D.A. Pearman, in: Preston et al. 2002).
Genetic variation
It has been known for almost 40 years that R. amphibia consists of two cytotypes, a rare diploid,known in B & I only from the Thames valley, and a very much more widespread tetraploid elsewhere inthese islands. The tetraploid form is also found throughout most of temperate Europe (Jonsell 1968;Jalas & Suominen 1994, Map 2318). A very powerful sterility barrier exists between these two geneticforms, such that no triploids have ever been found (Preston & Croft 1997). Although there are minormorphological differences between the two cytotypes, they are regarded as too similar to merit taxonomicdifferentiation (Jonsell 1968; Rich 1991).
Hybrids
While the species remains on the whole morphologically uniform (Jonsell 1968, p. 116), the variation thatexists may well be the result of hybridization. R. amphibia is known to form hybrids with threeother water-cress species, R. sylvestris (Creeping Yellow-cress), R. palustris (MarshYellow-cress) and R. austriaca (Austrian Yellow-cress), but only the first of these is at allwidespread in B & I, and its hybrid with R. amphibia (R. ×anceps (Wahlemb.) Rchb.) has just a few Fermanagh records, although here, as elsewhere in B& I, it is very much overlooked.
Jonsell (1968) raised the possibility that the curious dissected leaf-shape of Irish specimens of R.amphibia, might represent a character intergraded from R. sylvestris throughhybridization and subsequent back-crossing.
Irish occurrence
R. amphibia is better recorded in the New Atlas than in the earlier BSBI Atlas(Walters & Perring 1962). This is especially so in Ireland, where it is chiefly found scatteredaround Lough Neagh and Lough Erne, with a similar presence in the Irish Midlands along the Shannonbasin, but with only a sparse sprinkling of stations elsewhere.
British occurrence
In Britain, R. amphibia is common in C and SE England, but absent or a rare introduction north ofa line between Liverpool and Hull. It also appears to have retreated in recent years from some previoussites in the English West Country and in Wales (Preston et al. 2002).
European and world occurrence
The wider world distribution shows the species native from temperate W Europe and N Africa, to E Asia. InW Europe, it becomes rare and scattered southwards in Spain and Portugal and towards the Mediterraneanand is absent from many Mediterranean islands (Jalas & Suominen 1994, Map 2318). R. amphibiais also widely introduced and naturalised in areas including eastern N America and Australasia, forexample, North Island (New Zealand), where it was first recorded in 1870 (Webb et al. 1988). Inmost cases, these introductions are undoubtedly dispersed by human agencies, as for example in the StLawrence River in Canada, where the species was carried long distances with ships (Marie-Victorin 1930).Jonsell (1968, p. 111), concluded that R. amphibia in C Sweden shows a distribution whose bordersare probably not determined by environmental factors, but rather by historical causes, including pastpatterns of boat traffic.
Threats
None.
Introduced, archaeophyte, naturalised from cultivation, very rare.
July 1982; Northridge, R.H.; roadside at Granshagh, 6 km SSW of Enniskillen.
June to August.
Growth form, reproduction and preferred habitats
The large, erect, wavy, coarsely or bluntly toothed, shiny, long-stalked basal leaves of this perennialgive off a strongly pungent, quite unmistakable smell if bruised. Although occasionally it produces atall, leafy spike of numerous self-incompatible, sweetly scented, white flowers from May onwards, it isalmost always completely sterile and, if formed, the small globular seed pods never (or hardly ever)ripen in Britain and Ireland (Weber 1949; Rich 1991). Despite this, Horse-radish, which develops thelong, tapering, fleshy but hard, white, pungent-flavoured tap root for which it is culinary famed, plusnumerous deeply penetrating, thin, secondary roots branching from it, manages to reproduce quiteefficiently and vegetatively by means of these underground organs.
Very small fragments of secondary root material are sufficient for propagation. Uozumi et al.(1992) found that root fragments of A. rusticana with an apical meristem or a branch, couldefficiently regenerate to form whole plants, and that fragments more than 5 mm in length possessed ahigh shoot-forming ability. Root fragments are readily dispersed in transported soil, mud on tools andotherwise by man the cultivator. Once transferred in this manner, often to other cultivated ground,waste ground near habitation or, as in Fermanagh, to roadsides where soil has been dumped, stored orspread, individual established plants can prove extremely persistent, even for example, in overgrown,abandoned kitchen-gardens of long-derelict houses.
Many gardeners regard the plant as eternal, rather than perennial, for once introduced to a vegetablepatch it can become a pernicious weed, virtually impossible to get rid of without resort to repeatedapplications of powerful systemic herbicide. To give an indication of potential root penetration, Edlin(1951, p. 108) tells of excavating roots in a cutting in the English chalk, to a depth of 6 feet [2 m].The secret of cultivating it is to lift the plant every year and to store it over winter in sand.Alternatively, it should be grown in a closed container.
Fermanagh occurrences
There are just four records of Horse-radish, all in non-garden, roadside settings in the Fermanagh FloraDatabase and all recorded by RHN. Apart from the first, which was in a more rural setting, the remainderare all near habitation around Enniskillen and in two villages in the SE of the VC. The details of theserecords are: Lisnaskea, July 1988; by Silverhill Bridge, on the NW outskirts of Enniskillen, August1988; and, just S of Teemore village, June 2002.
Although A. rusticana is extremely persistent when established, it is not an invasivespecies when left unmanaged, at least in Ireland. Even well established plants tend either to besolitary or to display little vegetative spread (Reynolds 2002).
Variation
Being almost entirely sterile and clonal, it is not surprising that several distinct leaf forms of A.rusticana occur and are perpetuated by its asexual reproduction. The leaf forms include, forexample, a distinction between those that are crinkled in texture and markedly notched at the base (iecordate) and others which have smooth surfaces and taper gradually into the petiole (ie cuneate).Illustrations of both these Horse-radish leaf forms are found in 16th century herbals, proving that theclonal distinctions are old and maybe of ancient origin (Courter & Rhodes 1969).
The problem of disease transmission
Another more unwelcome consequence of continual asexual propagation in a crop plant is that microbialdiseases are readily transmitted in the propagated maternal tissue. Horse-radish is widely cultivated inthe USA and produces a root crop of good quality in spite of being very susceptible to White BlisterRust and Horseradish (= Turnip) Mosaic Virus (Weber 1949). Pound (1948) showed that 100% of the thenmost frequently cultivated clone in the USA was virus infected and a great desire arose to producevirus-free Horseradish seed and then to breed greater disease resistance into the crop.
Seed sterility, genetic cytotype and possible hybrid origin
The high level (but not absolute) seed sterility of A. rusticana has been shown to arise frommeiotic irregularities in the formation of male and female sex cells, including only partial pairing ofchromosomes and aneuploidy: the plant has a normal chromosome count of 2n=4x=32, but occasional plantshave only 2n=28 (Easterly 1963). Only twelve viable seeds were harvested following approximately 10,000crosses carried out by Weber (1949). Carrying this work on, Stokes (1955) found that failure to developviable seed following fertilization was due mainly to endosperm-maternal tissue incompatibility,resulting in endosperm failure and, less frequently, in embryo abortion. These facts point to thepossibility that A. rusticana may be of hybrid origin, or indeed it might itself be a hybridrather than a true species (Courter & Rhodes 1969).
While over the years Horseradish has been transferred by taxonomists through numerous related genera, e.gThlaspi, Raphanus, Cochlearia, Radicula, Nasturtium and Rorripa, if it were of hybridorigin, or an actual hybrid itself, no one has yet suggested the possible parents. Molecular isozymegenetic analysis using nuclear and chloroplast DNA by Franzke et al. (1998) has shown A.rusticana is closely related to the genus Rorippa.
There is a considerable body of active genetic research involving A. rusticana, much of itbiochemical and very technical, or associated with tissue culture.
Irish occurrence
Although A. rusticana has been found at least once in 35 of the 40 Irish VCs (Scannell &Synnott 1987; Reynolds 2002), it is really quite rarely recorded on this island, being found mainly onroadsides, waste ground or dumping sites near the larger cities and towns, or in coastal locations whereit may originate in fly-dumped household and garden refuse (a nasty, dirty habit, and unfortunately alltoo prevalent), or like many other members of the Brassicaceae it may just prefer the relatively open,sandy soil of such situations. Support for this latter suggestion comes from the fact that in otherparts of these islands it sometimes colonises sandy seashores, railways and river-banks (D.A. Pearman,in: Preston et al. 2002). De Candolle (1884) noted the same tolerance of salty soils in E Russia.Indeed, the very word 'armoracia', which has been applied to the plant both as a generic name andspecific epithet, is formed from the Celtic words 'ar' meaning 'near', mor, 'the sea', and rich,'against', ie a plant growing near the sea (Barton & Castle 1877).
In Northern Ireland, the relative scarcity of the species is indicated by the fact that it was firstrecorded in Fermanagh as late as 1982, in Down (H38) in 1913, but with no second station until 1946, andin Antrim (H39) in 1965 (Kertland & Lambert 1972; Hackney et al. 1992). The Northern IrelandFlora Website shows A. rusticana is now quite well represented along the coastal shores of Downand Antrim, but is much more rare and scattered inland.
British occurrence
While it is generally regarded as a hardy perennial, A. rusticana is essentially confined to thelowland situations in which it is or has been cultivated and, in Britain, it is so much more widespreadin England compared with Wales and Scotland (Preston et al. 2002), to the extent that one mighteven dare postulate a cultural divide in Horseradish sauce use by the populace!
Toxicity
The pungent aroma and flavour of the root and leaves of this table condiment are due to two glucosides,sinigrin (allyl glucosinolate) and 2-phenylethyl glucosinolate. These poisonous principles are similarto mustard oil and, when hydrolysed, are potent irritants of the eyes and skin and can cause lethalpoisoning of stock animals, including cattle, ponies and pigs (Cooper & Johnson 1998).
Uses
A. rusticana, under a variety of early names, but clearly referring to this plant, was familiarenough as a medicine to the 16th century English botanical Herbal authors Turner (1551) and Gerard(1597), and already both knew of it as an escape from cultivation (Grigson 1987). Thus the plant isnowadays regarded an archaeophyte in Britain and Ireland (ie an ancient pre-1500 AD introduction).Originally, it was a purely medicinal herb with a reputation as a very strong diuretic for treatingdropsy, sciatica, gout and stones, a usage stretching back to Dioscorides in the first century AD(Grieve 1931; Courter & Rhodes 1969).
The first mention of the plant's cultivation in a mediaeval European herbal is by Albertus (a physicianso highly regarded he was also known in his own lifetime as 'Albertus Magnus'), his work dating from1260 (Harvey 1981). However, Gerard (1597) reported that in Germany the grated root, mixed with vinegar,was then being put to its present culinary sauce use for meat and fish, a practice he appears to reportto his countrymen as something novel. By 1640, when Parkinson wrote his early gardening book cum herbal,Theatrum botanicum, he was aware of the plant's condiment use in England and he issued a warningon the strength of its flavour.
Making horseradish sauce from the raw vegetable can be a traumatic experience, as the fumes from thegrated root are much worse than very powerful onions. A modern electric food-processor is the answer,and Mabey (1972, p. 62) provides a good recipe.
European occurrence
Like almost all such early cultivated introductions, the original 'wild' distribution of A.rusticana has been obscured by its long association with man. In a masterly combination ofgeography and philological analysis, De Candolle (1884, pp. 34-6) pointed to the fact that the plant (hecalled it Cochlearia Armoracia) was widely spread over an area of N, W and C Europe (nowadaysmapped for Flora Europaea from Finland (c. 67°N) to C Spain and S Italy, but rare and veryscattered in the Mediterranean basin (Jalas & Suominen 1994)) and beyond this to the Caspian Sea,the deserts of Cuman and in, "several localities in Turkey in Europe, near Enos, for instance,where it abounds on the sea-shore". Nevertheless, as one moved westwards, De Candolle noted theplant became increasingly rare and more scattered; hence he concluded that A. rusticana was notindigenous in W Europe.
Possible area of origin
De Candolle (1884) also identified the word 'chren', common in the numerous Slavic languages of E Europe,as the most primitive name for Horse-radish, a name that was introduced into German dialects as 'kren','kreen' and in French as 'cran' and 'cranson'. Another name used in Germany is 'Meerretig', in Holland'meer-radys', which becomes 'meridi' in the Italian Swiss dialect, all literally meaning 'sea-radish',but these De Candolle regarded as not primitive like 'chren'.
On this basis he concluded that the most probable place of origin was the temperate region of E Europe,from thence its cultivation had been spread westwards for about a thousand years.
Due in part to their original vagueness, and to the numerous subsequent changes and introduced errors innames and uses perpetrated by copyists and later scholars upon the texts of ancient herbal authors overthe intervening centuries, it is often impossible to identify with certainty which of several relatedspecies the oldest works are referring to, the belief is that this plant was mentioned by Dioscorides(1st century AD), yet not by Theophrastus (372-287 BC) and thus it has only been cultivated for lessthan two thousand years (De Candolle 1884; Courter & Rhodes 1969).
Names
As noted above, there is a suggestion that the name 'Armoracia' is derived from Celtic wordsmeaning 'by the sea' (Barton & Castle 1877), but we regard this as doubtful (Deirdre Forbes, pers.comm.). Other authorities suggest the name is Greek or Latin in origin, and that it was first applied byPliny, not to this species which he called Persicon napy (Courter & Rhodes 1969), but toanother unidentified radish-like plant, possibly Raphanus raphanistrum (Gilbert-Carter 1964;Stearn 1992). The specific epithet 'rusticana' is derived from Latin, and like the word 'rustic',means 'of the countryside', or 'of wild places' (Gledhill 1985).
The English common name 'Horseradish' was first given by Gerard (1597). The word 'horse' commonly appliedto a name in this manner, means it is a larger, coarser or stronger variety of plant. Thus 'Horseradish'means 'a strong radish', which is of course apt in this case. Grigson (1974 & 1987) has alsosuggested that Gerard may have coined the name from a mistranslation of the German 'Meerettish',which means 'Sea-radish', in the sense of a radish or root from foreign parts, ie 'Overseas Radish'.Grigson seems to have thought that Gerard took the 'meer' element to mean 'mare', a female horse, oralternatively as 'mähre', which translates as 'an old horse', as if it referred to the rankness andtoughness of the roots (Courter & Rhodes 1969).
The name 'Radish' is derived from Latin 'radix' or the Italian 'radice', meaning a plant valued for itsroot (Prior 1879). It was spelt 'radyce', which is the Old English form, in Turner's (1538) 'Libellusde re herbaria novus' (Rydén et al. 1999, p. 80-81 & Table 1, p. 121).
Previous English Common names included 'Redcole' or 'Redco' (Turner 1551), or 'Redcoll', 'Redcoal''Radcole' and 'Rotcoll', 'Cole' commonly referring to cabbage and other members of the Brassicaceae(Britten & Holland 1886). Lyte (1578) called it 'Rayfort' and 'Mountain Radish', from the French'raifort', meaning 'strong root' (Grigson 1987; Courter & Rhodes 1969).
Threats
None.
Native, very rare, possibly mis-identifications. European temperate.
28 July 2006; ENSIS New Lake Survey; Lough Corry, Kimran Td.
Two records of this quite conspicuous bitter-cress with its rather easily recognised, largish whiteflowers with six distinctive violet anthers were reported to RHN & RSF as VC H33 Recorders by NIEnvironment Agency (NIEA). The species was recorded at two perfectly suitable acidic, peaty lakeshoresites in separate surveys mounted by NIEA in SE Fermanagh. However, we have not been offered vouchersfor these First County Records, and must assume none were collected. Unfortunately the recordinformation for this new species to the VC was so belatedly given we have not been unable to check itsaccuracy. We cannot accept the records at face value in part because the two wetland areas where thespecies is claimed to occur are already well surveyed. In addition, C. amara flowers early inspring from April to June, and thus the late July date of the first record made by visiting Englishbotanists suggests that the plant would most likely be in fruit. It is possible that the visitors wereunaware that this would be a significant extension of the known NI distribution of the species, thatthey expected to find the species, and therefore they did. The most likely error is either anon-flowering specimen of C. pratensis (Cuckooflower) or a Rorripa species.
The seasonal doubt does not attach to the second reported record, the details of which are: 7 May 2009;EHS Habitats Survey Team; Crom Td shore, Upper Lough Erne. However, we cannot confirm the speciespresence without a voucher.
Some credence is lent to both of these records by the fact that Large Bitter-cress has been reported fromsites not too distant from SE Fermanagh in neighbouring Co Tyrone (H36). In his 2010 Flora of CoTyrone, Ian McNeill records C. amara from Lough Nurchossy, SWE of Clogher (Grid referenceH5150), which lies just 5 km from the Fermanagh boundary. It may be significant that in Tyrone, McNeillnoticed that, apart from around the shore of Lough Neagh, C. amara appears to be much morefrequently a riverbank species than a lakeshore one.
Native, common, very widespread and locally abundant. Circumpolar wide-boreal.
1881; Stewart, S.A.; Co Fermanagh.
March to January.
Growth form and preferred habitats
A semi-rosette perennial, with pinnate leaves, shallow rooting and possessing a short horizontal rhizomewhich overwinters with a tiny, almost insignificant leaf rosette (Grime et al. 1988), C.pratensis is most frequent and locally abundant in damp to wet, open (ie well-illuminated) watermeadows (ie seasonally flooded ground) − of which around its lakes, Fermanagh has very many hectares,plus other forms of more or less damp pasture and grassy roadside verges. The species can also be foundmuch less frequently and growing very much more sparsely in shaded situations in drier sites, eg on woodmargins or in clearings, on hedgerows, as well as in wetter lakeside fens and marshy ground by streamsand rivers. It also occurs thinly scattered in upland rushy pastures and, again, in wet or damp flushesto a limited extent, but it is never found on steep slopes or on shallow, rocky ground. Neither wouldone find it in permanently wet ground.
C. pratensis is a plant closely associated with ± closed grassy turf rather than with bare orheavily disturbed ground, but obviously its seed (or plantlets − see below), must colonise gaps in thevegetation cover. Like the majority of small or less sturdy plants (with which it would definitely begrouped), Cuckooflower is only weakly competitive and cannot invade and establish itself among tall,rank, heavily shading and vigorously competing species in any form of vegetation. However, it cancolonise and become established when the environment is modified and plant competition limited byfactors which restrict dominance, including regular grazing, cutting, or moderate levels of other formsof disturbance (Grime et al. 1988; Rich 1991).
It seems to prefer moderately acid to neutral soil reaction (pH 5.0-7.5) and performs best in moderatelyfertile soils, although it is also found in calcareous, base-rich sites. Due to excessive amounts ofcalcium ion, these soils are generally mineral-poor or unbalanced for the plants' nutrientrequirements and are thus best considered nutrient-poor (Dale & Elkington 1974; Grime etal. 1988).
Fermanagh occurrence
Cuckooflower (or Lady's Smock) is one of the most familiar, widespread and welcome spring flowers toFermanagh residents, with whole fields being coloured by its charming, soft lilac blossom in early May.
C. pratensis is in fact by far the most commonly recorded member of the Brassicaceae(Cruciferae) in the Fermanagh Flora Database. There are a huge number of records from 405 tetrads, 76.7%of those in the VC. C. pratensis is well over twice as frequent and three times as widespread asthe second ranking species in the family, Rorippa amphibia (Great Yellow-cress).
Genetic variation
The flower colour of C. pratensis is easily observed to vary from a delicate lilac-mauve to purewhite, but in both genetic terms and in chromosome complement, the plant variation is so great that thespecies really comprises a species aggregate or complex, involving a wide spectrum of chromosome numbersfrom 2n=16-84, plus aneuploids and hybrid forms involving aneuploids (Hussein 1955; Allen 1981). Studiesin a wide range of habitats in C & N England found that almost all C. pratensis populationswere of the same chromosome complement (2n=56) (Dale & Elkington 1974). This study showed there waswide variation in numerous floral and vegetative characters and transplant experiments proved that muchof the variability was environmentally governed (ie the plants were phenotypically very plastic). Thiswork also showed that the form recognised as C. palustris (Wimmer & Grab.) Peterm. in lowlandNW Europe, does not fit English material and, consequently, in his BSBI Handbook, Crucifers of GreatBritain and Ireland, Rich (1991, p. 144) decided it was best to treat C. pratensis as asingle polymorphic species until such time as its taxonomy had been further studied.
Flowering reproduction
The pretty flowers of Cuckooflower are insect pollinated and show a high degree of self-incompatibility,but it is not absolute (Salisbury 1965). The incompatibility mechanism is of the 'di-allelicsprophytic type', similar to the genetic control of 'pin' and 'thrum' heterostyly in the genusPrimula (Richards 1997a, p. 206). Multi-allelic sporophytic incompatibility systems appear to belargely (but again, not entirely) confined to two large plant families, the Brassicaceae (or Cruciferae)and the Asteraceae (or Compositae).
While C. pratensis is extremely variable in numbers of flowers and their size and form, it has asurprisingly low seed output. This is due to a combination of factors amongst which soil moisture level,severity of competition and the intrinsic self-incompatibility, rank as most important. In a closedwater-meadow community, Salisbury (1965) reckoned that the average plant produced less than 200 seedsand, locally, the figure could be appreciably lower than this estimate.
Seed dispersal
Seeds are shed explosively, by the sudden splitting and coiling of the valves of the ripe fruit, amechanism which can fling the seed up to a metre or more. The seed coat is mucilaginous and becomessticky when wet, a feature which undoubtedly enables secondary distant dispersal by attachment to thefeet of animals and perhaps also to bird feathers (Salisbury 1965).
Seed longevity
Buried seed survival is a matter of some dispute, there being many measurements and estimates suggestingit is transient, or short-lived and a few indicating longer term seed persistence and viability for upto five years or more (Thompson et al. 1997).
Vegetative reproduction
It has been known since 1825 that C. pratensis can reproduce vegetatively by the production oftiny adventitious viviparous plantlets on the leaves (Smith 1825, 3, p. 190). Salisbury foundthat these plantlets developed from cells over the veins on the upper surface of the terminal leaflet ofradical leaves and to a lesser extent from other leaflets on the basal leaves. He also confirmed rootingfrom axillary buds took place (Salisbury 1965, p. 331). C. pratensis can carry out additionalforms of vegetative reproduction, by branching of the short rhizome, or from plant fragments (ie brokenstems and leaves), all of which are capable of rooting. So frequent and successful are these asexualprocesses in multiplying and dispersing the plant − particularly in the wetter habitats of the species −that Salisbury reckoned vegetative reproduction was probably the predominant mode of increase of C.pratensis in moist ground. The resultant clonal development could help explain the species' lowseed production, since it would be possible for all the plants in flower at one time to belong to thesame clone and thus self-incompatibility would block seed set.
The frequency of the vivipary is known to vary markedly with habitat conditions and Salisburyacknowledges that it is commoner in "the wetter situations". He quotes White's (1912) Floraof Bristol, where the latter comments that, "the plant multiplies freely by a process ofproliferation from leaflets of the lower prostrate leaves", and, "repeated examinations ofplants in the field have shown that over 70% are viviparous − more in bogs, less in dry places".
Plants have been examined in mown and unmown roadside wet grassland in Fermanagh, Down (H38) and WestCork (H3), none of which showed any evidence of vivipary whatsoever. It would be an interesting projectto examine throughout B & I the frequency of vivipary in this and in other species in which it isreputed to occur. Salisbury (1965, p. 335) lists vivipary as occasional in Drosera rotundifolia(Round-leaved Sundew) and more rarely occurring in D. intermedia (Oblong-leaved Sundew).Buds have been recorded as arising from leaves of various other Brassicaceae (Cruciferae) − see thespecies account of C. flexuosa for more details.
Hybrids
An extremely similar reproductive strategy, also heavily dependent upon specialized foliar and axialvivipary, is described for the triploid hybrid C. × insueta (C.amara L. × C. rivularis Schur), from hay meadows and open pastures in Central Switzerlandby Urbanska (1981). This and other studies emphasises that we should always remember that reproductivecapacity may be largely influenced by environmental conditions.
A hybrid between C. pratensis and C. amara (C. × ambiguaSchulz (C. × mixta Druce)) has been reported from Oxfordshire (VC23), but Stace (1975) commented that the little hybrid seed produced from many pollinations betweenthese species proved inviable and thus reputed wild hybrids reported both in the BI and on theContinent, require verification.
British and Irish occurrence
Reflecting its powers of seed dispersal, C. pratensis is found throughout the whole latitudinalrange of Britain, from the Channel Isles to the tip of Unst in the Shetland Isles. However, it is notomnipresent, being absent from some of the smaller western isles of both Britain and Ireland and alsofrom sections of the SW coast of Ireland and from parts of the English Wash (Preston et al.2002).
European and world occurrence
Due to the unclear pattern of variation and confused taxonomic position of the C. pratensisspecies group or aggregate in B & I, in their European treatment of its distribution, Jalas &Suominen (1994, Map 2350) mapped the seven subspecies of the C. pratensis group together as onespecies. They also mapped the component taxa separately − most of which are either absent or poorlyrepresented in B & I.
The European distribution of the C. pratensis group shows it as native and widespread throughoutthe Continent, but becoming scarce towards the S & SE and absent from most of Greece and all of theMediterranean islands. Hultén (1971, Map 72) shows the C. pratensis group continuouslydistributed around the northern hemisphere and also present as an introduction in New Zealand.
Trends
Populations of C. pratensis probably have declined or been destroyed in B & I during the last50 years, due to agricultural drainage and the increased use of agrochemicals, although there is somesuggestion that the species may contain genotypes resistant to some herbicides (Rich 1991). The NewAtlas (D.A. Pearman, in: Preston et al. 2002) suggests that there has been no overallchange since the earlier BSBI Atlas (Perring & Walters 1976) but this is the picture when working at'the one record in a 10 km square' level of discrimination.
Casual and garden forms
Cuckooflower does occasionally occur as a casual garden weed of damp, heavy soils and most especially oflawns. The latter is not really surprising, since recent mowing experiments in an abandoned fen meadowshowed this treatment had a positive influence on population survival of C. pratensis and othertypical wet meadow species (Jensen & Meyer 2001).
Double flowers are also sporadically found in the wild and in Fermanagh, as elsewhere, these are oftenbrought into garden cultivation. Small populations of 'normal' C. pratensis have been reportedfrom neglected damp grassy areas in suburban gardens in Dublin city and these plants are considered aconsequence of the increasing commercial trade in 'wild flower seed mixtures' (Scannell 1997).
Folklore and names
The fact that this species has so many English common names (Grigson (1987) lists no less than 53)suggests that it must have numerous traditions associated with it. Many of the local names havespringtime associations, eg involving the cuckoo which sings in April and May, or names which contain'May' as an element, or lambs. Other names involve milkmaids and their smocks, or the lady (ie our lady,the Virgin Mary), pigs, pigeons and their eyes. Grigson (1987) gives quite a detailed account of thesenames, their sexually suggestive connotations and the tradition of ill luck if they are picked (analternative name is 'Pick-folly'). Vickery (1995) also recounts several witnesses to the ill luck onflower gatherers. Several folk names include mention of 'blobs' and these and the references to 'milk'may readily be connected with the familiar sight of 'cuckoo-spit' on the plant − the white foamprotecting frog-hopper nymphs from predators. In his Dictionary of English Plant Names, Grigson(1974) enlarges on the origins of the two most frequently met names, 'Cuckoo-flower' and 'Lady's Smock'in similar vein. Another general name for the genus is 'Bittercress' and the young leaves do have apeppery taste and have been used as a Cress substitute in salads and sandwiches (Mabey 1996, p. 151).
There are two notions regarding the origin of the genus name 'Cardamine'. It is either derived from twoGreek words 'kardia' meaning 'heart', and 'damao' meaning 'subdue', a reference to its medicinalproperties as a heart sedative (Johnson & Smith 1946; Chicheley Plowden 1972), or it may simply be acarry over of an ancient Greek name given to some cress-like herb in the Dioscorides herbal(Gilbert-Carter 1964; Stearn 1992). The Latin specific epithet 'pratensis' simply means 'growing inmeadows' (Gilbert-Carter 1964).
Threats
None.
Native, common, widespread and locally abundant. European temperate, but also in C and E Asia andpossibly introduced in eastern N America.
1881; Stewart, S.A.; Co Fermanagh.
January to November.
Growth form, species origin and preferred habitats
Morphologically very similar and genetically very closely related to C. hirsuta (HairyBitter-cress), most Fermanagh records have been distinguished from the latter on the 'ji*z' of the plant(ie overall size, leafiness and hairiness of the stem), plus the number of stamens, ie plants having apreponderance of six anthers were taken to be C. flexuosa. While C. hirsuta is always anannual species, C. flexuosa is very much more variable in terms of life-form and it can be eitherannual, biennial (ie a short-lived monocarpic perennial), or even a polycarpic (repeat flowering)overwintering perennial (Grime et al. 1988; Webb et al. 1996). When it is behaving as anannual, C. flexuosa can germinate in the autumn − germination probably under day-length controland, in this case, it overwinters as a small wintergreen leaf rosette. Alternatively, the seed mayundergo chilling and vernalisation, so that germination is delayed until the springtime.
The flexibility in life-form which C. flexuosa can assume, allows it to exploit a variety ofecological situations, ranging from the colonisation of temporarily open, bare ground or unoccupiedlitter, to persistence in a rather more closed vegetation cover − where the species can persistas a biennial or as a perennial, always provided conditions such as grazing, seasonal shade or flooding,ensure that competition is not excessive, or sufficient to oust the plant.
Wavy Bitter-cress is a tetraploid (2n=32), which is believed to have arisen by allopolyploidy from asterile hybrid between C. hirsuta and C. impatiens (Narrow-leaved Bitter-cress) (both2n=16), followed at some stage by chromosome doubling to create the new, fully fertile species (Ellis& Jones 1969).
Since C. flexuosa and C. hirsuta are so closely related, they share many of theirecological requirements and tolerances. Thus they overlap in their many habitats and may occur together,making their distinction even more uncertain at times. However, having hybrid origin, C. flexuosatends to be the larger and more lax in growth of the two species, but it is not necessarily the morevigorous, or the more floriferous plant.
Comparative competitive ability
Neither of these two Cardamine species has much competitive ability, so they tend to occupy eitherbare soil or mud, or gaps in vegetation cover produced by some form of moderate disturbance thatminimises negative interaction with more aggressive, robust herbaceous species. Very early, rapidseasonal growth and the ability to self-fertilise their tiny flowers characterises both theseCardamine species, allowing them to successfully reproduce earlier than most associated speciespresent in the wide range of habitats they frequent.
C. flexuosa prefers, or is more restricted to, somewhat damper, deeper, moderately fertile andacidic, organic soils (but not in strongly acidic or permanently wet conditions), in comparison withC. hirsuta. Of the two species, it most typically frequents cooler, more sheltered, semi-shadedsituations, where the likelihood of desiccation is reduced or minimised, although in these habitats itsometimes has to compete with its close relative.
In the Sheffield area of England, C. flexuosa was categorised as intermediate between astress-tolerant ruderal and a ruderal species by Grime et al. (1988). In the current author's(RSF) view, however, this classification would exaggerate the level and frequency of disturbance thatC. flexuosa faces in its typical Fermanagh habitats which are more semi-natural than urban incharacter.
Fermanagh occurrence
C. flexuosa is much more frequently found than C. hirsuta in woods and bystream-sides, lakeshores and in shaded, sheltered, damp rural places in general – includingwaysides, cliffs and overgrown areas in old quarries. Since these are common situations inFermanagh, it is not surprising that C. flexuosa has been recorded more than 2.5 times asfrequently as C. hirsuta in the county and in very nearly twice as many tetrads, ie in 291tetrads, representing 55.1% of those in the VC.
Flowering reproduction
Quantitative data on the flowering and seed production capacity of C. flexuosa seemunavailable, but both are probably very similar to those of C. hirsuta. As in otherCardamine species, seeds are explosively released and may be flung up to a metre from the parentplant. The soil seed bank of C. flexuosa is less persistent than that of C. hirsuta, theseed being classified as either transient, or surviving at most one year in the ground (Thompson etal. 1997).
Vegetative reproduction
C. flexuosa is capable of vegetative reproduction by the rooting of shoot fragments broken off andtransported as a result of disturbance (Grime et al. 1988), but so far the extent or significanceof this appears to be unstudied. Salisbury (1965), writing of the viviparous plantlets produced onleaflets of C. pratensis, mentions that buds have been recorded arising from leaves of variousother members of the Brassicaceae (Cruciferae), and he lists six species, including C. flexuosa,C. hirsuta and C. impatiens (Narrow-leaved Bitter-cress). He points out, however, that,"... there is a suspicion that some at least of these may have been axillary buds that haddeveloped roots ... " (ie rather than leaf cells giving rise to plantlets), "... sothey are not comparable", ie with the viviparous situation he was describing in C.flexuosa (Salisbury 1965, p. 335).
British and Irish occurrence
While common and widespread throughout most of B & I, the New Atlas shows C. flexuosais less well represented in the Irish midlands and on the extreme W coast, while in Scotland it thinsout towards the NW, and in E England it is absent from parts of Cambridgeshire, S Lincolnshire and SEYorkshire (Preston et al. 2002). At the same time the distribution is not considered to havechanged since the earlier BSBI Atlas (Perring & Walters 1976).
European and world occurrence
In continental Europe, C. flexuosa has a markedly western distribution (not extending much furthereast than Poland) and thinning towards the south and absent in many of the Mediterranean islands. To thenorth, it is more extensively represented in Scandinavia than C. hirsuta but, unlike it, C.flexuosa is absent from Iceland (Rich 1991; Jalas & Suominen 1994, Map 2375). Although thetaxonomy is uncertain, five related forms (possibly subspecies) are believed to occur in the Himalaya, EAsia and the Philippines (Hultén 1958, Map 125; Grime et al. 1988; Rich 1991). Thus C.flexuosa is polymorphic, and the subspecies (or perhaps related species) are widely disjunct intheir distribution around the N Hemisphere (Hultén & Fries 1986, Map 934). Subsp. flexuosa isalso sparsely present in eastern N America. It is claimed by some to be native there, and thereforeamphi-Atlantic, but it is much more likely an introduction, as is definitely the case in Australasia.
Names
The Latin specific epithet, 'flexuosa' is derived from 'flecto' meaning 'bend' or 'curve'and refers to the rather wavy, flexuous flowering branch typical of the species (Gilbert-Carter1964).
C. flexuosa is not sufficiently distinguished from C. hirsuta to have any English commonnames associated with it.
Threats
Very possibly capable of further increase in fertile, disturbed ground, but probably not endangering anyother native species in doing so.
Native, common, widespread and locally abundant. Eurosiberian southern-temperate, but very widelynaturalised and now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Characteristically a winter-annual, perennating as a leaf rosette 4-8 cm across and perfectly capable ofsurviving hard frosts even as very small seedlings (Salisbury 1962, p. 349), C. hirsuta is a veryabundant and, indeed, cosmopolitan ruderal weed of disturbed and cultivated ground. The species isparticularly loathed by gardeners on account of its short generation time, fecundity and vast seed bank.
Hairy Bitter-cress is very closely associated with damp or dry disturbed ground and with shallow, verydry soils in crevices on walls, paths, car parks, in waste places and hedgerow bottoms. However, it doesoccur, although less frequently, in more natural settings, eg in the shade of woods and scrub and moreoften in open well-lit conditions, on cliffs, ledges and screes – particularly in limestone districts.It is also said to frequent sand-dunes (Rich 1991) but, being land-locked, Fermanagh is devoid of these,although there are some local sand quarries where the plant does occur.
Fermanagh occurrence
C. hirsuta has been recorded less than half as frequently in Fermanagh as C. flexuosa (WavyBitter-cress) – the latter plant favouring our wetter soils. The two species are closely related (seethe C. flexuosa species account above) and they can be difficult to distinguish, but there arerecords of C. hirsuta from 145 Fermanagh tetrads, 27.5% of those in the VC.
Identification
C. hirsuta and C. flexuosa are closely related (see the C. flexuosa species accountabove) and they can be difficult to distinguish. Useful field characters that separate them are stamennumber (C. hirsuta almost always has four, C. flexuosa almost always has six), theleaf-stalks of the basal rosette are decidedly hairy in C. hirsuta, while C. flexuosa hasa hairy, flexuous stem, usually bearing more, and more prominent, stem leaves than C. hirsuta.
Flowering period
C. hirsuta flowers very early in the season − in mild areas of the country from March onwards whentemperatures become favourable. In damper soils, however, it often completes two or more generations ofits brief life-cycle, flowering from March to May, and then again from fresh crops of plants throughfrom June to October (Salisbury 1964; Grime et al. 1988). Thus, as with other common garden weedssuch as Lamium purpureum (Red Dead-nettle) and Euphorbia peplus (Petty Spurge), floweringand fruiting of Hairy Bitter-cress may occur during eight months of the year in many parts of the B& I (Salisbury 1962, p. 350).
Seed production capacity and dispersal
The tiny seeds are released explosively and can travel up to 80 cm in still air whenever the ripe fruitis touched or disturbed even slightly (Salisbury 1964, p. 109). Although normally it is only a smallplant, 7-30 cm tall, flower and seed production are high (even on plants dwarfed by regular oroccasional disturbance). Typical small sized plants each produce an average of around 600 seeds(Salisbury 1964). Under ± ideal conditions, a single large, branched fruiting individual can shed hugequantities of seed.
As with other members of the genus, the seeds are coated with a layer of mucilage which becomes stickywhen wetted, so that secondary dispersal by attachment to animals, including ourselves, and in thegarden, by sticking to tools, wheels and so on, helps convey the species over longer distances(Salisbury 1964).
As is always the case, reproductive capacity is extremely variable and dependent upon the environment,especially upon soil conditions. For example, comparison of a sand dune population with an adjacent onegrowing on clay on a wall top, found the former produced c 98 seeds per plant, while the plants onshallow clay had a mean of 640 seeds (Salisbury 1942). In the same season, a well grown plant in amanured garden soil under little or no competition, produced an estimate of around 52,000 seeds. Underthese very favourable conditions this plant showed there were increases in both the number of seeds perpod, as well as in the number of pods per plant (Salisbury 1942, pp. 44-5).
Germination
C. hirsuta seed may or may not undergo a period of after-ripening or natural dormancy. Astudy by Salisbury (1962, p. 390) suggests there is, if any, a very brief delay − only two to threeweeks between seed release and first germination of this species, but according to Grime et al.(1988), after-ripening requires several months, and the main period of Hairy Bitter-cress germination isin the autumn. Roberts & Boddrell (1983) found some seedlings appeared in spring, but most did so insummer. Whatever the case is regarding dormancy, (and it may vary between populations), eventualgermination is characteristically intermittent, generally spread over six or more weeks, and theannual's plant growth in spring, or in summer, is rapid, but very dependent on suitable temperature andmoisture conditions.
Dwarfed plants and seed longevity
The plants have a shallow fibrous rooting system and very often in mid-summer, because of droughting, theplants and their flowers are so tiny that they can readily be overlooked. However, since even dwarfedplants are capable of flowering and the flowers automatically self-pollinate, unless the plants areheavily disturbed, they generally succeed in fruiting and releasing additions to the soil seed bank,where they can persist for five or more years (Roberts & Boddrell 1983; Thompson et al.1997).
Hybrids
A sterile triploid hybrid between C. hirsuta and C. flexuosa is known to occur. Said tomore closely resemble C. flexuosa, it is easily overlooked or very rare (the only BI record is inVC 47 (Montgomeryshire)), and nothing suspiciously like it has been noticed in Fermanagh (Stace 1975;Rich 1991).
British and Irish occurrence
The New Atlas shows C. hirsuta is common and widespread throughout almost the whole of B& I, becoming less frequent in NW Scotland and W Ireland. It is regarded as introduced in some ofthe most northerly Scottish islands.
Following the advent and increased use of herbicides from the 1960s onwards to control weeds and tomaintain parks, gardens and waysides, short-lived ruderals, including C. hirsuta, have tended toincrease in these types of artificial habitat, especially if they are sprayed in spring or early summer.The chemical kills off, or severely curtails, the vigour of perennial competitors, thus allowingseedlings of opportunistic, rapidly evolving, annual species like C. hirsuta the chance to fillthe liberated ecological space (Warwick 1991). In recent years, Hairy Bitter-cress has become a majorweed of horticultural nurseries and garden centres in B & I, since it so rapidly colonises thesurface layers of bare soil in plant containers.
In the garden, vigilance and regular (even weekly!) hoeing before the plants fruit is the answer, ormulching with 5 cm of grass cuttings; otherwise it requires a resort to herbicide (Roth 2001).
European and world occurrence
C. hirsuta is an originally Eurasiatic species that has spread widely around world (Hultén &Fries 1986). In Europe, C. hirsuta is very widespread in the W & C, from the Mediterraneanbasin northwards to around 62°N in Scandinavia and Iceland. However, it rapidly peters out in moreeasterly areas and is only present beyond longitude 30°E along the northern shore of the Black Sea(Jalas & Suominen 1994, Map 2376). Beyond this, it has spread widely across both hemispheres, sothat its distribution has become circumpolar southern temperate and, indeed, it is often now considereda cosmopolitan weed (Rich 1991; Hultén & Fries 1986, Map 935).
Having said this, ten plants collected in S Australia in the middle of the 19th century and previouslyregarded as native C. hirsuta (Bentham1863), all turned out after modern examination to be anendemic species (Kloot 1983). A large measure of scientific caution is required when assessing the worlddistribution of any weedy species, since very often closely related yet different taxa occur insimilar habitats around the globe. In this respect, we may be talking of anything from varieties tospecies, or possibly even genera. On the other hand, the pace of modern air transport means that seedcan all too readily hitch a ride, so that rapid invasion of new territory is a very real possibility.
Names and uses
The English common 'book-name' is 'Hairy Bitter-cress', but its flavour is not bitter at all. In fact itsfresh young leaves are pleasantly tangy and delicious and nutritious in salads and, not surprisingly,reminiscent in flavour of Water-cress (Rorippa nasturtium-aquaticum agg.), but a little sweeterand more fleshy (Mabey 1972). Carl Linnaeus named the related English wildflower Cardamine amara,the Latin specific epithet of which means 'bitter'. No country folk would ever have described theseplants as being bitter in flavour. However, despite this the Herbal and Flora writers copied oneanother and between them managed to transfer this inaccurate name to almost all the species of thegenus.
The Latin specific epithet 'hirsuta', of course, means 'hairy', which is rather overstating the case inthis instance, although the upper surface of the leaflets is slightly hairy.
Worldwide, C. hirsuta has numerous local English common names such as 'Spit-weed', 'Shot-weed','Flick-weed', 'Touch-me-not', 'Popping Cress' and 'Poppits', all of which refer to the verycharacteristic explosive seed release. Other names listed by Britten & Holland (1886) include'Lamb's Cress' and 'Land Cress' (as opposed to 'Water-cress') − but the latter name is also sometimesapplied to Barbarea vulgaris (Winter-cress). The name 'cress' comes from the Old English'cærse', 'cerse', or 'cresse', with cognate names in other Germanic languages, andall these words are derived from an Indo-European base meaning 'to nibble' or 'to eat' (Grigson 1974).
Whatever we call it, we can agree that C. hirsuta is generally very abundant and very widespread,easy to weed out in the garden, but virtually impossible to get rid of entirely. Perhaps we should eatmore of it in our salads!
Threats
None to it, but the species is spreading with the assistance of the horticultural trade and it is atorture for gardeners.
Native, occasional. Circumpolar boreo-temperate.
1881; Smith, T.O.; Ardunshin.
April to February.
Growth form and preferred habitats
The erect fruits, small white flowers and flat, overwintering basal rosette of bristly, hairy, dark greenleaves, make this a distinctive and easily recognised short-lived perennial or biennial species (Grimeet al. 1988; Ryser 1990). The typical habitats of this species in B & I are shallow, warm,well-drained soils on exposed limestone cliff ledges, rock crevices, scree slopes and fairly steep rockypastures. In Fermanagh, however, this tap-rooted species also colonises a few more obviously man-made ordisturbed sites, including limestone gravel foundations on the sides of forest tracks and rock faces andspoil in a limestone quarry. In other areas of B & I, it also occurs on sand dunes and in limemortar on old bridges and walls (D.A. Pearman, in: Preston et al. 2002). The species iscompletely absent from heavily or regularly disturbed sites, however, and also from wetlands.
A. hirsuta is an excellent example of what is meant by a 'calcicole' plant. This term meansthat it is closely confined to base-rich rocks (in Fermanagh – Carboniferous limestone) and theirderived, infertile (nutritionally unbalanced), unproductive, often open, well-drained soils, whichfrequently support a moderately species-rich plant assemblage. Vigorous, potentially dominantcompetitors are either scarce in these soils, or limited by other factors including grazing pressure anddrought. Occasionally, Hairy Rock-cress is also found on adjacent more acid, peaty ground, alwaysprovided this is percolated by base-rich drainage water (Grime et al. 1988; Rich 1991). At higherlevels of fertility, competition from more vigorous taller growing species quickly ousts rosette plantslike A. hirsuta, so it must be assumed that even in these percolated peaty areas, theproductivity of the vegetation remains low (Ryser 1990, p. 51).
Fermanagh occurrence
Currently there are records in the Fermanagh Flora Database from a total of 38 tetrads, 7.2% of those inthe VC. Apart from Smith's early record listed above, A. hirsuta is completely confined to thelimestone terrain west of Lough Erne. The isolated nature of the Ardunshin record in north-easternFermanagh (which has a voucher in BEL), strongly suggests that Smith collected the plantelsewhere in the county at a time when site location was not regarded as significant. Along with all hisother biological records, the site was simply given as his home address!
Flowering reproduction and genetic variation
Although usually Hairy Rock-cress is a polycarpic perennial, it has no means of vegetative spread orreproduction, relying entirely on plentiful seed production for the species increase, survival anddispersal. Numerous (often more than 50), small white flowers are produced on an erect, 5-60 cm tallstem from April or early May onwards, with the slender fruits ripening through into August or September(Grime et al. 1988). Just exactly how A. hirsuta carries out fertilization is still notfully resolved, since throughout its range the plant forms a polyploid complex of microspecies, four orfive of which occur in Europe (Jalas & Suominen 1994; Roy 1995).
An American study by Roy (1995) found that allozymes of A. hirsuta were variable at the populationlevel and that progeny arrays revealed fixed levels of heterozygosity, suggesting that apomixis (ieasexual seed formation) was taking place. However, Roy (1995) could not rule out the possibility ofselfing polyploids, since many European populations have been shown to be tetraploid and these can beeither apomictic or sexual, since the even chromosome number which they possess allows normal meioticpairing to occur.
Titz (1972), who has studied the species aggregate in great detail, found that European A. hirsutaplants require pollen for seed set. From his studies, he has concluded that the plants are sexual,normally self-pollinating and autogamous (ie self-fertilizing). Titz also believes that the species hasnormal meiosis and he made successful inter- and intra-specific crosses involving it. Data in Roy's(1995) study suggested that the populations he examined were pseudogamous, since there was nosegregation of alleles in progeny arrays. However, since these arrays probably were the product ofself-pollination of the wild collected seed he used, Roy feels that further study is required todistinguish between pseudogamy and autogamy in A. hirsuta.
Seed production and dispersal
Seed production is quite high for a relatively small plant, each pod or siliqua containing approximately40 seeds (Grime et al. 1988). Thus a minimum of around 2000 seeds per plant is quite usual, withlarger plants achieving production several times this figure. The seed is small, light, flattened andnarrowly but variably winged. While there are no special adaptations for its dispersal, we may assume itrelies on wind for transport (Titz 1972; Rich 1991).
Seed germination and buried longevity
The seed does not require chilling for germination and the vast majority of seedlings emerge in theautumn (86-96%). The exact period of the autumn depends upon growing conditions, but generally it occursbefore the end of September (Grime et al. 1988; Ryser 1990). The survey of soil seed banks in NWEurope found that A. hirsuta seeds were either transient (persisting less than one year), orshort-term persistent (surviving buried for between one and five years) (Thompson et al. 1997).
Population studies
In a detailed experimental population study of nutrient-poor species-rich limestone grassland in NSwitzerland, Ryser (1990) found that moderate cover or shelter provided either by the proximity ofneighbouring plants or by a relatively loose 1-1.5 cm moss layer carpeting the soil in vegetation gaps,slightly delayed, but actually enhanced the levels of germination and establishment of A. hirsutaseedlings. These same growing conditions also favoured spring and early summer survival of the youngA. hirsuta plantlets growing in a Swiss environment, since winter frost-heave of the soil surfacedisturbed their roots and exposure to subsequent dry, sunny April conditions resulted in desiccation andmajor population losses. Total mortality was high when compared with five other species in this study,only 23% and 15% of the autumn cohort of seedlings surviving through their first year of growth in 1986and 1987 (Ryser 1990, p. 24). The pattern of establishment shown by A. hirsuta in this study wasclearly very much governed by the harsh abiotic conditions of late winter and early spring inSwitzerland.
A similar investigation of A. hirsuta population behaviour and ecology carried out under themilder, oceanic growing conditions typical of Ireland or western Britain might well provide a totallydifferent analysis of the species' growth and survival strategy and such a comparison would be verywelcome.
British and Irish occurrence
Two varieties are recognised within A. hirsuta by Sell & Murrell (2014), of which var.hirsuta is the most widespread. This variety is locally concentrated in its British distributionon base-rich, mainly chalk or limestone derived soils. In Ireland, both varieties probably occur; var.brownii (Jord.) Titz being frequent on sand dunes and more rarely on rocks in the West. Overall,in Ireland, A. hirsuta is much more infrequent and sparsely scattered than in Britain and it ismainly (but not exclusively) westerly and coastal in its occurrence (Rich 1991; Preston et al.2002).
A. hirsuta has declined in Britain over the past 80 years, probably due to intensification ofa*griculture combined with the species low competitive ability, which prevents it colonising moredisturbed and/or artificial habitats (Grime et al. 1988).
European and world occurrence
Beyond the British Isles, the A. hirsuta species complex is widely represented throughout W, C andNW Europe from N Africa and N Spain to the W shore of the Black Sea and northwards to within the ArcticCircle in Norway, although it is absent from Iceland and the Arctic Islands (Jalas & Suominen 1994,Map 2397). A. hirsuta s.l. extends right across Eurasia and the whole complicated species complexspreads around almost the entire northern hemisphere in boreal and temperate continental latitudes,although it is absent from higher latitudes in most of Canada and from Greenland (Hultén 1971, map 152;Hultén & Fries 1986, Map 940).
Names
The genus name 'Arabis' is Greek and literally means 'of or from Arabia'. Since none of the plantscurrently referred to by the name occur in that country, most authors of books on plant names skirt thisembarrassing difficulty and refer to it as being 'of obscure derivation'. However, Chicheley Plowden(1972) suggests the connection might lie in the ability of the genus members to thrive in drysituations, which must be accepted as something of a truism! The Latin specific epithet 'hirsuta'translates as 'hairy' or 'rough-haired', the latter being appropriate in this case (Gledhill 1985).
The plant has not attracted any uses or folklore and the English common name 'Hairy Rock Cress' simplyinforms us that it is a hairy member of the cabbage family, that frequents dry, rocky sites.
Threats
None.
Introduction, neophyte, a very rare adventive, often associated with horticulture and gardens.
7 July 1988; Northridge, R.H.; wall of a church in Clabby village.
April to September.
Growth form and preferred habitats
Thanks to its well-developed taproot and spreading fibrous root system, this small winter annual istolerant of fairly dry soils in May and early June at what really is the end of its growingseason. In terms of its ecological characteristics, D. muralis is essentially a warmth-loving,Mediterranean or sub-Mediterranean, spring flowering, calcicole annual of open, winter-moist, butshallow, immature, sometimes rather unstable soils of base-rich Carboniferous limestone geology. Thepreferred soil has plenty of lime, is often stony, free-draining, rich in humus and has a pH of 6.5 orabove (Ratcliffe 1960).
D. muralis is a well-adapted winter annual and typical therophyte, very resistant to adversegrowing conditions and thus avoiding competition. While often persistent in its existing stations, itsdistribution is much confined by its limited powers of natural dispersal, a situation further aggravatedby its seed survival being transient. Only the fact that it is accidentally transported by man explainsits current wide and still spreading distribution.
Flowering reproduction
Following vernalization by winter cold, from mid to late April onwards plants flower freely irrespectiveof their size. Flowering and fruiting continues throughout May and June, or even into July provideddrought does not intervene by killing the plants (Ratcliffe 1960). The small white flowers areautomatically selfed, each resultant fruit containing around ten or eleven small, light seed, althoughthey can produce up to 16 in each oval, flattened pod. Salisbury (1964), who published these figures,reckoned the mean production per plant was over 500 seeds and Ratcliffe (1960) likewise estimated meansof between 500 and 1500 per plant.
The species has no specialised mode of seed dispersal, wind probably carrying the seed little beyond onemetre from the parent plant.
Germination
After seed dispersal, there is an after-ripening period of around two months which prevents prematuregermination in the summer months, even if conditions are mild and wet. The current author (RSF) has notlocated any information in the literature on seed longevity or persistence in the soil (eg in the surveyby Thompson et al. 1997), but presumably it is transient (ie less than one year). Germination ofthis winter annual can be described as ± continuous once it has begun. It stretches over a period ofaround 25 days, beginning in the September after seed production when the soil becomes permanentlymoist, although the process may exhibit a slight intermittence towards the end of this period. Up to 65%of the seed germinates in the autumn and there is a hiatus of growth in the months around the turn ofthe year, with germination and rosette growth resuming in late February or March depending, upon theseasonal weather (Salisbury 1964).
Fermanagh occurrence
D. muralis has persisted, for instance, on the wall of St Margaret's Church in Clabby since atleast July 1988. It has also been seen, along with Erophila verna (Common Whitlowgrass), ongravelly path sides on the Necarne Estate near Irvinestown in April 2000, but these two stations for thespecies, both discovered by RHN, are the only ones so far recorded in Fermanagh. The species istherefore not yet established in this VC, and is thus an adventive.
British occurrence
While D. muralis is a rare native with small, scattered colonies on lower-lying, warmed limestonerocks in SW England, the Peak District and the Pennines, it does manage to reach up to 490 m in theCraven Pennines in W Yorkshire − where the first discovery of the species in Britain was made by JohnRay in 1670 (Ratcliffe 1960; Rich 1991; D.A. Pearman, in: Preston et al. 2002).
It addition to this very limited native distribution, D. muralis is also widely but thinlyscattered elsewhere in Britain, mainly in mild, winter-wet, mild western districts, to which it"spread artificially", as it was quaintly put by Ratcliffe (1960).
The anthropogenic dissemination of the plant in Britain began early in the 19th century, for it wasreported from the old botanic gardens in both Edinburgh and Glasgow by 1824 and 1865 respectively. Manyother early county records beyond the native English range of the species, describe its discovery inhorticultural nurseries and in botanic or private gardens, nearly always growing on old stone wallswhere the lime mortar provided the preferred nutrient conditions, along with little or no competition.
Irish occurrence
D. muralis is also a very thinly and widely scattered alien throughout Ireland, having beenrecorded in 20 of the 40 Irish VCs at any date (Reynolds 2002). The New Atlas map displays itsoccurrence in 21 hexad squares with 1987 or later dates (Preston et al. 2002). Here again, as inBritain, dissemination along with garden plants from particular horticultural nurseries is the mostlikely source of records (Brunker 1950) and the plant is always found in man-made habitats involvinglimestone rock, or on old walls with lime mortar.
The earliest Irish record appears to have been a single plant on the walls of Blarney Castle, CountyCork, found by Mr James Drummond, a Scotsman who around 1809 was Curator of the short-lived Cork CityBotanic Garden and who emigrated to W Australia in 1829 (Praeger 1949). His undated find was publishedin the Catalogue of the indigenous plants of Ireland (Mackay 1825). The second published Irishrecord was by another Scotsman, George Dickie, who reported it growing, "On old walls aboutBelfast", in a supplementary list in his Flora of Ulster, recognising it as an introduction(Dickie 1864).
It appears that D. muralis is a well adapted winter annual and typical therophyte, very resistantto adverse conditions, and thus avoiding competition. While it is very persistent in its existingstations, its distribution is much confined by its very limited powers of natural dispersal, a situationfurther aggravated by its seed survival being transient. Only the fact that it is accidentally orotherwise transported by man explains its current wider distribution.
European and world occurrence
D. muralis is considered native in temperate areas of W, S & C Europe, extending eastwards toTurkey and the Caucasus. It is also native in NW Africa and Madeira (Rich 1991). It is absent from muchof NW France, the Netherlands and becomes more scattered northwards into southern Scandinavia (Hultén& Fries 1986, Map 963). It has been introduced in N America (Rich 1991).
Names
The genus name 'Draba' is from the Greek 'drabe', a Classical name given by Dioscorides to a plant of theCabbage Family, possibly Lepidium draba (Hoary Cress) which was supposed to have value inpoulticing whitlows, that is, wounds of the nails, and hence the English common name. The Latin specificepithet 'muralis' simply means 'growing on walls' (Stearn 1992).
Threats
Extreme rarity always represents a survival problem.
Native, occasional. Eurosiberian southern-temperate.
1900; Praeger, R.Ll.; Co Fermanagh.
February to November.
Growth form and preferred habitats
This is a small, rosette-forming, ephemeral, winter annual of dry, shallow, nutrient-poor, neutral tocalcareous soils. In much of B & I, Erophila verna s.l., or the polyploid species aggregateis generally common and predictable in any open, disturbed habitat having a high proportion of baresoil. It often occurs in a wide range of linear or urban situations including, in B & I, bothcoastal and inland examples, in base- or lime-rich, sandy, stony or rocky soils (Salisbury 1964; Prestonet al. 2002).
Man-made habitats include paths, or in mortared crevices of pavements and walls, plus alongside railwaytracks where these survive (Salisbury 1964; Sinker et al. 1985). E. verna is tolerant ofexposure and is widespread at low to medium altitudes throughout B & I and more rarely up to around2,400 ft (730 m). It is completely absent from Shetland and becomes increasingly rare or absent in themore acidic soils and the wetter conditions of W Ireland, NW Scotland and the English Midlands(Salisbury 1964; Preston et al. 2002).
Fermanagh occurrence
In Fermanagh, E. verna s.l. is occasional only, or infrequent, and has been recorded in 40tetrads, 7.6% of the total in the VC. It is found in a wide range of artificial and semi-naturalhabitats from urban waste ground, along roadsides (and previously along railway lines), plus lakeshore,wayside gravel, car parks, paths, slipways and quarries. As the tetrad distribution map clearly shows,although decidedly uncommon, the distribution of E. verna s.l. very definitely follows linearhabitats – both artificial (roads) and semi-natural (rivers). For a number of years, E. vernas.l. grew in long swathes on the gravel covered hard shoulder alongside the Enniskillen to Belfast roadfor some kms. In this site, the flowers in early spring were so dense they resembled a recently fallenshower of hailstones. Subsequent road works tarred over many of the areas where the plant previouslygrew so abundantly and to date it has not re-established in this site to anything like the same extent.
Flowering reproduction
The leafless stems produce a variable number of small white flowers with deeply notched petals from thebeginning of March until June. Growing as they do in open, often exposed and disturbed, shallowinfertile soils, plants of E. verna are typically very small or dwarfed, and they tend to produceonly a few fruits per plant. In Norfolk sand dunes, for example, Kelly (1984) found that plants wereseldom more than 4 cm tal and a many were less than 1 cm tall. Small plants produce very few fruits andKelly's work proved that so-called 'depauperate' plants bearing only one or two fruits also producereduced mean numbers of seeds per fruit. The mean number of fruits on the Norfolk dune E. vernaplants varied over three years between 1.54-2.39 per plant, yet despite such low figures the plantpopulations of E. verna and five other associated 'depauperate' species, appeared relativelystable in numbers.
E. verna flowers habitually inbreed, self-pollination automatically occurring when the outerstamens dehisce immediately adjacent to the ripe entire stigma (Rich 1991).
Seed dispersal, survival and ecology
The winter annual life cycle, which is so frequently and so well displayed in small, short-livedcruciferous species like E. verna, may be considered a very effective drought-avoiding syndrome.The small seeds of these little annuals facilitates wind dispersal of the species and, in thisparticular case, the seed also displays remarkable longevity, surviving five or more years in the soilseed bank (Thompson et al. 1997). Germination drops off very rapidly after the second year ofstorage in damp soil, however. In a Polish sand dune experiment, an average of only 5.7% of the E.verna seeds present in the surface layers germinated in the spring under natural conditions,although recruitment (referred to as 'natality', which really means 'birth rate' (Holmes 1979)) variedgreatly (as expected) with existing plant density (Symonides 1984).
The early spring flowering of E. verna has traditionally been used by farmers in some parts of theBritish Isles as a guide to when conditions are suitable for sowing spring barley (Vickery 1995).
Variation
Erophila verna s.l. is a species aggregate or complex which consists of a large number ofmorphologically different cytotypes, each exhibiting genetic constancy. The 19th century French botanistAlexis Jordan set out to distinguish and describe these local populations and as a result of hiscollections and their subsequent cultivation, he was able to describe 53 "elementary species"and recognise more than 200 distinct genetic "lines" or "morphs" in the genus(Briggs & Walters 1997, pp. 30-1). The situation is maintained by habitual, long-term inbreeding.This in turn has created hom*ozygous, true-breeding, pure genetic lines that are morphologicallydistinct, eg differing in fruit shape, size, seed number per fruit and pubescence (Proctor & Yeo1973; van Andel et al. 1986). This situation led Filfilan & Elkington (1988, 1998) followingthe earlier study of Winge (1940), to delimit four forms within the species complex with differingchromosome counts between 2n=14 and 2n=64, recognising each of them at the species level. One of thesecytotypes with 2n=24 (E. semiduplex Winge), appears confined to Germany, but the other threeforms occur in B & I. So far, only one of these segregates has been recognised in Fermanagh (seeE. glabrescens Jord. below) and their true distribution in B & I has yet to be properlydetermined (Rich 1991, pp. 256-9; Filifilan & Elkington, in: Rich & Jermy 1998, pp. 126-8;Preston et al. 2002).
While these new forms (species or otherwise) may prove useful constructs, Kelly's study should still beborne in mind. The fact remains that Symonides (1983, 1984) found that individuals of E. vernarespond in a plastic manner to seedling population density. Also, van Andel et al. (1986)discovered a strong correlation between plant morphology and seed weight in populations of E.verna and this in turn reflects multi-niche selection and adaptation to environmental'uncertainty'. This may reflect, for example, soil moisture and nutrient levels and, possibly, also theeffect of plant burial by blown sand in dune populations.
The existence of highly inbred annuals appears to contradict both the Darwinian principle that Natureabhors perpetual self-fertilization and the Mendelian notion that crossing, even if onlyoccasional, is essential in order to maintain genetic fitness. As with every situation in life, inreproduction there are costs and benefits to be analysed and the balance of outcomes closely depends onthe particular circ*mstances, rather than a simple 'one size fits all' scenario. Forms of E.verna with relatively large seed, for example, may enjoy a competitive advantage allowing rapidearly growth of seedlings and might result in these plants overtopping surrounding vegetation. Othermorphs with a smaller seed size may allow the fruiting plant to produce greater numbers for the sameoutlay of resources, thus conferring significantly greater powers of dispersal into bare ground, orenabling colonisation of sites where seedlings face less competition, or occupy a more mesic, lessecologically testing environment.
Habitual selfing is particularly common in polyploid plants such as E. verna s.l. and its closerelatives, in which the chromosome number has increased (usually doubled), often followinghybridisation. It has been suggested that the extra genetic material that this process creates, may wellreduce or dilute the effect of deleterious mutant genes that inevitably accumulate over time in thistype of ± continually selfing species (Proctor et al. 1996, p. 335).
European and world occurrence
Erophila verna s.l. or the species aggregate is considered native in large parts of Europe, N.Africa and W Asia (Hultén & Fries 1986, Map 966), but the complicated taxonomy and recentsubdivisions has led to a great level of uncertainty as to the distribution of species (or subspecies)and varieties, so they really are unknown at present.
Names
The genus name 'Erophila' comes from the Greek 'er' meaning 'spring', and 'philos' meaning 'loving'(Stearn 1992). The Latin specific epithet 'verna' means 'of the spring' and is very appropriate. TheEnglish common name 'Whitlow Grass' was first coined by Gerard (1597) for this medicinal plant, whichwas too inconspicuous to have acquired common names (Grigson 1974). The species was known to 16thcentury herbalists as 'Paronychia vulgaris', from its use in (supposedly) curing a whitlow (inLatin, paronychia), ie any puss* inflammation at the nail of a finger or toe (Watts 2000). The word'whitlow' is a 14th century modification of the previous 'whitflaw', thought to be derived from 'whiteflaw' (Hanks 1986).
Threats
None.
Native, very rare, but almost certainly under-recorded. The world distribution range is uncertain.
23 April 1999; Northridge, R.H.; hard shoulder alongside the Enniskillen-Belfast road near Coollane.
April to May.
Very little work has been done on separating the various species and subspecies of the Erophilaverna (Common Whitlowgrass) complex in the Fermanagh survey, a situation which so far is thecase throughout most of the British Isles (New Atlas).
E. glabrescens first appeared in a national Flora as a separate taxonomic entity in the thirdedition of the Flora of the British Isles by Clapham et al. (1987), a source ofinformation not consulted since it was quite rapidly and almost completely superseded by Stace's (1991)New Flora of the British Isles. During most of the Fermanagh flora survey RHN and the currentauthor (RSF) preferred to carry around and rely on as our field identification yardstick, the sixthedition of An Irish Flora (1977). Knowing now what the available alternative was (Stace 1991),this is quite a major confession.
The significant paper on the British Isles distribution of the constituent taxa of the Erophilaverna polyploid complex, based largely on herbarium records, (Rich & Lewis 1999, Table 1 andFig 4) showed that, in Ireland, E. glabrescens had at that time a total of 39 records from 32hectad grid squares. Fermanagh was not represented in the paper though a total of just three records didexist from N Ireland sites: one each from Cos Tyrone, Down and Antrim (H36, H38 & H39). Three yearslater, Preston et al. (2002) published a map with records from two additional N Ireland hectads,adding grid square J34 in Co Down and the first Fermanagh record given above, from grid square H33.There are now two additional Fermanagh records in the new century and they lie in contiguous gridsquares, H15 and H24. The sites are on roadsides along the lowland axis of the county, in or near theonly substantial conurbation, namely Enniskillen town. All three Fermanagh records were made by RHN andthe remaining details of the other two are: on Sligo road, Enniskillen town, 4 May 2000; abundant in thecar park at Tully Castle, Lower Lough Erne, 16 April 2001. The habitats involved are lowland, artificialand disturbed, ie roadside verges or hard shoulders, and a tarred public car park.
It is quite clear from this evidence and from the comment of D.A. Pearman and C.D. Preston in the briefspecies account accompanying the B & I hectad map in the New Atlas, that being a small andrather insignificant looking ephemeral spring annual and, furthermore, a polyploid taxon or species thathabitually selfs, E. glabrescens is difficult to reliably distinguish from E. verna s.s.It therefore remains seriously under-recorded throughout the British Isles. This is especially so inIreland in comparison with Britain, since a very much smaller population of sufficiently expertrecorders is resident and active.
Introduction, archaeophyte, a casual weed, now extinct. Native range uncertain, probably Europeanboreo-temperate, but widely naturalised.
14 July 1953; Moon, J.McK.; waste ground in Enniskillen town.
Growth form, reproduction and preferred habitats
Previously, this overwintering annual or biennial was a regular, accidentally introduced seed contaminantand arable weed of flax, corn and lucerne. In parts of Britain, this small-flowered yellow crucifer withyellowish, inverted pear-shaped fruits was also a wool-shoddy alien.
The quite tall plant flowers in midsummer and is either pollinated by bees or self-pollinates (Garrard& Streeter 1983). The 12-20 brown seeds produced per pod (around 900 to 4500 per plant) areshort-term persistent, surviving in the soil for about a year or so (Salisbury 1964, pp. 121-2; Thompsonet al. 1997).
Until the advent of efficient scientific seed cleaning in the 1940s, it was a frequent and widespreadpersistent weed of arable cultivation throughout B & I. At the same time, it was always rather localand generally casual in its appearance (Garrard & Streeter 1983; Blamey & Grey-Wilson 1989; Rich1991; Clement & Foster 1994).
As an oil-seed and fibre producing plant, C. sativa is known to have been either an importantcultivated crop in its own right and/or it was tolerated along with another oil-seed crucifer species,Eruca vesicaria (Garden Rocket) as a common contaminant of Linum usitatissimum (Flax) sowneither for linseed oil or for its fibre. These three species grow together in the Near East, beingharvested and processed far back into prehistory when the distinction between 'crop' and 'weeds' wasalmost or entirely irrelevant (Loudon 1829; Jones 1988; Rich 1991).
N Ireland was a major flax growing area in the 19th and early 20th century, the fibre being used in thejustly famous and still surviving Irish linen industry. The current author (RSF) suspects thatC. sativa probably persisted here in the Province of Ulster (the nine northern counties ofIreland), longer than it did in other parts of Ireland.
Fermanagh occurrence
Despite this historical connection with the Ulster linen industry, Gold-of-pleasure has only once beenrecorded in Fermanagh as listed above. In recent decades, C. sativa has been re-introduced tothese islands in a fresh context. It is again spreading as a local casual throughout B & I, the seedbeing a constituent of wild bird food mixtures, provided on garden bird tables (Hanson & Mason 1985;Clement & Foster 1994). Nowadays, it frequents various forms of disturbed ground, for instance nearfeeders in gardens, chicken runs, dockyards, waste ground and rubbish tips (D.A. Pearman, in: Prestonet al. 2002). We believe it is only a matter of time before it appears again in Fermanagh fromthis type of source.
Irish and British occurrence
The New Atlas records just five post-1987 10-km squares recorded for the species in Ireland, onein Co Dublin (H21) and the remainder in Co Limerick (H8) along the estuary of the River Shannon. TheCensus Catalogue of the flora of Ireland lists a further 14 VCs from which old records exist, butwhere it is now regarded as extinct (Scannell & Synnott 1987). Similarly, the New Atlas plotsjust 43 10-km squares with post-1987 records for the species in Britain (Preston et al. 2002).
European and world occurrence
As a consequence of its connection with Flax cultivation, the native range of C. sativa isuncertain. At present, it is widespread in C & SE Europe and in SW Asia and has also been(presumably accidentally) introduced to the Far East, N & S America and Australasia (Rich 1991).
Names
The genus name 'Camelina' appears to be derived from two Greek words meaning 'on the ground' or 'dwarf'and 'flax', that is, 'Dwarf Flax', which is one of the alternative English common names and obviously adirect translation. The Latin specific epithet 'sativa', applied to so many plants, as always means'planted' or 'cultivated' (Gilbert-Carter 1964). Other English common names include, amongst others inBritten & Holland (1886), 'Cheat', 'Dutch Flax' and 'Gold of Pleasure', all of which probably alludeto the fact that the seed appears like flax but is not! The rather poetic name 'Gold of Pleasure' issaid by C.P. Johnson in his 1862 book The useful plants of Great Britain to, "bear ironicalreference to the disappointment of its first cultivators here, who found their investment in it about asprofitable as gold squandered on 'pleasure' usually proves." (Watts 2000).
The very ingenious derivation of this English name given by Prior (1879) based on an account of anoil-producing plant he says is mentioned by 'Gerarde', involving the corruption of 'Oleo de Alegria' to'Oro de alegria' (Gold of Pleasure). Unfortunately, this passage does not appear to refer to Camelinasativa at all (see Gerard 1633, p. 273) and despite quite a diligent search of the latterreference, the current author has failed to locate the passage to which Prior refers.
Threats
None.
Introduced, archaeophyte, common. Eurosiberian wide-temperate, but widely naturalised and nowcircumpolar.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Although very variable in plant size, fruit and leaf-form, this small cosmopolitan annual, oroccasionally biennial weed with its distinctive and unique triangular, flattened fruits has to be one ofthe most familiar and best recorded ruderal species in the flora of B & I. Its reproductive strategyand dispersal is so effective that C. bursa-pastoris has become one of the most ubiquitous weedsin the world and thus it features in Holm et al.'s 1977 book, The world's worst weeds.However, being a small, non-competitive annual, definitely pernicious, but seldom present in largequantity and certainly not noxious, it hardly deserves such an extreme degree of notoriety. Widespreadhuman transport of the species is achieved by numerous methods, but Reynolds (2002) in her Cat AlienPl Ir proposed that it arrived in Ireland with animal foodstuff.
C. bursa-pastoris is a stress-tolerant pioneer colonist, ± confined to disturbed or bare soil inopen habitats where it can avoid competition. It prefers disturbed, fertile soil situations and,elsewhere in B & I, is most frequent as a weed of arable crops and gardens, particularly ofbroad-leaved vegetables, eg potatoes, cabbage, peas and sugar-beet (Holm et al. 1977). However,it is also very common in less fertile, disturbed habitats, including trampled or compacted ground,especially where the land is seasonally wet, eg around farm gateways, animal troughs, or in clayeydeposits at the base of walls. Only rarely is it found in gaps in pastures, meadows or in woods.
The plant rapidly develops a rosette of basal leaves and a tough branched taproot. It is the latter thatenables it to successfully colonise and survive in potentially dry, stony or calcium-rich soils, eg inbuilding rubble, cliff crevices, or in trampled and compacted ground. While tolerant of some drought,essentially C. bursa-pastoris is a mesophyte, preferring near neutral, moderate to base-richsoils. It is therefore absent from very acid, very dry, or permanently wet sites.
Fermanagh occurrence
C. bursa-pastoris is common and widespread in disturbed, open habitats throughout Fermanagh, beingrecorded in 256 tetrads, 48.5% of those in the VC. As the tetrad map indicates, it is least frequent orabsent on the wetter, more acid soils of the Western Plateau area of the county.
Flowering reproduction
The species is tetraploid, probably of hybrid origin
and reproduction is entirely by seed (Hurka & Neuffer 1997). Plants frequently branch from near thebase and stems produce numerous long racemes of flowers. As is the case with other ruderal therophytesin this family, C. bursa-pastoris flowers readily and profusely. It also is predominantly aself-pollinating species, often self-fertilising before the flowers open. Under cloudy and rainyconditions flowers are mostly self-pollinated before they open, but dry, sunny weather favoursoutcrossing. At low temperatures (c 4-10°C) the duration of flowering is prolonged up to five-fold, butpre-conditions favouring cross-fertilisation are strongly reduced (Hurka et al. 1976).
The usually rapid fertilisation process enables C. bursa-pastoris to complete its life-cyclewithin six weeks with a guaranteed full seed set. Greenhouse experiments suggest outcrossing is rareunder field conditions, but is somewhere between 0-20%, so that the breeding system is flexible (Hurka& Neuffer 1997).
The gradual shedding of ripe seed from the plant, together with a very wide range of phenotypicplasticity within the species in response to its environment, frequently allows two or more generationsto complete their development within a single growing season. The number of potential offspring producedin a year in this inbred manner is extremely high (Salisbury 1964) and the abundant seeds are long-livedin the soil, persisting for at least 30 years (Salisbury 1964).
Seed germination
Cool temperatures below 10ºC break seed dormancy and since like other cosmopolitan weeds C.bursa-pastoris is indifferent to day-length and photo-period control, subsequent germination isintermittent and can occur during any month of the year. Germination is favoured by seasonally higherand preferably naturally fluctuating temperatures, plus exposure to light (Salisbury 1963, 1964; Popay& Roberts 1970). These particular germination conditions produce a major burst of growth in earlyspring, while subsequent tillage or disturbance of the soil during the summer brings to the light afresh supply of non-dormant seed, which will then germinate given sufficient warmth and moisture.
Herbivory and ecology
The plant is eaten by stock and by wild animals, most notably rabbits (Crawley 1990), but slugs are alsopartial to the leaves, as are numerous insects (Aksoy et al. 1998). Being an insubstantialannual, C. bursa-pastoris is very much an opportunist colonist and pioneer species of open, bareground conditions and it has very little competitive ability, so that even if abundant amongst crops, ithas little effect on the ultimate yield.
European and world occurrence
C. bursa-pastoris is probably of Mediterranean origin and has spread far and wide as anagricultural weed to develop a worldwide distribution that avoids only the hot and the wet tropics(Hurka & Neuffer 1997). A related diploid taxon, C. rubella Reut. replaces it in southernparts of C Europe (Hultén & Fries 1986, Map 971).
Names
The genus name 'Capsella' is a diminutive of 'capsa' meaning 'a box', and refers to the small,notched-triangular or heart-shaped fruit capsule which opens downwards by means of two valves. The Latinspecific epithet 'bursa-pastoris' translates as 'shepherd's purse' from 'pastor' meaning 'shepherd' and'bursa', 'purse' (Gilbert-Carter 1964). This and the familiar English common name 'Shepherd'sPurse' derive from the shape of the small triangular fruit which resembles a miniature mediaeval leatherbelt purse and, as the seeds are yellowish to golden brown and oval and flattish in shape, they resembleminiature coins and thus reinforce the allusion (Salisbury 1964). In Europe, in ancient times, it wasused as a pot-herb and it is indeed quite nutritious, eg 100 g of the fresh basal leaves in the springcontaining one and a half times the recommended daily human requirement of vitamin C (which is 60mg/day) (Zennie & Ogzewalla 1977).
Threats
Possibly capable of further increase and spread in disturbed habitats, but unlikely to oust any but otherruderal species.
Introduction, archaeophyte, a very rare casual. Native origin obscured by long history of cultivation,widely naturalised.
24 July 1995; Northridge, R.H.; garden greenhouse in Enniskillen Town.
Growth form and preferred habitats
This annual is the original 'cress' of 'mustard and cress' seedlings, which was grown in gardens forwinter salad for at least a thousand years. Its seed germination and growth on dampened blotting paperby windowsills was watched with interest by many generations of primary school children.
The salad form of the species has been selected for its milder, peppery taste, which is sharp butpleasant when eaten raw and at the young cotyledon stage (Edlin 1951). In his brief note on the subjectin BSBI News, Rich (1988) helpfully provided an illustrated key to the seedlings of the plantsinvolved, but the curious and unusual three-lobed seed leaves or cotyledons of L. sativum reallyare absolutely unmistakable. Rich's excellent Crucifers of Great Britain and Ireland details thedistinguishing characters of the more adult plant (Rich 1991, p. 230).
L. sativum is frequently used as a plant bioassay in many kinds of laboratory and fieldexperimental situations and, for instance, a 'Google Scholar' internet search for"Lepidium sativum plant bioassays" on 11 April 2021 produced 3,930 literature 'hits'.
Fermanagh occurrence
L. sativum has been found only once in Fermanagh. As listed above, it was growing as a weed andproducing seed in an Enniskillen greenhouse, rather than in a truly wild or semi-wild situation.Nowadays it has been almost completely replaced by Brassica napus (Rape) both in the schoolcurriculum and in salads purchased in shops and in eating establishments (Rich 1988).
British and Irish occurrence
Elsewhere in the wild in B & I, L. sativum now occurs only rather rarely as a casual,non-persistent individual, or in small populations, chiefly as a ruderal on roadsides, or on disturbedor waste ground, in rubbish tips and occasionally in newly re-seeded grassland (Reynolds 2002). The seedsources nowadays appear to be as bird feed and grass seed mixture contaminants, as well as from culinaryand horticultural waste. In England, north of the Midlands, the New Atlas map shows L.sativum is very thinly and widely scattered and there are very few recent records of it at allfrom Scotland, Wales and Ireland (Rich 1991; D.A. Pearman, in: Preston et al. 2002).
Centre of origin
The species is considered to originate somewhere in W Asia and/or N Africa (Rich 1991), possibly in Iran(Edlin 1951), or Egypt (Jalas et al. 1996, p. 209), although being one of the more ancient saladplants it is so widely introduced, cultivated and naturalised worldwide that nobody can be certain whereit comes from, so that the latter reference decided not to map it.
Names
The English common name 'Cress' is descended from Old English 'caerse', 'cerse' and 'cresse' and hasequivalents in all other Germanic languages. These words all derive from an Indo-European base meaning'to nibble' or 'to eat' (Grigson 1974). The name 'Cress' is often loosely applied to many differentmembers of the Cabbage Family, but according to Prior (1879), when used absolutely, it properly refersonly to the genus Lepidium.
Threats
None.
Possibly introduced, very rare. Apparently oceanic southern-temperate, but the native range is uncertain.
1950; MCM & D; abundantly by the roadside south of Glenross.
Growth form and preferred habitats
L. heterophyllum is a rather weedy rosette-forming perennial, or more rarely a biennial plant witha stout woody rootstock and branching grey-green stems 10-50 cm tall. The basal leaves disappear beforeflowering, but they subsequently re-sprout after fruiting (Clapham et al. 1962). The numerousprostrate or more or less procumbent flowering branches are clothed throughout the season with manycrowded, short, stem leaves, both toothed at the side and with auricles at the base to varying degrees.
The species colonises dry, disturbed lowland ground, as well as arable fields and dry, open pastures. Itappears to avoid calcareous conditions (being something of a calcifuge) and often is a plant ofdry acidic soils on heaths and gravelly places by paths, roadsides verges, alongside railways and onwell-drained embankments. In other parts of the B & I, it is frequent on seashore shingle and, to amuch lesser extent, it also appears in arable fields and short-turf pastures, where we can assumecompetition is of a low order. The established strategy of L. heterophyllum is categorised asSR/CSR, ie intermediate between a stress tolerant ruderal and a competitor-stress tolerator-ruderal(Grime et al. 1988 & 2007).
Flowering reproduction
Terminal clusters of numerous small, white flowers are produced from May to August or September and theviolet or reddish anther colour (prior to their bursting) is a useful distinguishing character fromL. campestre (Field Pepperwort) which is otherwise similar, but which has yellow anthers (Rich1991; Webb et al. 1996). American common names for the plant are 'Purple Antherfield Pepperweed'and 'Variable leaved Pepperweed', both of which are helpful reminders of significant identificationfeatures of the species. The flowers appear to be self-fertilized and the fruits contain just one 2 mmdiameter seed in each of their two compartments (Fitter 1987).
In ecological terms L. heterophyllum appears to be a fairly undemanding plant of dry, sunny,infertile, open, more or less disturbed, stony conditions where competition is fairly minimal orrepressed (Sinker et al. 1985). L. heterophyllum is also said to be tolerant of grazingpressure (Rich 1991) − probably because, like L. latifolium (Dittander), its roots andleaves taste burning and bitter (Gerard 1597; Grigson 1987).
Just as there appears to be little or almost nothing written on the ecology or biology of this species,there is likewise no mention of its dispersal mechanism, seed longevity or germination in any of theliterature the current author (RSF) has consulted (eg Ridley 1930; Grime et al. 1981; Thompsonet al. 1997). Clearly, if we are ever to understand what governs the occurrence of widespreadplant species like this one, we must have some more basic biological research data to fill such obviousgaping holes in our knowledge. However, the ecological behaviour of L. heterophyllum and L.campestre (Field Pepperwort) appears rather similar and their British distributions appear socomplementary (L. heterophyllum declining in the SE, which is the principal area of L.campestre), that one might suggests that the latter could possibly be competitively excludingthe former (Preston et al. 2002).
Fermanagh occurrence
Although very rare in Fermanagh, L. heterophyllum is by far the most common Lepidiumspecies in NI. It is generally reckoned to be a native species in Ireland, as is also the case inBritain. However, there are only four Fermanagh records, three of which date from 1950. It has beenfound by RHN in only one site recently. Apart from the first record given above, the remaining detailsare: sand pit near Killadeas, 1950, R. Mackechnie; roadside 1.5 km W of Tempo, 1950, MCM & D; oldQuarry at Lisbellaw, 3 July 1994, RHN.
Questionable native status of the species
While in Fermanagh this species is very rare and occupies, or once did, disturbed ground on roadsides,quarries and sand-pits, RHN thinks it might possibly be an introduction, or an escape from arablecultivation. Elsewhere, in quite a wide portion of B & I, L. heterophyllum is a fairlyfrequent or local, uncompetitive species of dry, acid, sandy, gravelly or heathy soils. It is also afrequent species of coastal shingle, railway ballast and embankments and less commonly appears in arableground and in dry, open pastures. The current author (RSF) finds it difficult to accept that the statusof this grey-green crucifer species is different in Fermanagh from everywhere else in the British Isles.However, we might well ask, what real evidence is there for regarding this species as indigenous inthese islands? There does not appear to be any fossil record of the species (Godwin 1975), so we areleft to consider circ*mstantial evidence to determine its status (Webb 1985).
Rich (1991) suggests that L. heterophyllum is native in Europe from Spain to Czechoslovakia and is"occasionally introduced elsewhere in Europe". Jalas et al. (1996), in Atlas FloraeEuropaeae, 11, map the species as being native only in W Europe (Spain, Portugal,France and the British Isles), but they believe the species is probably or more definitely introduced inregions from the Netherlands northwards into Scandinavia, and entirely absent from Italy and moreeasterly and more continental European regions (Jalas et al. 1996, Map 2787). It is introduced inN America and Australasia (Rich 1991).
There is no space here to further argue the case, but the current author (RSF) concludes that in commonwith other widespread weeds of disturbed ground, the native distribution of L. heterophyllum isbasically unknown and remains a disputable subject. The status of all such species in botanicalliterature owes more to tradition than to science. Currently developing genetic techniques should,however, eventually allow an analysis of species cytotypes and this may then provide definitive answersto questions of native occurrence of weedy species such as this one.
British and Irish occurrence
In view of its weediness, it is rather surprising that L. heterophyllum is not more generallydistributed in B & I than it is. The New Atlas shows it most commonly distributed along theIrish Sea coasts of both islands, plus the S coast of Ireland and NE Scotland, but only scattered andmuch rarer elsewhere across these isles. It is either entirely absent or very rare in the Midlands andthe west of Ireland (Preston et al. 2002).
Names
The genus name 'Lepidium' is a Greek name used for some plant by the classical botanist Dioscorides andis the diminutive of the Greek word 'leptis' meaning 'a scale', the fruits of Lepidium beingconsidered scale-like (Gilbert-Carter 1964; Stearn 1992). The Latin specific epithet 'heterophyllum'means 'other leaves', or 'diverse leaves', that is the plant has leaves of different forms (Stearn1992).
Previously, the botanical name of the plant was L. smithii, called thus in honour of Sir JamesEdward Smith (1759-1828), author of the influential English Botany and much else (McClintock1966, pp. 82-4). This connection is retained in its current recommended English common name, 'Smith'sPepperwort', sometimes also given as 'Smith's Cress' (Melderis & Bangerter 1955). Other moredescriptive common names include 'Downy Pepperwort' and 'Hairy Pepperwort', the former the more accuratesince the whole plant is usually (but not always) well furnished with a soft pubescence of short, grey,simple (unbranched) hairs. The American name 'Purple Antherfield Pepperwort', provides a reminder of thesimple differential character between this species and L. campestre, with which it probably ismost often confused.
The name 'Pepperwort' was first used by Turner (1568) for L. latifolium (Dittander), which in the16th century was cultivated for its roots and leaves that were used in making hot pungent sauces untilpepper and Horse-radish drove it out (Grigson 1987). Gerard (1597, 1633, p. 240) says "the root issharp and biteth the tongue like pepper, whereof it tooke the name pepperwort".
Threats
None.
Introduction, neophyte, established, but very rare. Native origin unknown, possibly S American.
11 August 1989; Northridge, R.H.; flower tub in Enniskillen Town.
May to October.
Growth form and preferred habitats
A small, sprawling, soft, deeply pinnatisect, feathery-leaved, alien weedy ruderal annual or biennial isgradually spreading across these islands. In view of the colonising behaviour of the species in Europeand elsewhere, its gradual spread to all regions, while not spectacularly rapid, remains inexorable.
Flowering reproduction
C. didymus forms a disc-like mat of prostrate shoots spreading out around the original longcentral tap-root bearing finely divided pinnate leaves which smell strongly foetid or pungentlycress-like when bruised. The very small, generally petal-less flowers are crowded on severalinconspicuous inflorescences amid the leaves. However, the flowers are actually borne opposite theleaves, or less often in the leaf axils, or completely terminal. Flowering takes place from mid- tolate-summer and automatic self-pollination is the norm, although it has been suggested that ants maysometimes be involved (Rich 1991). A study in India found that four species of ant plus two of aphidswere attracted to the flowers by their unpleasant scent and by nectar (Chauhan 1979).
Salisbury (1964) estimated that, depending upon size, individual plants could produce between 1,600 and18,000 seed per season. The small doubly notched, twin-valved fruits (which give the plant its specificname, Latin 'didymus' meaning 'double' or 'twin' (Gilbert-Carter 1964)) contain a single seed in eachlobe. The two fruit halves split apart when mature and are probably distributed in mud by animals,including man. Dispersal may perhaps also involve ants as vectors.
Seed of Lesser Swinecress persists in the soil seed bank for more than five years and while germinationoccurs chiefly in April and September, flushes of emergence following soil disturbance can occur in anymonth except December and January (Roberts 1986). Growth of the plant occurs all year round in milderareas of New Zealand (Popay et al. 1995; Roy et al. 1998). Since the oceanic climate inour part of Ireland is rather similar to that in lowland New Zealand and as so many 'Kiwi' plants growexceptionally well in our gardens, we may expect the same sort of year round growth and seed and weedpopulation behaviour by C. didymus if or when it becomes fully established in Fermanagh and otherparts of western Ireland.
Centre of origin unknown
The origin and native distribution of this small ruderal plant is unknown. There is disagreement amongFlora writers as to whether it is from Eurasia (Hickman 1993), or S America (Clement & Foster 1994;Webb et al. 1996; Stace 1997). Whatever the truth of the matter, it is now a very successfulcosmopolitan weed (Baker 1972).
Fermanagh occurrence
The first Fermanagh record for this species was in a planted urban flower tub as detailed above. Thereare just four additional records as follows: Lakeland Forum, Enniskillen town, 1993, I. McNeill; ingreat profusion in a garden at Magheranageeragh, 1 October 1994, RHN; beside Dunnes' store car park,Enniskillen town, 25 May 2001, RHN; in quantity at Brockagh Sandpit, to the W of roadway, 6 October2001, RHN.
Although thankfully C. didymus is still only a very rare casual plant in Fermanagh, it seems verypossible that like Arabidopsis thaliana (Thale Cress) and Cardamine hirsuta (HairyBitter-cress), this crucifer has the potential to become a common weed in horticultural nurseries andgiven the popularity of local garden centres it could then spread throughout the VC with purchasedplants.
British and Irish occurrence
In some cases, as in Fermanagh, C. didymus appears to be spread by the horticultural trade but, ofcourse, this is only one example of human assisted dispersal. Since its first introduction to Britain(probably as ships' ballast), from the early 18th century onwards, C. didymus has continued tospread slowly. In B & I, Lesser Swine-cress does appear to be more actively spreading in recentyears. Salisbury noted this fact in 1964, stating then that, "at one time rather uncommon, it isnow widespread in the southern counties [of England] and would seem to have become appreciably moreabundant during the last thirty years".
Lesser Swine-cress has now become an established weed of disturbed situations, including in arablefields, open pastures, waysides, waste land and coastal shingle and the New Atlas map shows it tobe very common in southern parts of B & I. Although present as far north as Inverness, C.didymus becomes much more scarce and coastal in its occurrence northwards from the EnglishMidlands. Similarly, in Ireland, it declines above a line across the map between Dublin and Galway(New Atlas). The New Atlas editors reckon that the species is "now frequent in urbanand industrial areas, and is still spreading into rural areas where it is widespread but scattered"(D.A. Pearman, in: Preston et al. 2002).
European and world occurrence
Represented as an introduction "probably from S America", Jalas et al. (1996) inAtlas Florae Europaeae 11 indicate that C. didymus is widespread, but notcontinuously so, in W Europe from S Spain to N Germany, becoming much rarer and strictly coastal furthernorth into Norway and Sweden. The species also extends discontinuously eastwards along the Mediterraneanreaching the Greek Isles and Crete.
Toxicity and grazing
The unpleasant smelling constituent of the feathery leaves has been identified as benzyl isothiocyanate,which is capable of tainting the milk of grazing cattle (Shimoda et al. 2000). Fortunately, thespecies is avoided by stock animals given adequate pasture and while tainting has been noted in NewZealand and studied in Japan, to date it is rare for the plant to be sufficiently abundant in grasslandin B & I to create this problem (Cooper & Johnson 1998).
Names
The genus name 'Coronopus' is a name used by Theophrastus, possibly for Plantago coronopus(Buck's-horn Plantain), from two Greek words 'corone' meaning 'crow' and 'pous' or 'pus' meaning 'foot'(Gilbert-Carter 1964). One has to suppose that 'crowfoot' here refers to the divided leaves of theplant, as with the aquatic species of Ranunculus. The English common name 'Lesser Swinecress' isgiven in relation to C. squamatus, 'Swine's Cress', a 16th century derogatory name referring tothe taproot of the plant which may be grubbed up by pigs, the plant being a cress fit only for pigs(Grigson 1974). The New Zealand common name 'Twin Cress' referring to the characteristic shape of thetwo-notched fruit capsule is a much more useful name in the current author's view (RSF). An alternativeEnglish name for C. didymus is 'Lesser Wart Cress', the fruit of C. squamatus, 'WartCress' being here regarded as wart-shaped (Prior 1879).
Threats
None as yet, but is likely to spread, increase and become a persistent weed.
Introduction, archaeophyte, occasional. Native range obscured by long cultivation, but probably it isEurasian southern-temperate.
1900; Praeger, R.Ll.; Florencecourt.
April to December.
Fermanagh occurrence
Turnip has been recorded in a total of 78 tetrads, 14.8% of the total in Fermanagh. It is an occasionalspecies on riverbanks, waste ground, roadsides, quarries and sand pits. The records are rather widelyscattered across the VC but, as would be expected, this tall annual or biennial is most frequently foundin the more fertile, lowland, farming area lying to the east of Lough Erne.
RHN and the current author (R.S. Forbes) feel that B. rapa is probably under-recorded and thespecies more common than the tetrad map indicates. We make this suggestion knowing that thistaxonomically complicated group of plants naturally leads to identification confusion for all concerned.We also believe that, perhaps more so in the past than now, some field workers in Fermanagh choose toignore obvious escapes from cultivation.
At the same time, due to an easily understood confusion between several very similar yellow-flowered taxaof the genus Brassica (plus a plethora of names and complicated synonymy), it is quiteconceivable that some of the B. rapa records in the Fermanagh Flora Database might really referto one or other of two forms of B. napus L., ie subsp. oleifera (DC) Metzg. (Oil-seedRape) or subsp. rapifera Metzg (Swede) (Rich 1987c, 1991). The situation regarding names is nothelped by there also existing a variety of B. rapa subsp.oleifera called 'Turnip-rape'. This is another oil-seed and fodder crop plant, which occurs inBritain as a casual alien (Clement & Foster 1994; Stace 1997). B. napus subsp. oleifera has been more widely grown in Ireland in recentyears, although not to any great extent in Fermanagh in comparison, for instance, to the more arableareas of south-eastern NI and especially in Co Down (H38).
The Swedish turnip is an orange-yellow fleshed swollen Rape root, usually referred to simply as 'Swede',and botanically named B. napus subsp. rapifera. It is cultivatedfor animal and human consumption and is usually eaten in the wintertime. It may also occasionally escapeinto disturbed sites, especially where farm, garden or kitchen material is used or dumped. Swede isoften sold under the incorrect name of turnip in NI, adding to any existing confusion the reader mayalready be experiencing!
In Fermanagh, we consider both Irish forms of B. rapa to be present (ie Wild Turnip and thecultivated Turnip), either as occasional recent escapes from cultivation, or older, feral derivatives orreversions from such escapes. B. rapa subsp. campestris (L.) A.R.Clapham (Wild Turnip) is sometimes locally abundant as a casual ruderal weed on disturbed ground,roadsides and waste ground, but it also occurs as a larger, biennial, semi-persistent and apparentlynaturalised form on unstable river banks, winter-flooded lakeshores, quarry ponds, sand-pitsstream-sides and ditches. Clapham et al. (1962, 1987) considered it doubtful whether these twoforms of subsp. campestris are genetically distinct.
Wild Turnip appears in urban situations too, especially in neglected ground in and around countryvillages and also in church grounds in a few instances. Quite where the latter form of the plant springsfrom remains something of a mystery, but perhaps soil disturbance in the form of grave digging andchurch yard tending may be all that is required!
Irish occurrence
The New Atlas map shows that B. rapa s.l. is a very frequent and widespread archaeophytethroughout most of lowland Ireland, with the exception of the most acidic peaty and mountainous areas,eg in Co Donegal, the Mourne mountains in Co Down, and in Connemara. Reynolds (2002) also reckons thatR. rapa is common and locally abundant in Ireland both as a ruderal and in more natural habitats.She also considers B. rapa subsp. rapa with its swollen 'root' tobe less common than subsp. campestris with normal tap roots.
British occurrence
In Britain, likewise, B. rapa is widespread in most of England and Wales, but its distributionbecomes increasingly fragmented and more coastal northwards into Scotland (Preston et al. 2002).
Plant and seed ecology
The seed of B. rapa s.l. is long-persistent in the soil seed bank, surviving at least five yearsand possibly very much longer (Thompson et al. 1997). Some form of regular disturbance of theupper soil horizons is necessary to enable germination and the maintenance of a recurring population ina particular site or area and subsp. campestris, in particular, requires and must have sufficientcompetitive ability in order to maintain its local presence. Information on the ecological requirementsand tolerances of the subspecies does not appear to exist and further study is clearly required.
An excellent summary of the history of B. rapa cultivars can be found in Simmonds (1976) and inZohary & Hopf (2000).
Threats
None.
Introduction, archaeophyte, occasional and declining. Origin is probably Eurosiberian temperate, butwidely naturalised in both hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
May to November.
Growth form and preferred habitats
A coarsely hairy, summer or more rarely winter annual, previously a widespread and abundant weed of farmcultivation throughout B & I, but which nowadays has hugely declined thanks to its susceptibility toselective herbicides. Together with the almost total absence of arable agriculture in Fermanagh, thismeans Charlock is now only occasionally found in the area, usually as solitary plants, typically inregularly disturbed, open, fully illuminated, lowland sites. The result of such a solitary or minimalpresence is that the once ubiquitous weed is readily overlooked nowadays and is very probablyunder-recorded.
Having said that, Charlock, throughout B & I, appears to tolerate a wide range of soil types,preferring a medium texture clay of neutral or alkaline reaction, with a high to moderate lime content.At the same time, it can also be found on peaty agricultural soils of low lime content with a pH around5 or so. What it abhors, in this respect however, are waterlogged conditions and both very light, dryterrain subject to regular drought, or very acid, cold, heavy clay or bog soils (Fogg 1950). In any soilor site, the disturbance associated with the activities of man are of primary importance and Charlocktherefore has always been virtually confined to artificial habitats.
Well established S. arvenis individuals not subject to excessive competition will eventuallyproduce a deeply penetrating tap root up to 74 cm deep and the species has a very rapidly developing andextensive fibrous secondary root system. Vegetative growth of the seedling plant is very vigorous in theearly stages, so that it often outgrows the seedlings of other species, including crop plants (Mulligan& Bailey 1975).
Competitive ability
In general, S. arvensis is only weakly competitive and prefers open sites with bare, or disturbedsoil available for colonisation. The competitive success and size of Charlock plants depends verygreatly on the particular cohabiting species (singular or plural) involved, the density of the competingplants and the fertility of the soil, but in general Charlock has a pronounced depressing effect on thegrowth and yield of cereal crops (Mulligan & Bailey 1975, p. 179).
Identification
Identification features, variation and characters distinguishing the most similar yellow-floweredcrucifer species with which S. arvensis might be confused are clearly spelt out and keyed byMulligan & Bailey (1985), Rich (1991) and also by the latter author in the Plant Crib 1998(Rich & Jermy 1998).
Variation
The vegetative growth of S. arvensis plants, their reproductive performance and even characterssuch as the depth of flower colour, all display a very wide degree of plasticity with respect toenvironmental conditions (phenotypic plasticity). For example, flowering stems frequently range fromdwarfs only 8 cm tall, to comparative giants up to 90 cm in height (Fogg 1950).
Fermanagh occurrence
The Fermanagh Flora Database has records from a total of 42 tetrads, representing 8.0% of those in theVC. Thirty-seven tetrads have post-1975 records. The tetrad map shows that S. arvensis is widelybut very thinly scattered throughout lowland Fermanagh, being slightly more frequent around Enniskillen.Charlock appears in both urban and rural disturbed waste ground and also on regularly visited, disturbedlakeshores, on field margins, waysides and in quarries.
When considering a local map displaying occurrences of a species like this one, we must remember that dueto the scarce, occasional visits of recorders and the solitary nature of most of the plants discovered,the map may well underestimate the plant's true distribution. However, the map symbol size and thelength of the period it represents can equally create a greater mental impression of the speciespresence than the recorder experiences in the field. This is illustrated rather well with S.arvensis by comparing the standard tetrad map and symbol size (2-km square) used in thisFlora, with the equivalent 1-km square representation. The scarce and scattered nature of thespecies and its real decline in frequency is much more readily appreciated from the latter.
Many, or indeed most, of the recent occurrences of S. arvensis in the VC probably derive fromburied seed emerging from the species long-persisting seed bank after local soil disturbance, eg inroadworks, by lakeshore jetties, on building sites and on waysides, especially those near sand andgravel quarries. Since so much of Fermanagh's ground is regularly covered with flood water or consistsof damp to wet habitats such as acidic pastures and meadows, wet peat or scrub woodland, it is reallynot terribly surprising that suitable sites are scarce in the county for a ruderal, short-lived plantlike S. arvensis which prefers open, unshaded, fertile, regularly but not excessively disturbedground, with a high proportion of exposed bare soil (Grime et al. 1988).
Flowering reproduction
Individual Charlock plants normally require about two and a half to three months growth to order tomature and flower. The species is phenotypically extremely plastic, however, and almost irrespective ofindividual size plants flower very freely from March or April through to July. Occasional very latedeveloping individuals can be found flowering even into December, although as with other species whichbehave in this way, these individuals are not likely to set much seed, if any (Mulligan & Bailey1975). The yellow flowers form a tight corymb, which together with the sometimes dense clustering of theplants, makes them conspicuous to both man and insect. Additional insect attractants are a slight, sweetperfume, nectar and abundant pollen. The flower reflects not only yellow wavelengths but also UV, so theinsect eye sees more detail and pattern in the flower than humans do. Each flower lasts just two daysand Fogg (1950) reported numerous insect types visiting to collect nectar and/or pollen foods, includinglong-tongued bees, butterflies, flies and beetles.
Until recent years, it was generally but incorrectly assumed that S. arvensis was both cross- andself-pollinated and was self-compatible (Knuth 1906-9; Fogg 1950; Salisbury 1964, p. 144). During mostof the last half of the 20th century, however, it has been realized that the species is completelyself-incompatible (Bateman 1955; Mulligan & Bailey 1975). The tissue recognition mechanism centralto the incompatibility was shown by Ford & Kay (1985) to involve sporophyte rather than gametophytetissue − ie the proteins on the coat of the pollen grain are sampled by the female stigma (Proctor etal. 1996, pp. 324-5). In fact, S. arvensis has a single-locus, multi-allelic sporophyticincompatibility system similar to that found in almost all other self-incompatible members of theCabbage family. Ford & Kay (1985) reckoned that there were as many as 24 allele forms of the onegenetic locus controlling the incompatibility in Charlock. The efficiency and benefit of this type ofbreeding system is that it allows any individual plant to breed successfully with most other members ofthe population, but not with itself or with some of its nearest neighbours (Proctor et al. 1996;Richards 1997a, pp. 224-30). Since cross-pollinated plants rely on external agents for pollination andthus cannot actively choose their sexual partner, this form of last-minute mating selection carried outon the stigma gives the plants which possess it a considerable evolutionary advantage in the long term.
One important consequence of flower self-incompatibility in S. arvensis is that when the plantbecomes rare and appears either solitary or in very small numbers as it currently does in Fermanagh,then seed set likewise becomes impossible, or less frequent and of a lesser order. Inevitably, anysignificant decline in seed production will hasten local population extinction and this is particularlythe case with annual species which, by their very definition, are completely dependent on seed for theirsurvival.
Seed production and dispersal
After fertilization the elongated, beaked fruit pods develop between 1-24 seeds, the average plantproducing 1,000-4,000 seeds (Salisbury 1964, p. 186). Seed is released by the splitting of the pod and,in addition to the consequent adjacent scatter, numerous birds including Greenfinch and Bullfinch aswell as stock animals such as cows dine on the pods and subsequently disperse the still viable seed intheir dung (Ridley 1930, p. 440; Salisbury 1964, pp. 102, 104). In the past, seed was also accidentallylong-range dispersed by man, as a crop seed contaminant.
Seed survival and germination
It has been appreciated for many years that Charlock seed have a remarkable capacity to survive for longperiods in the soil seed bank (Roberts & Boddrell 1983), and viable seeds up to 60 years old havebeen recovered at depths down to 30 cm (Brenchley 1918). This pronounced seed longevity permits thespecies' survival in sites which are cultivated or otherwise disturbed, on both a regular and avery infrequent basis. It has been shown that some S. arvensis seed is ready to germinateimmediately upon ripening in the autumn after it is formed, but the remainder possess innate dormancy,which is triggered, enhanced or enforced by even very shallow burial. Continuous variation was found inthe germination progeny of individual plants, and segregation of single genotypes with reduced dormancyoccurs, indicating that dormancy in Charlock is very much under genetic control (Witcombe &Whittington 1972).
The majority of seeds remain dormant in the soil for about 2 years, but there are also seasonal changesin the dormancy status of buried seeds. The capacity for germination is lowest in summer and earlyautumn, and it reaches a peak in the spring. Young plantlets of S. arvensis are sensitive tofrost and except in areas of mild climate and in sheltered sites that are slightly disturbed, it appearsthat few seedlings of summer or autumn germinating Charlock survive overwinter in most parts of B &I (Fogg 1950; Edwards 1980). Their mortality is due either to frost, or to excessive disturbance,e.g., early winter or spring cultivation, or competition with more vigorous species. Thus itappears that the autumn generation of S. arvensis appears to have little influence on thecapacity for survival of the species (Edwards 1980).
Population dynamics
In a population study in cereal fields in Leicestershire, climate was the major factor influencingCharlock population dynamics and reproductive capacity. While the persistent soil seed bank permitsspecies survival in sites which are cultivated or otherwise disturbed only on an infrequent basis,Edwards (1980) estimated that at the particular sites she studied, local population maintenance requireda minimum of one successful reproductive cycle every eleven years in order to top up the seed bank.Edwards also analysed a survey of published results from cultivated ground elsewhere and comparing thisto her own findings, found the half-life of Charlock seed is usually around three years. However, thereis considerable variation within S. arvensis from year-to-year in seed germination, dormancy andlongevity.
Population outcome is often determined by factors such as competition from the crop, or from other weeds,in which case the frequency and degree of soil disturbance is important in favouring established, butnot very competitive, S. arvensis plants. The effect of competition on Charlock plant size andreproductive capacity was shown to be more important in controlling the population than individual plantsurvival. This is undoubtedly due to the exceptionally prolonged seed longevity of S. arvensisand the large scale of the buried seed population − numerically much larger than the actively growingplant population; in this case, species survival depends mainly on the seed population (Edwards 1980).
Even more frequently than the influence of competition and soil disturbance, however, Charlockpopulations in B & I are limited by climatic effects including winter frost, early- or mid-summerdrought, or unfavourable conditions for germination such as a late spring. Edwards found that theprimary effect of climate was on the timing of germination and seedling emergence and establishment inthe springtime. In her study, seedlings began to emerge when the mean weekly temperature at 10 cm soildepth was above 4.4°C and emergence always coincided with rainfall (Edwards 1980, p. 157; Roberts &Boddrell 1983, p. 306).
British and Irish occurrence
On the basis of the BSBI Atlas (Perring & Walters 1962), Rich (1991) suggested that Charlockis "virtually ubiquitous" in B & I. However, this overstates the real position. Somewhatsurprisingly, the same major underlying habitat limitations seen in Fermanagh are well displayed on theNew Atlas hectad map of these islands, which indicates (even at such a large scale) that S.arvensis is unrecorded in many western and upland areas of B & I where wet acidic soilssupporting pasture grasslands, bog, heath or woods are the predominant forms of natural or semi-naturalvegetation (Preston et al. 2002).
European and world occurrence
S. arvensis has been a common weed of cultivation in Europe including B & I since earliesttimes (Fogg 1950). Godwin (1975) lists three fossil sites in B & I from the Neolithic, Roman andMediaeval periods. As with many such cereal weeds, the species is so closely associated with man andwith disturbed ground, that it is not really possible to be certain about its geographical centre oforigin or its native habitats. It is considered 'probably native' throughout most of Europe, especiallyin the Mediterranean basin, but it becomes rarer and more coastal further north in Scandinavia (Jalaset al. 1996, Map 2874). It has a similar status in N Africa, Asia Minor, SW Asia to the Himalayaand eastwards to Siberia. It is also introduced and widely naturalised in N & S America, S Africa,Australia and New Zealand (Clapham et al. 1962; Tutin et al. 1993). S. arvensis isalso a casual in many of the Atlantic islands, including Iceland, the Faeröes, Madeira and the CanaryIsles (Mulligan & Bailey 1985).
Historic uses
Young Charlock plants are palatable and were boiled and eaten as a green vegetable by poor people in thepast from Classical times onwards. This practice continued in B & I until the early decades of the20th century (Fogg 1950). Vickery (1995) quotes several Irish sources which describe the use of this'famine food', one of which explicitly testifies just how nauseous the plant tasted. In Scotland,likewise, seeds were eaten in hungry times, being ground and used to make bread. The mustard oil wasalso sometimes used as a fuel or 'burning oil' (Grieve 1931, p. 570).
Toxicity
The plant is toxic, but only becomes so when it is mature and forms fruit pods. There are three poisonousprinciples, a volatile mustard oil, Allyl-iothiocyanate, which is released from Sinigrin, plus thealkaloid, Sinapine and an alkaloidal glycoside called Sinalbin (Fogg 1950; Cooper & Johnson 1998).Toxicity has been shown in a number of animals to which seed has in the past been fed, includingchickens and, in pastures where mature Charlock was present, sheep, cattle and horses all suffered. Whenlarge amounts of either the plant or the seed were eaten the animals died (Cooper & Johnson 1998).
Names
The genus name 'Sinapis' is an ancient classical appellation given by Theophrastus to the mustard plant.Collaterally, it is sometimes spelt 'sinapi' and 'sinape' (Gilbert-Carter 1964). There is no apparentderivation of the name. The Latin specific epithet 'arvensis' is derived from 'arvum solum' meaning'arable land' and thus conveys 'growing in or pertaining to cultivated fields' (Stearn 1992).
The plant has a plethora of English common names, of which Grigson (1987) lists 31 and Britten &Holland (1886) 56! The most widely known name 'Charlock' is also applied to Rhaphanusrhaphanistrum, although it is more often qualified as 'White Charlock', or 'Jointed Charlock'(Watts 2000). The name 'Charlock' comes from the Old English 'cerlic', or 'cearloc', which Grigson(1974) says is, "an old name, as befits a weed of farm land, but of unknown significance". Itmust be a name that has been around a long time since it has accumulated a very large number of dialectforms ranging from 'Carlock' and 'Chedlock', through 'Harlock' and 'Skedlock' and 'Skellocks' to'Warlock'.
In addition, there are a number of forms beginning with 'k', such as 'Kecklock', 'Keblock' and 'Kerleck'(Grigson 1987; Watts 2000). 'Keck' or its variants refers to the hollow dry stems of members of theUmbelliferae, so these names do not meet this requirement and therefore must simply be variations of'Charlock'. The names 'Runch', 'Runches', 'Runch-balls', 'Runchik', 'Runchie' and 'Rungy', from variousparts of the country north of Yorkshire, also appear to refer to dried flower stems (Britten &Holland 1886).
Other more local names include 'Bread-and-Marmalade' from Somerset, a double name which perhaps suggeststhe fact that Sinapis arvensis seed was used in some districts to make bread in times of famine,plus a reference to the yellow colour of the flower. Four other common names listed by Grigson (1987)include 'yellow' as a word element.
Threats
Once a serious noxious weed of both cereals and broadleaved crops in lowland arable farmland,Charlock is so sensitive to modern herbicides that their widespread use since the 1970s has eliminatedthe species from cultivated ground throughout B & I. Plants are also sensitive to mechanical damagesuch as provided by grazing and trampling and it persists on wayside and waste ground sites that aresubject to only moderate levels of disturbance. Otherwise plants appear, usually in small numbers, fromlong buried seed in sites undisturbed for many years, eg roadworks and building plots.
Introduction, archaeophyte, an extinct casual. Native distribution has been obscured by long cultivationand wide naturalisation in both hemispheres, but it is probably European southern-temperate.
1903; Praeger, R.Ll.; Belleek village.
Growth form and preferred habitats
A tall (20-100 cm), yellowish-green, annual with deeply lobed upper stem leaves, fibrous roots and paleyellow flowers in dense, crowded racemes. The fruit pod has a flattened terminal segment or beak, 10-30mm long. There are usually two to three seeds in each swollen loculus of the fruit (Rich 1991).Previously a common weed of arable cultivation, it has declined to become a rare casual of waysides,disturbed ground and gardens, where it is used in wild bird seed feeding mixtures.
Flowering reproduction and physiology
S. alba flowers from May to September and is pollinated by bees and flies. It has been used inphysiological experiments on flower induction and, as far as is known, it is unique in being a long dayspecies in which flowering can be induced by just a single exposure to a day-length greater than 10hours. Under the specified experimental conditions, flowering is maximised by an exposure to an 18 hourday. It was also found that the timing of exposure to light has a considerable influence on the extentof the flowering response and that a single short 8 hour day could also induce flowering provided itfell within the 20 hour circadian period when the species was receptive to a light stimulus promotinganthesis (Kinet 1972).
Fermanagh occurrence
There have been no further records of White Mustard in Fermanagh since Praeger's 1903 find on wasteground in the village of Belleek on the county boundary. While the seed can survive hidden in the soilseed bank for many years, after a century of no sightings we may safely conclude that what was always inearlier days, a casual plant, is now locally extinct in the VC.
British and Irish occurrence
Elsewhere in Ireland, where White Mustard still occurs, it is an even more infrequent or rare casual thanwas formerly the case (Cat Alien Pl Ir). In Britain, S. alba has also declined over fouror more decades (the New Atlas change index is -0.90). However, the species remains very widelyscattered as a casual ruderal in lowland areas of these isles. S. alba is more persistent incalcareous soils and in B & I it is only really frequent and abundant in SE England.
In the past, White Mustard was occasionally grown and used by farmers as a green manure, being ploughedin to improve soil fertility. Nowadays, many, and possibly the majority of records of this species, areassociated not with arable farming, but with its use in wild bird seed mixtures, used to attract andfeed wintertime avian garden visitors. Perhaps this is why it tends to be found mainly on disturbedwayside and waste ground, where it may have been discarded, or escaped from gardens by its own means ofdispersal – including by the assistance of feeding birds (Reynolds 2002). Seed of this long cultivatedannual is associated with Roman sites in Britain, making it of archaeophyte status (Salisbury 1964, p.29).
European and world occurrence
As with S. arvensis, the native distribution of White Mustard is entirely obscured by its longhistory of cultivation and association with man, to the extent that Jalas et al. (1996) do noteven attempt to map it in their Atlas Florae Europaeae, 11. In its world distribution, wemay say that S. alba is primarily a southern temperate zone European species, possibly native inthe Mediterranean basin (Rich 1991). However, S. alba is so very widely introduced andnaturalised it has become almost circumpolar in the N Hemisphere (ie it is missing only in E Asia).S. alba is also naturalised in New Zealand (Hultén 1971, p. 6) and in S America and Australia(Rich 1991).
Uses
Grown chiefly for use as a green salad, S. alba was once the mustard of 'mustard and cress' fondlyremembered by many people from primary school biology experiments or early 'Nature Table' experiences.Nowadays, however, even this decidedly minor vegetable role has been replaced by Brassica napus(Oil-seed Rape) (Rich 1988). However, commercial mustard preparations do still include the use of seedsof Sinapis alba (White Mustard), mixed along with those of Brassica juncea (BrownMustard), or rarely with B. nigra (Black Mustard) (Rich 1991; Vaughan & Geissler 1997).
Names
The English common name 'Mustard' is of 13th century origin and is derived from the Old French'moustarde' and 'mostarde', which referred to seeds of S. alba and Brassica nigra togetherground up and mixed with 'moust de vin', that is, with 'must', new wine, a term itself derived from theLatin 'mustum', making a condiment for eating with meat (Grigson 1974).
Threats
None.
Introduction, neophyte, a very rare casual. Native of the Pyrenees and Central Europe, but widelynaturalised including in N America.
15 October 1995; Northridge, R.H.; disturbed ground on the recently widened roadside, SE of Lisbellaw.
Growth form and preferred habitats
This casual winter or summer annua introduction can sometimes behave as a biennial. Hairy Rocket grows20-60 cm tall, flowers from May to November and can develop on a range of mineral soils from sand tochalk. E. gallicum often appears as solitary individuals, apparently remote from any obvioussource of introduction. Generally it does not persist long, although vegetative growth and seedproduction are very plastic with respect to local environmental conditions and thus populations can varygreatly from year-to-year (Rich 1991).
Fermanagh occurrence
There is a solitary record in the Fermanagh Flora Database. The details of the record are: site about 200m along the Maguiresbridge road from the Presbyterian Church, SE of Lisbellaw. RHN has a voucher and theidentification has been confirmed by T.C.G. Rich. Is this change correct?
Irish occurrence
At present, the New Atlas map shows that apart from the solitary Fermanagh record listed above, inIreland E. gallicum is entirely restricted to, and rather thinly scattered within, an area S of aline between Dublin and Limerick. Reynolds (2002) in A Catalogue of Alien Plants inIreland lists around 35 records, mostly dating from the late 1980s and the 1990s, in tensouthern Irish VCs in the area mentioned. Rich (1991) considered the species to be currently spreadingrapidly in Ireland but, despite this, Reynolds regarded E. gallicum as an infrequent casual, withno evidence of it spreading (Reynolds 2002).
British occurrence
In Britain, the species is a weed of disturbed lowland roadside, railway, dockland and waste groundhabitats, occasionally more established and persistent in quarries and along track-ways on chalky soilsin S England. The plant is sometimes sown to bind and stabilise steep roadside cuttings, for instancethrough chalk in parts of Hampshire (Brewis et al. 1996).
European and world occurrence
The plant is considered native to the Pyrenees and Central Europe, but is widely introduced in many otherparts of Europe, across N America and the Urals, as animal feed grain and as bird seed (Hultén &Fries 1986, Map 993; Rich 1991; Clement & Foster 1994).
Introduction, archaeophyte, a rare or very rare casual weed of cultivation.
1900; Praeger, R.Ll.; Florencecourt.
July to October.
Growth form and preferred habitats
This weedy annual form of the Wild Radish is not at all common in the north of Ireland and it never hasbeen. The cultivated Garden Radish is a completely different species (R. sativus L.), but anotherform of the current species, common in particular on Irish Sea coasts, is subsp. maritimus (Sm.)Thell. (Sea Radish). The latter is a robust biennial as opposed to this annual form. Fruit charactersclearly distinguish the subspecies (Stace 1997).
While the flower colour of Wild Radish can vary from white to pale yellow or pale lilac (rarer), thepetals often being conspicuously veined dark violet beneath (Hackney et al. 1992), the fruit ismuch more clearly distinctive. It has a long, tapering, persistent beak and, when mature, the poddevelops distinct but not very deep constrictions between the bead-like seeds (Clapham et al.1962; Webb et al. 1996). In April and May, when not yet in flower, the rough bristly hairiness ofthe plant makes it easily mistaken for the extremely variable Sinapis arvensis (Charlock). Whenflowering from June to September, the two can be distinguished by the deeper yellow petals of Charlockand by the orientation of the sepals: erect in Wild Radish, spreading widely in Charlock. Undoubtedly,however, the mature fruit of R. raphanistrum is its most distinctive feature: the brown podsdevelop bead-like swellings separated by internal cross-walls and they break between the 1- or 2-seededbeads, unlike most crucifer pods which split downwards along their length (Rich 1991). Subsequently, thesegments of the broken pods gradually decompose with age to release the seeds.
Wild Radish appears to prefer disturbed, moderately fertile, fairly dry, sunny soils on waysides, gardensand waste ground, including around docks (Sinker et al. 1985; Rich 1991). Previously, however,before the major switch in Irish land use towards pastures and meadows in the 1950s, Wild Radish wasquite a common noxious weed colonising arable fields and their margins, especially those with sandy orpeaty-loam, mildly acid soils.
Fermanagh occurrence
In Fermanagh, over the years, subsp. raphanistrum has only been recorded a total of nine times,six of the records belonging to the post-1975 period. Locally, therefore, it is a rare or very rarecasual annual weed and is very thinly and widely scattered.
Additional to the first record given above, the details of the other eight are: field at Cranbrooke,Colebrooke estate, near Fivemiletown, 1946, MCM & D; sandy fields below Gortaree, Slieve Rushen,1949, MCM & D; waste ground behind St Michael's Church, Enniskillen, September 1986, RHN;Derrychara, Enniskillen, 29 July 1987, RHN: four plants still there 16 July 2010, RHN; Tempo, 31 July1987, RHN; Belleek village, 1 August 1987, RHN & RSF; roadside at Mullaghmore Bridge, NW of Ederny,19 October 1996, RHN.
British and Irish occurrence
This noxious weed is still quite common, although often merely casual, throughout all latitudes inBritain and especially frequent south of a line between Hull and Liverpool (New Atlas). In theFlora of NE Ireland, 2nd edition, Praeger (ever the optimist) went so far as to describethe occurrence of subsp. raphanistrum in the three NE counties as being, "not rare". In1938, Praeger may have been recollecting times when the plant previously was a common weed of arablecultivation and indeed 'not rare' to Irish naturalists of his generation.
The plant appears to have spread as a seed contaminant of grain and in Ireland was most frequent in the Sand E of the island. The Cen Cat Fl Ir 2, for instance, lists Wild Radish from 37 of the 40 VCson the island, the exceptions being S Tipperary, SE Galway and Roscommon (H7, H15 & H25).
With the advent of much improved, scientific seed cleaning in the 1940s and especially following thedevelopment of selective herbicides during the last 50 years, Wild Radish is now easily controlled andsubsp. raphanistrum has significantly declined, both as an agricultural weed and in the widercountryside. The New Atlas map shows that although there has been a major decline in Wild Radishacross both B & I (a Change Index of -1.39), it is certainly not on the verge of extinction inIreland as was imagined or predicted in Flora of Connemara and the Burren. Indeed Reynolds (2002)in A Catalogue of Alien Plants in Ireland declared the plant, "fairly common".
European and world occurrence
While Clapham et al. (1962) considered R. raphanistrum as "doubtfully native" inthe flora of B & I, it is now recognised as an archaeophyte which almost certainly originated in theMediterranean basin. It spread widely from there among agricultural seed and possibly the young plantwas also used as an edible pot herb, as well as in herbal medicine. It has been spread throughout Europeand N Africa, although rarer towards the east and throughout Asia. It was also carried by man in thesame way almost world wide, to N & S America, the Cape of Africa, Australia and New Zealand. It haseven been transported to E Greenland and to numerous remote oceanic islands, eg Kerguelen in the SouthIndian Ocean (Hultén & Fries 1986, Map 996).
Threats
None.
Introduction, archaeophyte, a very rare casual.
Eurosiberian temperate, but widely naturalised in both hemispheres.
1884; Barrington, R.M.; E shore of Lower Lough Erne.
Growth form, origin, preferred habitats and uses
This tall (50-150 cm), robust, hairless biennial possesses a strong tap-root plus many side-roots and, asusual in plants of this biotype, produces a leaf rosette in its first growth season and a stiffly erect,ribbed, hollow flowering stem in the second. The tiny (4-5 mm), 4-petalled, yellowish-green flowers areproduced in long, slender racemes, and the basal rosette of linear leaves withers after the firstseason. R. luteola typically grows in lowland areas on disturbed, neutral or base-rich, oftencalcareous, stony or sandy, neglected waysides, waste ground and disused quarries.
A definite native of the Mediterranean region and W Asia, R. luteola, 'Weld', 'Yellow-weed'or 'Dyer's Rocket' as the latter names imply, provides a natural yellow dyestuff and from Neolithictimes it has been introduced and cultivated for its yellow flavone in many parts of the world, includingB & I. Every part of the plant except the roots can be used for dyeing.
British and Irish status
Webb (1985) included R. luteola in his list of 41 species previously considered native in B &I, which he believed were probably introduced by man. The plant is very robust and produces a large cropof seed which is extremely long-persistent in the soil seed bank. In 1931, an archaeological dig at afirst century ditch at Cirencester resulted in a crop of Weld appearing on three occasions. Thesuggestion that the seed remained viable for 1,800 years was described at the time as being, "notincredible" (Rees 1931, quoted by Salisbury 1964, p. 323). However, until the editors of the NewAtlas (2002) recognised the species as an archaeophyte, all B & I Floras and other botanicalworks regarded R. luteola as native to these islands (eg Cen Cat Fl Ir 2; Clement &Foster 1994; New Flora of the BI 1997; Cat Alien Pl Ir).
Irish occurrence
The New Atlas map clearly shows that R. luteola is relatively frequent, more widespread andestablished in S & E Ireland than is the case in Fermanagh and neighbouring VCs. The hectad mapshows that the species distribution lies chiefly in the Midlands and the SE of the island and the plantbecomes increasingly rare and more confined to the coast as one moves in a north-easterly direction, ieit is most frequent E of the River Shannon and NE of Lough Neagh (Preston et al. 2002).
Fermanagh occurrence
There are just four records of R. luteola in the Fermanagh Flora Database, from waste groundspread across separate lowland tetrads. They date from 1884 onwards and only one of them is post-1975,but the latter, too, is now 30 years old! Clearly this is a very rare, only casual species in Fermanagh.
The details of the other three Fermanagh records are: quarry near Donagh Crossroads, 1950, MCM & D;old Ulster Canal near Gortnacarrow Bridge, 1950, MCM & D; Muckross near Kesh, 1976, Miss N. Dawson.
British occurrence
R. luteola is widespread and common in lowland Britain as far N as the Scottish urban areasof Edinburgh and Glasgow. It also has outliers further N around the Inverness area (New Atlas).In the past, the principal growing areas for the Weld dye crop were in Yorkshire, Lincolnshire, Essexand Kent and relics of these plantings may still persist in these areas alongside 'wild' plants (Sell& Murrell 2014).
European and world occurrence
R. luteola is considered native to the Mediterranean region and W Asia and, after a long period ofwidespread introduction and cultivation, it is a widespread weed in most of Europe northwards to around66°N in Sweden and Finland. It extends eastwards to W & C Asia, south to N Africa and SW to theCanary Islands. It is introduced in N & S America, Australia and New Zealand (Hultén & Fries1986, Map 997).
Native, very local and uncommon. Circumpolar boreo-arctic montane.
1882; Stewart, S.A.; Legland Mountain, SW of Knockmore.
Throughout the year.
Growth form and preferred habitats
This distinctive evergreen, small-leaved, often almost prostrate, creeping, much-branched subshrub hassmall, inconspicuous flowers and black, drupe, berry-like, fleshy fruits. It is found on windy, peatymountain summits, in exposed places on cliffs and upland slopes, and very occasionally in drier spots onlowland bogs, or on shallow raw peat over limestone rock.
Fermanagh occurrences
While E. nigrum is definitely uncommon in Fermanagh, it is locally frequent and very rarelydominant on some of the more exposed peaty mountain summits, cliffs and upland slopes. It has beenfrequently recorded in 31 tetrads, 5.9% of those in the VC, but as the distribution map shows it is verylocal and apart from several isolated stations on Slieve Beagh mountain in the east is almostexclusively confined to high ground in western Fermanagh.
Although listed by Meikle and his co-workers in the card index that formed the basis for theRevised Typescript Flora for the lime-flushed blanket bogland site of Erica vagans(Cornish Heath) at Black River (ie at or near the Carrickbrawn ASSI) and elsewhere in 'District IV', itis occasionally found growing in thin, raw peat over hard, crystalline Carboniferous limestone in placeslike Trien Mountain above Florencecourt and around the Monastair Gorge. The vast majority of thepost-1975 records are from wind-exposed, damp, but always well-drained, acid, nutrient-poor peatysummits, cliffs and upland rocky blanket bog slopes. These conditions are most often met on N-facingslopes in the Lough Navar, Reyfad and Cuilcagh Plateau areas. Here, Crowberry is often associated withVaccinium vitis-idaea (Cowberry) and other more common heathers – V. myrtillus(Bilberry), Erica tetralix (Cross-leaved Heath) and Calluna vulgaris (Heather), plus theclubmosses Selaginella selaginoides (Lesser Clubmoss) and Huperzia selago (Fir Clubmoss).Occasionally, Crowberry fruits in profusion and it does so on the exposed summit of Cuilcagh, whereindeed it is the locally dominant species.
The lime tolerance of E. nigrum that RHN and the current author have noticed in Fermanagh is oftenrepeated in the famous Burren district of Co Clare (H9) (Flora of Connemara and the Burren) andthis ecological property is also remarked upon in the Biological Flora account of the species byBell & Tallis (1973, p. 298). E. nigrum tolerates a soil reaction ranging from pH 2.5 to atleast 7.7 and while it is generally considered a calcifuge species, clearly this is not always the case.
E. nigrum has been found only twice growing on somewhat drier, better drained spots on peatyhummocks on lowland raised bogs in Fermanagh. This habitat type has rapidly declined due to drainage andpeat cutting and indeed it has disappeared in many other areas of B & I. The 1952 record fromClontymullan bog (Revised Typescript Flora) has not been confirmed recently, while Moninea bog inSE Fermanagh, where the species was first recorded in 1986, is now a protected ASSI site. The latter boghas been described as one of the top ten examples of surviving lowland raised bogs in NI (Leach &Corbett 1987).
Fossil history
Fossil pollen and macrofossil remains, such as fruit stones and leaves, indicate that in Ireland towardsthe end of the last glacial period approximately 13,000-10,000 years BP, Empetrum nigrum was oneof the first species to colonise de-glaciated ground in the cold, wet, oceanic (but gradually warming)climate of the period. Along with dwarf creeping Salix herbacea (Least Willow) and Juniperuscommunis (Juniper), it appears to have dominated the tundra heath vegetation widespread overIreland at the time. This was especially so in the wetter west of the island, during much of the LateGlacial and Early Post-Glacial periods, Zones I to IV of the developing Woodgrange Interstadial and theLittletonian Interglacial, as these warm stages are referred to in an Irish context (Jessen 1949, p.222; Mitchell & Watts 1970, p. 19). The dominance of Empetrum nigrum is so pronouncedin the fossil record that Jessen (1949) initiated what soon became the common practise of using thelevel of its presence as an indicator of the degree of 'oceanicity' of past climate phases.
However, it is important to realise that between these two warm stages of different duration whichwere suitable for plant growth and vegetation development, there was a return to Polar climaticconditions during the Nahangan Stadial, which lasted for around a thousand years, from 11,000-10,000 BP.This relatively brief, but very significant, cold stage meant that newly arrived and establishing plantspecies would have again been destroyed, and vegetation forced to retreat to more suitable warmer,sheltered sites, which it is generally imagined probably lay to the south and west of Ireland (Mitchell1986). The representation of Empetrum nigrum fossil pollen dropped suddenly from abundance torarity and disappearance over much of Ireland in Zone III; however, as Watts (1963) has shown, Crowberrygave place to Calluna vulgaris and other heathers common nowadays, all of which appeared in theBetula peak zone accompanied by Cladium mariscus (Sword Sedge).
A very concise summary of the fossil history of Empetrum heaths in Britain is provided by Bell& Tallis (1973) and an expansive account is given by Godwin (1975, pp. 300-5). The basic message ofthese accounts, which is true for all of B & I, is that Calluna vulgaris finally oustedEmpetrum nigrum from most heathlands at the end of Pollen Zone IV. Fossil records of Empetrumnigrum in the later Post-glacial Period are sparse and it becomes progressively restricted tohigher ground in the N & W, very much the pattern that pertains to the present day (Godwin 1975).
Variation and two subspecies
Nowadays E. nigrum is recognised as a Circumpolar boreo arctic-montane species of cool, dampclimates of the N & W in terms of its distribution in B & I, and indeed beyond our shores,although the distribution is not restricted to areas with true oceanic climates, ie regions wherethe climate is governed by the proximity of a very large body of water. Circ*mstantial evidence suggestsinstead, that while the distribution of E. nigrum is without a pronounced continental or oceanicemphasis, in areas away from the oceans it is controlled by environmental features which are analogousin their effects to a true oceanic climate (Meusel 1943; Brown 1971). At the same time, climatic controlof the species distribution is more strict than the effect of soil type (ie edaphic control is weakerthan climate).
The overall picture of E. nigrum distribution is made more complicated by the existence of atetraploid form which has mainly (but not entirely) perfect (ie hermaphrodite), bisexual flowers andwhich is regarded as taxonomically distinct at either subspecies level, as subsp. hermaphroditum(Hagerup) Böcher (as Stace (1997) and Sell & Murrell (2014) prefer it), or at species level. Asindicated by the use above of the word 'mainly', the dioecious (two separate sexes) versus hermaphroditeflower character is not absolute; hermaphrodite forms of subsp. nigrum have been reported inBritain (Blackburn 1938) and monoecious forms (ie, having flowers of different sex on different branchesof the same plant) of subsp. hermaphroditum can also occur (Danielsson 1988).
Both these E. nigrum subspecies exist in Britain, the tetraploid being heavily concentrated inScottish mountain sites, but all Irish material belongs exclusively to the monoecious subsp.nigrum (Preston et al. 2002). Due to considerable variation and overlap, the twosubspecies are vegetatively indistinguishable until they flower and fruit (Bell & Tallis 1973). ThePlant Crib 1998 gives useful advice for spotting subsp. hermaphroditum (Rich & Jermy1998, pp. 131-2).
Both subspecies (or species) flourish only under the low light levels associated with predominantlycloudy skies, combined with constant high relative humidity in the atmosphere, low summer temperaturesand small annual air temperature fluctuations near ground level where the plant resides (Brown 1971).These features are very characteristic of the oceanic, cool temperate climate current today in NW Europe(Miller 1961) and thus, in the Fermanagh area at least, the presence of Empetrum nigrum plantsand their pollen in sediments records an oceanic climate. Elsewhere, the presence of the plant and itspollen records only a direct analogue or substitute for this type of climate, eg through shade andrelative humidity provided by snow cover or the canopy of ericaceous and other subshrubs.
British and Irish occurrence
The present-day Irish distribution of E. nigrum in the New Atlas shows it widelyscattered throughout the island of Ireland, but chiefly represented in the upland, acidic terrain of theN & W. In Britain, subsp. nigrum is almost entirely confined to oceanic moorland heaths andmountain slopes and cliffs, NW of a line drawn on the map between Whitby and Plymouth.
At the same time, the altitudinal range of this subspecies in B & I is considerable, it being foundfrom near sea level to over 1270 m in the Scottish Cairngorms and reaching a height just over 1000 m inKerry (G.T.D. Wilmore & D.A. Pearman, in: Preston et al. 2002).
The rarer subsp. hermaphroditum (Hagerup) Böcher has never been found in Ireland, but in Britainit can overlap with subsp. nigrum, although generally occurring at higher altitude (>650 m)and often in drier places than the latter.
Although it is not a rare, or even a scarce species in B & I at present, the advent of 'GlobalWarming', ie a rapid increase in overall mean annual temperature, represents a real threat to specieslike E. nigrum which compete much better in cooler conditions (Anon. 1991).
European and world occurrence
A number of other related taxa (either forms or varieties, depending upon which taxonomist is followed),possessing differing leaf characteristics or fruit colour, have been described from distant areas inJapan and eastern N America, but if we take the broad view of the species as E. nigrum s.l., thenit has a distribution which is circumpolar (Bell & Tallis 1973; Hultén & Fries 1986, Maps 1463,1464).
The diploid form, E. nigrum subsp. nigrum, is present today inboreal N, W & C Europe, from Iceland in the NW, stretching across Scandinavia and Russia and rangingsouthwards to middle European latitudes. At its southern margin the distribution becomes progressivelyconfined to higher altitudes in the French Massif Central, the Alps, the Italian Apennines and mountainsranging further east to reach the Caucasus. Subsp. nigrum is very probably close to its southerngeographic limit in the current climate of B & I and it may have retreated northwards and upmountains in historic times in these islands.
Subsp. nigrum also extends through boreal Asia and the coastal mountains of western N America,although the Japanese plant is distinguished as var. or subsp. japonicum (Hultén & Fries1986, Map 1463).
Competition between the subspecies
In all of the cooler areas of subsp. nigrum's distribution (including the more southerlyEuropean mountains), it is overlapped and accompanied by subsp. hermaphroditum and clearly thetwo subspecies must compete strongly for territory. Leaf longevity is one character in which it is knownthere is a slight, but perhaps significant difference between these two subspecies. The evergreen,peinomorphic, needle-like, reduced, ericoid leaves of subsp. nigrum persist on average for1.5-1.9 years, while those of subsp. hermaphroditum survive longer, averaging 1.9-2.1 years(Karlsson 1992).
Winter frost protection by snow
At both higher latitudes and altitudes, E. nigrum, like other dwarf woody shrubs includingRhododendron ferrugineum (Alpenrose), requires the winter frost protection of a moderately thickblanket of snow cover. The association of snow patches with rocks, hummocks, hollows or piles of stones,probably helps account for the small-scale pattern of Crowberry in the more extreme, wind-exposed sitesit typically occupies. In common with most arctic-alpine plants, E. nigrum is adapted to lowtemperature regimes and re-commencement of its growth begins in the early spring. E. nigrumnormally flowers in April and May, so the snow must disappear quickly in springtime, or else the alreadybrief growing and reproducing season in these regions and habitats would be significantly shortened,which could prevent fruiting (Bell & Tallis 1973). Snow is a rarity in Fermanagh, never lying longerthan a few days, but fortunately its frost-shielding and protective blanketing properties are scarcelyrequired either, since frosts are seldom hard in the hyper-oceanic climate of NW Ireland.
Reproduction
E. nigrum subsp. nigrum is dioecious, both male and female plantsflowering freely early in the growing season, with dates in Britain ranging with latitude and altitudefrom March to May. The pinkish-purple flowers are wind-pollinated and although it is often few-berriedor sterile, occasionally the glossy black berry-like fruits (drupes) are borne in profusion from earlyJuly onwards. In Fermanagh, they are especially abundant on the exposed, elongated summit of Cuilcaghmountain, where very locally Crowberry is the dominant heath species. The berries are described byGrigson (1987) as "eatable, but poor eating, suitable for crows or crakes", and hence theEnglish common name 'Crowberry' and the alternatives 'Crawberry', 'Crawcrooks' and so on. Grigson listsa total of 17 English name variants, some of which are also applied to Calluna vulgaris and otherheather species (see also below under 'Names').
Fruit dispersal
Grouse and other moorland fruit-eating birds, including thrushes, fieldfares and crows are considered themain agents of dispersal, although foxes and deer are also said to eat them. On Danish moors, Hagerup(1946) describes commonly finding dark blue excrements derived from various animals, the colourgenerally indicating fruits of Vaccinium spp. or Empetrum nigrum. He says that thefruits are eaten in large amounts, not only by birds, but also by men, foxes and bears. Thus the'stones' and seeds of these genera are easily recovered on Danish moors, and the blue dung is socommonly observed that in many places that it has given rise to numerous vulgar names for E.nigrum!
Seed ecology
Each berry contains up to nine small seeds, which pass through the animal vectors undigested (Bell &Tallis 1973). However, the fruits often persist on the branches over winter, many of them failing todisperse and eventually being dropped around the parent plant (Lang 1987).
The fate of seeds is controversial; according to Hagerup (1946), under natural conditions only very lowrates of seed germination are reported in any one year. Passage through an animal's gut is notabsolutely necessary in order to break dormancy and seed may persist for four or more years beforegerminating. On the other hand, the survey of soil seed banks in NW Europe lists seven relevant studies,five of which regarded E. nigrum seed as transient, while the other two could not determine thelength of persistence (Thompson et al. 1997). Overwintering low temperature exposure is requiredto break seed dormancy and low numbers of seedlings, if any, emerge throughout the year (although theymainly appear in spring), over a period of several years.
Seedling development
Seedling development and plant establishment by E. nigrum plants is remarkably slow, anotherfeature shared with members of the genus Vaccinium (Flower-Ellis 1971). As one mightexpect, seedlings are generally observed in gaps in the vegetation, although in truth the species tendsto occupy habitats with little in the way of bare soil (Grime et al. 1988). Perhaps as a resultof this, in common with Vaccinium myrtillus (Bilberry), V. vitis-idaea (Cowberry) and mostother ericaceous dwarf shrubs, E. nigrum seedlings are rarely observed in the field anywhere in B& I.
Vegetative reproduction
Vegetative increase and dispersal by layering of procumbent or prostrate shoots is undoubtedly the mostsignificant current means of reproduction in these islands, although unlike most of the Ericaceae, E.nigrum clones are long-lived, some individuals surviving for up to 150 years (Bell & Tallis1973). The overall reproductive strategy of E. nigrum subsp.nigrum is thus parallel to that of other berry-producing, large-seeded, long-lived species, suchas Vaccinium vitis-idaea and V. myrtillus, and quite different from the rather more seedorientated reproduction and the shorter lifespans, observed in Calluna vulgaris and someErica species (Grime et al. 1988).
Grazing pressure
A study of Bilberry-dominated heathland in the Derbyshire Peak District found that E. nigrumincreased significantly when the mixed shrub heath was fenced and protected from sheep grazing pressure,yet while a regime of summer grazing allowed Crowberry cover to spread, there was a negligibletrend either way in winter-grazed plots. It was concluded that longer term studies would be required todiscover whether Empetrum nigrum would continue to replace Vaccinium myrtillus under thegiven conditions (Welch 1998).
Crowberry is very unpalatable to sheep and it is only ever grazed by them between January andApril when the animals are most likely to go hungry, or might otherwise starve (Welch 1984).
Fire and trampling
Crowberry can tolerate moderate, well-controlled burning of heaths, but it takes a long time to recoverfrom severe, hot, deeply penetrating fires. There is some suggestion that it can recover more quicklyfrom some Scottish muirburns than can Calluna vulgaris, Erica spp. and Vacciniumspp. However, this is probably seldom the case, or it represents only a temporary situation, since theseEricaceae are very much more often than not the dominant heath and bogland species that E. nigrumhas to compete with for space and survival (Gimmingham 1964; Bell & Tallis 1973).
Most dwarf shrubs suffer considerable damage if trampled and this is also true of E. nigrumalthough the only experimental evidence the current author (RSF) has come across is rather contradictoryof itself! A study was made in four sites in the Scottish Cairngorms of damage caused by simulated humantrampling around skiing areas (Bayfield 1979). This suggested firstly that the E. nigrum agg. (iethe two subspecies occur here together and were not differentiated) is among a group of three species,the cover of which recovered fairly rapidly after damage: the other two plants studied wereTrichophorum caespitosum (Deergrass) and Vaccinium spp. (Bilberry). Each of these replacedmore than half the loss of plant cover within two season's growth and, as the only evergreen among them,the rapid recovery of E. nigrum agg. was speculatively put down to stored food reserves in theextensive subsurface root system of the plant (Bayfield 1979).
However, the particular site where E. nigrum was studied possessed a thick carpet ofRachomitrium moss, the cushioning feature of which was possibly very significant in helpingCrowberry absorb the physical impact of the trampling. The situation is greatly complicated by the factthat when discussing the experimental treatments that meted out the most severe trampling and therecovery over an eight year period, Bayfield included E. nigrum agg. along with Callunavulgaris, Arctostaphylos uva-ursi (Bearberry) and Sphagnum rubellum in a list of speciesdescribed as "most susceptible − with high initial damage and poor recovery", while T.caespitosum and the Vaccinium spp. were classed as "moderately susceptible totrampling − moderate to high initial damage followed by fairly good recovery" (Bayfield 1979, p.175).
Names
The genus name 'Empetrum' is derived from two Greek words 'en' and 'petros' meaning 'on a rock', ie areference to the habitat of the plant. This name was first given by the Classical botanist Dioscoridesto a quite different, unrelated widespread Mediterranean species, possibly Frankeniapulverulenta, or another closely related evergreen prostrate species of that genus (Johnson& Smith 1946; Gilbert-Carter 1964). The Latin specific epithet 'nigrum', refers to thecharacteristic shiny black berry-like fruit.
Threats
The current steady increase in mean annual environmental temperatures we are experiencing in Britain andIreland in recent years may further reduce the competitive ability of this species and thus the area ofhabitat suitable for it and, indeed, this is also the case for other 'Arctic-alpine' species at theselatitudes.
Introduction, neophyte, uncommon, widely scattered, but locally dominant.
1946; MCM & D; near Colebrooke Park.
Throughout the year.
Growth form and preferred habitats
Originally planted around B & I in the larger estates and gardens probably around the early to middleperiod of the 19th century as a decorative plant and for upland game cover tolerant of exposure andpeatland soils, R. ponticum soon became firmly established, locally dominant and then spread toadjacent areas of acid terrain. This large, vigorous evergreen shrub is especially frequent on peaty andsandy soils, but really in terms of soil conditions it avoids only limestone.
A study of invasive spread in N Wales by Thomson et al. (1993) found that whenever a seed sourceof R. ponticum existed, factors favouring colonisation were a northerly aspect, steep slopes anda habitat where soil is occasionally disturbed by some form of management, eg tree planting or thinning,scrub clearance, moor burning or grassland ploughing. Despite the toxins it contains, intensity of sheepgrazing appears to help inhibit Rhododendron establishment, possibly by removing or trampling smallseedlings. However, ungrazed vegetation may simply provide a more favourable microclimate forRhododendron seedlings. Spread of R. ponticum is further inhibited by excessive wind exposure onhigher ground and it avoids permanently boggy ground at all altitudes (Thomson et al. 1993).
Fossil history and native distribution
The fossil pollen record preserved in peat bogs shows that R. ponticum was widespread in southernC Europe and extended northwards into Ireland in the Gortian interglacial warm period, 250 to 200thousand years ago (equivalent to the Hoxnian Period in England) (Jessen et al. 1959; Watts 1959;Mitchell & Watts 1970). In the current interglacial the native distribution of the species is muchfurther south in the Mediterranean basin and it is very disjunct − the main region being the Pontic andAnatolian regions of Turkey, Bulgaria and the Russian Republics (W Caucasia) on the S & E shores ofthe Black Sea and the immediately adjacent areas, plus a small outlier in the Lebanon (Chamberlain &Cullen 1982, p. 315, Map 94). The secondary native area of the species is comprised of three smallTertiary relict populations, surviving, but now vulnerable, in C and S Portugal and the Gibraltar areaof S Spain (Cross 1975; Bean 1976; Mejias et al. 2002).
Introduction to B & I and origin
The original introduction of R. ponticum to B & I either took place around 1763 (the dateusually quoted eg by Elton (1958) and Harvey (1988)), or more probably (at least for the commercialintroduction) between 1775 and 1780 (see D.L. Clarke, in: Bean 1986, p. 742). The original material wasalmost certainly imported via Gibraltar − thanks to the Rock's strategic importance and its colonialstatus. Confirmation of the Iberian origin of most B & I R. ponticum, rather than from thePontic or other eastern region, has been obtained from a recent genetic study of chloroplast DNA andnuclear ribosomal DNA haplotypes (Milne & Abbott 2000). Examining a total of 260 samples from sitesaround B & I, these workers found that 10% were of a unique Portuguese haplotype and 89% were of atype almost exclusively from S Spain.
Interestingly, the remaining 1% of accessions examined possessed chloroplast and nuclear ribosomal DNA ofother introduced Rhododendron species that are cultivated in B & I, but which have never escapedinto the wild. Two American species are undoubtedly involved, R. maximum and R.catawbiense, the latter closely allied to R. ponticum despite it being a plant of EasternUSA origin. R. catawbiense was first implicated as being introgressive along "with otherinfluences here and there", by Cox & Hutchinson (1963) when they inspected the wide range ofR. ponticum variation in Turkey and compared it with naturalized material growing wild inBritain.
The DNA study showed that introgression involving R. catawbiense was significantly more abundantin E Scotland, Britain's coldest region. Milne & Abbott (2000) suggested that the B & Inaturalised R. ponticum may have increased cold tolerance conferred on it as a result of thisintrogression, a factor which may have allowed the plant to colonise colder parts of these islands towhich the Iberian material was not preadapted (Abbott et al. 2003). Further study is required toeliminate the possibility of this particular introgressed material having been introduced to E Scotlandby mere chance, rather than the distribution reflecting such an advantageous adaptive feature.
The introgressive genetic material that the evidence suggests must have originated from crosses R.ponticum made with other introduced Rhododendron species, or with hardy hybrids duringtheir joint garden cultivation and these genetic pairings probably date from the early years of the 19thcentury (D.L. Clarke, in: Bean 1986, p. 743). Nelson (1994) has reported that R. ponticum was incultivation in Ireland by at least as early as 1800.
Naturalization and spread
The species appears to have been first recorded in the wild in Britain by at least 1894 and it appears tohave spread particularly rapidly in the early years of the 20th century, both by seed and by layeringitself (Cross 1975). Little detail is known about the dates of naturalization and the rate of invasivespread into the wild in either Ireland or Britain, due to a general lack of documented evidence, orperhaps to its current obscurity. It is however known that the plant was recognised as having"strong regenerative ability" by the middle of the 19th century, for this was reported to J.D.Hooker by his correspondents in Hampshire and S Wales while he was still in India collectingrhododendrons and writing his major work on the genus in the Himalaya (Hooker 1849; Desmond 1999).
Flowering and conditions for seedling establishment
Unlike some Rhododentron shrubs, R. ponticum flowers when it is still very young. It isself-compatible and like other members of the heather family it produces massive quantities of tinyseed. Each Rhododendron inflorescence produces around 5,000 seeds and Cross (1981) estimated that inMuckross, Co Kerry, a small bush 11 m in circumference and just 2 m high in a semi-shaded position couldproduce over one million seeds.
In producing vast numbers of small, readily dispersible seed, the reproductive strategy of theRhododendron shrub resembles that of species in the Pyrolaceae and the Orchidaceae. This is very unlikeother B & I woodland shrubs, which produce seed on average 100 times heavier than those of R.ponticum (Salisbury 1942; Cross 1981). R. ponticum seeds are short-lived, surviving onlya few months. They require light to germinate and in favourable conditions do so about five or six daysafter release. Seedlings survive best on slopes where there is a thin carpet of moss orliverwort, since when small they are extremely susceptible to drought. Other factors limiting thespecies involve the facts that young seedlings are unable to survive burial by shifting soil litter andthey do not have much competitive ability against taller herbaceous vegetation (Cross 1981).
Once the shrub has become established, grown taller and branched to form a domed shape, the heavyevergreen shade that it casts and the toxins it contains allow the species to dominate the vegetation.Indeed, typically R. ponticum excludes all other plants from beneath it. In suitable growingconditions in woodland and with the physical support of trees, naturalized specimens of R.ponticum can reach a height of 7 to 8 m and form an impenetrable thicket of branches (Cross1975, 1982).
Fermanagh occurrence
In Fermanagh, R. ponticum has been recorded in 74 tetrads, 14% of those in the VC, chiefly aroundUpper and Lower Lough Erne. Here it often forms very dense evergreen thickets from 3 to 5 m tall.Although, as the distribution map indicates, it is widely scattered throughout the county but, in highdensity, it remains centred on the old estate woods where it was first planted and in a few other uplandpeaty areas, eg within the Lough Navar Forest Park.
While the species has reached significant weed status in some lowland woods in Fermanagh, it is stillcomparatively rare on both the lowland bogs and in most upland woods and peatland areas of the VC.
Toxic properties
The leaves, flowers, pollen and nectar all contain several very poisonous diterpenoids (or grayanotoxins)and the shrub has been recognised as poisonous since ancient times (Cooper & Johnson 1998). In 400BC, Xenophon reported poisoning of Greek soldiers from honey made by bees from wild rhododendrons − athing virtually unknown since then due to better apiary management (Allison & Day 1997).Fortunately, stock animals are only likely to graze evergreen shrubs, including R. ponticum,under very adverse conditions when other food is extremely scarce. Poisoning is most common in sheep forsome unknown reason (particularly in rams) and losses are annually reported in the N & W of Britain(Cooper & Johnson 1998). Roots of R. ponticum also poison the soil, releasing short-chainaliphatic acids which deter the roots of other plants − the phenomenon known as allelopathy. Itsown roots are mycorrhizal, having an association with soil fungi and in nutrient poor acidic soils theymay starve competing plants of vital nutrient requirements.
Invasive nature and control
In Fermanagh, while the species has reached significant weed status in some lowland woods, it is still acomparatively rare species on lowland bogs and in most upland woods and peatland areas of the VC. Inview of the invasive behaviour of the species elsewhere in these islands in wet, cool oceanic climaticconditions, eg in W Ireland, N Wales, the Lake District and the W Highlands of Scotland (Cross 1975,1982; Abbott et al. 2003), every effort should be made to keep these types of habitats inFermanagh and elsewhere completely free from R. ponticum, by the immediate uprooting and burningof any plants that are found spreading.
Numerous studies around the world have proven how difficult or impossible control is once Rhododendron(often R. ponticum) gains a real hold, particularly in a wood or forest situation where itspresence significantly decreases tree regeneration and growth (Esen & Zedaker 2003). A comparativefield study of various manual and herbicide 'control' measures in Turkey by the latter authors foundthat low rates of foliar-applied Arsenal SL (imazapyr) (ie 3.0 kg active ingredient/ha), sprayed whentranslocation is downward to the roots (ie between mid-July and mid-September), gave significantlygreater Rhododendron control than other herbicides including Garlon 4 (triclopyr ester). Physicalgrubbing (ie manual uprooting from the uphill direction) also gave a great potential for sprout crownreduction. From what has been said above, grubbing up bushes and burning them on site provides an idealseed bed if other Rhododendron bushes remain nearby to provide a source (Thomson et al. 1993).
Names
The genus name 'Rhododendron' comes from two Greek words meaning 'Rose tree' and is thought to haveoriginally been applied to the rose-flowered form of Oleander, Nerium oleander, now transferredto this genus (Gilbert-Carter 1964). The Latin specific epithet 'ponticum' is geographical, meaning'belonging to the south shore of the Black Sea' (Stearn 1992).
Threats
Although R. ponticum is a serious invasive weed of woodland and peatland, it has limited powers ofdispersal and thus its spread can be predicted. Once established, it is very persistent, steadilyincreasing in density. Control is best achieved by preventing spread, which requires constant vigilanceto restrict the shrub to existing sites by avoiding soil and vegetation disturbance near seed sources(Rotherham 2003). In 2003, the fungal pathogen Phytopthora ramosum responsible for a 'Sudden OakDeath' epidemic in N America was introduced to a garden plant nursery in Cornwall, very probably on arhododendron cultivar. The pathogen has quickly spread across most of southern Britain and all ofIreland, and is now capable of attacking a wide range of woody species including bilberry (Vacciniummyrtillus), larch (Larix spp.), ash (Fraxinus excelsior) and chestnut (Castaneasativa).
Native, extremely rare, but appears to be increasing. Circumpolar boreal-montane.
1995; Hamill, B. & Corbett, P.; bogs west of Clontymullan Td, near Arney.
Growth form and preferred habitats
This small, trailing, hairless evergreen ericaceous subshrub, 10-25 cm tall, with a creeping woodyrhizome and attractive pinkish urn-shaped flowers is a characteristic species of the lowland raised bogsof Central Ireland. A. polifolia is usually confined to relatively undisturbed bogs and hasalways been considered very rare in N Ireland (FNEI 3). Richards described A. polifolia asthe raised mire specialist par excellence, growing amongst living Sphagnum mosses in a substratelargely composed of sphagnum peat in the pH range 2.8-4.8 (Stewart et al. 1994). Undisturbedraised bogs, or indeed undisturbed bogs of any kind, are now extremely rare in B & I.
In Britain, favourable conditions for Bog-rosemary are found at low altitude, between 10 and 530 m, withbetween 800 and 1200 mm precipitation and around 140-180 wet days per year (Rodwell 1991(b)). Nutrientanalysis of A. polifolia tissues of plants growing in subarctic peat bogs showed that the speciessurvives there with extremely low values of inorganic nutrients (N, P and K) in the shoot parts, theseminerals being transported to the actively growing region of shoots with green leaves (Malmer &Nihlgard 1980). Flower buds are produced the previous season and open in April or early May.
Bogland ecology
In addition to the widespread acidic raised bogs of the Irish central plain, A. polifoliais locally frequent on raised bogs in other, more southern parts of Ireland, yet it also has arestricted distribution on Atlantic blanket bogs (Doyle & Moore 1980). The two father figures of theclassical school of European Vegetation Classification, Braun-Blanquet and Tüxen during their jointinvestigative tour of Irish vegetation in July 1949, reported Atlantic Blanket Bog only in theirsub-association with A. polifolia within the overall Vegetation Association Pleuroziapurpurea-Erica tetralix, where it was strongly dominated by E. tetralix (Cross-leavedHeath) and Narthecium ossifragum (Bog Asphodel) (see Braun-Blanquet & Tüxen 1952, Table 48).
In Finland, however, A. polifolia is rather common in poor fens and in transitional rich fens andit also grows less frequently in nutrient-rich, eutrophic Pine and Spruce swamps (Jacquemart 1998). Thisfundamentally different ecological behaviour by the species forms an interesting parallel with that ofSchoenus nigricans (Black Beak-sedge, or Black Bog-rush), a species of eutrophic fens in Englandwhose widespread occurrence on blanket bog in Ireland has puzzled British ecologists for generations(Sparling 1968).
Flowering and vegetative reproduction
Flower buds are produced the previous season and open in April or early May. Bog-rosemary flowers aresmall, more or less spherical, pendulous, pinkish in colour and borne in a very short inflorescence of2-7 flowers. It usually flowers from April to June. Nectar is secreted by swellings at the base of thesuperior ovary, but insect visitors are often too late, the stamens and stigma ripen simultaneously (iethe flower is hom*ogamous) and self-compatible, so that self-pollination and self-fertilisation regularlyoccurs, sometimes while still in bud, thus displaying cleistogamy (ie closed marriage) (Hagerup 1954).
Some open pollinated flowers are pollinated by bumblebees and butterflies and achievecross-fertilisation. In lowland sites in Europe, some plants may have a second flowering period fromSeptember to October. In C Europe, about 27% of ovules develop to maturity, representing about elevenseeds per capsule, but in B & I fruits are seldom observed (Jacquemart 1998; M.C.F. Proctor, in:Preston et al. 2002). Most reproduction in B & I is probably restricted to vegetative spreadthrough active growth and layering of horizontal stems to form clones, although seed productionobviously remains essential for any degree of dispersal between bogs.
Fermanagh occurrence
A. polifolia was completely unknown in Fermanagh until the above listed solitary site wasdiscovered in 1995 by members of the EHS Habitat Survey Team, on the same bog near Clontymullan which isthe only known station for Rhynchospora fusca (Brown Beak-sedge) in the whole of NI. The moreaccurate name for this piece of lowland valley bog, lying just S of the Arney River where it entersUpper Lough Erne, is 'Drumanacabranagher Td', since this is the name on the current 1:50,000 map nearestthe grid reference given by the recorders. RHN who has visited the newly discovered A. polifoliasite on five occasions, described it in July 2002 as being, "scattered over 30 paces by 2 paces,parallel to a narrow drain".
A joint visit to the bog by RHN and the present author in 2009 showed A. polifolia had spread toform two largish clonal patches. Parts of the bog surface have been disturbed by drain cutting and peatremoval, but a relatively small area of intact surface remains, approximately 6 ha in size. Themicro-topography is subdued, and consists mainly of low peat hummocks and a few small pools, besidewhich the A. polifolia grows. Apart from the two rare vascular plants mentioned, this site alsocontains several rare bog mosses, including Sphagnum fuscum and S. imbricatum.
In B & I, A. polifolia is usually confined to relatively undisturbed bogs, a habitat typewhich is under extreme pressure from drainage, afforestation, heather burning and peat extraction.Several A. polifolia sites in NI are known to have been lost to these operations, andClontymullan (or Drumanacabranagher) bog is disturbed and continues to be exploited for peat,despite the presence of two exceptionally rare plants growing on its surface (ie Rhynchosporafusca and Andromeda polifolia).
British and Irish occurrence
In NI, Bog-rosemary is a rare and slowly declining species as its very specialised bogland habitatbecomes modified or destroyed. Apart from the one Fermanagh site described above, there are between oneand three recent records in each of Cos Tyrone, Armagh, Antrim and Londonderry (H36, H37, H39 &H40). Two previous sites in Co Down (H38) near Donaghadee were destroyed by development around 1927 andpre-1938 (FNEI 2, FNEI 3).
Although A. polifolia is very well represented on bogs in the more central counties of Ireland,and remains present to a very much lesser extent in NI, in hectad maps both in the first BSBIAtlas (Perring & Walters 1976) and in the map in the Biological Flora species account(Jacquemart 1998), the 1987-8 BSBI Monitoring Scheme survey showed (within its limitations) a9.0% decline in occurrence in the RoI, and a 18.0% reduction in sites overall in B & I (Rich &Woodruff 1990). In direct contrast with this, after a complete survey of B & I, the New Atlasmap indicates numerous fresh finds of Bog-rosemary during the last 60 years, especially in Wales and inthe Clyde Basin (M.C.F. Proctor, in: Preston et al. 2002).
The almost total absence of A. polifolia from the Scottish Highlands is quite the most strikingfeature of its B & I occurrence. While it may be possible to explain this absence by suggesting thatA. polifolia is purely a raised bog species, and that it cannot tolerate the differing, morevariable and more testing conditions of shallow blanket bog peat, its widespread and common occurrencein Scandinavia would immediately refute this contention. In her review of the species, Jacquemart (1998)indicates that while the chief habitat of the species in NW Europe is on the main dome of raised bogs,it does extend into wet hollows, wooded bogs and occasionally onto blanket bogs. The isolated station onMount Keen in NE Scotland, discovered in summer 1979, is on a high-level blanket bog sitting at 735 m(Birse 1980).
In view of its isolated and very rare occurrence in the N of Ireland, the abilities the species has toself-fertilize and tolerate prolonged inbreeding are clearly advantageous, allowing even solitary plantsto set seed and create populations. Dispersal of the small seed from the fruit capsule is presumably bywind and must be sufficiently effective for the plant to manage jump-dispersal to Mount Keen in Scotland(Birse 1980) and, of course, to the isolated station at Drumanacabranagher or Clontymullan Bog inFermanagh.
European and world occurrence
The distribution map for W Europe shows A. polifolia is most frequent in countries around theBaltic and less often found in S Sweden, Denmark, the Swiss-French Alps and the Pyrenees (Jacquemart1998). It is widespread in the boreal zone of Europe, Asia and N America making it circumpolar. Itoccurs throughout Scandinavia, making its near total absence in the mountains of Scotland verysurprising (Hultén & Fries 1986, Map 1456).
In view of its isolated, and very rare occurrence in the N of Ireland, the abilities to self-fertilizeand tolerate prolonged inbreeding are clearly advantageous, allowing even solitary plants to set seedand create populations. Dispersal of the small seed from the fruit capsule is presumably by wind andmust be sufficiently effective for the plant to manage jump-dispersal to Mount Keen in Scotland (Birse1980), and of course to the isolated station at Drumanacabranagher or Clontymullan Bog in Fermanagh.
Threats
Drainage and peat cutting.
Native, very common, frequently dominant. European boreo-temperate, but also widely naturalised beyondits native range.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This evergreen subshrub is the characteristic dominant species of the W European Lowland Heath Zonevegetation communities which were delimited by Gimingham (1972, p. 10, Fig. 2). The extent towhich Ling is often dominant is illustrated by Webb's (1986) statement that, "the ecology ofCalluna is, to a large extent, the ecology of the European heathlands".
Ling is generally associated with rather better drained areas of oligotrophic, strongly acidic,nutrient-poor bogland, heaths, upland woods and moorland. However, it also turns up in much drier, butstill acidic, nutrient-starved, canopy gap situations, including locally on some of the wooded islandsof Lough Erne and Lough Melvin. The shrub also colonises very shallow raw humus formed directly overhard, crystalline limestone in open situations, eg above Florencecourt and also in the Burren, Co Clare(H9). This latter situation allows C. vulgaris, normally considered a strongly calcifuge ('limehating') species, to grow intermingled with definite calcicole ('lime loving' or, rather better, 'limetolerant') species, in an exposed, low-growing community. This fact is indicative of the strong leachingof soils associated with very high rainfall levels in Western Ireland.
The characteristic high atmospheric humidity throughout the whole year associated with a cool, mild, dampOceanic or Atlantic type of climate is highly significant, permitting Ling to survive on very shallowpeaty soils. The one thing that all ericaceous species cannot tolerate, however briefly, is fortheir roots to completely dry out − and indeed this absolute drought restriction also applies to otherericoid shrub species, including those grown in gardens (Gimingham 1960, 1972, p. 11).
Variation
Considering the wide geographical and ecological range and the abundance of the species in many of itshabitats, apart from the normal expected plastic variation in size and performance associated withfavourable and unsuitable environments, Calluna vulgaris is incredibly constant in its growthform, to the extent that, apart from over 1,000 selected mutant cultivars of garden importancemaintained by the horticultural trade, taxonomists recognise only one named wild variety, a denselydowny grey form, well named 'var. hirsuta S.F. Gray' (Gimingham 1960; Griffiths 1994).
Growth and vegetative reproduction
Although C. vulgaris is a shallowly rooting shrub and does not possess a rhizome or stolons, theprocumbent lower branches produce adventitious roots and layer themselves on damp soil or peat, enablinga very limited amount of annual lateral spread by vegetative means. Aerial stems are sclerophyllous,possessing evergreen, overlapping scale leaves on long and short shoots, the former of which branch veryfrequently producing a dome-like growth form in vigorously growing 'building' plants. Leaves persist fora year on long shoots, but up to three years on the dwarf side shoots.
Individual unmodified C. vulgaris shrubs are surprisingly short-lived for an evergreen woodyspecies, normally reaching maturity between 11 and 15 years of age (Marrs 1986). However, shrubs oftenbegin to senesce and gradually lose physiological vigour when they are over 15 years of age, althoughthey can survive for 25, or perhaps very exceptionally for up to 30 years. As they age, plants becomeincreasingly woody and growth greatly slows, the dense crown of the young shrub gradually opening up asthe branches diverge and becoming more and more 'leggy', typically forming a central gap which exposesthe rather deep layer of litter accumulated beneath the shrub canopy. Watt's (1947) original descriptionof the growth cycle of the heather plant forming four characteristic stages, ie pioneer, building,mature and senescent, and their properties with respect to light, litter, soil moisture, pH, competitiverelations with neighbouring species, are summarised by Gimingham (1972) and more concisely by Webb(1986) and by Grime et al. (1988). These references should be consulted for more detail on themorphological and physiological properties of the species during its growth cycle. However, see belowfor limits on the application of the ecological concept of 'cyclical vegetation succession'.
Flowering reproduction
From July to September, C. vulgaris flowers freely and densely on the current year's long shoots,with zones of short or dwarf shoots produced both below and above them (Gimingham 1960, 1972; Mohamed& Gimingham 1970). Pollination can be achieved by wind, bees and other large insects, or by tiny (1mm long) slender insects called thrips that live their lives in and on the flowers. Depending oncirc*mstances, the flowers, which offer nectar and perfume to attract visitors, are pollinated by pollenfrom the same or other flowers, but direct contact between the anthers and stigma of an individualflower is impossible. From a genetic point of view, however, the pollination of C. vulgaris inmany cases must be regarded as autogamy, and the vast superabundance of pollen released is easilyappreciated by anyone who has walked through a stand of flowering heather and got it coating theirtrouser legs (Hagerup 1950a; Proctor et al. 1996).
Seed dispersal and behaviour
Huge quantities of very small, wind-dispersed seed are produced from September onwards, some of which,given damp, open soil, may germinate in the following two months, or else in the following spring. Thedeep litter layer beneath heather bushes is not conducive to germination and seedling establishment ofC. vulgaris, or indeed of any ericaceous species, but disturbance of the litter layer exposingbare soil or peat and, especially, the disturbance associated with fire which releases essential plantnutrients, greatly assists the process of Ling regeneration from seed. The remainder of the seedproduced becomes part of the vast soil seed bank of the species, in which individual seed may persistfor over 40 years (Gimingham 1972; Webb 1986, p. 110).
Fermanagh occurrence
Calluna vulgaris is very widespread in Fermanagh, occurring in 303 tetrads, 57.4% of those in theVC. This demonstrates just how much acid, peaty or podsolic ground of poor fertility there is across thecounty!
British and Irish occurrence
Calluna vulgaris is the most common and widespread member of the Ericaceae in both Irelandand Britain. The New Atlas species map indicates it being recorded in almost 86% of the hectadsin these islands and on this rather crude scale of presence measurement, it is well ahead of the secondranking ericoid species, Erica tetralix (Cross-leaved Heath) represented by map symbols in a mere70% of hectads (Preston et al. 2002).
Habitat loss and species decline
Having said this, there has been a decline in suitable heath, bog and moorland habitats throughout theseislands over the last 60 years affecting all the characteristic bogland species, so that the percentageof hectads in the New Atlas with the most recent date class (1987-99) has dropped substantiallyfor all ericoid species. The respective figures for C. vulgaris and E. tetralix are 79%and 62%.
The pressures on these infertile habitats have come from a very wide variety of changes in thecountryside and its management. The primary influences come from increasing forestry, intensification ofa*griculture − including movement between arable and pastoral that depend upon regional climate.
Additionally there has been over-grazing by sheep, abandonment of previously worked land and consequentrecolonisation by scrub and trees, and change of land use for building, or for mineral or peatextraction (Tudor & Makey 1995; M.C.F. Proctor, in: Preston et al. 2002). It is estimatedthat around 18% of heather dominated moorlands in Britain have been lost over the last 70 years, andconsiderably more than this since the late 18th century (Webb 1986, p. 183; Simmons 2001).
Since about the early 1980s there has been a growing awareness in B & I that C. vulgaris andother ericaceous subshrubs are not only losing ground through the landscape developments mentionedabove, but are also suffering reduction in their cover values and competitive dominance, even in areasof extensive heathlands that have been managed for long periods and remain unthreatened by changes indevelopment or land use. Indeed, it is becoming apparent that the contraction of heather species is nota new phenomenon, as the palaeoecological evidence from peat studied suggests that the species losseshave been operating over a prolonged period, one measured not in decades, but in centuries (Webb 1986).
In addition to the pressures already mentioned, there are fears that long-term global climatic warmingand increasing levels of nitrogen oxides pollution might already be adversely influencing the extent ofheather-dominated vegetation directly through changes in the productivity of moorland soils andvegetation, and indirectly through increased opportunities for cropping and the extension of the grazingseason. A survey in England and Wales found that there was approximately 44,000 km2 of landwith suppressed heather having less than 25% subshrub cover. Suppressed heather is defined as thatsuffering from, and showing growth forms affected by heavy grazing, neglect or inappropriate management(Bardgett et al. 1995). Although such heaths are damaged, they are potentially capable ofrecovery. In 1995, the Government introduced a subsidy Moorland Scheme, designed to regenerate moorswith more than 25% heather present, by restricting sheep stocking density − in effect an uplandequivalent of the Environmentally Sensitive Area scheme Fermanagh farmers enjoyed for some years(Simmons 2001, p. 280).
European and world distribution and recent changes
The range of C. vulgaris covers most of N, W & C Europe stretching from Iceland and NScandinavia southwards throughout the continent, but reaching only as far as Morocco and the Azores. Itis most abundant in the moist, mild, chiefly lowland, heathland region on the western seaboard of thecontinent, while in the much drier climate of the Mediterranean basin, the species becomes limited tomore humid parts of the coasts and mountains of NE Spain, S France, N Italy and W Dalmatia. Corsica isthe only Mediterranean island on which it is naturally found. Eastwards the species stretches in a verydisjunct manner into C Russia and China and it is also a scattered introduction in E & W NorthAmerica and in New Zealand (Hultén & Fries 1986, Map 1447).
In Europe, in common with B & I, the species has suffered a decline in habitat area and localabundance during the last five decades or more due to pollution and, particularly, the effect ofatmospheric nitrogen deposition, which reaches a critical point of heathland species tolerance at 15 kgnitrogen per hectare. Beyond this threshold, grasses such as Deschampsia flexuosa (WavyHair-grass) and Molinia caerulea (Purple Moor-grass) replace ericaceous subshrubs in dry and wetconditions respectively (de Smidt 1995).
Origin and history of Calluna vulgaris heaths and bogs: If we considerpresent day heathland and many areas of bogland in the longer term, measured in thousands of years, theyalmost certainly originated following disturbance of woodland by early farmers (Mitchell 1986; Simmons2001). While this is definitely the case for most lowland inland heaths, especially those in S England,a growing body of opinion believes that at least some ancient heaths and bogs may represent 'natural'climatically governed climax vegetation communities, rather than degraded woodland or nutrient-depletedprevious Neolithic farmland, an example of the latter being the Céide Fields of West Mayo (H27), anarchaeological site in coastal W Ireland.
Palaeoecological evidence in Britain points to the occurrence of Calluna and Empetrumspecies communities somewhat comparable to those of today (opinions differ as to the degree ofsimilarity), as long ago as 10,000 BP (Birks 1988), and maybe up to 1,000 years earlier than this in WIreland during the peak in Betula pollen at the end of the Early Post-glacial (Mitchell &Watts 1970; Pilcher & Hall 2001). Having said this, the period of extensive typical blanket bogdevelopment in Ireland did not occur until a great deal later: in the Killarney area, Co Kerry (H2),this occurred only around 3,000 years ago (Jessen 1949; Mitchell & Watts 1970).
Different patterns of geological prehistory, topography, changing land use (including grazing pressureand other factors exporting plant nutrients), plus soil and vegetation development process obviouslyexisted around these islands. The type and age of heath or bog vegetation communities that have formedare in response to such factors, as well as to overarching dynamic climatic conditions measured overmany centuries. Information on all of this must be collected, examined and integrated in orderfor us to interpret any particular local instance of current vegetation on the ground. It is thereforeunrealistic to generalise too widely on this matter, especially when we are dealing with a species likeC. vulgaris which possesses a broad ecological amplitude, and which occupy differing roles innumerous habitats and their plant communities.
This does not prevent us from drawing broad regional distinctions, as has been done for Britain betweenupland and lowland, and between wet northern and western situations, contrasted with drier eastern andmore southerly heather moorlands (Nolan & Robertson 1987; Thompson & Miles 1995). Many wetheaths possibly date back to the early post-glacial and some of them may have natural, rather thananthropogenic origins (Gimingham 1995). On the other hand, fire (naturally occurring or managed by man)and grazing might have tipped the balance from woodland trees towards dwarf shrubs, sedges, heaths andblanket bogs during climatic conditions differing from today.
If the change in vegetation type towards heathland was initiated by man, do we really want to preserveand maintain the status quo? Conservation is necessary for us to appreciate the complex processes ofvegetation change, since this should enable the development of practical guidance and interventionskills in order to better manage diversity in related vegetation types in future.
Natural change in heather-dominated vegetation
In terms of our understanding of dynamic vegetation processes, the cyclical succession of speciesdescribed by Watt (1947, 1955) for heather-dominated vegetation, involving the age of stands andeventual opening of the canopy to permit seedling establishment, probably applies better in wet,more species-rich conditions, compared with drier, more productive heathlands. This type of heatherregeneration cycle might not occur at all in some Scottish and Irish moorlands, simply due to a lack ofsufficient viable seed (Gimingham 1995; Thompson & Miles 1995; Welch et al. 2000).Heather-dominated ground below the tree-limit, (wherever that might lie in our much modified landscape),would naturally be colonised by sapling trees, especially birch and rowan, but possibly also by oak orby introduced species such as pines and larch (Khoon & Gimingham 1984).
The use of fire to prevent tree invasion and to regenerate heather and other dwarf shrubs and maintainthem in the building phase of their age structure has been widely practised as a management tool forheathlands over many generations. The frequency and severity of burns must be carefully related toprevailing grazing pressure and, if unskilfully executed, firing could have contributed to the gradualloss of heather cover across B & I throughout the last 200 to 400 years now observed from maps andfrom ground, aerial and satellite surveys. Khoon & Gimingham (1984) found birch was virtuallyexcluded by dense C. vulgaris stands in the building and mature phases of its growth cyclemaintained by carefully controlled regular burning of heath vegetation. However, occasional fires werefollowed by slow or patchy heather regeneration and this provided ideal conditions for birch invasionand establishment.
The need is obvious for more research on patterns of heath and moorland change and the relationshipsbetween vegetation types, including a re-examination of past results for possible causal factors whichcould then be field tested to discover which communities are the most stable when left alone, ie themost nearly 'natural' under the conditions of minimal or nil interference. When stable or almost stableunmanaged heathland stands are found, they tend to gradually develop into uneven-aged clones of C.vulgaris forming a mosaic pattern of the different growth stages described by Watt (1947, 1955).Even here, a minimal level of intervention will eventually be required, involving some use of grazing,burning, occasional herbicide treatment, or manual removal of taller woody invaders in order to maintainthe vigour of heather species and prevent succession to woodland (Marrs 1986).
Cyclical succession in heathland − does it happen?
Reappraisal of the model of cyclical succession involving Calluna vulgaris and other small-scaleheathland species proposed by Watt (1947, 1955) suggests it is an oversimplification. The basicassumption of long term heathland stability seems something of a myth. Yet the model has been useful inapproaching the interpretation of C. vulgaris dominated heaths with respect to a wide range ofenvironmental influences and pressures and it remains a helpful tool for this purpose, provided weaccept its limitations (Gimingham 1988). For example, if we consider grazing pressure, Ericacinerea (Bell Heather) is less productive and more unpalatable to browsing animals than C.vulgaris, but it is more palatable than E. tetralix, so in vegetation containing anymixture of these common heath subshrubs, a browser ranking order exists whenever the most favouredspecies becomes scarce. This needs to be appreciated and applied in the context of the dynamicinstability of the particular heath under review, its history of past management (essentially thepattern of grazing and burning) and the current spectrum of environmental pressures it exists under.
The growth phase patches of the ericaceous shrub species present often comprise a mosaic-like pattern, inwhich dominance may temporarily, and perhaps very locally, switch between one heather plant clone andanother. This will depend upon their relative age, individual history of shoot damage and the differentspecies inherent physiological vigour and morphological powers of recovery. However, other stands mayprove to be more even aged, and the older these are, the more vulnerable they become to catastrophicdeath events, since natural senescence affects the physiological ability of the subshrub to recoverafter any form of damage. Thus Watt's model of cyclical change in heather-dominated ground may or maynot operate, or may do so to an extent which perhaps is geographically (or climatically) determined.Operation of cyclical processes is especially doubtful where larger areas of ericaceous shrub vegetationsuffer damage at the same time, eg the catastrophic effects of a severe out-of-season frost, drought,insect attack, disease, major landslip, or an excessively hot fire event. Under this stochastic, lesspredictable or more randomly occurring event scenario, the more death suffered by C. vulgaris andits subshrub associates, interrupting or cutting short their endogenously governed life-cycles, thegreater the acceleration of directional succession leading to their replacement by either birch woodlandor acidic grassland (Marrs 1986).
Foliage damage
C. vulgaris growing in wet sites is commonly observed to suffer winter damage known as 'frosting',although in fact it is not directly caused by low temperatures, but rather it is drying damage, due tothe low levels of atmospheric humidity the plant encounters during cold windy spells of weather. This'frosting' effect undoubtedly reduces the productivity of mature evergreen plants, and it may kill offor adversely affect seedlings and young plants growing in bare, unshaded and, therefore, unshelteredpatches (Loach 1968; Webb 1986, p. 111). Similar foliage damage, reddening and sometimes killing theplant may occur during summer drought, although this does not happen very often in the mild, dampclimate of the west of Ireland.
Names and uses
The genus name 'Calluna' is from the Greek 'kalluno' meaning 'to cleanse', alluding to the factthat the branches of Ling and of numerous less common associated ericaceous subshrubs were often usedfor making rough sweeping brooms or 'besoms' (Johnson & Smith 1946). Apart from fodder for stockanimals, the plant has been used in rural areas until quite recent years to make a springy bed foreither man or stock, as roof thatch, besom brushes for the hearth, house and yard, for baskets, aspacking material, fuel (especially for kindling), strong rope (strong and suitable for mooring boats),as structural material for wattle-and-daub walls, as a dye plant and, in common with the otherericaceous subshrubs, for herbal medicine (for the latter see the current author's Erica cinereaspecies account).
Another possibly very important past use of C. vulgaris was as an ingredient of Heather Ale, amythical Scottish Highland brew, the recipe for which was lost, but which has recently (1990s) beenrecreated on the basis of supposed archaeological evidence. Basically it simply substitutes floweringheather shoot tips as flavouring instead of hops (Mabey 1996, pp. 158-61). Not least amongst the uses ofheather is the wearing of a sprig for luck, thankfully still a widespread practice at appropriatemoments in life!
In the light of such extensive use, it is not surprising that many local folk and English names haveaccumulated for the species, of which Grigson (1987) lists 16, and Britten & Holland (1886) as manyas 26! The name 'Heather', together with 'heath', derives from the Anglo-Saxon or Old English 'Hæddre'(Grigson 1974), which Prior (1879) says originates from a word that means the open, untilled andun-forested ground on which heath grows, and its even older root is shared with the word 'home', sinceopen ground provided sites for dwellings, even in very early times, ie the Neolithic. Thespelling changed from the 14th century 'hathir' to 'hedder' and, eventually, to 'heather' during the18th century. The other fairly widespread and well known English common name 'Ling' comes from Old Norse'lyng' and it is still most frequently found in areas of B & I where Vikings (Danes or Norwegians)settled (Grigson 1974).
Threats
As already mentioned, the species is affected by atmospheric and groundwater pollution which enrichesnitrogen levels in particular. It cannot effectively compete with grasses under these conditions, norwhen heavily grazed, or after severe or frequent burning or heavy trampling.
Native, common, but very probably declining in area. Suboceanic temperate.
1881; Stewart, S.A.; mountains west of Lough Erne.
Throughout the year.
Growth form and preferred habitats
E. tetralix is readily distinguished from E. cinerea (Bell Heather) by its greyer, hairierleaves, occurring in whorls of four, rather than in threes and its larger, more inflated, paler pinkflowers held in terminal umbel-like clusters of 4-12. E. tetralix occupies much wetter, moreoften waterlogged, poorly aerated, acid, peaty habitats than does either of the two other common heatherspecies, E. cinerea and the usually dominant Calluna vulgaris (Heather), or indeedany of the other ericaceous subshrubs found in the wild in these islands. While E.tetralix is most commonly found on, and is characteristic of, very wet or waterlogged,nutrient-starved, peaty, organic soils of acidity in the range pH 4.0-5.0 (Bannister 1966), the speciescan sometimes manage to compete and survive in more mesotrophic or even eutrophic habitats, alwaysprovided that some factor or group of factors other than nutrient level is constantly acting to limitthe competitive growth of any potentially dominant species present, thus preventing the species beingousted.
Fermanagh occurrence
In Fermanagh, E. tetralix is almost three times more frequently recorded than E. cinereaand it is also distributed across twice as many tetrad squares as the latter (ie there are E.tetralix records from 236 tetrads (44.7% of those in the VC), compared to just 98 (18.6%) forE. cinerea). Further numerical comparisons of these two species in Fermanagh and elsewhere in B& I are made in the current author's E. cinerea species account.
Apart from around wet hollows, in flushes and on the lower slopes of hummocks of deeper peat on bogs, onpasture moors, exposed, sunny mountain slopes and summits of lower hills, regular habitats of E.tetralix in Fermanagh also include roadsides, riverbanks, lakeshores and under canopy gaps inupland damp mixed deciduous woodland, including eg that in the Correl Glen NR.
In W Ireland, E. tetralix reaches an altitude of 550 m in Donegal and around 520 m in Fermanagh.This means it is absent from Cuilcagh ridge and its higher scarps and screes. In continental Europe,apart from the Pyrenees, C France and W Norway, where it can reach 2,200 m, E. tetralix isusually confined to low-lying areas (Bannister 1966).
Flowering reproduction
Robust plants produce flowers from the second or third year onwards and seed is set annually. Floweringbegins in June and reaches a peak in July and August, a little before E. cinerea and C.vulgaris. Both self- and cross-pollination probably occur, the flowers producing abundant nectarattracting a range of insects including bees, butterflies and flies as pollinators.
Another insect, the thrip, Frankliniella intonsa (Tryb.), is very intimately associated with E.tetralix as it lives and breeds inside the flowers. While thrip species inhabiting flowers tendto be specialised feeders on pollen, piercing the grains with their asymmetric mouth-parts and suckingout the contents, in a few cases and, especially in circ*mstances where other insect visitors are rare,they may also assist the host plant by being significant pollinators. For example, in the Faeroe islandswhere the climate is even cooler and wetter than ours, and where flying insects are naturally scarcebecause of this, flowers of Erica tetralix and Calluna vulgaris harboring thrips werefound to be both cross- and self-pollinated by the small insects living and sheltering inside them,pollen being unintentionally transported on the hairs on their bodies (Hagerup 1950; Hagerup &Hagerup 1953).
The normal pollinators of E. tetralix are, however, bees and bumblebees, some of which, inaddition to collecting nectar, may operate the technique of 'buzzing' the flowers to extract theirpollen (Haslerud 1974). To do this they hold onto and vibrate the flowers by rapid contractions of theirindirect flight muscles (Knudsen & Olesen 1993), so that the anthers resonate and almost explodepollen (Proctor et al. 1996, pp. 125, 179).
Nectar is often stolen from E. tetralix flowers by a variety of insects after bumblebees havevisited and drilled a convenient hole in the corolla tube near its base. Insects stealing food in thismanner avoid the legitimate entrance to the flower and thus they fail to service the evolved pollinationmechanism of the plant (Hagerup & Hagerup 1953). It remains to be seen how the plant species willevolve a mechanism to counter this thieving, evidence of which appears very frequent when Ericacorollas are examined.
Reproductive capacity
Numerous seeds, between 60-100 per capsule, are produced, although up to about 60% of ovules fail to set(Bannister 1966). The ability to colonise a much wider range of habitats than the physiologically severeenvironments for which it is particularly well adapted, helps E. tetralix to produce a largeannual seed crop. In moist conditions, seeds germinate readily with little need of pretreatment,although germination is much better in the light. Seeds can survive periods of at least three monthswaterlogging without loss of viability. Germination may be poor, however, on very acid peat and oncalcareous soils. Seed longevity can stretch to 33 years or more in the soil seed bank (Thompson &Band 1997).
The performance of Calluna vulgaris is significantly better with regard to the scale of seedproduction and its longevity in comparison with the two common Erica species, E. tetralixand E. cinerea (Barclay-Estrup & Gimingham 1994; Thompson & Band 1997). The persistentcorolla characteristic of the two Erica spp. undoubtedly to some extent hampers seed dispersalfrom their ripe capsule from October onwards, resulting in the observed rarity or absence ofErica seedlings in all except disturbed (usually burnt) ground. The seed dispersal limitation,together with the ease with which prostrate branches adventitiously root in wet habitats spreading andreproducing the species vegetatively, means that the latter is really the effective means ofreproduction of E. tetralix in most situations (Bannister 1966).
Response to herbivory
Sheep, cattle and grouse will all graze E. tetralix to a limited extent, but it is ratherunpalatable and the animals much prefer C. vulgaris if it is available. Like E. cinerea,Cross-leaved Heath is tolerant of this mild degree of browsing and it can recover quickly, pruned shootsresprouting from their base (Bannister 1966). Like all dwarf shrubs, however, including E.tetralix's most vigorous competitor C. vulgaris, heavy grazing such as has been thewidespread practice in recent years due to an ill-advised and short-sighted European Commission farmsubsidy involving animal headage payments that encouraged high levels of stocking, can exterminate thewoody heath and moorland vegetation and replace it with grassland. Whenever this occurs, continuedheavy, selective browsing of the pasture results in it very quickly becoming dominated by the mostunpalatable and aggressive grass species, Nardus stricta (Mat Grass).
Response to burning
As E. tetralix is mainly associated with wet ground and its crown and lower stems tend to beburied in wet moss or peat, whenever the heath is subjected to 'muirburn' management by planned,controlled firing, it experiences lower temperatures and/or shorter burns than other subshrubs that growon drier soils. As a result, E. tetralix is often less damaged and better able to survive thanother evergreen ericaceous species managed in this manner. E. tetralix also regenerates well fromits shoot base, so that it may become temporarily dominant after fire until C. vulgaris or othermore aggressive plants recover their normal vigour and size. Burnt ground appears to be the onesituation where seedling regeneration by E. tetralix may be reasonably frequent and obvious,although experimental results demonstrate that germination and seedling establishment is poor and ratherslow in wet peat conditions in comparison to that on well-drained mineral soils (Bannister 1964a).Excessive burning leads to the extermination of Cross-leaved Heath and other dwarf shrubs and theireventual replacement by grasses, sedges and rushes.
British and Irish occurrence
Both common Erica species are abundant and widely distributed throughout the N and W of both B& I, corresponding with where the wetter, often milder, climate conditions apply and the most acidicgeology and soils occur. However, while E. tetralix is still very well represented in the inlandcounties of C Ireland which was, until vast quantities of peat were extracted, the major area of deepand widespread raised bogs in the country, in this same region E. cinerea is scarce, orcompletely absent.
In S England both Erica species were previously associated with rather drier lowland heaths andbogs, habitats which have contracted sharply in the last 60 years due to pressures from farming andother land use developments. In the English Midlands, neither species has ever had much of a presenceand both are now increasingly rare, extinct or absent over a considerable area (Preston et al.2002).
European and world occurrence
The distribution of E. tetralix in B & I and in W Europe is very similar to that of E.cinerea, but rather surprisingly, despite its better competitive performance in wetter habitatsituations, E. tetralix is less rigidly oceanic and coastal, and more continental − or suboceanicin its occurrence than E. cinerea (Hultén & Fries 1986, Map 1445). It penetrates both furthernorth and east, eg to near Narvik, around 68°N on the coast of Norway, east to Latvia and C Finland andscattered further eastwards through south C Europe towards the Carpathian mountains (Hultén & Fries1986).
On the other hand, of the two species, E. cinerea is alone present in the Faeroes, Scillies and NAfrica (Hulten & Fries 1986, Map 1446). Although the map just mentioned also shows E. cinereapresent in Madeira, the form of the plant previously known as E. cinerea var. maderensis Bent.,is now regarded as an endemic species, E. maderensis (Benth.) Bornm, and indeed was firstdescribed as such as long ago as 1904 (Press et al. 1994). Bannister (1964 (b)) has suggestedthat E. cinerea has only a very limited tolerance of winter and spring reduction in the watercontent of its tissues, compared to E. tetralix, and to an even greater when compared toCalluna which can tolerate large reductions, and that this factor might well determine thecontinentality of the distributional range of these three heather species.
Established introductions
In some of its more continental sites in C Europe remote from the oceanic climate influence, E.tetralix is very probably a relatively recent introduction, eg in S Germany, where it isbelieved to have arrived along with conifer saplings imported from further north (Bannister 1966). E.tetralix is a rare introduction in eastern North America (Bannister 1965, 1966; Hultén &Fries 1986). E. tetralix has been cultivated in New Zealand gardens along with other introducedheathers since the early period of European settlement. While seedlings of most heathers are not oftenseen there, even in gardens, escapes from cultivation and deliberate attempts to naturalize C.vulgaris and E. cinerea have occurred there and these particular heathers are nownaturalised in a few stations on both NZ islands. On the other hand reports of E. tetralixoccurring with the other two species in the wild in NZ are discounted as being identification errors forE. cinerea (Webb et al. 1988).
Uses
Similar to Calluna vulgaris and Erica cinerea; see the species accounts of these.
Names
The Latin species epithet 'tetralix', is a name first used by Theophrastus for a thistle-like plant whichalso had leaves in whorls of four, forming a cross-like arrangement as in E. tetralix; over thecenturies the name became a technical term to describe this leaf state and as a name it was transferredto this heather by the Swedish taxonomist, Carl Linnaeus (Gilbert-Carter 1964; Gledhill 1985).
The English common name 'Cross-leaved Heath' is a simple translation of 'tetralix' and it is the onlyname given by Prior (1879) in his dictionary of popular names. In their similar but much larger work onthe subject, Britten & Holland (1886, Supplement 1896), list about 17 English common names fromaround B & I, many of them also regularly applied to C. vulgaris and to E. cinerea, eg'Bell-heath', 'Broom-heath', 'Besom-heath', 'Heather', 'Carlin Heather' and 'Ling'. One name fromSomerset and Hants specifically given to E. tetralix is 'Honey Bottle', but really this would bemuch more appropriate if applied to E. cinerea. Another N Yorkshire name is 'Father of Heath',while both this species and E. cinerea are often locally referred to by shepherds as'She-heather', by which they 'ungallantly' distinguish them as inferior grazing to 'He-heather', whichis Calluna vulgaris (Britten & Holland 1886).
Threats
Like all plants of heath and bogland habitats in B & I, there has been a decline in both area andabundance of E. tetralix during the last 60 years or so due to a combination of changing land useand pollution, both atmospheric and water-borne. The increased drainage involved in land improvement foragricultural production has probably more severely affected E. tetralix than the other ericaceousspecies, since they have a preference for drier soils than it does. Where ground that supported E.tetralix has not been reclaimed for agriculture, planted with conifers, built over or otherwisedeveloped, drainage and soil nutrient enrichment have together led to a one way shift in vegetation fromwoody dwarf shrubs towards pasture grasslands.
In common with other ericaceous subshrubs, E. tetralix cannot effectively compete with grasses andherbaceous plants under the developed/improved agricultural ground with its less nutrient-starvedconditions, nor can it survive heavy grazing, severe or frequent burning, or much human or heavy animaltrampling. E. tetralix seems set to decline further unless active conservation managementintervenes.
;
Native, frequent and quite widespread. Oceanic temperate.
1882; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
E. cinerea is a much more scarce plant in Fermanagh than E. tetralix (Cross-leavedHeather), usually occurring only in small patches, upright or sprawling in form, but almost alwaysstraggly in appearance, perhaps mainly due to the very slender, semi-prostrate woody branches, but alsoto the often rather sparse, dark, purple-tinged, needle-leaved evergreen foliage being more or lessrestricted to the ends of shoots.
On heaths, E. cinerea is associated most closely with Calluna vulgaris (Heather), oftenaccompanied by varying amounts of Erica tetralix (Cross-Leaved Heath), Vaccinium myrtillus(Bilberry), Ulex europaeus (Gorse) and other heath species, including occasionallyEmpetrum nigrum (Crowberry) and Juniperus communis (Juniper) on the most exposed heathsand cliffs. This subshrub is most frequently found on drier, well-drained, shallow, acid, peaty podsolpastures, including some formed locally over limestone. Bell Heather can also occur in openings inupland woodland and it is frequent on the drier tops of hummocks on otherwise very wet raised andblanket bog surfaces. It is especially frequent on cut-over turf banks, which by their nature areexposed, better drained and considerably drier than their boggy surroundings. A constant feature of allthe habitats E. cinerea occupies is their moderate to severe acidity (pH 3.6-6.6) and theconsequent nutrient-starved nature of their soils (Grime et al. 1988). This is reflected in theevergreen habit and reduced, peinomorphic or xeromorphic, sclerophyllous, needle-like leaves of thespecies. (For discussion of this important topic, see the Vaccinium oxycoccus (Cranberry) speciesaccount below.)
Fermanagh occurrence
E. cinerea usually forms only small patches of subshrub and it is a much more scarce plant inFermanagh than E. tetralix (Cross-leaved Heath). E. cinerea has just over one thirdthe number of E. tetralix records and it is represented in 98 tetrads, 18.6% of those in the VC.While it is quite widespread, in quantitative terms it remains concentrated in the peatlands of theWestern Plateau. E. tetralix is much more widely distributed across 228 Fermanagh tetrads.
Although E. cinerea is chiefly a plant of open habitats, growing in small quantity, Bell Heatheralso accompanies Calluna vulgaris (Heather) and Vaccinium myrtillus (Bilberry) in canopygap situations in upland, damp, mixed deciduous woodland, eg in the Correl Glen NR. Here, Bell Heatheris often forced to grow in the shade of other shrubs, often forming a semi-prostrate understorey to thedominant C. vulgaris. Shading produces the straggly, weak-stemmed, adventitiously rooting andhorizontally spreading, few-flowered or non-flowering, easily overlooked shoots of E. cinereathat we frequently meet in these situations. Bell Heather is much more tolerant of shade than C.vulgaris, so it penetrates deeper into coniferous or other evergreen or dense deciduous canopythan the latter, thus avoiding competition with this otherwise very dominant subshrub species.
E. cinerea also grows in Fermanagh in crevices on rock outcrops, including the damp sandstonescarps of the Western Plateau and in the more sheltered spots where winter snow might lie on otherwiseexposed cliffs and on mountain summits, including, eg on Cuilcagh the highest mountain in the VC.
Calluna vulgaris and E. cinerea also co-exist in Fermanagh on dry, grassy, low-growing,sheep-grazed heath, rooting in shallow, raw humic peat 'ranker soils' formed directly over hardcrystalline Carboniferous limestone, eg above Florencecourt and around the Knockmore area. Thisvegetation and habitat is the local equivalent of the acidified and leached lowland grassland limestoneheath or chalk heath of S England, where E. cinerea commonly occurred until the habitat losses ofthe last 50 years (M.C.F. Proctor, in: Preston et al. 2002).
Under these drier growing conditions, E. cinerea appears better able than other common ericaceoussubshrubs at maintaining a favourable water balance. It may even become the dominant species, orco-dominant along with Calluna vulgaris during some of the stages in heathland succession. Thisis especially so after any form of disturbance that reduces the vigour and dominance of C.vulgaris.
Finally, in addition to almost innumerable vegetation communities of heaths at all levels, boglands,woods and open montane communities, Rodwell et al. (1992) lists E. cinerea occurring insix NVC calcifuge, base-poor grasslands and non-heathy, grazing-sensitive, bracken-infested orrelatively unstable substrate montane communities in Britain. It is not a constant species in any ofthese situations, however, and tends to be of low frequency and cover/presence.
Flowering reproduction
Like Calluna vulgaris, Bell heather is a surprisingly short-lived subshrub. Individuals matureearly and flower in their second or third season. The majority of individuals appear capable ofsurviving for only around 20 summers at most (Bannister 1965).
Between July and September, the small urn-shaped, vivid-purple flowers, borne in elongate, terminalracemes or panicles are very conspicuous and are recognisable even from long distance. The plants flowerbefore C. vulgaris, so their presence amongst or partially beneath the canopy of the latter ismost noticeable in July. Plentiful nectar is produced at the base of the ovary in the pendulous,urn-shaped flowers which attracts various bees, butterflies, moths and flies as pollinators. It isprobable that unvisited flowers can self-pollinate, if necessary.
Some insect visitors, including some bumblebees, have a proboscis (or 'tongue') that is too short toreach the nectar through the narrow corolla tube. The thwarted insects often bore a small hole near thebase of the flower and extract the sugar without disturbing and operating the anthers and stigma at all,thus robbing the flower (Proctor & Yeo 1973, p. 143). Nevertheless, in addition to the nectar energysource, bees do require pollen for their protein and mineral nutrition, and therefore some of themmust operate the floral mechanism legitimately and carry out cross-pollination. In B & I,hives of honey bees are often transported in July to upland areas with plenty of C. vulgaris andE. cinerea, so that bee keepers can harvest 'Heather' honey (MacLennan 1995).
Seed production, dispersal and longevity
Fruit capsules examined in Britain by Bannister (1965) contained between 18-34 seeds, about one third ofwhich were abortive. A moderately sized 10 year old plant produced around 1,000 capsules; younger plantsproduced fewer capsules. The estimated annual seed production based on these figures was c 450,000potentially viable seed/m². Ripe capsules split open and vast quantities of seed are shed from Octoberonwards (Barclay-Estrup & Gimingham 1994). Dispersal, however, tends to be over very shortdistances, travel on the wind being greatly hampered by the fact that the withered, urn-shaped corollapersists around the capsule well into the winter months.
Seedlings are most commonly found in the immediate vicinity of the parent plant (Bannister 1965). Despitethe heavy seed rain and their known long survival in the soil seed bank (up to at least 33 years(Thompson & Band 1997)), generally seedlings are only observed colonising and establishing newplants in areas of relatively dry, bare mineral soil in vegetation gaps. Seedlings either do not occur,or do not survive for long, in wet, peaty, or litter- or moss-covered soil situations (Mallik etal. 1984).
Competitive ability and vegetation dominance
Temporary dominance in dry parts of heaths or moors may sometimes be achieved by E. cinerea, egafter ground is burnt, since its stumps recover more quickly from the effects of fire than those of thenormal dominant species, Calluna vulgaris. While both these heathers have large seed banks buriedin the soil and litter layers, that of E. cinerea is greatly stimulated by a brief heattreatment, and its germination is both more rapid and more uniform than that of Calluna, givingthe species at least a short-term competitive advantage (Bannister 1965; Mallik et al. 1984;Rodwell 1991(b), p. 383).
A study of threatened lowland heaths in England by Britton et al. (2000) found that whileCalluna is highly competitive over much of its life cycle, its competitive ability is reducedduring the regeneration phase that follows the death of stands, or which results in response to normallyapplied management activity including burning and grazing. These workers also discovered that sitegeographical location had the greatest effect on the pattern of regeneration of any of the four otherfactors they studied (ie dominant species, management techniques, gap size and seed sourceavailability). One thing becoming clear from this and other studies, is that heath regeneration dynamicsmay be extremely variable, and that conservation management must be tailored to the environmentalconditions found on each individual site (Britton et al. 2000).
Vegetative reproduction
Observed rapid seedling colonisation of burnt bare ground indicates that reproduction of E.cinerea from seed does occur and this is significant both in maintaining genetic evolutionarypotential and in transporting the species to fresh sites. However, vegetative spread by rooting ofprostrate branches is very common, and shoot recovery by resprouting of rootstocks and older branchesafter grazing, burning and trampling damage is also prevalent, so that vegetative reproduction isprobably very much more frequent than successful establishment from seed, and it is vitally important inmaintaining the species in its existing sites.
British and Irish occurrence
Erica cinerea is native and widespread throughout both B & I, but is much less common in CIreland and the English Midlands and S & E, especially in calcareous areas. Being characteristic ofoceanic heathland communities, it is most abundant in the N & W of both islands. E. cinereais closely associated with the occurrence of well-drained, acid, bare mineral soils of dry heaths andmoorland, but has declined in S England due to a general loss of lowland heathlands and changes in chalkheath vegetation associated with a decline of grazing pressure from sheep and rabbits (Bannister 1965;M.C.F. Proctor, in: Preston et al. 2002).
European and world occurrence
Unlike the other widespread B & I heathers Calluna vulgaris and E. tetralix, E.cinerea is very much more restricted to the moist, mild, chiefly lowland, oceanic and southwardsuboceanic heathland region of the western seaboard of Europe. It stretches from S Portugal to SW Norwayand is also present in the Faeroes, Scillies and N Africa. It has been introduced to at least onelocation in coastal eastern N America (Nantucket Island) (Hultén & Fries 1986, Map 1146). Bannister(1964) has suggested that E. cinerea has only a very limited tolerance of winter and springreduction in the water content of its tissues compared to E. tetralix. This is true to an evengreater extent when it is compared to C. vulgaris which can tolerate large moisture reductions.This property might well determine the very limited continentality shown in the distributional range ofE. cinerea.
Variation
There appears to be very little variation in E. cinerea apart from the existence of rare whiteflowers. A single endemic variety, var. maderensis D.C. from Madeira, is now recognised as aseparate species E. maderensis (Benth.) Bornm (Press & Short 1994). Having said this, thereis some possibility that a wet peat-tolerant ecotype of the species may exist in Ireland and in similarclimatically hyper-oceanic regions of NW Scotland. Plants of E. cinerea over much of the species'range cannot survive prolonged waterlogging and they certainly are much less tolerant of this thaneither E. tetralix or C. vulgaris. Nevertheless, Bannister (1965) reported E.cinerea growing on wet rankers in vegetation communities of northern affinities. Many of the wetheaths and Molinia-Calluna dominated vegetation types in N & W parts of B & I develop onthe slightly better-drained patches of peat banks and hummocks, on peaty podsols, or over shallow peatyrankers. This form of somewhat better-drained, damp to wet vegetation does have northerly, sub-alpine,or even possibly sub-arctic affinities, so that the local tolerance of E. cinerea to wet peatsoils may demonstrate wider variation than the species norm on the N & W margins of its overallgeographical range.
Limiting effects of temperature and drought
While there is some evidence from oceanic W Ireland and NW Scotland suggesting the existence of awet-soil ecotype within E. cinerea, or that the species demonstrates a wider ecological amplitudein the west of its range, Bell Heather definitely does not have an arctic tolerance of low temperature.The N and E of the species distribution conforms closely with the limit of the 2°C January isotherm.Also, since its main absorbing roots are shallow and normally form a mat at soil depths around 10 cm,Bell Heather cannot tolerate great summer heat or prolonged drought. Bøcher (1940) found that the S andE limit of its range correlated well with the 22°C June isotherm. Despite considerably higher figuresfor absolute altitudinal occurrence in various regions of B & I, the upper limit at which E.cinerea forms a regular component of the vegetation in these islands is around 620 m (2,000 ft)(McVean & Ratcliffe 1962), a fact which ties in with the 'temperate-thermic oceanic' classificationof the species given it by Bøcher (1940) on the basis of the limiting isotherms quoted above.
An additional limiting factor is the susceptibility of E. cinerea to late spring or early summerfrosts. Plants suffered very severe frost damage on exposure to a ground minimum temperature of -3.5°Cin late May and June in NE Scotland in 1964, an event which, in comparison, left C. vulgaris, themuch more continental species of the two in its distribution, totally unharmed (Bannister 1965). Interms of seasonality or phenology, Bell Heather, being evergreen, shows little winter growth, followedby a rapid burst of development in late spring and it flowers earlier than C. vulgaris,beginning in July and continuing through until September.
Dynamic succession processes
To a very great extent, the vegetation types in which E. cinerea grows are almost entirelysemi-natural and anthropogenic in nature, being produced by mans' destruction of pre-existing forest andsubsequent management for grazing or other uses of the heathers present (Bannister 1965; Gimingham1972). Except in otherwise severe environmental conditions, heathland below the tree limit dominated byCalluna vulgaris and other dwarf or semi-dwarf heathers and subshrubs is not a 'Climatic Climax',nor is it even a particularly stable form of secondary vegetation in these islands. There is always thelikelihood of invasion of such ericaceous heath and moorland by species typical of later successionalchanges, most notably by taller, longer-lived shrubs and trees, amongst which birch, having abundant,very light mobile seed, provides the most obvious threat in many instances (Khoon & Gimingham 1984;Gimingham 1995).
Response to grazing
Sheep are the principal browsers of heaths and moorland in B & I, followed by grouse and cattle,although numerous insects are also listed by Bannister (1965) in his Biological Flora account ofthe species. Larvae of the Heather beetle, Lochmaea suturalis (Thoms.), can cause localdefoliation of the shrub in some years. The foliage appears to be relatively unpalatable, since mostherbivores avoid it and show a preference for other available species, including Calluna vulgaris(Bannister 1965).
Experimental work on sheep grazing has shown that the season and period of browsing, as well as theweight of stocking density, are significant in terms of maintenance of heather species. In Europe, as inB & I, the extent of heathland has been very considerably reduced in the last 50 years andespecially so in the two most recent decades, due to cessation of previous management for agriculturaluse, ie a set regime of ± regular burning and grazing designed to maintain the vigour of the heaththrough keeping the subshrubs in the building phase of their growth cycle. Neglect of active managementhas led to encroachment by trees and shrubs on many heaths. Other heathlands have been destroyed byoverexploitation of the subshrub resource, giving rise instead to nutrient impoverished, species-poor,acid grasslands (Gallet & Roze 2001).
Sheep grazing at a high stocking rate (10-15 sheep/ha/year) on plots of dry and mesophilous heaths inBrittany where E. cinerea was an important component species found that, as expected, heavygrazing led to a decrease in vegetation cover, reduction in canopy height and also in the production offlowers (Gallet & Roze 2001). In this French study, E. cinerea appeared to suffer largedecreases in its frequency under several seasonal grazing regimes, but it always recovered to itsinitial level of cover after the livestock was removed. Following winter grazing, which tends to removeboth branch apices and flower buds, thus having a negative effect on flower production, subsequentdevelopment and full recovery of vegetative canopy was most probably due to the stimulation after theremoval of apical dominance, of the numerous lateral dwarf shoots which are such a characteristicfeature of E. cinerea shoots. Grazing in summer removes flowers, but in addition it also appearedto affect the plant physiologically, since the effect was still apparent in the following summer'sreduced flower production.
It has been suggested that heavy grazing towards the end of summer causes a reduction in carbohydratereserves, which could have detrimental effects on the important spring regrowth. It may also causewilting and death of branches during the winter and, in more general terms, grazing not only has animmediate effect, but it also seems to weaken heathers and other woody plants, decreasing their abilityto cope with winter cold (Grant et al. 1978, 1982; Bayfield 1979; Gallet & Roze 2001).
Response to trampling
Experimental human trampling of heathers has shown that even occasional trampling severely disrupts allthe plant communities studied and causes structural deviations in them. The presence of the ericaceousspecies in "humid heaths" over dry sandy soils nevertheless enhanced the vegetationresistance, at least to modest intensities of trampling in comparison for instance, to the damagesuffered by the more herbaceous floor layer of mesophilous woodland. The woody habit of heathers confersresistance to trampling, but also mitigates against the ability of the plants to recover from tramplingdamage. In respect to this, there are definite differences between species, eg V. myrtillus has amore limited wear and tear tolerance compared to the other species studied (Roovers et al.(2004).
Another much longer term experimental study on four sites at montane levels in the Scottish Cairngormmountains involved trampling treatments (ie two men wearing hiking boots!) applied over a short initialperiod followed by no further disturbance. It examined the responses of heathers and graminoid speciesin terms of initial damage and subsequent recovery over an eight year study. As expected, damageincreased with the level of trampling, but some woody species showed delayed damage; substantialdie-back occurred during the following winter, or even later. Again there were major differences betweenthe performances of ericoid species. Calluna vulgaris and Arctostaphylos uva-ursi(Bearberry) proved most susceptible − suffering high initial damage and poor, exceedingly slow recovery.Erica cinerea and E. tetralix were among the species of low susceptibility, showing low ormoderate levels of initial damage followed by an increase in relative cover compared to untrampledcontrol plots, although again their recovery process was slow, undoubtedly on account of the altitude.The deciduous species, Vaccinium myrtillus and V. uliginosum (Bog Bilberry), were placedin an intermediate grouping in terms of susceptibility − moderate to high initial damage, followed byfairly good recovery, most of which occurred in the first year − due perhaps to their being able toproduce a complete new set of leaves the following spring (Bayfield 1979).
Uses
Apart from the dark brown 'Heather Honey', Bell Heather flowers have been used in Scotland for ayellow-brown dye for wool. Many of the uses listed in the Calluna vulgaris species account applyalso to E. cinerea, and all three common and widespread heathers in B & I (ie C.vulgaris, Erica tetralix and E. cinerea) were used interchangeably in herbalmedicine for the treatment of a wide range of ailments. These heathers provided almost a panacea,through faith in a relaxing or mildly soporific effect the plants were believed to possess. Heather hasbeen used in the Scottish Highlands to sooth nerves and as a sedative to counter insomnia. Itsastringent properties have also been used for stomach upsets and for diarrhoea, and it has some diureticuses too. In Ireland uses of 'Heather tea' have varied from the treatment of coughs and asthma, and torelieve the pain of rheumatism (Allen & Hatfield 2004).
Names
The genus name 'Erica' is from the Greek 'ereike' meaning a 'heath' or 'heather', probablyoriginally a species of this genus (Johnson & Smith 1946; Gilbert-Carter 1964). The Latin specificepithet 'cinerea' means 'ash-coloured', which is not at all apt with respect to the whole plant,but might be a reference to the colour of the undersurface of the needle-like leaves. The currentwidespread English common name 'Bell Heather', appears to be a recent transfer from E. tetralix,since Britten & Holland (1886) list it among a total of 19 names for that species on theauthority of Jamieson's Dictionary of the Scottish Language (1867 edition). In their laterpublished Appendix (1896), however, Britten & Holland do list E. cinerea as 'Bell-ling', alocal Yorkshire name, along with a total of ten alternative names for the species. The name 'Blackheath' from Hants the current author feels is quite an appropriate one, in view of the very darkpurplish tinge of the typical leaf colour. According to Grigson (1987), the unusual Yorkshire name'Carlin Heather' is from the Old Norse 'kerling', meaning a witch or hag.
Threats
Like all plants of heath and bogland habitats, there has been a decline in both area and abundance ofE. cinerea during the last 60 years due to a combination of changing land use and increasingpollution, both atmospheric and water-borne. Where ground that supported the species has not beenreclaimed for agriculture, planted with conifers, built over or otherwise developed, drainage and soilnutrient enrichment has led to a shift in vegetation from woody dwarf shrubs towards grassland. As withCalluna vulgaris, Bell Heather cannot effectively compete with grasses under the changingconditions, nor when the subshrub is heavily grazed, severely or frequently burnt or heavy trampled.
Protected as native, very rare and vulnerable, but could possibly be an introduction. Oceanicsouthern-temperate.
1936; Dickie, Major; blanket bog near Black Bridge, above Belcoo.
June to December.
Growth form and preferred habitats
An evergreen, dwarf or tall shrub up to 80 cm tall with numerous ascending, erect branches without shortaxillary shoots. The glabrous leaves, 7-10 mm in length, are borne in whorls of four or five. Thesub-terminal inflorescence is a dense, cylindrical raceme often terminated by leaves. The four-lobedflower corolla is bell-shaped, either pink or white and the purple anthers are fully exerted beyond thepetals. E. vagans is a very rare, sometimes locally abundant and co-dominant shrub of heathlandover ultrabasic rocks (serpentine or gabbro), or moist gleyed peat flushed with alkaline, base-richgroundwater.
E. vagans is long regarded as native on the Lizard Peninsula in Cornwall where John Ray found it350 years ago, and it has also been recorded in most of the southern and midland counties of Britain,plus in a handful of widely scattered sites further NW into N Scotland and further east in Holland,generally appearing in these other areas as isolated bushes representing casual garden escapes(McClintock 1971; New Atlas).
Fermanagh occurrence
This really is a puzzling plant since it was first reported in Ireland in 1834, not in its extant remoteW Fermanagh station, but rather from a coastal site in Co Waterford (H6), about which there arose anerror in communication with J.D. Hooker, who had first published the find in his British Flora of1834. Eventually this station was given in More and Moore (1866) as, 'Cliffs in Islandicane townland,west of Tramore'. The station was never confirmed and the original voucher specimen apparentlydisappeared after it was sent to Dr Ball, the referee of the time. However the location in the far southof the island is precisely where the species might most feasibly appear, mirroring the native occurrenceof the plant on the Lizard peninsula in S Cornwall and minimising the disjunct distribution between thetwo stations. The Waterford population or individual plant was never refound so the report becameregarded with scepticism (eg in Colgan & Scully 1898). The assumption grew that the recordwas a garden escape, and gradually the record was discounted. At the turn of the century Praeger totallyignored the Waterford record in his influential and important contribution to distribution studies,Irish Topographical Botany (Praeger 1901).
Two other subsequent Irish appearances of E. vagans have occurred: one plant found in 1899 onsandhills at Dundrum, Co Down (H 38) (Praeger & Megaw 1938), which was still present in 1978 (Nelson1979); and a stony hillside west of Lough Swilly, Co Donegal (H35), where it was definitely planted andonly fleeting grew (Browning 1928; McClintock & Rose 1970).
Imagine the surprise when a substantial established population of entirely white-flowered plants wasdiscovered by an army Major from Enniskillen when out shooting! The shrub was found on a remote and atfirst glance apparently undistinguished moorland hollow and on a slope above a small nearby stream,referred to locally as 'the Black River', near Belcoo in W Fermanagh (Praeger 1938; Webb 1954b;McClintock & Rose 1970). It remains the solitary Irish site where this species appears wellestablished, and it might possibly be indigenous at this site (but see later argument below on thistopic).
Contrary to first impressions, the Fermanagh E. vagans site is ecologically quite distinctive.Investigations carried out between 1952 and 1970 by ecologists and taxonomists has shown that almost allthe ground now known as Carrickbrawn ASSI is flushed by a spring that seeps alkaline, lime-rich waterthrough the rather sandy mineral substrate underlying the shallow moorland peat. The soil here isderived from sandstone and shale, overlain with a thin layer of sedge and moss peat (Webb 1954b). Theflushed vegetation here is comparatively species-rich with 56 flowering plants species and 26 bryophytesrecorded by McClintock & Rose (1970).
Within its very restricted, constantly moist, but not waterlogged site, E. vagans waspreviously abundant and co-dominant with Molinia caerulea (Purple
Moor-grass), Carex spp. (Sedges) and Juncus acutiflorus (Sharp-flowered Rush). Beyond thespecies-rich, lime-flushed ground, apart from a few outlying clumps of the shrub, some of which are onthe stony 'cliff' above the stream within a short distance of the main E. vagans area, thisunusual heather species is surrounded by completely ordinary Calluna-dominated moorlandvegetation containing just 22 flowering plants species and 19 non-flowering species (McClintock &Rose 1970).
Growth in garden cultivation
The fact that Cornish Heath is such a widely appreciated garden subject, with 15 horticultural varieties,pink, white or cream flowered being listed in the RHS Index of Garden Plants (Griffiths 1994),highlights the reality that the species does not require or prefer base- or lime-rich flushedsoil conditions in order to grow, but simply that it tolerates such conditions better than otherheathers. In the wild, it is essentially this property which enables Cornish Heath to compete andpersist on shallow, relatively dry or well-drained, somewhat lime-flushed peatland surrounded byCalluna vulgaris and its associated species. Basically, E. vagans avoids competition withthese more aggressive species by tolerating conditions which they cannot. In garden cultivation, E.vagans actually prefers and grows best on a heavier, peaty soil and Flora Europaea alsodescribes it as a calcifuge (ie lime hating or lime avoiding) species (D.A. Webb & E.M. Rix, in:Tutin et al. (eds), 1972, p. 7)). E. vagans also tolerates shade better in the gardensetting than most other heathers, continuing to flower profusely, even under significantly darkenedconditions (Grey-Wilson 1989).
Growth and ecology at the Fermanagh site
The Carrickbrawn E. vagans colony has definitely contracted, perhaps by 75% during the last 60years, very possibly due to changes in the grazing regime. Sheep replaced cattle at the site 30 yearsago and the thus a modified trampling and grazing regime could well be responsible for the observedlosses. Early reports of the colony by Praeger and by Webb both refer to vigorous regenerationafter fire and grazing, involving both re-sprouting of stem bases and the appearance of seedlingsbetween the somewhat elevated clumps of mature Cornish Heath (Praeger 1938; Webb 1954b). Both thesevisitors recognised the limited population variation at Carrickbrawn and absence of pink flowers −unlike at the Lizard in Cornwall, where there is an almost equal mix of lilac to pink coloured flowersand white ones; the entire Fermanagh population is white flowered. The white flowers suggest that theIrish colony may be derived from one (or more) original hom*ozygous plants, which would mean that ifsexual reproduction has taken place (and elsewhere the species is considered self-fertile), the colonywould retain its uniformity (Nelson & co*ker 1974).
At present there is no evidence of recruitment from seedlings at the Fermanagh site. Prof Webb, inparticular, would have been very alert to the possibility of population sterility, since at the time ofhis visit he had just discovered this to be the case in Erica mackaiana (Mackay's Heath) in bothConnemara and Donegal (Webb 1954a). At Carrickbrawn, he described finding seedlings of Cornish Heath,"frequent on the intervening stretches of cattle-grazed turf", ie between the peaty hummockson which the larger bushes of E. vagans then grew (Webb 1954b, p. 216).
In terms of genetics, small isolated populations inevitably lose their vigour over extended periods oftime as they gradually lose genetically variability (a process sometimes referred to as 'geneticerosion'). This is a consequence of their restricted gene flow and 'genetic drift' (ie the tendency forgene allelles to fix within small populations at random, or even somewhat against selectiveforces)(Richards 1997, pp. 46-49). In small populations, these processes associated with inbreeding,lead to the accumulation of both hom*ozygous gene alleles and deleterious recessive mutants, but inpolyploid species, such as e.g., most Pteridophytes, these weakening processes limiting sexualreproductive success, may be very slow indeed to operate.
The lack of seed germination in the limited samples collected by Nelson and co*ker (1974) at Carrickbrawndoes not provide sufficient evidence to enable us to conclude that the population is totally sterile.These authors themselves remarked that small plants growing on the cliffs overhanging the stream couldhave been produced only from seed, although just to be safe they qualified this verdict by commenting,"unless very small fragments of parent plants had successfully produced adventitious roots andbecome established''. We believe the improbability of vegetative reproduction by fragmentsrather than seedling establishment in the river 'cliff' environment merits very little seriousconsideration, and in the light of Praeger's and Webb's observations, and with our own knowledge of theoccasional nature of heather seedlings - typically associated only with fire or with other major habitatdisturbance -, we believe that a low frequency of successful sexual reproduction probably stillis involved in the maintenance of this population.
This opinion is strengthened and coloured by the fact that dwarf heath plants of any ericaceous speciesare generally not long-lived, at least in comparison with other woody plants. Dwarf heather shrubsnormally survive only 15, 20 or 25 years, or very exceptionally up to 30, perhaps 50 years maximum.Regular grazing and occasional burning helps keep woody crowns and underground parts of ericaceoussubshrubs rejuvenated, but in any heath, older individuals will inevitably die from time to time, andthey must be replaced if the population is to maintain itself. The contraction of the overall colonysize might well be a reflection of such inevitable deaths, but the density of the remaining shrubsargues otherwise. Close study of our autumn 1978 photographs of E. vagans at Carrickbrawn showsthat there is some die-back of tall, leggy stems within otherwise vigorous, actively flowering clumps ofthe heather.
Another consideration is the possibility of a genetic reservoir of Cornish Heath seed persisting in thesoil. However, while seed of other heather species is known to remain viable for 30 years or more,unfortunately we have not been able to trace any mention of E. vagans seed longevity in theliterature (Thompson et al. 1997).
A detailed investigation is urgently required of the reproductive ability of E. vagans and itscurrent powers of regeneration at this site, perhaps including a controlled burn of a portion of thecolony.
Contraction in size of the Fermanagh colony
When examined by Praeger in August 1937, shortly after its discovery, the main area of the E.vagans colony in Fermanagh was closely grazed, apparently chiefly by cattle, which must alsohave damaged it to some extent through trampling, although sheep, goats and donkeys have also been seenat times in the vicinity (Nelson & co*ker 1974). The owner of the site has stated that cattle grazedthe area up until 35 years ago, when sheep were substituted and only they have browsed the ground since.
During his visit Praeger (1938) reckoned the patch of E. vagans covered an area of 75 × 50 yards[69 × 46 m] plus a few outliers. In the early 1950s, the size of the colony was roughly estimated byProf. Webb to cover a 50 m square (Webb 1954b), but by 1966 and 1970 it had contracted to occupy just 50× 30 m plus the small outliers (McClintock & Rose 1970; Nelson & co*ker 1974). Curtis &McGough (1988) in The Irish Red Data book of Vascular Plants estimated that there were thenaround 500 plants covering 45 × 30 m.
A visit by the current author and RHN in 2003 found that the plot where E. vagans was stilldensely present measured approximately 40 × 30 m, but the individual clumps of the plant were smaller insize than the two of us had previously noticed them, and Molinia caerulea (Purple Moor-grass) andMyrica gale (Bog-myrtle) appeared to be more obviously dominant and overgrowing them (Northridge& Northridge 2004). The fact that the colony appears to have shrunk by around half its reported sizeduring the 70 years it has been known is undoubtedly significant. The reduction is too large to beimaginary or the result of careless estimation and we can be certain that eminent naturalists likePraeger and Webb made no such mistakes. A survey conducted in September 2004 by conservation staff ofEHS, using satellite positioning (GPS), found the colony covered 1577 m2, slightly largerthan a previous estimate they had made and very similar to the autumn 1970 estimate made by Nelson &co*ker (Nelson & co*ker 1974), although the plants currently appear very overgrown and less vigorousand, indeed, had to be searched for (P. Corbett, pers. comm., December 2004).
The importance of herbivory and trampling
It appears to from the above observations that the role of herbivory in maintaining E. vaganspopulation competitiveness and co-dominance with Molinia caerulea, Juncus acutiflorus andMyrica gale on the Fermanagh site has probably been underestimated (Hulme 1996). The replacementof cattle by sheep could well be responsible for the observed decline of the E. vaganspopulation. Direct effects such as changes in selective browsing and in the level and timing of overallgrazing pressure, could lead to increased competition from more unpalatable shrubs and graminoidassociates. The heavyweight trampling disturbance which previously created vegetation gaps and thusassisted E. vagans seedling establishment has been greatly reduced by the change from cattle tosheep. The vegetation gap deficit would certainly limit the species' successful sexual reproduction. Inaddition, cattle are more likely to graze new spring shoots of the deciduous grass Moliniacaerulea, a serious competitor of Cornish Heath on this stretch of bogland. On the other hand,sheep are more likely to graze young heather seedlings than cattle, and it is at the seed and seedlingstages that herbivory principally influences plant mortality, both directly and indirectly (Harper 1977;Watt & Gibson 1988).
A variety of animals, including birds, insects, molluscs and mammals forage on seeds and seedlings in afrequency-dependent manner and, indirectly, this may affect interspecific competitiveness (Hulme 1996).Cattle were probably also responsible for more breakages of older woody tissues, which might bothstimulate and assist vegetative regeneration of Cornish Heath on the site, by fragmentation and bylayering of surviving trampled stems. Undoubtedly there are further indirect effects of a major changein herbivory involving differences in soil nutrient cycling, the significance of which would be moredifficult to quantify, but one only has to consider cow and sheep manure for a moment to realise thatsuch differences really can be significant (Crawley 1983; Hulme 1996).
Status of the Fermanagh colony
Nobody knows for certain how long the E. vagans colony has been at its Fermanagh site. It might beless than a century, although this would be greatly stretching the imagination in view of the observedlow competitive ability of the species, the narrow range of ecological circ*mstances where it achievesestablishment and persistence, and its probable, consequent slow growth rate. Alternatively, like otherericaceous subshrubs in Ireland, it might prove to be an ancient, genetically attenuated remnant of alarger prehistoric population of wider variability, ecological amplitude and geographical range,currently surviving in this one very isolated locality. If this is the case, it has become very disjunctfrom its Cornish nearest neighbours. Whether it arrived naturally or with the assistance of man, cannotyet be decided. What is certain is that Carrickbrawn is an extremely unlikely place foranyone to choose to deliberately plant any species.
British occurrence
Although E. vagans has been recorded at around 20 scattered sites in Britain in the wild, eitheras a garden escape or deliberately planted out, it occurs as an established species only on the Lizardpeninsula in S Cornwall. Due to widespread losses of lowland heaths in England, nowadays it is gone fromtwo of the Cornish hectads and has become restricted to just four of such squares and a total of lessthan ten sites (Perring & Farrell 1977; D.E. Coombe, in: Wiggington 1999; Preston et al.2002). Despite the reduced distribution, it is still co-dominant here with Molinia caerulea andSchoenus nigricans (Black Bog-rush) on silty clay and on neutral to slightly acid gley soilsderived from serpentine and gabbro rocks. The vegetation it occupies is described locally as 'TallHeath', since on the Lizard Peninsula shrubs of Cornish Heath often grow up to 80 cm high − at least 20cm more than is normally the case in Fermanagh.
European occurrence
E. vagans is a member of the small group of plants (15 at most), with disjunct geographicaldistributions strictly confined to mild oceanic climatic conditions spanning some or all of N Spain, W& NW France, SW Britain and S & W Ireland. The Latin specific epithet 'vagans' is the presentparticiple of 'vagor' and translates as 'wandering' or 'of wide distribution', and the former seems alot more appropriate in this case (Gilbert-Carter 1964; Stearn 1992). From an Irish perspective, thisgroup is often referred to as either the 'Hiberno-Lusitanian', the 'Hiberno-Cantabrian' (Praeger 1934,paragraphs 35, 37), or sometimes simply as the 'Atlantic' element in the Irish flora, although thelatter encompasses a more northerly extension than is appropriate to this discussion and is thereforebest avoided (Perring 1962; Webb 1983).
Alternatively, like other disjunct ericaceous species such as Arbutus unedo (Strawberry Tree) inKerry and Sligo (Mitchell 1993), Daboecia cantabrica (St Dabeoc's Heath) and E. erigena(Mediterranean Heath) in Connemara, E. mackaiana (Mackay's Heath) in Connemara and Donegal,Erica ciliaris (Dorset Heath) in Connemara and Cornwall (Webb 1966; Webb & Scannell 1983;Rose et al. 1996), E. vagans might prove to be a fairly ancient, genetically attenuatedremnant of a larger population of wider variability, ecological amplitude and geographical range,currently surviving in Ireland in just this one highly disjunct Fermanagh locality.
The extremely restricted NW Irish distribution of E. vagans is mirrored to various extents by allthe other ericaceous species in the W & N of Ireland listed above, and indeed the population ofE. ciliaris near Clifden in Connemara is very much smaller than that of E. vagans atCarrickbrawn. It consists of just five entirely vegetatively reproducing plants, which did, however,recover very well after a severe bog-fire in May 1966 (Webb 1966). This tiny Irish population of E.ciliaris, so small that it was entirely lost for 119 years, is distinct from native Englishpopulations of E. ciliaris in Dorset and Cornwall (vcs 1, 2 & 9), in that it never hasglandular tips on the stout marginal hairs (the cilia) of its leaves, a feature which only about 10 % ofEnglish plants display (Webb 1966; McClintock 1968; Nelson 1989). However the fact that this tiny bogroadside population of E. ciliaris has neither increased nor decreased over the 160 years sinceits first discovery, does suggest to some botanists at least, that it probably is a human introduction,transported by unknown means (Matthew Jebb, pers. comm., Dec. 2004).
In common with all these other 'Lusitanian' plant species except Arbutus, there is no fossilevidence to prove, indicate, or even as much as suggest the presence of E. vagans anywhere inIreland in the Late-glacial or the early Post-glacial periods (Mitchell & Watts 1970). This is notterribly surprising since the flowers are chiefly - but probably not exclusively - insect-pollinated,and although the anthers are exerted from the corolla tube in E. vagans, and the plantflowers profusely, the chances of pollen or fruit and seed preservation and recognition of them atspecies level, are not very great, even if it were once a much more common species than it is atpresent. While not of course conclusive in itself, the lack of fossil evidence tends to suggest that atleast the more frost-sensitive members of the Lusitanian plant group such as Arbutus andDaboecia could not have survived peri-glacial conditions in the vicinity of Ireland during thelast cold spell, and that species perhaps including E. vagans must have survived further southand arrived here more recently during the current warm interglacial period, called the 'Littletonian' inIreland (Mitchell 1965).
In addition to E. vagans, five other members of the Heather Family belong to the'Lusitanian' group, Erica ciliaris (Dorset Heath), E. erigena (Irish Heath), E.mackaiana (Mackay’s Heath), Daboecia cantabrica (St Dabeoc’s Heath) and Arbutusunedo (Strawberry-tree), the last four of which are oddly absent from Britain. The'Lusitanian' distribution pattern is followed by E. vagans in all the regions listed above,except that in Ireland it occurs in Fermanagh, unlike the other Ericaceous members that are mainly orentirely found in Connemara or Kerry. It is also the case that in N Spain and in France, E.vagans extends further inland than do most other species of the Lusitanian group (Praeger 1938;Webb 1983; for a map see Nelson & co*ker 1974, Fig. 1).
Threats
The population is vulnerable, especially to fire, but also to overgrazing by sheep and to any changesreducing an already shrinking population with only limited genetic variation. Myrica gale,Molinia caerulea and other common graminoid plants appear to be exerting considerable competitivepressure upon E. vagans, severely limiting its reproduction, or perhaps completely preventing it.Careful monitoring and active conservation management directed at encouraging the species survival,based on detailed scientific research of the local population, is urgently required.
Native, occasional to frequent, but rather local. Circumpolar boreal-montane.
1902; Abraham, J.T. & McCullagh, F.R.; Feddan Bog.
April to January.
Growth form and preferred habitats
A tiny loose, spreading subshrub whose wiry stems need to be actively searched for, often onSphagnum tussocks around the margins of pools in the wettest parts of raised bog surfaces. V.oxycoccos appears to need strongly acidic, wet growing conditions, but it cannot tolerateconstant immersion. Thus it colonises wet bog hollows, only after they have already begun to fill withSphagnum moss, principally the common yellow-green species, S. cuspidatum (Lusby etal. 1996).
Fermanagh occurrence
V. oxycoccos and Andromeda polifolia (Bog-rosemary) are the two dwarf ericaceous shrubswhich best manage to colonise the transition zone between hummock and hollow on acid peat bog surfaces.However, while Cranberry is quite frequent, though easily overlooked because of its small size andinconspicuous growth form, the more upright Bog-rosemary is a much rarer and declining species, at leastin NI (FNEI 3). In the whole of Fermanagh, there is just one relatively small, recentlydiscovered Andromeda polifolia population on a lowland Fermanagh bog, but V. oxycoccos hasbeen recorded over 130 times in the VC, in a total of 48 tetrads. This presence represents Cranberry in9.1% of Fermanagh tetrads and 40 of them contain post-1975 records. As the distribution map indicates,Cranberry is thinly and quite widely scattered in boglands across the county.
Bog plant leaf adaption to nutrient starvation
The relatively long, slender, wiry stems of Cranberry allow it to keep pace with the rapid growth of theSphagnum moss on which it sits around bog pools. It bears small, hard, evergreen leaves (iesclerophylls) which are covered with wax. Like Calluna vulgaris (Heather), Erica spp. andother genera of ericaceous subshrubs, Cranberry is apparently drought resistant (ie the sclerophyllsshow 'xeromorphic' adaptation). However, these types of leathery, reduced leaves are now regarded in awider ecological context than desert conditions and they are understood to be a reflection of nutrientstarvation, an adaptive structural feature called 'peinomorphism'. This term meaning, 'a hunger orstarvation induced form', was introduced by Weissenböck (1969) in connection with salt-toleranthalophytes, but it is becoming more widely applied to all nutrient-starved sclerophylls (Seddon 1974;Ellenberg 1988; Lusby et al. 1996).
Raised bogs by their nature and development are isolated from underlying mineral soils by a layer of fenor sedge peat. Thus plants growing on them receive only an extremely limited supply of mineral nutrientsfrom blown dust and materials brought down onto their surface by rain. There is strong evidence fromnumerous very different ecological situations (including Tropical Rain Forest in Australia) suggestingthat the apparent xeromorphic anatomy of this and many other bog plants, and especially the sclerophyllyof their leaves, reflects tolerance of very low levels of available phosphate (Müller-Stoll 1947;Loveless 1961, 1962; Beadle 1966; Small 1972).
The small, hard, thick-cuticled, waxy, evergreen leaves of V. oxycoccus have down-rolled marginsand their pores (stomata) are confined to the lower surfaces. The leaves also persist for two years, sothat altogether, the species provides a classic example of the extremely stress-tolerant, highlyconservative strategy of acquisition and use of mineral nutrients which typifies peinomorphicsclerophylly.
Root system
V. oxycoccus has a very shallow fibrous root system, hardly penetrating more than a fewcentimetres below the living parts of the moss layer on which it grows, so that it must have to competewith the Sphagnum for scarce nutrients. Like other ericaceous bog subshrubs, all species ofVaccinium produce dense root systems which end in fine absorbing rootlets, termed 'hair roots',ie rather than the normal, but anatomically different, 'root hairs'. Hair roots merely consist of one tothree layers of cortex cells around a narrow central conducting stele (Pearson & Read 1973). Becauseof its very shallow root system, V. oxycoccus relies on the excellent water-conducting andretaining capacity of the Sphagnum mosses it grows over for its water supply (Malmer etal. 1994). Thus the growth of Cranberry can be limited by either the prevailing chronic shortageof mineral nutrients (especially nitrogen and phosphate), or by a lowered or fluctuating water table.Surface drying of the bog is very deleterious to V. oxycoccus and it has been shown that growthof the subshrub is best when the average groundwater level is 25-30 cm below the root-stem junction(Gronskis & Snickovskis 1989).
Flowering reproduction
The tiny, pink, waxy flowers resemble a miniature Turk's-cap Lily or a Cyclamen sp. and have asimilar pollen release mechanism to the latter (visited by bumblebees). The open flowers are actuallyslightly less conspicuous than the flower buds, which display a more vivid crimson colour when stillclosed. Flowering begins in June and extends into August, the individual flower being unusuallylong-lived, an average of 27 days. Fruit development is very slow, taking between two and three monthsafter pollination (Jacquemart 1997).
Occasionally, V. oxycoccus can be found fruiting in great abundance and the bright red, or redspeckled with brown spots, very sharp-tasting berries, which slowly ripen over the winter period, areeither transported locally by streams, or are eaten and dispersed in the spring by birds such as robins,thrushes, blackbirds and red grouse and probably also by mammals. The most important Cranberrycollecting mammal in this part of the world is possibly the fox, but voles and mice may also feed onthem if they can negotiate the wetter parts of the bog (Jacquemart 1997; Mundell & Povey 2002). Thenumber of seeds per berry averages 7-8 and the estimated annual seed production per m2 isregarded as low, at around 480 ± 510 (Vander Kloet & Hill 1994).
Vegetative reproduction
Studies of successful recruitment from Cranberry seed in several parts of the Northern hemispherestrongly indicate that this type of plant establishment must be a rare, or even an extremely rare event.The dominance of vegetative over sexual reproduction is very prevalent in this genus, as it is in otherericaceous subshrubs, including most common Erica spp. (Jacquemart 1997).
British and Irish occurrence
Widespread in suitable bogland in Britain NW of a line between Cardiff and Hull, north into Scotlandsouth of the Great Glen and with a few thinly and widely scattered sites beyond these limits. InIreland, it is widespread but has declined in the Midland bogs due to very extensive, industrial peatextraction and in the W due to drainage and afforestation (M.C.F. Proctor, in: Preston et al.2002).
European and world occurrence
V. oxycoccus is widespread in N & C Europe extending locally to south-central France, NItaly and SE Russia. It then extends through northern Asia to Japan and N America. It is circumpolar inthe Boreal region of the N Hemisphere although there are distribution gaps in N Atlantic and Pacificareas (Hultén & Fries 1986, Map 1458; Sell & Murrell 2014).
Threats
Peat cutting and forestry and associated drainage operations still pose real threats to many Irishpopulations of Cranberry, including some in Fermanagh.
Native, occasional, locally frequent. Circumpolar boreo-arctic montane.
1881; Stewart, S.A.; Cuilcagh Mountain.
Throughout the year.
Growth form and preferred habitats
Cowberry can very easily be identified throughout the year by its broad, oval, evergreen leaves, darkgreen above and paler with black glands below. In Britain, and locally also in Fermanagh, this creeping,evergreen, rhizomatous, up to 30 cm tall subshrub has a remarkably similar distribution to that ofanother, even more prostrate species of the same habit − Empetrum nigrum (Crowberry). Inphytogeographical terms, both belong to the circumpolar boreo-arctic montane element and, in B & I,they are chiefly confined to the well-recognised 'edge communities' on upland wind-exposed scarps(including sea cliffs), rocky acid heaths on upland moors and mountain summits (New Atlas).However, in addition to such wind-dried heathy sites, in Fermanagh, V. vitis-idaea also growsover rock outcrops on damp-but-drained heaths around lakeshores. This happens across the Western Plateauand especially in the Lough Navar area of the VC.
The soils in which Cowberry grows range from pH 3.5-6.8 (Ellenberg 1988; Grime et al. 1988) but,in Fermanagh, the species is confined to the extreme acid end of this spectrum and it is seldom observedin soils over pH 4.0 and then only when the substrate is overlying limestone. The humus-rich soils itfrequents are typically in half-shade, constantly damp or wet throughout the year, although from theirposition, always quite porous and free-draining. Soils in the more wind-swept, exposed sites will alsobecome air-dried to some extent whenever the sun shines.
In every site where it occurs in Fermanagh, V. vitis-idaea co-exists and competes with two verymuch more common ericaceous subshrubs, V. myrtillus (Bilberry) and Calluna vulgaris(Ling). Grime et al. (1988) have described V. vitis-idaea as being, "intermediatebetween a stress-tolerator and a stress-tolerant competitor". This summarizes very well thesituation we observe in the field, in that Cowberry appears more tolerant of wind and drying soilconditions than V. myrtillus in exposed sites, and considerably more so than the most competitivespecies of the three, C. vulgaris.
The ecological tolerances of Cowberry suggest that in B & I it might eventually join V.myrtillus in colonising the less shaded areas of pine and spruce plantations in upland blanketbog peatlands, especially when such coniferous forestry is continued into a second generation of trees(Ellenberg 1988, p. 531).
Fermanagh occurrence
In terms of the frequency of the three B & I Vaccinium species which occur in Fermanagh, avery small margin (just three records), separates V. vitis-idaea from V. oxycoccus(Cranberry), making the former the least frequent of the Vaccinium species trio. The localdistribution of Cowberry is more upland and restricted of these two mentioned species, however, it beingrecorded in just 32 Fermanagh tetrads (6.1% of the total in the VC), whereas V. oxycoccus isknown from 48 tetrads.
Vaccinium myrtillus and V. vitis-idaea are the most abundant ericaceous dwarf shrubs in themountainous regions of C Europe as well as in the boreal and sub-arctic regions of N Europe, where theycommonly co-exist and are widespread both in the understorey of coniferous forests in montane andsubalpine zones and in treeless open 'tundra' habitats at higher altitudes (Ritchie 1955b; Renato etal. 2004). However, in NW Ireland, where truly high mountains do not exist and where in WFermanagh the Atlantic Ocean is within 15-20 km distance, the only site in the VC where Cowberry occursin woodland is in the Correl Glen NR. Here, red sandstone scarps occur under the canopy of humid, uplandoak-birch mixed woodland festooned with bryophytes and filmy ferns at the lower end of a very damp glenon the Western Plateau. V. vitis-idaea grows here both over rocks on the woodland floor and, veryrarely, also as an epiphyte on tree trunks.
Locally, Cowberry has one other interesting, very isolated Fermanagh outlier, on a scarp of rather loweraltitude at Drumskinny, which lies to the north of Kesh and Ederny, to the east of Lower Lough Erne.Finally, V. vitis-idaea appears in small quantity on eroded peat on one stretch of raised bog inGlen West Td, in the far west of Fermanagh.
Mechanisms of co-existence of similar species
While the results of many studies indicate areas of similarity and overlap between V. vitis-idaeaand V. myrtillus in terms of their plant structure, biology and habitat ecology, equally thereare sufficient differences in these areas plus in their reproductive systems, ecological requirementsand tolerances, or they could not co-exist in the same community and site and remain separate species.Ecological theory suggests there are two principal explanations for co-existence of similar or closelyrelated species in a community: a. habitat differentiation − meaning that species utilize differentportions of the available habitats; or b. resource differentiation − in which species partition thelimiting resources in such a way that each is limited by a different component of the availableresources or inherent environmental pressures (nutrients, light, temperature/shelter, water, fire,grazing pressure, disturbance and so on).
Shmida & Ellner (1984) found that differences in 'life-history strategy', such as availability ofseed from nearby habitats, differences in demographic response to environmental fluctuations andturnover in species composition between different habitat patches (ie 'patch dynamics') were allrelevant to the co-existence of similar or closely related species.
Significance of the evergreen habit
The most immediately obvious differences between the two Vaccinium species here underconsideration is in their morphology and life-form: V. vitis-idaea has thick, glossy, leathery,evergreen, two or three ranked leaves, while V. myrtillus has much thinner, more herbaceous,deciduous (or very occasionally semi-evergreen) leaves, which are much shorter lived, each leaftypically lasting less than one year. By definition evergreens are species that maintain a cohort ofleaves for more than one year (Chabot & Hicks 1982).
Using a mathematical model, Monsi (1968) showed that plant growth is largely determined by thepartitioning of the net photosynthetic gain into new photosynthetic and non-photosynthetic tissues, andby the turnover rate of the photosynthetic parts of the plant. The deciduous leaf habit requires areplacement of the photosynthetic system after every production term (ie the annual cycle of growth).This necessity slows down the growth of deciduous species in comparison with plants of evergreen habit,as demonstrated for instance by Beech (fa*gus sylvatica ) versus Spruce (Picea spp.)trees, and in this case, V. myrtillus growth slows in comparison with V. vitis-idaea(Schulze et al. 1977).
A study by Karlsson (1992) of leaf life span measurements for 16 evergreen shrubs in the Ericaceae andEmpetraceae, made in the European Alps and in C & N Europe, found that mean leaf longevity for eachshrub species varied between 1.4 and 3.8 seasons and that leaf persistence was consistently greaterfurther north. Evergreen leaves of V. vitis-idaea were retained for between 2.0 and 3.8 seasons,and at each of three latitudes studied, Cowberry leaves functioned the longest of all of the 16shrub species that were examined. However, the figures observed for shrubs are much lower than for theneedles of evergreen conifers such as Pinus and Picea, which often survive for 4-10 years(Karlsson 1992).
Plant nutrition and evergreen leaves
Leaf characteristics have been shown to be influenced by different nutrients. One of the earliestsuggestions was that soils low in potassium supported a more evergreen community (Harper 1914). Loveless(1961, 1962) showed a proportional increase in the degree of sclerophylly occurred with a decrease inleaf phosphorus below 0.3%. Sclerophyllous leaves are hard in texture and have cells with thickcuticles, small lumens (ie cell interior volumes) and reduced intercellular spaces, allowing the plantto be much more resistant to drought.
Several hypotheses have been raised to explain the ecological significance
and benefit to the plant of the evergreen habit in terms of nutrient conservation and improved carbonbalance − both of which give advantages to plants in environments where low nutrient levels limit leafgrowth. The evergreen habit might also be an adaptation to more general environmental stress. Chabot& Hicks (1982) reviewed this topic, listing and discussing eleven hypotheses along these lines, mostof which are not mutually exclusive.
The evergreen habit has long been regarded as an adaptation to nutrient-poor habitats, conferring anecological advantage over deciduous plants, or over those species with less persistent leaves. Theadaptive makeup of the evergreen plant involves: 1. low capacities to photosynthesize and to absorbnutrients; 2. a slower turnover of plant parts, combined with a high re-absorption of minerals fromsenescing leaves; and 3. storage of carbohydrates and nutrients in old evergreen leaves (Gerdol etal. 2000). A further advantage of evergreen species that reduces loss of minerals from theecosystem is year-round leaf drop. Through a gradual leaf fall, combined with a subsequent slower decayrate than for deciduous leaves, small amounts of nutrients may be made available to the roots ofevergreen species throughout the whole year. This will be particularly important in a mild, temperateclimate like that of Fermanagh, with moderate to heavy rainfall, where the growing season is relativelylong and leaching of soluble minerals from decaying litter and from the soil is rapid (Monk 1966).
The potential ecological benefit of the evergreen life-form was demonstrated by Karlsson's (1992) studythat found leaf life span of evergreen shrubs increased with decreasing soil nutrient status, ie leaveswere longer-lived in wet bogs than in drier heaths, and likewise at high altitudes compared with lowermontane levels. Evergreen shrubs also tend to dominate dry heathlands, being well adapted to droughtstress, probably through being better able to maintain a positive net carbon dioxide balance at lowwater potentials. Paradoxically, evergreen shrubs also dominate many waterlogged soils, since theseconditions can lead to conditions of physiological drought. This feature of evergreens has been knownsince Schimper's days in the late 19th century, resulting in the recognition of the 'peinomorphosisadaptation' typical of evergreen bog species, which develop plant growth forms and leaf types similar tothose of desert perennials (Gerdol et al. 2000).
Different types of evergreen leaves
At the same time, it clearly emerges from many studies that all evergreen leaves are notthe same, eg xeromorphs are drought resistant, while the superficially similar life-form, sclerophylly,may relate more to herbivore deterrence and the reduction of nutrient leaching in wet, infertile soilsituations. Thus the various anatomical and morphological structures that relate to shrub leaf life spancannot necessarily be explained by a single hypothesis. The same may also be said of herbs; no generalpatterns of leaf longevity appear in them either (Diemer et al. 1992).
Experiments in subarctic shrub communities
A number of studies lasting 5 or more years have been made examining annual variation in growth andreproduction of subarctic shrub communities dominated by co-existing V. vitis-idaea, V.uliginosum and Empetrum nigrum, and in which V. myrtillus is often also present.These investigations found that responses to experimental manipulation of temperature, water, nutrientlevels (in some cases only N, or N & P), and species composition (ie the selective removal ofabove-ground parts of same or neighbouring species), produced results that were highly complex andextremely difficult to predict. The complexity arose through varying species-specific patterns ofgrowth, great year-on-year variation, and high numbers of interactions between the four factorsmentioned (Parsons et al. 1994; Shevtsova et al. 1995 & 1997; Leith et al.1999).
Elevated temperature and increased nutrient level both produced the expected increase in totalabove-ground biomass, canopy height and rate of nutrient cycling, but there was little or no consistencyin the effect this had on the species competitive outcome and dominance. Nutrient addition did lowerspecies richness, but mainly through its negative impact on the mosses and lichens of the ground flora,a consequence of increased shading by the canopy of growth-stimulated dwarf shrubs (Press et al.1998). These latter workers also noted that their measure of plant cover revealed an accumulation oflitter and standing dead material in response to increased nutrient and temperature levels, operatingboth singly and in combination, suggesting a faster rate of turnover of plant material in the dwarfshrub community under these treatments.
Although temperature appeared to have a greater stimulating effect on dwarf shrubs than nutrientadditions in these higher latitudes and altitudes studies, it is likely that under warmer, more mesictemperate conditions, nutrients will become proportionally more important, affecting comparative growthrates and perhaps also the competitive relationships between these dwarf shrubs. The effect ofadditional water produced surprisingly little response in the subarctic dwarf shrub vegetation, despitethe experiment being carried out in one of the driest parts of Scandinavia (Press et al. 1998).
Comparative evergreen/deciduous nutrient-use efficiency: A similar study in a subalpine heath inthe Dolomites in N Italy comparing the water- and photosynthetic nutrient-use efficiency in co-existingV. vitis-idaea and V. myrtillus. This found as expected that deciduous V. myrtillusproduced the higher rates of net photosynthesis, and that this was positively correlated with leafnutrient-status and with carbon dioxide concentrations within the leaf. The percentages of N (nitrogen)and P (phosphate) pools reabsorbed from senescing leaves was also somewhat higher in the deciduousspecies. Lower concentrations of P in senescing evergreen leaves showed however, that V.vitis-idaea was more proficient at re-absorbing this element (but not N), when comparedto the deciduous species, and the evergreen shrub had a higher carbon gain per unit foliar N and P, dueto a longer mean residence time of both nutrients in the plant tissues (ie it conserved these nutrientsbetter).
The study did not detect any differences in water-use efficiency between the two shrub species, either onan instantaneous or a long-term basis, but this is probably because there was no sign of any waterdeficiency in the habitat throughout the growing season studied; there were no appreciable dry periodsexcept at the very end of the season when V. myrtillus had already shed most of its leaves butV. vitis-idaea was still active (Gerdol et al. 2000). However, in drier, perhaps morewind-exposed, or in semi-arid or arid conditions, it is highly likely that possession of evergreen orsclerophyll leaves may serve a different ecological role, functioning then in awater-conservation mechanism. In north-central Florida, for example, deciduous species predominate inmesic fertile sites, while evergreen plant communities segregate off to occupy dry, sterile sites, afamiliar enough pattern worldwide (Monk 1966).
Gerdol and co-workers (2000) concluded that evergreen V. vitis-idaea is competitivelyadvantaged over deciduous V. myrtillus, but only in extremely nutrient-poor habitats, andespecially so when the latter are phosphorus-limited. However in terms of their carbon economy (iephotosynthetic assimilation and storage), the co-existence of the two species in mixed communitiessurely must also reflect some degree of niche differentiation with respect to their light regime, apartthat is, from the differing duration of their assimilation period mentioned above.
A further study in treeless subalpine heath on three differing soils carried out in the Italian SE Alpsby Gerdol and co-workers (2004), looked at both above-ground and below-ground biomass of V.vitis-idaea and V. myrtillus in relation to manipulated soil moisture and nutrientcontent. Results indicated that V. myrtillus was primarily P-limited and V. vitis-idaeaprimarily N-limited. Water content affected the distribution of the two shrubs in a similar way, bothspecies producing the lowest biomass of the experiment when growing on peat, possibly due to a toxiceffect of waterlogging in wet substrates. Higher P-availability in the soil enhanced V. myrtillusrather than V. vitis-idaea, the presence of which is less distinctly related to soil nutrientcontent.
Carbohydrate cycle phenology
Deciduous species generally have higher photosynthetic capacities than evergreen species and requirehigher photon flux densities (ie light intensities) before photosynthesis in individual leaves becomeslight saturated. When photosynthetic performance was examined in situ in a subarctic dwarf-shrubheath, evergreen V. vitis-idaea assimilated about one fifth of its yearly carbon gain duringperiods when the related deciduous species V. myrtillus and V. uliginosum were leafless(Karlsson 1989). Although it would need to be investigated, it seems very likely this fraction could beeven greater under temperate conditions, and might be sufficient to significantly affect the competitiveoutcome between evergreen and deciduous species at least at the beginning of the growing season.
On the other hand, carbon lost from evergreen leaves through respiration during the winter months willundoubtedly reduce the gain from the longer growing season in more temperate latitudes. Another findingwas that the seasonal photosynthetic productivity of the deciduous V. myrtillus was more lightlimited than that of the evergreen V. vitis-idaea under most natural conditions (Karlsson 1989).
A Scottish Vaccinium phenology study: Carbohydrate content measured insamples of three species of Vaccinium collected from an exposed Scottish cliff ledge at 700 m onBen Lui, Perthshire at intervals of 2 to 8 weeks over a period of least 16 months found that all threesubshrubs followed a similar cycle to one another in both their aboveground and underground parts(Stewart & Bannister 1973). A rapid, substantial increase in their carbohydrate reserves occurredearly in the spring. In deciduous V. uliginosum for instance, this 'spring rise'occurred between the end of April and the third week in May, before its leaves were open. The springrise occurred even earlier in V. vitis-idaea and V. myrtillus, both of which possess atleast some photosynthetic tissue at this time of year - ie V. myrtillus has wintergreen, barklessstem tissue, while V. vitis-idaea has evergreen leaves. This was followed by a rapid drop incarbohydrate content later in the spring, or the early summer in the case of V. uliginosum, andin the below-ground parts only of V. vitis-idaea. A general increase in carbohydrate levels thenoccurred during the months when all three plants bear leaves (referred to as 'the summer rise'),followed in V. uliginosum by 'the winter fall' after the end of October, when respirationcontinues but temperature and other conditions no longer favour photosynthesis.
Carbohydrate levels in V. vitis-idaea and V. myrtillus actually begin dropping earlier thanin V. uliginosum, beginning to occur around the end of August. In Cowberry this continued onlyuntil the first week in November and the fall in stored carbohydrate is occasioned by an autumn flush ofgrowth (Ritchie 1955 b). In V. myrtillus the dropping levels of carbohydrate reserves ("thewinter fall") continues until the "spring rise" in the second week in February, and thecauses are an autumn flush of growth followed by subsequent respiratory losses during the winter months(Stewart & Bannister 1973).
Leaf decomposition and nutrient release
Against the trend favouring possession of evergreen leaves is the considerably greater length of timerequired for leathery leaf litter to decompose and release nutrients bound up in the dead tissues. Thisincludes time for the water-soluble nutrients to leach into the soil and become re-available to livingroots, ie to recycle, and also time for the mass of such dead residues to physically disappear from thesoil surface, rather than accumulate in a deepening litter layer that might inhibit other species andaffect moisture soil relations and root nutrient uptake (Monk 1966).
Sexual and vegetative reproduction
As discussed in detail for V. myrtillus (see species account), over a wide geographic area of thenorthern hemisphere seed of V. vitis-idaea is either absent from the soil, or is present only invery small quantity, so that reproduction must primarily be vegetative. This is accomplished bybranching and fragmentation of its shallow, underground rhizome (Vander Kloet & Hill 1994).
The shrub usually flowers sparingly in B & I, only one to three blossoms appearing on each pendulousraceme cluster, although the species is capable of producing up to twelve flowers on each (but morenormally five or six). It is reported that flowering occurs in spring and again in early summer (Ritchie1955b; Grime et al. 1988), but the plant is much too rarely recorded in Fermanagh to determine ifthis is the case. The N American form of the species, subsp. minus, differs in flowering onlyonce, as one might expect of a plant with a largely subarctic, short season distribution (Hall &Shay 1981).
According to Ritchie (1955b), Cowberry plants do not produce many flowers until they are somewherebetween 5 and 10 years established. Russian studies put the time of first flowering considerably laterthan this, most plants requiring 14 to 20 years growth before reaching sexual maturity (Hall & Shay1981). The flowers are said to be hom*ogamous, or almost so, the sexual parts ripening more or lesssimultaneously. Pollination is either by insects (commonly bees and butterflies), or involves selfing,since the species is partially self-compatible. Like V. myrtillus, Cowberry is reported tohabitually inbreed, but in V. vitis-idaea this is largely or entirely due to the scarceoccurrence of many of the over-dispersed populations in B & I, making cross-pollination a relativelyrare event (Ritchie 1955b).
V. vitis-idaea suffers a drastic decrease in fertility after self-pollination when compared tocross-pollination, another limiting property it shares with V. myrtillus. Partial self-sterilityin both these Vaccinium spp. is due to embryo abortion early in seed development (ie earlyinbreeding depression). Guillaume & Jacquemart (1999) hypothesized that this is based on theexpression of partially recessive lethal alleles during embryo development.
Fruit and seed production
In respect to fruit production, toxic properties of the berries, seed transport (mainly by birds),germination, rarity of seedlings and their slow rate of establishment, there appears to be very littlephysiological or ecological difference between V. vitis-idaea and V. myrtillus (Ritchie1955b, 1956; Hall & Shay 1981). Obviously since the former subshrub is much rarer,particularly further south towards the limit of its range in England, Wales and throughout Ireland,under the prevailing environmental conditions, opportunities for successful sexual reproduction andgenetic recombination are even more unlikely in Cowberry than in Bilberry. Nevertheless, in parallelwith other arctic-alpine plant species surviving changing conditions in B & I from the Late-glacialperiod to the present day, V. vitis-idaea demonstrates the characteristic tenacity all thesespecies share, being able to maintain existing, but perhaps shrinking populations, even when sexualreproduction and the ready transport which seed provides for the colonisation of fresh sites becomesseverely limited and they become increasingly dependent upon vegetative reproduction for their survival.
Hybrid
In Britain, a very rare intermediate hybrid (V. × intermedium Ruthe)is formed with V. myrtillus, but it has never been recorded anywhere in Ireland.The hybrid is widespread in C & N Europe (Stace et al. 2015).
Irish occurrence
In the rest of Ireland apart from Fermanagh, the distributions of V. vitis-idaea and Empetrumnigrum do overlap, but the latter species is very much better represented throughout theRepublic of Ireland. In comparison, south of the International border V. vitis-idaea is confinedto sites in six mountainous regions in the Donegal, Sligo-Leitrim, Connemara, Wicklow, Limerick andWaterford areas, although there are old records in the Irish Census Cat. from three other VCsfurther south (H7, H10 & H13). In Co Limerick (H8) for example, Cowberry was found by Stelfox in thewestern portion of the Galtee mountains (Praeger 1946), and one small, very inaccessible colony survivesthere (Reynolds 2013). Elsewhere none of these southern records have been refound for many years andthey may well be extinct (Scannell & Synnott 1987). In the northern province of Ulster, V.vitis-idaea has a very much wider, although still a rare and mainly upland occurrence, which isstrikingly coincident with that of Empetrum nigrum (Hackney et al. 1992).
British occurrence
Although managing to persist in single isolated hectads in S Devon and S Somerset (VCs 3, 5), furthernorth V. vitis-idaea is widespread in much of upland Britain and has its greatest presence inScotland. In England, it is most abundant in the SE Pennines and the Pennines in Cumbria. Cowberry isdescribed as "not infrequent" on mountains in the Lake District and it descends to nearsea-level close to Morecambe Bay (Halliday 1997). V. vitis-idaea is described as rare, sparse anddeclining on the Pennine Uplands of Co Durham (23 tetrads mapped; Graham 1988, p. 155), while on theother hand in Northumberland the species is described as being scattered and frequent and plentiful onhigher parts of The Cheviot (being mapped in 93 five km squares, plus three pre-1968 (Swan 1993, p.188)). The New Atlas gives the species a low change index value of ‑0.18, which suggeststhat overall, at the hectad level of discrimination, the distribution has not significantly alteredsince the 1962 BSBI Atlas (Preston et al. 2002; Walters & Perring 1962).
European and world distribution
Hultén (1971, Map 69) illustrates the distributions of the two closely related subspecies, thecircumpolar subsp. minus overlapping the more southerly subsp. vitis-idaea in Scandinavia,where introgression between the subspecies occurs; they co-exist again in Asia in parts of E China andJapan. In W Europe, V. vitis-idaea is widely distributed in arctic, boreal and temperate areasand towards the south of its range has montane and alpine outliers in France, N Spain and N Portugal.The same form of the plant also has further outliers east of the Black Sea (Hultén 1971; Hultén &Fries 1986, Map 1460).
The overall picture of the total species range fits the familiar pattern of the arctic-alpine floraelement of Matthews (1955), the plant having a significant portion of its geographical range lying northof the tree limit, although this itself is a very variable and difficult concept to define. For goodaccounts of the problems, historical and otherwise, in delimiting arctic and alpine timberlines, seeBarry & Ives (1974), Larsen (1974) and Wardle (1974).
Uses
Cowberries are very sharply tart until they are subjected to frost, so they become somewhat morepalatable later in the winter period. Despite their strongly astringent nature, in N Europe, andespecially in Scandinavia, there is a tradition of collecting them by combing them from the branch ends.The berries are used to make jelly, or mixed with other wild fruits, such as rose hips, to make jam.Cowberries are also widely processed and marketed in Japan and are commercially harvested in parts ofScandinavia, Russia, Alaska and Canada. Considerable amounts of fruit are imported into the UnitedStates annually. Much of this imported fruit is consumed by people of Scandinavian descent who use theso-called 'Swedish lingenberry' in traditional dishes.
V. vitis-idaea s.l. has the potential for more extensive commercial development and some nativestands in the subarctic could be managed with a minimum of cultivation, as are those of Low sweetblueberry. The feasibility of expanded commercial operations has being trialed in parts of North America(Tirmenstein 1991).
Porsild (1937) pointed out the valuable antiscorbutic properties of Cowberries due to their high vitaminC content. In comparison with V. myrtillus, V. vitis-idaea appears to have very littletraditional use in herbal medicine, the only reference to it uncovered by Allen & Hatfield (2004)being as an ingredient of an inhalant for a blocked nose or for treating blocked sinuses in Cumbria.
Names
The Latin specific epithet, 'vitis-idaea', is a name first used by Theophrastus and it means 'vine ofMount Ida or Idaea', a reference to a Greek mountain where presumably the plant grows, or once did(Gilbert-Carter 1964). The Mountain Flora of Greece, 1 (Strid 1986, p. 741) mentionsseveral sites for the species, but it does not appear to include any Mount Ida!
Twelve English common names are listed by Britten & Holland (1886) including 'Flowering Box','Brawlins', 'Clusterberry', 'Ling-berry', plus some names shared with other species of the genus. Themost familiar and entirely inappropriate name, 'Cowberry', is said to have arisen from a 19th centuryblunder which confused the genus name 'vaccinium', the fruit of the whortle, with 'vaccinum', the Latinfor 'what belongs or pertains to a cow' (Prior 1879, p. 55; Grigson 1974).
Threats
Since V. vitis-idaea is already very local and sparse, only occurring in out of the way sites, itis somewhat protected by these factors. The lack of effective seed regeneration however, means that oncepopulations are lost for any reason, they are not naturally replaced and thus the species is vulnerableto any change in the environment.
Native, common and locally dominant. Eurosiberian boreal-montane.
1818; Walford, T.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This very variable calcifuge, semi-deciduous subshrub forms large clonal patches by horizontal vegetativegrowth of its branching underground rhizome. Frequently, but locally, this enables Bilberry to becomethe dominant or subdominant species of the under-storey field layer in damp, upland, acidic, mixeddeciduous woodlands and also, to a certain extent, in more open areas within coniferous plantations(Ritchie 1956). Since the plant can tolerate more shade than both Calluna vulgaris (Heather)and Erica spp., to a limited extent it can occupy darker areas in conifer plantations thancan those shrubs (Ritchie 1956).
The breadth of the species woodland tolerances is demonstrated by the fact that Rodwell et al.(1991a) lists its presence in nine different woodland communities in the British NVC classification (notall of which, of course, occur or are relevant in an Irish context); it is considered a 'constantspecies' in five of these communities. While the subshrub grows largest and most bushy (c 60-90 cm tall)in fairly open woodland, it noticeably tolerates more shade than the other ericaceous species with whichit almost always closely associates and competes.
Small plants of it are frequently present in an often very subsidiary role in heaths, heathy grasslandand bogs dominated by other species, so that Rodwell et al. (1991b) lists it occurring in sixtypes of NVC mires (bogs) and as many as 17 forms of heath vegetation, and as a constant speciessufficiently significant to feature in the community name, in six of the latter. It is therefore notsurprising that Grime et al. (1988) described it as a "stress-tolerant competitor".
Bilberry is common on peat bogs where it tends to dominate the very tops of the characteristic vegetationhummocks. It also becomes abundant around the margins of isolated rock outcrops in bogland and likewiseits presence increases on the somewhat better drained sloped margins of the peat dome of lowland raisedbogs. On bogs that are cut for turf (peat fuel) on an irregular, occasional basis, Bilberry oftenbecomes prominent on the drier edges of peat banks or those of cut surfaces after the spade or 'turfslipe' is used for peat sod extraction (Evans 1942, p. 136-7). The steadily increasing use nowadays ofmodern mechanised turf extractors which tunnel below the surface of raised and blanket bogs(appropriately enough nicknamed 'sausage machines') can, at least temporarily and very locally, increasedrainage in the peat and thus favour the growth of Bilberry over C. vulgaris and Ericatetralix (Cross-leaved Heath), at least in the short term. Unfortunately, this method of peatextraction eventually causes the collapse and frequently also the rupture of the thin layer of surfacevegetation, so that erosion (either cyclical or longer term) often results.
On and below cliff scarps in upland areas, especially as in the Lough Navar Forest Park where many of theslopes are covered with a relatively well stabilised, overgrown block scree, V. myrtillus mayalso be locally abundant, competing well in these generally completely ungrazed circ*mstances withCalluna vulgaris, Erica cinerea (Bell Heather), Pteridium aquilinum(Bracken) and Luzula sylvatica (Great Wood-rush). Bilberry is often closely associated with V.vitis-idaea (Cowberry) growing under its canopy in this type of habitat.
Variation
V. myrtillus is phenotypically extremely variable and 'plastic' in its response to growingconditions in its wide variety of habitats. It generally performs best however in well-drained, somewhatsheltered situations. At the same time it is able to persist and is often quite abundant (albeit in amuch dwarfed form, c 5.0 cm tall) in sheltered spots near exposed mountain summits. Dwarfism and reducedstem rigidity are genetically controlled, and both are correlated with increasing total leaf nitrogen athigher altitudes. This in turn controls maximum rates of photosynthesis and levels of stomatalconductance, assisting absorption of carbon dioxide as the concentration of the gas thins with altitude,thus compensating the plant for a shortened growing season. The thinner, less rigid stems enableincreased levels of leaf nitrogen to occur and the stems also survive wind buffeting better. However,they do so at the expense of sexual reproductive capacity, each stem being only able to bear the weightof one or two fruits at the most (Woodward 1986).
Fermanagh occurrence
In Fermanagh, V. myrtillus is frequent or common and locally abundant, recorded in 244 tetrads,46.2% of those in the VC. Here, as elsewhere in B & I, Bilberry grows in a very wide variety ofstrongly acidic, peaty conditions, ranging from sheep pasture grasslands, even on those formed over veryshallow raw humus, or on heathy podsols formed directly over hard Carboniferous limestones. Examples ofthe latter occur around Knockmore, above Florencecourt and in the Cladagh River Glen (= The MarbleArch), as well as on ombrogenous raised and blanket peat bogs (ie Atlantic mires). Bilberry also thriveson drier, steeper, heathy areas of moorland, especially in sheltered pockets of ground around rocks, orin hollows and on, or close to, mountain summits, cliff scarps and on stabilised, talus scree slopesbelow cliffs.
Flowering, fruit and seed production
When in full sun, V. myrtillus flowers freely with two peaks of anthesis between April and earlyJuly. Flowers are either insect-pollinated by bees and wasps, or self-pollinated by gravity (Richie1955a). Inbreeding is described as 'habitual' (Richie 1956) and a recent study has shown a drastic'early-inbreeding' depression in fertility takes place in Bilberry due to embryo abortion in the earlystages of seed development following successful self-pollination. It has been plausibly suggested thatthe observed partial self-sterility results from the accumulation of lethal recessive gene alleles, ierather than any form of self-incompatibility system existing in the plant (Guillaume & Jacquemart1999).
Masting behaviour
The familiar small but plump, blue-bloomed, purple-black berries (called 'Fraughans' locally, a GaelicIrish name of the plant) are produced from July to September in numbers which generally appear, or whichare assumed, to vary with the prevailing seasonal weather conditions. In Fennoscandia, however, Bilberryhas been reported to produce large berry and seed crops at three to four year intervals, ie it isobserved to follow a masting cycle synchronised within the population (Myrberget 1982; Selås 1997). Mastyears in S Norway have been followed by temporal population increases of a large number of animalspecies of all trophic levels and an analysis of a 50 year record of seed production found that both theprevious degree of Bilberry reproduction and weather helped to explain the pattern. At the same time,regularity was commonly interrupted by unfavourable weather conditions and Bilberry production wasdepressed by, amongst other factors, low temperatures in spring and when flowering was taking place, andlow or high levels of precipitation during berry ripening in summer (Selås 2000).
Seed population behaviour
Each berry may contain up to 20 small yellow seeds, but the calculated average is just over 16 seeds perberry. In the shade of woodland canopy, or in the more exposed upland habitats over c 300 m, far fewerflowers are produced by comparison with well lit, sheltered conditions on open hillsides. In these morestressful environments, berries tend to be scarce or rare, some of them only partially formed or filledand, in view of the above revelation regarding self-pollination, we may assume that most seed containedin the poorly formed berries are sterile (Guillaume & Jacquemart 1999).
Studies in Canada, albeit further north in latitude than we lie, involving six species ofVaccinium in heathland, showed seed production is variable in time and space and varies from yearto year. Even where Bilberry and V. vitis-idaea are dominant in the vegetation, their seed iscuriously rare or absent in the soil seed bank. This is often the case despite the fact thatexperimental measurements of V. myrtillus seed longevity showed that their survival variesbetween two years and more than six (Vander Kloet & Hill 1994). Fungal rot appears to play a part indetermining, or rather limiting, seed viability; Cippollini & Stiles (1992) have suggested that theseeds of summer-ripening berries in the Ericaceae generally have ineffective defence againstfungal invasion and rot. This often overlooked topic has previously been reviewed by Janzen (1977) andby Herrera (1982), both of whom concluded that the rot factor affecting both fruit and seed could be assignificant a limitation on reproductive performance as seed predation, or it might be of far greaterconsequence.
A parallel study on seed production and its fate on moorland in Scotland found that even whenV. myrtillus produced many berries containing highly viable seed, collection of the berries bybirds and mammals was minor. Most berries fell into the mossy ground under the bushes that produced themand the soil seed bank was small (c 0-274 seeds/m²). Observations of seedlings were extremely rare andthen only in bare peat in disturbed ground conditions. Furthermore, the viability of buried seeddeclined from 90% to 20% in three years, due to a combination of decay in the soil and prematuregermination, ie germination soon after dispersal, in or on top of the litter zone in conditionsinimical for seedling growth, establishment and survival (Welch et al. 2000).
Contrary to this, Ridley (1930) quotes numerous reports of birds of the thrush, crow, pigeon and grousefamilies feeding on Bilberries in areas of Britain and Europe, while on the other hand, in their bookBirds and Berries, Snow & Snow (1988) make no mention of this happening, and these authorsalmost totally overlook Bilberry. On the basis of these reports we might conclude, as have some otherstudies of British moorland soils, that the seed bank of V. myrtillus is virtually non-existent.It certainly contrasts very strongly with the huge seed store of Calluna vulgaris that existsunder similar habitat conditions eg 52,900 viable seed/m² (Hester et al. 1991).
Seed dispersal and seedling rarity
Despite the above, it appears that Bilberry does achieve occasional long-distance jump-dispersal to new'vacant' sites by being transported by frugivores (ie fruit-eating birds and mammals). Guitian etal. (1994) reported migratory flocks of Redwings (Turdus iliacus coburni) consuming wildfruits of Empetrum nigrum (Crowberry), Vaccinium uliginosum (Bog Bilberry), V.myrtillus, Arctostaphylos uva-ursi (Bearberry) and Rubus saxitilis (Stone Bramble)in SW Iceland. This form of transport occurs at an enormous expense in terms of lost seed (Flower-Ellis1971). Low levels of seed persistence in soil and the observed rarity of seedlings in the field in siteswhere the species grows are very probably due to poor defence of the propagules from pathogens andpredators in the soil and, rather unexpectedly, a poor competitive ability after germination (Welchet al. 2000).
The most likely fruit and seed consumers in the Fermanagh area are sheep, small rodents ie mice and voles(probably very frequent) and, much more occasionally, birds of the thrush family. The habit of rodentspecies storing food in caches may apply to bilberries in moorland and, since this could assistdispersal, it deserves further study (Price & Jenkins 1986). However, some of the potential animalvectors are probably as much seed predators as transporters (Janzen 1969, 1971); the predators includeinsects, as well as birds and mammals. None of the mammalian dung examined in two studies in NE Scotlandproduced any seedlings of V. myrtillus and it therefore appears likely that passage of the seedthrough the gut of animals as different as cattle and voles most of the time effectively destroys embryoviability (Welch 1985; Welch et al. 2000). On the other hand, game bird studies of Red Grouse andCapercaille have reported viable seedlings growing on their dung in parts of England and Scotland (Welshet al. 2000).
Another significant observation which helps explain the rarity of V. myrtillus seedlings on moorsand in woodland is the extremely slow growth that Bilberry seedlings are capable of achieving in damp,cool, acidic, nitrogen-limited and other nutrient-deficient soil, which means that a rhizome does notdevelop until at least the third year of a plant's life (Flower-Ellis 1971). The result of thislimitation is that the young seedling cannot extend laterally to escape competition from other morerapidly growing ericoid or graminoid seedlings, or from the established moorland or woodland groundlevel sward surrounding it.
Vegetative reproduction and clonal spread
It is clear from the above that in both shaded and more exposed conditions, the photosynthetic resourcesand growth of Bilberry is often directed not towards sexual reproduction, but rather to vigorousvegetative spread achieved by underground horizontal extension of its sympodial branching, woodyrhizome. Thus many Bilberry plants form large clonal patches of indeterminate, but presumably oftenconsiderable age, in stable woodland, scrub or exposed heath situations (Richie 1956). Numerousobservations indicate that recruitment from seedlings is infrequent in many species of clonal plants,perhaps especially in those which, like Bilberry, advance using a 'phalanx growth form' (ie tightlyaggregated shoots or ramets, with clonal extension growth confined to the perimeter); V.myrtillus appears to be a good exemplar of this type of primarily vegetative reproductivebehaviour, which directs the long-term growth and dispersal of the species in a locality (Eriksson1989).
Response to sheep grazing
The phenology of Bilberry is unusual for a shrubby plant in B & I in being deciduous yet havingangular stems which stay green and relatively free of woody tissue during the winter months, so thatconsiderable winter and spring utilisation by browsing herbivores may be expected. Bilberry growth alsostarts earlier in the year than in most other moorland plants (Welch et al. 1994) and so it isliable to be heavily grazed by sheep or cattle when nutritious, digestible fodder is typically at itsmost scarce in February and March. In mixed stands with Calluna vulgaris and in ± pure Bilberryheaths in the Derbyshire Peak District, sheep given all year access consistently chose to graze theBilberry swards much more heavily in autumn (August-October) than in the rest of the year, while forC. vulgaris the increased grazing pressure fell between October and January.
Autumn peaks in carbohydrate (Stewart & Bannister 1973) and fructose and glucose (Pakonen etal. 1991) contents have been found in Bilberry and this, coinciding with a sharp decline in thenutrient content and digestibility of moorland grasses in autumn, appears to shift sheep feedingpreferences towards ericoids at this season (Powell & Malcolm 1974). In mixed stands, it is oftennoticeable that Heather and Bilberry exist in distinct patches, each of which has one species dominantand the other absent or very subsidiary. Under three different experimental grazing regimes, these mixedheaths showed a sharp decline in Bilberry cover and an equally marked increase in C. vulgariscover and height (Welch 1998). Flowering and fruiting will be more depressed (at least in the shortterm) by grazing in Bilberry than in Heather, since V. myrtillus flowers develop on the previousyear's shoots, whereas C. vulgaris flowers appear on the current year's shoots, which generallyare not sheep browsed until wintertime.
Insect herbivores
Many insects feed on Bilberry, predominantly the larvae of geometrids, tortrids and sawflies. Theselarvae utilise leaves, bark, buds and reproductive parts of the plant as their sole food. High larvaldensities in late June and early July coincide with the hatching period of many woodland breedinginsectivorous birds which rely, at least in part, on the larvae to feed their nestling broods. Exclusionof birds from experimental Bilberry plots in Swedish boreal Pine, Spruce, Birch forests found that thetotal density of larvae was 63% lower outside the enclosures where the birds had access and that thescale and type of damage to Bilberry annual shoots by the insect larvae was significantly affecting thecompetitive potential and reproductive performance of the shrub (Atlegrim 1989).
Other damage and comparative recovery potential
In common with the woody rhizome, branching of aerial ramets in V. myrtillus is sympodial and,since apical dominance is weak and dormant buds are abundant, branching is both frequent and flexible,allowing compensatory growth to occur in the different habitat-related light levels which bushesexperience after environmental stress caused by eg herbivory, late frost, burning, trampling or fungalattack (Tolvanen 1995). The woody stems of Bilberry and other ericaceous subshrubs, however, are verylimited in their ability to tolerate trampling wear and tear and their recovery is slow to negligible(Cole & Spildie 1998; Malmivaara et al. 2002). Since the rhizome typically grows at depths of15-20 cm, which is somewhat deeper than C. vulgaris roots, it is generally believed that regularburning of heathland favours Bilberry over Heather, but further studies to monitor the composition ofmixed Bilberry-Heather stands over the whole cycle of burning are required to test thishypothesis scientifically (Welch 1998). In comparison with its evergreen relative and associate, V.vitis-idaea, the deciduous growth form of V. myrtillus increases its regrowthpotential after damage by the various factors under consideration here, since its photosyntheticand growth rates are both greater than those of the former shrub (Tolvanen 1995).
British and Irish occurrence
The New Atlas hectad map shows that in Britain, V. myrtillus has a very marked N & Wdistribution, being almost entirely absent from south central and eastern England, which broadlycorresponds with the major areas of chalk and limestone geology. In Ireland, by comparison, V.myrtillus is shown at the hectad scale as being much more widespread throughout, although it isundoubtedly better represented in the N & S of the island. Elsewhere, it is much more restricted tohigher ground. In Fermanagh, Bilberry is much more frequent and abundant on the higher ground on theWestern Plateau than elsewhere, but it is well scattered throughout the county and lowland sites includelakeshore woods, scrub and peatlands. In the wet oceanic climate of W Ireland, regular, heavy rainfalland long-term leaching is known to induce shallow, acid peaty ranker soil formation directly on top oflimestone rock and, in Fermanagh, some of these profiles support at least stunted Bilberry bushes.
European and world distribution
Widespread in the boreal Eurosiberian region from Iceland eastwards through Fennoscandia to C Siberia andbeyond to N & C China. The range also reaches southwards more or less continuously to the Alps andthe other middle latitude mountain alpine and subalpine forest zones in Europe, from N Portugaleastwards to the Caucasus. It has also been reported from Alangorssuaq Island in S Greenland (Böcheret al. 1968), but there is a suggestion that it may be introduced here (Hultén & Fries1986, Map 1462).
Uses
In the past, berries were often collected and eaten raw − preferably with cream, or used to make pies,puddings, jam and jelly, or to add to wine or spirits to manufacture liqueurs, practices which at leastlocally appear to have declined or died out. Grigson (1987) reports that the berries were used fordyeing in Scotland and Ireland and that in the past the branches were fashioned into brooms and whisksto use in spraying the potato crop with Bordeaux mixture against blight. In the Hebrides, Bilberryleaves were used as a tea substitute and, being a strong astringent and diuretic, it was usedmedicinally for dissolving kidney stones and the treating of other ailments of the urinary tract. Allen& Hatfield (2004, p. 123) catalogue numerous other medicinal uses which closely mirror those of'heather' (ie the more common species of Erica plus C. vulgaris). As a result V.myrtillus has accumulated many local names.
Names
The origin of the genus name 'Vaccinium' is something of a mystery. It appears to be a Latinisedname of great antiquity said to date back to the same prehistoric Mediterranean language (ieThraco-pelasgian) as the Greek 'Hyakinthos', ie commonly known as 'Hyacinthus'. Hyakinthos is thepre-Hellenic name of a boy god beloved of Apollo in Homer's poems and Greek myth, accidentally killed bya discus which was diverted to hit him by the jealous wind god Zephyrus. From his blood the poets say aflower sprang marked by Apollo's cries of grief, 'AI AI' (Radice 1973). Alternatively 'hyakinthos' maybe a word in the ancient language describing the blue colour of water! It is thought that perhaps, oreven probably, 'Vaccinium' is a corruption of that name, but it depends upon the authority oneconsults (Gilbert-Carter 1964; Gledhill 1985; Stearn 1992). From the current author's low level ofawareness of the linguistics, that explanation of the name and its supposed transition to 'Vaccinium'does appear to be a very long shot indeed! The Latin 'vaccinus' is an adjective meaning 'dun in colour'(Stearn 1973), but this is not regarded as relevant, and neither is 'vacca', meaning 'a cow'(Gilbert-Carter 1964). The Latin specific epithet 'myrtillus' is a diminutive literally meaning'a small myrtle', ie likening the blue berry to a small myrtle-like fruit (Gilbert-Carter 1964; Stearn1992).
The English common name 'Bilberry' dates from the 16th century, and as with the Latinised genus name, itsorigin is uncertain. The first element 'bil' is probably of Scandinavian origin, as is 'blae', 'blea'(or 'blaa' in Old Scandinavian) meaning 'blue-black' of the 15th century English common name 'Blaeberry'(Grigson 1974). The widespread alternative name 'Whortleberry' is very probably a corruption of'Myrtleberry', and 'Hurtleberry', 'Hurtberry', 'Hurts', 'Whorts', Huckleberry', 'Hartberry' are alllikewise corruptions of one another linked to 'Myrtleberry', and used as local names around theseislands (Prior 1879, p.123). 'Wimberry' on the other hand may derive from 'Wine-berry', a reference to amore sociable use of the plant.
Threats
Although currently still widespread and common throughout B & I in suitable habitats, the limitedability of the species to reproduce sexually, especially the low level of representation in the soilseed bank and the virtual lack of successful seedling establishment, resulting in a nearly totalreliance on vegetative growth for maintenance and spread of existing populations, makes V.myrtillus vulnerable to local extinction and fragmentation of its distribution following anyexcessive environmental stress or stresses it encounters. The fungal pathogen Phytophthoraramosum, that recently arrived in B & I from N America and is rapidly spreading north andwest, has a very wide host range which includes Rhododendron and other ericaceous shrubs includingV. myrtillus. The disease has not yet appeared on this genus anywhere in Ireland, but wemust not be complacent.
Native, rare. Circumpolar boreo-temperate.
1904; Praeger, R.Ll.; scarp south of Carricknagower Lough.
Throughout the year.
Growth form and preferred habitats
An evergreen, mycorrhizal, patch-forming perennial herb with a creeping rhizome and thick, ratherleathery, roundish-oval, radical leaves borne in a rosette at the base of the very short, woody stem. Itoccurs in damp woods, both deciduous and coniferous, including plantations, on a variety of humus-richsoils, usually covered with a deep surface litter. It can also occur on damp upland heaths and on shadyledges on wet, rocky scarps. Members of the genus all have mycorrhizal roots and, although they arecapable of photosynthesis, they can be thought of as semi-saprophytic, being able to tolerate or endureconditions of very deep shade (Salisbury 1942). Unlike P. media (Intermediate Wintergreen) andOrthilia secunda (Serrated Wintergreen) which in Fermanagh are generally either completelyvegetative or very shy flowering, most populations of P. minor in the VC usually contain quite afew flowering spikes, which aids both the discovery and the identification of the plant.
Flowering reproduction
The pinkish-white, pendulous, globular, 6 mm flowers are borne in a dense terminal raceme around a 10-30cm scape from June to August. Although there is no nectar, a sticky fluid that insect visitors lick isexuded by the five lobed stigma. The straight 1-2 mm style is included within the five quite free,rounded petals of the corolla and is shorter than the stamens and ovary. The flowers are either insect-or self-pollinated and produce large numbers of very minute, lightweight (orchid-like, but even smaller)seeds, shortly tailed at each end, in a globular capsule that eventually splits to release them on anypassing breeze (Melderis & Bangerter 1955; Clapham et al. 1987). As the seeds havepractically no food reserves, successful germination and seedling growth appears to require the obligateassistance of a saprophytic fungal partner from the soil flora, in the same manner as terrestrialorchids do (Salisbury 1942, p. 94).
Fermanagh occurrence
In Fermanagh this pretty flowering herb grows in quite sizeable clonal patches in damp, old, mixeddeciduous woodland, in a coniferous plantation and on wet, shaded scarps. Colonies can still beimpressive, some producing hundreds of flower spikes and covering several square metres of shadedground. P. minor has been recorded in a total of six tetrads in Fermanagh, all of which containpost-1975 records. Of the three pre-1951 stations, two survive, ie Florencecourt plantation and theCorrel Glen woods, and four completely new sites have been discovered since 1985. The population in theCorrel Glen mixed oak wood recorded in 1946 by Meikle and his friends was only rediscovered by RHN &HN in 2004, although this excellent site has been exhaustively worked botanically and has for many yearsbeen protected and recognised as one of our most important Nature Reserves. Since the plant produceshuge numbers of tiny, dust-like seeds which allow the possibility of jump dispersal effected by wind,the species could very well turn up at more new sites.
In a paper entitled Among the Fermanagh hills describing five days in July 1904 spent botanisingin the county, Praeger (1904) conveys the excitement he felt on finding three Pyrola speciesgrowing together on the wooded scarp south of Carricknagower Lough. The species were Pyrolasecunda (= Orthilia secunda) (Serrated Wintergreen), P. media (IntermediateWintergreen) and P. minor. Praeger knew he was only the second botanist in Ireland to ever havefound such a site, the other fortunate recorder having been Dr Moore of the Glasnevin Botanic Garden,Dublin when the latter visited Co Londonderry (H40), 70 years previously! Unfortunately, despiteprolonged, careful searching, we have not been able to rediscover P. minor at Praeger'sCarricknagower site.
With the exception of Dryas octopetala (Mountain Avens) and P. minor, all the associatedrare or scarce plant species which Praeger mentioned in his impressive 1904 paper, including P.media, Orthilia secunda and Asplenium viride (Green Spleenwort), are still to befound today where he described them. Indeed, we find the absence of Praeger's station for P.minor at Carracknagower so odd, that despite his achievements, reputation and towering statureamong Irish naturalists, it is tempting to suggest he might have been mistaken – not in hisidentification of the plant, but rather in his recollection of the location of his discovery.
The record details additional to the first record are: Correl Glen, 1946, MCM & D; Florencecourtplantation, 1950, MCM & D; conifer plantation, Cassidy Td, Necarne estate, 28 June 1987, D. Irvine& RHN; Pollaphouca Waterfall, 26 October 1992, RHN & F. Carroll; Derryvore Oakwood, oppositeCrom Castle, Upper Lough Erne, 4 January 1995, RHN, HJN & I. Herbert; wooded scarp, Pollaphoucacliffs (different grid reference from waterfall), 24 May 1998, RHN.
Irish occurrence
The New Atlas hectad map indicates that in Ireland, Common Wintergreen is a rare and slowlydeclining species, increasingly confined to widely scattered sites in the northern quarter of theisland.
British occurrence
P. minor has a strongly marked northern distribution in Britain, being rare, widely scattered andin decline in most of SW, S & C England, East Anglia and Wales since before the 1930s. It is muchmore frequent and widespread in N England and Scotland, although even here it is absent from the OuterHebrides, Orkney and Shetland (Garrard & Streeter 1983; Preston et al. 2002; Sell &Murrell 2014).
European and world occurrence
P. minor is widespread and circumpolar in the boreal regions of the N Hemisphere and is alsocommon in montane situations further south, reaching the mountains of C Spain, C Italy and Greece and inN America stretching from Alaska south to California and east to Labrador and New England (Hultén &Fries 1986, Map 1436; Sell & Murrell 2014).
Threats
Forestry operations and overgrowth by brambles and Rhododendron at the coniferous plantation site nearNecarne.
Native, very rare and probably facing extinction. Eurosiberian boreal-montane.
1904; Praeger, R.Ll.; on the shore of Lough Fadd.
Throughout the year.
Growth form and preferred habitats
Intermediate Wintergreen is a rare, rhizomatous, mycorrhizal, evergreen, perennial herb with a basalrosette of fleshy ovate or rounded leaves. While the plant produces a creeping underground rhizome thatenables some degree of vegetative reproduction and clonal dispersal, this is a delicate organ of veryslender dimensions, rather aptly described by the famous Victorian alpine explorer, gardener and plantcollector Reginald Farrer as, "long, fibreless runners like fine macaroni, that never stops torest" (Farrer 1930, 2, p. 203). P. media can produce a flowering scape up to 30 cmtall bearing a terminal raceme of 10-15 pendulous, white, globular flowers that are larger than those ofP. minor (Common Wintergreen). However, it often fails to do so, the P. media plantrosettes remaining stubbornly vegetative.
The preferred growing conditions are semi-shaded, damp and mossy, yet well-drained, mildly acid toslightly basic, leached and nutrient-poor soils. On these types of substrate, it usually grows under thesecondary canopy of ericaceous sub-shrubs on heaths (often upland) and in woods (especially pinewoods)(Stewart et al. 1994; Hill et al. 1999; F.J. Rumsey, in: Preston et al. 2002).
With regard to distinguishing the two Pyrola species that occur in Fermanagh (the otherwintergreen present is now in the genus Orthilia), experience to date indicates P. mediaalways occurs in very small populations of discrete leaf rosettes. It grows in crevices or ledges onsandstone cliff scarps, or under dwarf ericaceous sub-shrubs on the steep, overgrown talus slopesbeneath these cliffs. This contrasts strongly with P. minor (Common Wintergreen) which occupiesmuch more horizontal ground in light to medium density woodland shade, generally under a canopy ofbirch, rowan, ash and hazel. P. minor usually appears in considerable quantity, forming eitherdiscrete patches or wide carpets of growth and it flowers very much more freely than does either P.media or Orthilia secunda (Serrated Wintergreen).
Fermanagh occurrence
Praeger found this very rare species on the rocky shore of Lough Fadd on 3 July 1904, where it has notbeen seen since; he also recorded it among steep-sloping, sub-montane ericaceous heath on four or fivenorth-facing dolomitized sandstone scarps (see below) in the Western Plateau, around what is now theLough Navar Forest Park, on subsequent days in the same week (Praeger 1904). His separate sites, whichhe described as, "Shean North, 1,135 feet [340 m]" and, "Half a mile [0.8 km] south ofShean North", are very difficult to locate on old or modern Ordnance Survey maps and in a mostunsatisfactory arrangement they are currently combined into an extant site for the species, the 'RadioMast Scarp' at grid ref. H066572.
Despite this difficulty, the current author (RSF) and RHN are able to identify a total of seven stationsfor P. media in Fermanagh, four of them with post-1975 records. All seven are situated on theWestern Plateau and they feature in a total of five tetrads, four with post-1975 records. Apart from thefirst record, all the sites are on the heathy scarps mentioned above or are associated with other namedmap features. The first five that follow are Praeger's sites from July 1904. The others were discoveredby RHN & HJN on the dates given: scarp S of Carricknagower Lough (refound 2 November 1995); E end ofMeenameen Lough; NE of Lough Anaban (refound 30 April 1995); at Shean North; half a mile [0.8 km] S ofShean North; scarp at Radio Mast, 7 August 1981 & 17 July 1982 (a solitary flower photographed); atDerryvahon Td, 500 m S of Sillees River, 9 March 1996, and monitored at the latter several times since.
The preferred ecological requirements of P. media of damp but drained, shaded, moderately acid,nutrient-poor conditions are very well met by the steep, heath-clad, north-facing, rock strewn slopesbelow the Lough Navar scarps, the rock of which is a weathered Old Red Sandstone in which mineralreplacement has been effected by percolating lime-rich water seeping down from overlying Carboniferousstrata.
Despite P. media possessing a slender, creeping rhizome, an organ enabling both energy storage andvegetative propagation, at its Fermanagh stations Intermediate Wintergreen is only present in extremelysmall plant numbers. As it grows beneath an evergreen heath canopy, P. media is made even moredifficult to locate by the fact that many of the individual leaf rosettes are very small, often bearingonly one or two leaves. Very rarely we have found individuals with up to ten fleshy evergreen leaves. Atthe four current Fermanagh stations we have never found more than 19 plant rosettes on any occasion andoften only six or fewer small vegetative plants are present.
As in other areas of B & I, P. media is extremely shy when it comes to flowering in theFermanagh sites' particular environmental conditions. We have seen the plant in flower only once (on 17July 1982), when it was photographed. The infrequency of flowering in Fermanagh obviously severelycurtails the production of the huge seed numbers essential to the success of the reproductive strategyof this species. We really must be observing a species declining towards inevitable local extinction.
Flowering reproduction
While flowering is extremely rare in the Fermanagh area at present, the flowering season of P.media is generally quoted as running from June to August, the leafless scape bearing betweenfour and twelve pendulous white hom*ogamous flowers. Pyrola flowers contain little or no nectarbut are said to attract bees, flies and beetles which collect and feed on plentiful pollen (Proctor& Yeo 1973; Fitter 1987). In some species, including P. minor, the stigma produces a stickyfluid that insect visitors lick. Sell & Murrell (2014) state that nectar is secreted by thebase of the petals in P. media. However, whatever the real situation is regarding an insect foodreward, several important authorities on pollination agree that insect visits to Pyrolaceae are rare(Knuth 1903-1909; Hagerup 1954; Knudsen & Olesen 1993) and wind-pollination, or more likely,selfing, must be a common feature since the majority of flowers do form fruit capsules and set seed(Salisbury 1942; Hagerup 1954).
A detailed study of insect pollination ecology in Pyrolaceae carried out in Denmark and Sweden includedthree species of Pyrola, plus Orthilia secunda (Serrated Wintergreen) (which does producenectar and scent) and Moneses uniflora (One-flowered Wintergreen) (Knudsen & Olesen 1993).These workers found that bumblebees were the principal visitors and although insect numbers were low inthe habitat of these species, the fact that two or more species of Pyrolaceae often grow together mayallow sharing of pollinators to the mutual benefit of the species. The anthers of Pyrola speciesrelease their pollen through restricted openings or pores and the bee visitors carry out'buzz-pollination'. This involves the bees gripping an anther and using rapid contractions of theirindirect flight muscles to vibrate or 'sonicate' them; they then harvest the pollen shower this processreleases (Knudsen & Olesen 1993).
Seed and seedling ecology
All species in the Pyrola family possess very small, light seeds which are comparable to those ofterrestrial orchids, but they have an even smaller central region containing the actual embryo(Salisbury 1942, p. 94). Members of the genera Pyrola and Orthilia may be regarded assemi-saprophytes, since although they possess evergreen leaves and are capable of photosynthesis, theyrequire damp, humus-rich habitats and can endure deep shade. The diminutive food reserves of the seed,allied to a near-obligate relationship with a mycorrhizal fungal partner for germination as is common inmost or all terrestrial orchids, reinforces the idea that the plants are probably best consideredsemi-saprophytic.
Measurements of fruit and seed production by Salisbury (1942) found that fruiting stems of P.media bore a range of between four and twelve capsules (mean 8.75), each containing an averageof c 1,400 seeds. Salisbury calculated mean seed production lay between 10,000 and 14,000 per plant.
If flowering was a regular seasonal occurrence in the majority of leaf rosettes, then P. mediacould survive in the long term perfectly well, since the small seed allows very efficient dispersal andthe mycorrhizal connection permits the tiny seed size without undue detriment to seedling establishment.It is obvious, however, that there is a smaller margin of safety against the risks of mortality in theearliest stages of development.
British and Irish occurrence
The New Atlas hectad map shows that in Britain P. media is very much more frequent in thehighlands of E Scotland than elsewhere. The species has declined and all but disappeared from Englandand Wales. In Ireland, it has likewise declined to near extinction in the Republic (one recent recordfrom Co Clare (H9), only) and it only appears to be maintaining itself in NI. Here, apart fromFermanagh, it is known from eleven hectad squares spread across three VCs (Cos Tyrone (H36), Antrim(H39) and Londonderry (H40)).
In view of the exceptionally shy flowering habit of P. media in our area, distinction of it fromP. minor (Common Wintergreen) might appear difficult. Even when they do flower, the two speciescan appear similar enough to be confused. Locally in the FNEI 3 and more generally in the NewAtlas, the editors recognise that these species have been mistaken for one another in the past.This makes it impossible to know for certain whether they are declining overall in the British Isles,although observers can be more certain of local changes in regularly visited sites (FNEI 3; F.J.Rumsey, in: Preston et al. 2002).
In Scotland and elsewhere in Britain, P. media is a rare member of the NVC H16 Callunavulgaris-Arctostaphylos uva-ursi heath community. It is especially concentrated within andcharacteristic of a herb-rich version of this vegetation, the Pyrola-media-Lathyrus montanussub-community (Rodwell et al. 1991b, p. 528). In Fermanagh, the heath vegetation where P.media occurs is much less herb-rich than Rodwell describes and in the total absence ofArctostaphylos the woody element is composed of some or all of Vaccinium myrtillus(Bilberry), V. vitis-idaea (Cowberry), Calluna vulgaris (Heather) (generally dominant) andErica cinerea (Bell Heather). On the other hand, here in Fermanagh, Intermediate Wintergreen isregularly accompanied by its very rare relative (at least in an Irish context), Orthilia secunda(Serrated Wintergreen) and very occasionally by the rare and inconspicuous orchid Listera cordata(Lesser Twayblade). Nowhere in the GB-based NVC do we find P. media consorting withOrthilia secunda and we may probably take it that this particular rare form of Irish vegetationhas yet to be investigated and described (Braun-Blanquet & Tuxen 1952; Ivimey-Cook & Proctor1966; White & Doyle 1982). It may perhaps be a transitional stage from calcicole to calcifugevegetation, due to prolonged leaching of the soil which is derived from a dolomitized sandstone.
European and world occurrence
In Europe, P. media is widespread in Northern boreal areas, extending S to the Maritime Alps and Eto Continental Russia and boreal Asia. The locations for it in the British Isles form the westernextremity of the core area of the species distribution and it is absent from immediately adjacentregions of the European continent including France (Hultén & Fries 1986, Map 1437).
Names
The name 'Pyrola' is the Latin diminutive of 'Pyrus', meaning or the name referring to the 'pear' treeand refers to the pear-like leaves of the species in the genus (Stearn 1992). The Latin specific epithet'media' simply translates as 'intermediate', that is, a reference to the scale of this species betweenP. rotundifolia (Round-leaved Wintergreen) and P. minor (Common Wintergreen). The plant istoo rare to have any English common names except the inevitable translated 'book name', 'IntermediateWintergreen'.
The English common name 'Wintergreen' itself was first given by Turner (1548) in his The Names ofHerbes, a straight translation of the German 'winter-grün', taken from the Ortussanitatis or the Hortus sanitatis, also called the German Herbarius or perhapsbest amongst further alternatives, just simply Cube's herbal, a medical text only partly reworkedfrom earlier material by Dr Johann von Cube of Frankfort, dating from around 1485 or 6 (see Arber 1938,p. 22). Prior (1879) points out that in Danish 'winter-grönt' refers to the ivy, Hedera helix andhe suggests that this is the rightful claimant of the name, being so conspicuously green when themajority of trees are bare of leaves.
The aromatic oil rub called 'Wintergreen' that is in everyday medicinal use by games’ coaches on theathletic field and which magically enables games’ players to get back on their feet, just moments afterapparently life-threatening muscular sprain, was originally distilled from the low, creeping Americanericaceous shrub, Gaultheria procumbens (Checkerberry). It has nothing whatever to do withthe members of the Pyrolaceae. The original G. procumbens plant oil can cause severe skineruptions and a synthetic replacement (methyl salicylate) is used nowadays to avoid the problem. Jokingaside, Wintergreen really does relieve rheumatic, joint and muscle pains (Grieve 1931, p. 849).
Threats
Very few and very small, isolated populations vulnerable to any of type habitat change, especially thoseassociated with forestry operations and heathland fires.
Native, very rare and conservation protected under Schedule8 of the Wildlife (NI) Order 1985.Circumpolar boreal-montane.
26 June 1901; Tetley, W.N.; N-facing sandstone scarp at the SE end of Correl Glen.
Throughout the year.
Growth form and history in Ireland
This is an extremely rare, evergreen, rosette-forming, creeping rhizomatous perennial in Ireland and ithas been so for many years. It became extinct at its previous Co Offaly (H18) bogland site (destroyed bypeat cutting in the 1950s), and has not been seen at a previous site in Co Londonderry (H40) since 1888.There were five stations in Co Antrim (H39) with dates in the early 19th and early 20th centuries and ithad not been seen at any of them since 1920 until RHN rediscovered it on the bank of the Cranny Water in1996 (Irish Red Data Book; Northridge & Northridge 1997).
Fermanagh occurrences
In Fermanagh, O. secunda sometimes occurs in dense clonal patches and considerable quantity, increvices and on ledges on dolomitized sandstone scarps and scattered in steep, submontane, mossy,Calluna-Vaccinium-dominated heathy slopes below these cliffs on the Western Plateau (ie inthe Lough Navar Forest Park area). When one has got one's eye in for it, the rather pale, grey-green,wintergreen, serrated leaves can be picked out amongst other foliage and moss all year round. All 59Fermanagh records are concentrated within one 10-km square (H05) and they can be mapped as nine tetrads,16 1-km squares or 31 100 m squares!
The plant still occurs in Tetley's original 1901 site on drier sandstone ledges on the lower half of thesandstone cliffs at the bottom end of the Correl Glen NR (Praeger 1901b). This is the only spot inFermanagh where the species grows under trees (ie in upland mixed deciduous woodland with oak andbirch), rather than under subshrubs, Calluna vulgaris (Heather) or Vacciniummyrtillus (Bilberry) (Praeger 1903c). Since the whole area where O. secunda occurs is onthe Western Plateau, it should be understood that what Praeger described as, "drier ledges in theCorrel Glen sandstone scarps", still represents very damp, humid conditions – given the prevailinghigh rainfall figures in the locality and the sheltered, wooded, gully-like nature of the lower (ie thesouthern) end of this deep wooded glen.
Flowering reproduction
Serrated Wintergreen flowers very sparingly in Fermanagh, somewhat less than 10% of rosettes bearing aninflorescence. On the evidence of Praeger's descriptions of it when it was first discovered by Tetley,West and himself (Praeger 1903b, 1904), RHN and the current author (RSF) reckon there has been a markeddecline in flower and rosette frequency during the last century. The account of O. secunda by DrD.A. Ratcliffe in Scarce Plants in Britain (Stewart et al. 1994), very closely matches theFermanagh situation described here, both in terms of habitats occupied (apart from rocky stream banks)and marked species decline in abundance and flowering capacity.
The characteristic secund (ie one-sided) raceme inflorescence develops in July and August and consists ofbetween six and twelve cream or whitish-green, bell- or funnel-shaped flowers, each of which persistsfor up to a week. The flowers do not have any scent (at least none perceivable to the human nose!), butthey attempt to attract insect visitors by providing food in the form of both nectar and relativelylarge amounts of pollen.
In a study of flower biology carried out in Denmark, Knudsen & Olesen (1993) found that O.secunda attracted bumblebees both as nectar-collectors and to a lesser extent as pollen-buzzers.The anthers of Pyrola and Orthilia species release their pollen through restrictedopenings or pores and the bee visitors carry out 'buzz-pollination'. This involves the bees gripping ananther and using rapid contractions of their indirect flight muscles to vibrate or 'sonicate' them andthey then harvest the pollen shower this process releases (Knudsen & Olesen 1993)
The flowers are strongly protandrous, the anthers opening a few days before the flower fully opens, thestigma only becoming receptive one or two days after anthesis. The Danish study found that in untreatedbagged flowers (ie autodeposition of pollen) only 0.5% of their ovules set seed, while underuncontrolled unbagged conditions (ie open-pollination) 84% of ovules set seed. Unfortunately, however,the experiment did not investigate self-compatibility, since the ability to self-fertilize is probablyimportant in O. secunda as is the case in many of its relatives. In addition, general observationindicates that insect visitors are infrequent to rare in the damp, shaded, upland habitats in whichO. secunda grows (Knudsen & Olesen 1993).
Seed production
About mid-September the ripe fruit capsules open to release the numerous tiny seeds (Helenurm &Barrett 1987). Measurements made in Britain by Salisbury (1942, p. 95) found that O. secunda seedproduction was of the order of 3,500 (± 500) per inflorescence. Since the species normally has abranched rhizome, however, each genome bears several inflorescences. In species of the closely relatedgenus Pyrola, the branched rhizome is more fragile than in O. secunda and the undergroundconnections between the leaf rosettes usually do not persist, so that separate individual ramets arise.This does not happen in O. secunda. O. secunda seed is even smaller than that of Britishorchids and it is light enough to be carried aloft by the slightest breeze.
Little or nothing appears to have been published on the reproductive ecology of O. secunda and thecurrent writer could uncover absolutely no information on seed germination requirements, season orseedling establishment and nothing on the competitive ability of the species − all clearly important inreaching an understanding of the ecological future of the species in B & I. There are two referencesregarding seed survival in the soil seed bank, both Russian and indicating that it is only transient(Thompson et al. 1997).
Seed germination and lack of establishment
A mycorrhizal fungal partner is required for successful germination and establishment, but observationsof the behaviour of the species in the last century in both Scotland and Ireland, strongly indicatesthat little or no colonisation of fresh sites by seed is taking place (D.A. Ratcliffe, in: Stewart etal. 1994). However, O. secunda can reproduce vegetatively and it has managed to survive,although certainly not in the same abundance as Praeger described, at or very close to all but one ofits early 20th century Fermanagh stations. The exception is Praeger's 1904 site on the cliffs E ofGlencreawan Lough (Praeger 1904) described in IN 13: p. 239, as, "North of the pointmarked 1,033 (ft) [315 m] on OS map (abundant)".
Variation
A recent study of Fermanagh O. secunda populations found there was very little geneticvariation present in any of them. All the patches examined, including one with 140 ramets, provedmonoclonal, ie they consist of a single genome. Reproduction involving extreme levels of clonalbehaviour and a lack of flowering and fruiting probably represents a means of conserving energy in anunfavourable environment. The fragmented population stands of O. secunda in Fermanagh are nearthe southern margin of the species distribution in B & I, which is centred in the ScottishHighlands. As such these Irish stands represent relict populations left over from cooler periods inEarth's history. Thus range-edge effects, the availability of suitable habitat, plus grazing pressurefrom sheep and goats may well be factors determining the strong predominance of clonal growth observed.Under Irish temperate conditions, populations are found on exposed scarps, or less commonly in deciduouswoodlands, rather than in the coniferous forests that represent the optimum boreal habitat of O.secunda. Thus what appears to be a sizeable population of Serrated Wintergreen may consist ofonly one or a very few genomes. Therefore the species lacks genetic diversity and is very much rarerthan it looks (Beatty et al. 2008).
British and Irish occurrence
By far the greatest presence of O. secunda in B & I is in the Scottish Highlands and northernregions of Britain. Further south in Britain, and especially in Wales and Ireland, it appears to be arare or very rare and declining relict species, although it is not clear if it has declined furthersince the 1970s (D.A. Ratcliffe, in: Stewart et al. 1994; F.J. Rumsey, in: Preston et al.2002).
European and world occurrence
The overall distribution of O. secunda (taking the taxon in the wider sense, since Hultén (1971),Tutin et al. (1972) and Hultén & Fries (1986) recognise two subspecies), is a circumpolarboreal montane species, characteristic of and widespread within boreal forests and upland heaths aroundthe N Hemisphere. It occurs throughout most of Europe and appears in a very much wider range of woodlandcommunities than it does in B & I (Ellenberg 1988), but is rare in the Mediterranean region andquickly becomes scarce to rare south of the boreal zone and in lowland areas (Hultén 1971, Map 129;Hultén & Fries 1986, Map 1441). Serrated Wintergreen is widespread and frequent to abundant insubalpine heaths and coniferous woodlands at all levels throughout Switzerland where, indeed, it is themost widespread and frequently met member of the Pyrolaceae (Welten & Sutter 1982, 1, Map1210).
Names
The genus name 'Orthilia' is of rather obscure origin, but the first part of the name comes fromthe Greek 'ortho' meaning 'straight' and probably refers to the female style. The Latin specific epithet'secunda' in Classical language means, 'following', but in a botanical sense it means, 'arrangedon one side only' (Gilbert-Carter 1964). The plant is too rare to have accumulated much in the way ofEnglish common names, apart from the obvious 'book names' or translations, for example, 'SecundWintergreen' and 'Serrated Wintergreen'. The one exception is 'Yevering Bells' (Prior 1879), or'Yavering Bells' (McClintock & Fitter 1956; Watts 2000), which the former helpfully explained is socalled, "from a lofty conical mountain near Kirk Newton in Northumberland, where the plant wasfound growing wild". However, it turns out that all these gentlemen were mistaken, taking thegeographical name of the site to be a local English common name. The correct name of the site is'Yeavering Bell', the solitary station for the plant in VC 68, where it was discovered in1834 and survives to the present day (Swan 1993).
Conservation status
Orthilia secunda is a protected species in Northern Ireland under Schedule 8 of the 1985 WildlifeOrder by the Environment and Heritage Service.
Worth looking for additional sites
Since such a high proportion of Serrated Wintergreen plants exist in a purely vegetative state and theyoften hide underneath sub-shrubs such as Vaccinium spp., it is possible the species might yet bediscovered in other suitably damp, reasonably undisturbed heathland sites elsewhere in the north ofIreland. It is likely that heath and heather moorland fires in the past have destroyed other populationsthroughout both B & I. Such losses have drastically restricted the distribution of this lovelylittle plant, which now, under the further threat of global climate change, really teeters on the brinkof extinction in Ireland.
Threats
Forestry operations, including fire and fertilizer spraying.
MONOTROPACEAE – Bird's-nest family
Native, very rare, conservation protected under Schedule8 of the Wildlife (NI) Order 1985.Circumpolar temperate.
July 1904; Carrothers, N.; Ely Lodge Forest, Lower Lough Erne.
July to October.
Growth form and preferred habitats
This small, pale yellow (sometimes tinged with red or brown), waxy-looking perennial has a few-floweredraceme, or rarely a solitary flower that droops at first but which later is held erect in fruit (Olson1990). It appears amongst leaf litter, generally but not always in deep woodland shade between July andOctober. The plant contains no chlorophyll and until recently was considered saprophytic (derivingnutrients from decaying leaf litter by means of mycorrhizal roots and fungal partners). Recent researchhas shown it is actually epiparasitic, using Tricholoma fungi to extract nutrients from the rootsof trees living in its vicinity (Leake et al. 2004).
In Fermanagh, where the species is at the extreme edge of its European range, it is typically associatedwith hazel or beech in woods or scrub growing on shallow limestone soils. It also occurs in the mixedoak-birch woodland of the Correl Glen NR where the geology is sandstone (although some of it is lessacidic than normal it having been dolomitized). In other parts of its B & I distribution, M.hypopitys more usually grows on more acidic soils under oak or pine. In damp coastaldune-slacks, it is occasionally found growing under Salix repens (Creeping Willow) (F.J. Rumsey,in: Preston et al. 2002).
Variation
M. hypopitys occurs as two subspecies in B & I and the 1957 Flora authors stated that allplants examined by them belonged to subsp. hypophegea, rather than subsp. hypopitys(Meikle et al. 1957), the occurrence of which is very scattered in England and unknown elsewhere(Stace 1997; Sell & Murrell 2014). Subsp. hypophegea produces fewer flowers (ie eight orless), they are less hairy inside and the style is shorter than in subsp. hypopitys (eleven orfewer flowers) (Stace 1997).
Fermanagh occurrence
Overall, M. hypopitys is currently known from nine tetrads in Fermanagh and, as the distributionmap shows, seven of them are scattered along the Lough Erne basin, three of them on the Castle Caldwellpeninsula in the NW corner of Lower Lough Erne. M. hypopitys has been known from the Ely Lodgesouth shore of Lower Lough Erne since 1904, but two stations nearby have been destroyed in recent years;Praeger and Carrother's 1945 site, "opposite the gate of Ely Lodge" (BEL voucher sheet)was planted with conifers, and the 1980 site of c 120 plants beside Castlehume Lough wasdestroyed by the development of a golf course around 1991. Two other moderately stable stands of up to adozen spikes survive in the Ely Lodge lakeshore lowlands, in mainly coniferous plantation. Happily, twoadditional Fermanagh sites have been more recently discovered: the first on the limestone Knockninnyhill overlooking Upper Lough Erne in 1995 and the second in the Correl Glen NR in 2003. Both are quitedistant from the Lower Lough Erne stations and the latter is much more upland.
The number of aerial flowering shoots in stations of the plant is typically very variable from season toseason (Söyrinki 1985; Olson 1990) and the population extent is thought by some to correlate withmaximal and minimal annual rainfall. However, while observed numbers are distinctly variable it shouldbe remembered that the aerial shoots are often small and retiring, variable in the timing of theirappearance and quite difficult to find and, therefore, easily overlooked. The edge-of-path locations ofmany of our discovered plants may simply reflect the rather inconspicuous, late-season appearance of theplant and the fact that even when we are exploring a woodland area for plants, we tend to followpre-existing tracks or paths.
Despite the welcome addition of the two new sites to the Fermanagh listing for the species, CastleCaldwell on the shores of Lower Lough Erne still seems to be the M. hypopitys stronghold in theVC. This always inconspicuous plant has been independently found there by three people in the lastdecade, a fact which encourages RHN and the current author to believe it is probably quite widespread inthat particular estate.
Flowering reproduction
The inflorescence is nodding at first but becomes erect later in development. It is a raceme of usually3–8 flowers, the terminal one with parts in fives and the remainder with parts in fours. Above the ovaryis a single, short, stout style expanding into a bright yellow, disk-shaped stigma. The flowers arefragrant and produce nectar from swellings at the base of the five-lobed ovary. The petals are expandedand saccate at their bases to hold the nectar. An insect visitor must push its proboscis (oftenerroneously referred to as its 'tongue') down between the edge of the stigma and the 8-10 anthers toreach the food reward, and thus pollen can be transferred between flowers.
Self-pollination is frequent, however, and close examination by median microtome sectioning showed veryclearly that the flower has contact autogamy (Hagerup 1954). The cavity of the flower is narrow and thepetals press the long stamens tightly against the thick ovary. The anthers are pressed inwards againstthe broad stigma, whose large mucilage plug retains a quantity of the pollen, with the result that thecanal of the style is soon densely filled with pollen tubes (Hagerup 1954).
A recent study of genetic variation in the Fermanagh populations at Castle Caldwell, Ely Lodge andKnockninny found they were fertile and reproducing sexually, although the plants are self-compatible andinbreeding is rife. In contrast with the closely related boreal species Orthilia secunda(Serrated Wintergreen), surprisingly low levels of clonal patch development were found in Fermanaghpopulations of M. hypopitys, confirming that the predominant mode of reproduction is seedproduction. It is likely that the high levels of inbreeding observed are the result of self-pollination,particularly given the small numbers of genetic individuals in most of patches of the plant. Beingself-compatible means, however, that M. hypopitys does not face the same problems of completeloss of sexual reproduction and/or rapid population extinction that can threaten populations ofobligately outcrossing clonal plants (Beatty & Povan 2011).
Seed dispersal
In view of the restricted, and, in Fermanagh and in Ireland in general, sporadic, isolated nature of thepopulations of this rare species, it is puzzling to contemplate just how M. hypopitys achievesdispersal and establishment. The globular fruit capsule opens by means of five vertical, slit-likevalves to release the numerous slender, white, extremely lightweight seeds it contains (Butcher 1961,2, p. 117). The contents of two capsules counted by Salisbury (1942, p. 94) contained 1,953 and1,980 seeds and since the number of flowers per raceme ranges from one to twelve, he reckoned that themean seed output would be around 16,000 per shoot. The mean seed weight is just 0.000003 g, very similarto those of native terrestrial orchids and we may assume, therefore, that wind provides the chief methodof seed dispersal.
However, seeds or roots might also be transferred inadvertently by man in soil, a circ*mstance mostreadily envisaged when imported tree material is being planted. Oddly enough, since the two most recentfinds in NI were both in already established nature reserves where tree planting is not a feature, itmay be necessary when searching for a vector to consider transfer on the muddy boots or clothing ofhabitual nature reserve visitors, a suggestion supported by the edge of path locations of many (but notall) M. hypopitys plants!
Germination and seedling establishment
The embryo is tiny, consisting of only three cells, and the nutritive endosperm is also reduced to justnine cells, so that even with the assistance of its mycorrhizal fungus, the odds against successfulgermination and establishment must be absolutely enormous (H.L. Francke (1935) quoted in Salisbury(1942)). The huge number of very small, biologically reduced seeds is undoubtedly responsible for thelow reproductive capacity in terms of seed germination and seedling viability found in M.hypopitys. However, since these biological properties are also the case with saprophytic orchidspecies, this reproductive strategy obviously gives greater assurance of efficient dispersa and thusfacilitates colonisation of very scattered, specialised sites suitable for growth and reproduction ofthese species.
Irish occurrence
In the rest of NI, M. hypopitys is known from only two other sites. One was discovered in 1997 atStraidkilly NR near Glenarm, Co Antrim (H39) and the other was in Co Londonderry (H40), although it hasnot been seen since 1944 and presumably has become extinct. In the RoI, Yellow Bird's-nest has been seensince 1970 at just two sites in Co Galway (H15, H16), two in Co Donegal (H34, H35) and one each in CosClare (H9), Wexford (H12) and Leitrim (H29) (New Atlas). If this is the true situation, itrepresents a long continuing decline of this always rare species in the RoI since before the turn of the19th century (Praeger 1901, p. 209). However, the authors of the Irish Red Data Book believedM. hypopitys is, "clearly under-recorded" due to its inconspicuous nature (Curtis &McGough 1988). In its Wexford site, Yellow Bird's-Nest was very abundant in a pine plantation and Curtis& McGough (1988) suggest that it was very probably introduced along with the tree material.
Conservation status in NI
On account of its rarity and vulnerability, M. hypopitys is listed for special conservationprotection on Schedule 8 Part 1 of the Wildlife (NI) Order, 1985. For wider based scientifically-drawnbiodiversity aims, it is also included on the NIEA list of Priority Species of special concern requiringlocal conservation action.
British occurrence
In Britain, Yellow Bird's-nest is prevalent in south central England and Anglesey, being rather thinlyscattered south of a line between Preston and Hull and very rare northwards to the Scottish Highlands.Even within the southern England area of most records, many sites have disappeared both before 1930 andduring the last 50 years, presumably due in the main to woodland disturbance and habitat destruction(F.J. Rumsey, in: Preston et al. 2002).
European and world occurrence
M. hypopitys s.l. (consisting of a total of four subspecies) is widely distributed in temperateregions of the Northern Hemisphere across Europe, Asia and North America, although throughout its rangeit is generally found scattered at low abundance (Hultén & Fries 1986, Map 1444; Beatty & Provan2011).
Threats
Woodland clearance through ignorance of the species' rarity and interest.
PRIMULACEAE – Primrose family
Native, common. European temperate.
1858; Brenan, Rev S.A.; by Ardunshin Bridge on the Colebrooke River.
Throughout the year.
Growth form and preferred habitats
Primrose is an extremely common and widespread rhizomatous, rosette-forming perennial throughout B &I, typically frequenting woods, scrub, hedgerows, grassy wayside banks and damp moderately acid toalkaline, preferably lime-rich pastures. It is, however, principally absent from four types of soil andhabitats: regularly flooded ground; deep, strongly acid peat; shallow, dry ranker soils formed over hardcrystalline limestone; and very light strongly acidic soils, particularly where these are likely to dryout in summer (Richards 1989, 1993).
Under the often cloudy, overcast skies of the most western regions of B & I, shade-tolerant and hotsun avoiding perennials like Primrose and Hyacinthoides non-scripta (Bluebell) are not confinedto woodland canopy or to north-facing cliffs and slopes as they generally are in drier parts of SEEngland with a more continental-style of climate. Rather here, in the western edge of the British Isles,these spring-active vernal herbs frequent mesic pastures, hedge banks, wayside embankments and vergesthat offer similar moderate levels of soil moisture, fertility and reaction, but which are completelyunshaded (Whale 1984). In well-lit yet humid grasslands of these types, the wrinkled, evergreen leafrosettes of P. vulgaris and their familiar luminous, buttery yellow, long-lasting flowers areespecially plentiful and conspicuous on the steeper grassy banks, quite irrespective of their aspect.The ecological strategy of established plants was summarised and described as S/CSR by Grime etal. (1988), meaning P. vulgaris has life strategy characteristics that lie intermediatebetween a stress tolerator and a balance of competitor, stress-tolerator and ruderal species.
Fermanagh occurrence
This familiar perennial is very common and widespread throughout Fermanagh in all kinds of damp grasslandand in woods, scrub and other shaded habitats. It is the 19th most frequently recorded vascular plant inthe county and has been recorded in 456 tetrads, 86.4% of those in the VC. While almost ubiquitous,P. vulgaris is most noticeably absent from the blanket bog area to the north of Upper LoughMacnean and around Little Dog and Big Dog, where deeper peat and extensive conifer plantations exist.
Herbivory and possible toxicity
On steep sites, Primroses encounter reduced grazing pressure from both stock animals and smallerherbivores, especially rabbits, the animals being discouraged by gradient and gravity. The currentauthor (RSF) has observed that horses leave perfectly accessible Primrose clumps and their leaves andflowers completely alone, even when the accompanying vegetation has become very well-grazed, indeedalmost entirely depleted. In coastal Wales, Knight (1998b) noticed that rabbits consumed only theflowers and appeared to find the leaves unpalatable. Slugs and snails are also fond of Primrose,nibbling leaves and flowers, often to destruction.
Flowering reproduction
The large, tubular, pale yellow flowers have their parts in fives and are usually borne on separate,long, hairy stalks (pedicels) arising from the base of the plant. Flowers rarely appear either white orreddish-pink. A rare variant that bears an umbel of flowers on long, hairy pedicels attached to a scape(leafless stem), called forma caulescens (W.D.J. Koch) Schinz & Thell, sometimes occurssingly amongst normal forma vulgaris flowering plants (Sell & Murrell 2014).
Plants usually flower from early March into May or early June and are well known to be of two types inroughly equal numbers, 'pin' (long style) and 'thrum' (short style), clearly differing in the length ofthe female style and the relative positions of the anthers and stigma. This heteromorphism (orheterostyly) is an apparently simple, but actually rather complex, genetic mechanism designed to achievea high degree of cross-pollination by means of an insect visitor inserting its elongated proboscismouthparts (often erroneously referred to as its 'tongue') into the tubular blossoms to reach for nectarsecreted at the base of the ovary (Richards 1989). Pin flowers can also occasionally self-pollinate, butthrum flowers never do, the thrum pollen tube failing to penetrate its own style (Richards 1989, 1993).In either case, seed is rarely set in the absence of cross-pollination (Richards 1989).
Many flowers contain sheltering thrips and small beetles, but these rarely carry pollen between flowers.Pollen transfer is carried out by insect visitors with long proboscis mouthparts, such as hive bees,bumblebees, butterflies and moths. Seed is rarely set without cross-pollination. All the flowers on asingle plant are of the same type, the heterostyly character being under genetic control (Richards1989). Experimental emasculation experiments that removed the anthers while still in the bud proved thatheterostyly does promote pollen travel between pins and thrums, so that more pin pollen is found onthrum flower stigmas and thrum pollen on pin stigmas, than would be expected by randomcross-pollination. Thus Charles Darwin was probably correct that the reciprocal positioning of theanthers and stigma in pin and thrum flowers helps pollen to travel legitimately to the stigmas of theother flower type, where it can successfully carry out cross-fertilisation (Richards 1989).
Seed dispersal
The very commonness of P. vulgaris and its widespread distribution in B & I is rather odd whenone recognises that the seedling is slow-growing, making establishment more hazardous, and the speciesis low-growing, rendering seed dispersal by wind difficult or unlikely. Ripe seeds are dark brown,irregularly shaped and about 1.5 mm in diameter. There is no specialized release mechanism: they simplydrop out of the weak-stemmed capsule which typically flops to the ground in summer, only reaching a fewcentimetres from the parent leaf rosette (Knight 1960). However, the resultant small pile of seedsattracts ants, since the seed-coat has an attached nutritive elaiosome oil-body, rich in fatty acids andsimilar in type to that of Viola odorata (Sweet Violet). Ants avidly collect and transport theseed and its sticky attached reward towards their nests. Many Primrose seeds will no doubt end up buriedin ant nests, but some are simply dropped en route and others have their oil-body bitten off and arediscarded outside the nest, some distance from their site of origin (Ridley 1930, pp. 520-1).
Study on the activity of European ants by Sernander (1906) quoted by Ridley (1930) indicated seeds ofvarious types being transported distances from 15 to 70 m and although this did not specifically involvePrimrose seeds, it gives some idea of the possible effectiveness of the mechanism. On the other hand,possession of a nutritive seed appendage can increase their depredation and destruction. Crosby (1966)found that seed capsules of P. vulgaris were heavily predated by mice, voles and slugs. Seedlingsof the plant are also sometimes found outside the burrows of voles and mice and these mammals mayinadvertently assist Primrose dispersal by attempting to store surplus capsules for their own furtheruse (Richards 1989). It would be interesting to know if any further work on this rather involved topichas been completed, since it has implications for other common woodland species including Violaspp.
Despite the observation quoted above that horses avoided grazing P. vulgaris, viable seedhas been recovered from the faeces of both horses and cows, indicating that at least the fruit capsulesare grazed and seed is internally transported (Ridley 1930, pp. 360-1).
The dispersal pattern of P. vulgaris appears to differ from other British Primula species,plants occurring either as relatively distant individuals or clumped in clones, rather than morerandomly distributed. Richards (1989) commented that the sites to which ants and rodents carry the seedsmay prove relatively favourable for seedling establishment.
Regeneration from seed
The regeneration strategy of P. vulgaris involves the survival of buried seed, either short-term(1 to 5 years), or long-term (longer than 5 years) (Thompson et al. 1997) and their eventualgermination and establishment after some form of disturbance bringing them back to the soil surface inan ecologically suitable environment. Seedlings are seldom found in sites occupied by other plants orwhere there is an accumulation of leaf litter. Most frequently they are found on ridges, sloping banks,disturbed ground and other bare earth sites, usually close to the parent plant (Knight 1960).
Weakening effect of drought
P. vulgaris is so very common and widespread in these islands, it is really absent only from peatyand/or heavily waterlogged situations and from very light, strongly acidic soils, particularly wherethese are likely to dry up in summer (Richards 1989, 1993). The latter observation is all the moresurprising since the plant does possess a sturdy, often near-vertical rootstock, which might beconsidered an adaptation allowing the species considerable drought resistance. It is a commonobservation that even severely wilted leaf rosettes whenever they are rewetted generally reviveremarkably quickly and, to the naked eye, they appear completely undamaged. Obviously drought survivaldepends upon the degree and duration of water depletion and any amount of wilting may physiologicallyweaken the individual plant, leaving it more vulnerable to competition from neighbours, less able torespond to depredations by vertebrate and invertebrate herbivores and, most likely of all, susceptibleto attacks by disease organisms. All Primula species are vulnerable to an aphid-dispersedCucumber mosaic virus, to a Botrytis root rot and, as gardeners know to their cost, slugs canalso do devastating damage to both leaves and flowers overnight.
British and Irish occurrence
P. vulgaris is widespread throughout B & I, still locally abundant in many parts, but innumerous other areas it has declined, becoming local and scarce. The reasons for this are not welldocumented, but in SE England the decline may be due to a sequence of dry summers (Rackham 1999) and, inall urban areas throughout the land, decline may be due to centuries of collecting for horticulture(Richards 1993).
Conservation
In Britain, Primrose is protected by a law passed in 1975 and in NI protection was given under theWildlife (NI) Order 1985, in being listed on Schedule 8 Part 2 which allows the flowers to be picked,but the plant must not be uprooted.
European and world occurrence
P. vulgaris is a polymorphic species with three subspecies in Europe (subsp. balearica(flowers white and very fragrant, an endemic, confined to the mountains of Mallorca), subsp.vulgaris and subsp. sibthorpii (flowers usually red or purple, confined to the E part ofthe Balkan peninsula, Krym (in the Black Sea), Turkey, Iran and locally naturalised in C Europe)(Flora Europaea 3, Tutin et al. 1972)). Richards (1993) recognises a fourthsubspecies, subsp. heterochroma (Stapf) Smith & Forrest loc. cit., which has flowers violet,purple, red, pink, white or yellow and occurs on the S shore of the Caspian Sea (Elburz Mountains), Iranand Azerbaijan, in a disjunct area.
The plant that occurs in B & I is subsp. vulgaris, which belongs to the Europeantemperate element and occurs in W, SC & S Europe, extending to Denmark and N Ukraine. It alsostretches to adjacent parts of N Africa and Asia (Hultén & Fries 1986, Map 1465).
Hybrids
P. vulgaris forms hybrids with both P. veris (Cowslip) and P. elatior (Oxlip) in thewild in Britain but, as the latter does not occur in Ireland, only the cross with Cowslip is relevanthere and it is very rare in Co Fermanagh (see the separate account).
Cultivated forms
Unlike its relatives, the primrose has given rise to a vast range of cultivated variants quite apart fromits hybrids (Richards 1993). Cultivated varieties of P. vulgaris include mutant forms withdouble, semi-double, 'hose-in-hose' (petaloid calyx), 'Jack in the green' (leaf-like calyx) and numerouscolour breaks such as gold- and silver-laced petals and a wide range of colours. (Richards 1993;Griffiths 1994). Old plants are susceptible to virus, root-aphis and weevils and clones can therefore bedifficult to keep in the garden setting. Plants are most successful when they seed around andsuccessfully self-sow.
Medicinal use and folklore
Since P. vulgaris is a lot more common and widespread in comparison with P. veris, Primrosehas been put to much more use in traditional herbal medicine than Cowslip. In England, Primrose leaveshave been made into an ointment to heal cuts, bruises and chapped hands or chilblains, or combined withbramble tops to clear up spots and sores on the face. The ointment was made with pork lard and primroseleaves, or else the rootstock was used with beef or mutton suet (Vickery 1995). The ointment has alsobeen smeared on ringworm and the leaves have also been applied (presumably as a poultice) to treat boils(Allen & Hatfield 2004). Primrose was also regarded as a suitable cure for yellow jaundice, theroots being boiled in water and a wine glass of the resultant fluid drunk each morning (Vickery 1995).The plant is also regarded as a relaxant and sedative in some parts of these islands, being used as atea for a wide range of complaints including muscular rheumatism, paralysis, gout, insomnia, nervoushysteria and headaches. The whole plant has also been used as an expectorant (Grieve 1931).
In folklore, Primroses were often associated with poultry keeping, less than 13 flowers brought into ahouse on the first occasion of doing so in the year, being unlucky and limiting the fertility and egghatching ability of hens and geese for the season to the number of flowers collected. Thirteen was thetraditional number of a clutch of eggs placed under a 'clucking' hen during the spring to hatch, eachyellow primrose therefore being the analogue of a young chick that would eventually emerge from the egg(Vickery 1995). A similar belief on farms in Ireland was that throwing a primrose into the byre door onMay Day prevented the fairies from taking away the milk from the cows for the year (Vickery 1995).
Names
The Latin genus name 'Primula' is the feminine of 'primulus' the diminutive of 'primus', meaning 'first'and refers to the idea that this is the first (or one of the first) flowers of spring (Gilbert-Carter1964). The English common name 'Primrose' refers to the first (prime) rose of spring, again referring tothe early flowering of the species.
Threats
Habitat destruction and collecting for horticulture.
Very rare.
1954; MCM & D; pasture near Roosky Turlough.
April and May.
Growth form, habitats and biology
Primrose (P. vulgaris) and Cowslip (P. veris) usually prefer rather different habitats fromone another, but they can occur together in the wild along woodland paths and edges, in clearings andscrub and in hedgerows, 'unimproved' meadows and on roadside verges where the more open ecologicalconditions exclude neither species. This perennial, rosette-forming Polyanthus is one of the mostcommon of all wild hybrids in B & I and is a very popular garden plant with a wide variety ofselected forms. It can arise fairly readily anywhere in the wild that primrose and cowslip occurtogether.
Experiments show that only the cowslip can act as the seed parent (ie female), the cross providing goodseed with up to 37.5% germination. In comparison, when P. vulgaris is the seed parent, onlynon-viable seed are produced (Valentine 1955). As the cowslip is the more ecologically restricted andless frequent of the two species, the hybrid is rarer than might otherwise be the case and usuallyrepresents only 1-2% of any mixed population (Woodell 1965). P. vulgaris flowers much earlierthan P. veris, shortening the duration of flowering overlap to some extent. However, both speciesattract the same range of pollinating insect visitors, despite the flowers emitting distinctly differentscents. The hybrid is partially fertile, but markedly less so than both its parents and seed germinationis only around 20% (Stace et al. 2015).
In addition to the native F1 hybrid, garden Polyanthus, in a range of flower colours,do occasionally occur as escapes in the wild and have been known to backcross with both their parents.The latter is a rare occurrence and the vast majority of P. ×polyantha records in B & I are of F1 crosses between native populations of thetwo species (Stace et al. 2015).
The F1 hybrid resembles a primrose, but has at least some flowers held at the top of a commonstalk like those of the cowslip (Richards 1989). It can easily be distinguished from the rare and localtrue Oxlip (Primula elatior), which is ± confined to East Anglia, by it having flowers whichpoint in all directions and lacking crisped hairs (Richards 1989). Hybrids are vigorous and regularlyform large, long-lived clumps which often are bigger than those of either parent. The hybrid isintermediate in most characters and, although it is fully fertile, back-crossing is rare and only inunusual circ*mstances of close proximity is there any evidence of introgression (measured in terms ofpollen fertility), even when the ecological barriers that normally separate the species are lowered.Thus the two species remain almost entirely distinct as the internal isolating barriers preventingbackcrossing and subsequent introgression are strong (Woodell 1965; Richards 1989).
Fermanagh occurrence
This hybrid has been persistent in the Roosky area for over 40 years. It seems to almost always occurwhere both parents have been established for a considerable number of years. However, on the newroadside embankment of the Enniskillen-Belfast road near Tamlaght, RHN and the current author (RSF)believe less than 15 years elapsed before both parents and their hybrid were discovered. Locally it isfound in meadows, scrub and roadside embankments, almost entirely confined to the limestone area SW ofLower Lough Erne.
In Fermanagh, P. × polyantha is present in six lowland tetrads. Thefull record details are: in rocky pasture between Monea and Drumcose, near the Green Loughs (ie RooskyTurlough), sparingly with both parents, 1954, MCM & D; roadside embankment, 2 km W of Tamlaght, May1976, RHN & HJN; Caldrum Glebe Td, Caldrum Hill, N of Derrygonnelly, 31 May 1986, I. Rippey; meadowat Beagh Big Td, 25 April 1987, I. Rippey; four plants by Main Drive, Castle Coole estate, Enniskillen,3 May 1992, RHN & HJN; scrub at Roosky Turlough, 3 May 1993, RHN; Tullynagowan/Beagh Big ASSImeadow, 18 April 1995, RHN & HJN (also 12 April 1997, RHN); twelve plants at knoll, 100 m SE ofconifer plantation at Roosky, 12 May 1995, RHN, HJN & B. Morwood; small population by main drive,Castle Coole, 26 April 2005, RHN; persisting here 2010; one plant by track S of Crom Castle near deerenclosure, 29 April 2005, RHN.
In view of the rarity of P. veris (Cowslip) in NI, it is unsurprising that this hybrid is alsoextremely rare. Apart from the three hectad squares plotted in Fermanagh, the New Atlas mapdisplays just one further square in NI, in Co Armagh (H37). As with P. veris, the hybrid is verymuch more frequent and widespread in the RoI, than in NI, as is also the case in England and Wales,although becoming rarer and more coastal northwards into E Scotland (Preston et al. 2002).
Threats
Meadow improvements and excessive mowing of verges.
Native and/or possibly accidentally introduced, rare. Eurosiberian temperate.
1881-2; Barrington, R.M.; Belleisle House estate, Upper Lough Erne.
April to August.
Growth form and preferred habitats
Cowslip is a rosette forming, early flowering, shortly rhizomatous, potentially long-lived, patch-formingperennial herb of damp but well-drained, mesic to calcareous, species-rich grassland soils in the pHrange 5 to 8 (Grime et al. 1988).
Although often reputed to favour constantly moist, lightly-shaded sites on the margins of woods, alongpaths or other openings, or by open scrub, none of the Fermanagh P. veris records are associatedwith such habitats. In many areas of B & I and Europe the restriction of Cowslip to relativelyunproductive, nutrient-poor, neutral or mildly acidic to lime-rich soil and to light- or half-shade, maysimply be a mechanism of avoiding excessive competition(Tamm 1972). Apart from this list of relatively open inland habitats, P. veris also occursin coastal sites on stabilised sand dunes and sea cliffs and shows a greater tolerance of drought thanP. vulgaris (Whale 1984; Grime et al. 1988). Prof Grime and his Sheffield co-workersclassified P. veris as a stress-tolerator with some competitive ability.
As is the case for the common Bluebell (Hyacinthoides non-scripta) and Bracken (Pteridiumaquilinum), the grey cloudy skies of the NW of Ireland may provide sufficient shade for Cowslip,so that it can compete better and survive in fully-lit, unshaded sites.
Fermanagh occurrence
The very pretty Cowslip has certainly declined in well-drained, species-rich 'unimproved' limestonepastures in Fermanagh since 1970, when it was found in the Doagh Lough, Knockmore and above the CladaghRiver Glen (ie the Marble Arch) areas. RHN and the current author (RSF) no longer know of it in thissemi-natural habitat anywhere in the county. However, P. veris does also occur in the seasonallyflooded, more or less organic soils in water meadows around sections of the shore of both Upper andLower Lough Erne, which is an occasionally noted habitat for this species, provided it is notpermanently water-logged. Of the three yellow Primula species that occur in Britain (P.vulgaris (Primrose), P. veris and P. elatior (Oxlip)), P. veris is theleast shade tolerant and the most drought tolerant (Whale 1984).
There are records from a total of 23 Fermanagh tetrads (4.4%). As the distribution map shows, records arescattered across the VC, although the species is mainly found around the Lough Erne basin and on thelimestones of the Western Plateau. Only 16 of the 23 tetrads have post-1975 records, indicating a verydefinite decline in Fermanagh populations. The map indicates that the most widespread loss occurred inthe Clyhannagh and Coaghan Tds, above and to the west of the Marble Arch caves (ie in the Marlbankregion), where Lynn Farrell recorded them in August 1970 and where the species has not been seen since.
Flowering reproduction
Unsurprisingly, Cowslips appear to flower more profusely amongst rank grasses andscrub than amongst large flocks of spring grazing sheep, but easing of grazing pressure tends to lead tomasses of flowers reappearing on English downland, eg in Hertfordshire (VC 20) (Mabey 1996). P.veris bears up to 30 flowers in a 1-sided, drooping umbel on top of a tall (10-20 cm), erect,solitary scape (leafless flowering branch) in April, but chiefly in May (Richards 1993; Sell &Murrell 2014). The flowers are heterostylous as in Primrose (P. vulgaris), hang slightlydownwards, are somewhat cup-shaped, about half the diameter of the Primrose and are clear golden yellowwith an orange to reddish spot at the base of each petal. Like P. vulgaris, the flowers aremostly self-incompatible and are dependent on 'long-tongued' insect visitors (principally honey bees andbumblebees) for their cross-pollination (Richards 1989). However, it has been found that even relativelylow levels of shading (41%-36% of full sunlight), can drastically reduce seed set in P. veris(Whale 1984).
In many areas of B & I, seed-set is good despite the requirement for cross-pollination. Plants withlong-styled 'Pin' flowers can, however, self-fertilise (14%) in the absence of 'Thrum' pollen, whileonly 0.6% of thrums can do likewise (Lees 1971; Richards 1993). After fertilisation, the ovoid fruitcapsule containing around 50 seeds (as large as those of P. vulgaris) is retained within theenlarged calyx, which must delay, restrict and limit seed dispersal distances achieved. Seed dispersalby gravity was found to be normally limited to around 50 cm from the parent plant (Richards &Ibrahim 1978). The seed is dormant when shed and primary seed dormancy is overcome by coldstratification, so that germination should occur in spring following dispersal, or later, if theysurvive soil burial (Milberg 1994).
Seed dispersal and its survival in soil
In 1992, Cowslip occurred as a 'weed' in a newly sown lawn at the Erne Hospital, Enniskillen and four orfive of its previous Fermanagh sites were on established grassland on roadside verges, where its originwas something of a mystery. The seed is without a gelatinous nutritive elaiosome to attract animalvectors and thus dispersal must be entirely wind-, water- or soil-borne. Wind is likely to be animportant means of seed dispersal, although the capsule is not elevated, which must limit distancetravelled. Experiments indicate few seeds disperse more than 1 m (Richards 1989).
One possible explanation of the rather transient occurrences observed in Fermanagh is that viable buriedseed might have persisted in the soil seed bank and have been induced to germinate and establish afterroad making or repairing disturbance brought them to the surface. However, the seed of P. verisis generally regarded as being only transient or short-term persistent in the soil, which rather knocksthis hypothesis on the head (Thompson et al. 1997). However, experimental work in Sweden showedthat P. veris seed can survive in soil for more than one year and might germinate in the secondor later spring after release from the parent plant, ie following two winter cold stratificationperiods. P. veris could easily be overlooked in soil bank studies if a cold treatment is notincluded (Milberg 1994).
Another totally unsupported suggestion is that seed or rhizomatous material may have been imported withfresh topsoil, or on the tyres of vehicles involved in the road works at some stage.
Role of vegetative reproduction
Field observations in England in fully illuminated, short-sward meadows and pastures suggest seed set ofCowslips is normally good, but can be variable from year to year, sometimes poor or sporadic. In theselatter circ*mstances, existing plants must rely on their limited potential for vegetative spread andrhizome growth for some of their increase and regeneration. In stable, favourable growing conditions,such as a meadow or pasture on deep soil that is capable of retaining moisture and with some marginal,light tree shade, P. veris can survive for many years without great change in number. In suchsituations, vegetative propagation by the rhizome branching and extending, compensates for the fewindividuals that die. Regular removal of vegetation by mowing or grazing keeps a sward low, reducescompetition and helps P. veris survive (Tamm 1972).
In open, dry, limestone sites, Cowslip plants remain small and are normally short-lived, seedlingssurviving an average of only two years. In closed grassland vegetation on a fertile, heavy soil theaverage lifespan of surviving seedlings is much longer, up to ten years and allows plants to developlarge clumps. Naturally, seedling colonisation and establishment is much more difficult in closedvegetation than in open or disturbed sites (Richards 1989).
Herbivory, toxicity and vegetation disturbance
As is also the case with P. vulgaris, the leaves of P. veris, which are often appressed tothe ground, tend to be ignored by browsing stock animals, very probably due to their saponin content(Grime et al. 1988; Cooper & Johnson 1998). Flowers and fruits are eaten by various animalsand the latter must assist in seed dispersal. Regular or occasional disturbance, such as cattletrampling and dung, certainly affects pasture vegetation and may provide spots favourable for seedlingdevelopment (Tamm 1972). Seedlings most frequently establish in small pockets of bare ground such asaround molehills, anthills and hoofmarks, often clustered around the parent plant (Richards 1989).
Possible introduction?
In more recent years, Cowslip has become a popular constituent of so-called 'wild flower' seed mixtures,increasingly sown by contractors to re-vegetate disturbed verges after roadworks. However, while thishas occurred in recent years, RHN and the current author (RSF) do not believe that the situation arosein Fermanagh at the earlier dates in question when records were made. The possibility of deliberateintroduction is strongly supported by the fact that the New Atlas distribution map for P.veris quite accurately plots the major road system in NI at least and it may do likewise inother regions of B & I (Preston et al. 2002). The fact that a high proportion of theFermanagh sites for this species are in or near large landed estates, eg Belleisle, Castle Coole, Cromand Florencecourt, might also be significant in this respect. There again, the occurrences might onlyprove what we already know, that the early planters (ie human settlers), recognised good, well-watered,fertile ground when they saw it!
In terms of conservation protection, the Beagh meadow site is an ASSI, and Cowslip is well established onthe main drive at the Castle Coole estate, a National Trust property, while the meadows at ClonattyBridge deserve protection. At the Roosky turlough, Cowslip clings on in a meadow which has already been'improved' − at least in terms of NPK fertility, if not of biodiversity.
Variation
Three subspecies are recognised by Sell & Murrell (2014), distinguished by the near absence ofglandular hairs (subsp. veris), or an abundance of glandular hairs, and then either a cordateleaf base (subsp. columnae (Ten.) Lüdi), or a gradually decurrent leaf base (subsp.canescens (Opiz) Hayck ex Lüdi). Subsp. veris is the form native to B & I and whichhas the widest range of the three, while subsp. columnae is a plant of the mountains of C Spain,C Italy, N Greece and NE Turkey, and subsp. canescens occurs in the Alps, Pyrenees and mountainsof N Spain.
Richards (1993) and Flora Europaea 3 (Tutin et al. 1972) add subsp.macrocalyx (Bunge) Lüdi in Hegi, loc. cit. to these three, it being a plant of SE Russia, theCaucasus and SC Asia that extends to E Siberia (Richards 1993). This form is possibly regarded as arelated, separate species by some taxonomists, which might account for its neglect or discard by Sell& Murrell (2014).
Although there are four recognised subspecies, the authors of the Flora Europaea account (D.H.Valentine and A. Kress) warn that there are many transitional plants (Tutin et al. 1972).
Irish occurrence
P. veris is very much more frequent and widespread in the RoI (especially on the Central Plain),than is the case in NI. Apart from Fermanagh, the few other occurrences in NI are all consideredprobable introductions or garden escapes (FNEI 3). Having said that, on account of the fewperhaps indigenous Fermanagh stations, plus one site in Co Armagh (H37), P. veris retainsProtected Species status in NI under Schedule 8 of the Wildlife (NI) Order, 1985. As BSBI VC Recorders,RHN and the current author (RSF) are sceptical as to the scientific basis for this protection.
British occurrence
The New Atlas hexad map shows that P. veris subsp. verisremains frequent and widely scattered throughout much of lowland Britain, especially in chalky,light-soiled areas, becoming much rarer and more coastal and eastern in distribution in Scotland(Preston et al. 2002). Many populations have survived the widespread destruction of calcareousmeadows and pastures that resulted from the intensification of farming that took place over a halfcentury in the 1930-1980 period, presumably persisting on adjacent marginal land. Pressure on Cowslippopulations, especially near urban conurbations, from flower picking to satisfy deep-seated folkcultural traditions around Easter and other celebrations, also took its toll until 1975 whenconservation legislation made clear the need to refrain from such collecting. There has been aconsiderable degree of population recovery from the 1990s onwards as a result of public awareness andconservation efforts.
Some of the currently mapped Cowslip populations may be derived from gardens and from deliberateplantation of 'wild seed mixtures' in more recent years. Plants of the other two subspecies from Europeare occasionally grown by specialist gardeners and some of these may escape into the wild, although theyare likely to be rare and probably not long persisting (Preston et al. 2002; Sell & Murrell2014).
European and world occurrence
P. veris is a polymorphic species with three or four subspecies depending uponauthority consulted. It is mainly restricted to Europe, but extends eastwards into Asia. Hultén &Fries (1986, Map 1467) choose to map just two subspecies. Subsp. veris is shown to be widespreadin much of Europe, becoming more coastal further north in Scandinavia and the Baltic area. Subsp.macrocalyx occurs in more eastern parts of C Europe and adjacent Asia, but occurs in adiscontinuous manner. The distributions of the other two subspecies is given above under 'Variation'.
Uses and folklore
The flowers have a very distinctive, fresh fragrance similar to Anise or to some volatile oil likeMannite. The flowers contain narcotic juices, including saponin, which encouraged their use inwine-making. Cowslip wine was given to children in small doses as a medicine for various ailments and itwas also prescribed for both jaundice and measles (Vickery 1995). Young cowslip leaves were used inspring salads and mixed with other herbs to stuff meat (Grieve 1931). The main medicinal uses were as asedative and anti-spasmodic, their special value being for strengthening the nerves and the brain,relieving restlessness, giddiness, nervous excitement and insomnia. It was also greatly valued forrelieving pains in the head and was formerly administered for palsy and all forms of paralysis, andnames like 'Palsy Wort' or 'Herba paralysis' were given. The root was also called in old herbals 'Radixarthritica' from its use as a cure for muscular rheumatism (Grieve 1931).
Cowslip balls were made of flower stalks tied together and thrown up for love divination, the names ofpossible suitors recited until the ball landed (Vickery 1995). Cowslips were scattered on the altar inchurches on May Day (the first Sunday in May) (Vickery 1995). This use in church may somehow connectwith plant names that refer to the keys of St Peter (an allusion to the pendulous umbel of flowers), ora Welsh name that translates as 'Mary's tears' (Grigson 1987). Previously Cowslip must have beenextremely common and abundant in at least some parts of England, as folklore has it that flowers wereworn as necklaces on May Day and were scattered in front of newlyweds emerging from church, likeconfetti, practices that could not happen nowadays even without conservation legislation (Mabey 1996).
Names
The English common name 'Cowslip' is a polite euphemism for 'cow-slop' or 'cow-pat' which hints at thekind of company the plants often keep in pastures (Mabey 1996). In old herbals, the plant is referred toas 'Herb Peter' or 'Key Flower', the pendulous flowers being likened to a bunch of keys and hence withSt Peter and his keys to the Kingdom of Heaven. They are also called 'Our Lady's Keys' and 'Keyflower'for similar reasons (Grieve 1931).
Threats
Further improvement of meadow fertility, causing decline of species diversity.
Introduced, neophyte, rare and now locally extinct. In its native range, European temperate and protectedin NI.
1939; Carrothers, E.N.; Whinnigan Glebe Td, near Farnaght.
Fermanagh occurrence
This submerged, shade-tolerant, truly aquatic member of the Primulaceae has not been seen since about the1950s at its only reported Fermanagh station, the bog holes and adjacent drains at Whinnigan Glebe Td,about 7 km from Enniskillen. This brief habitat description does not sound very suitable for thesurvival of the plant, since H. palustris really prefers clear, mesotrophic, base-rich, usuallycalcareous waters (Preston & Croft 1997; A.J. Richards, in: Preston et al. 2002). In theRevised Typescript Flora, Meikle et al. (1975) followed Praeger (1939) in statingquite unequivocally that the plant was introduced from the Bog Meadows in SW Belfast, some time around1920. The site at Whinnigan Td lies just beside the Carrothers' family farm at Farnaght, so we believethat the plant was most likely introduced there by N. Carrothers, ie the older Carrothersnaturalist. Praeger (1939) listed the next generation family member, R.D. Meikle's co-worker, E.N.Carrothers, as his source for the record of the Fermanagh introduction.
The Bog Meadows in SW Belfast, from which the Fermanagh plant derived, was itself an introduction of thisspecies. It is known that John Templeton, one of the most talented early Irish naturalists, transferredthe plant to his garden at Cranmore in south Belfast in the early 19th century. He collected hismaterial in the River Quoile marshes around Downpatrick, Co Down (H39), where the first Irish discoveryof this wintergreen aquatic perennial was made by a Mr Richard Kennedy in June 1810. The original sitewas described as, "in the marshy ditches on the right side of the road at Everogue bridge, nearDownpatrick" (FNEI 1).
The flowers and submerged leaves are attractive so that H. palustris is nowadays commonly used todecorate and oxygenate garden ponds and fish-tanks. The species is heterostylous and requires crossingbetween pin and thrum flowers to set seed. Being stoloniferous and capable of vegetative reproduction byfragmentation, the plant often produces single-morph colonies which fail to set seed. H.palustris should be capable of long-distance jump dispersal through the movements of water fowl,as this is presumed to have occurred elsewhere in these islands (McCallum Webster 1978; Hackney etal. 1992). An interesting summary of the biology, ecology and distribution of the species isprovided in Preston & Croft (1997).
Irish occurrence
In Ireland, this very attractive stoloniferous perennial is a very rare species which has been reportedin a small number of other scattered Irish counties, all of which stations are known or deemedintroductions. A small (but presumably sexually reproducing) population survives, totally submergedapart from the flower spikes, in deep, peaty, ditches, dug in a wood at Hollymount NR near Downpatrickand in the nearby marshes (VC H38). These sites are the only candidates for possible Irish nativestatus. In NI, it is protected under Schedule 8 of the Wildlife (NI) Order, 1985.
British occurrence
In England and Wales, H. palustris is quite frequent and widely scattered south of a line on themap between Sunderland and Holyhead and locally common in eastern England. However, with the exceptionof the Bristol area, it is poorly represented west of a line between Chester and Worthing on the Scoast. Isolated occurrences in C Scotland and the Isle of Man are regarded as definite introductions(New Atlas).
Sites have been lost in Britain due to drainage, vegetation clearance from waterways, eutrophication andboating disturbance beginning prior to the 1930s, but continuing to the present, especially in SEEngland (A.J. Richards, in: Preston et al. 2002).
European occurrence
H. palustris is mainly restricted to middle latitude areas of Europe, although it reaches north asfar as C Sweden and southwards to S France, C Italy and Romania (Hultén & Fries 1986, Map 1475).
Threats
Drainage, eutrophication and collection.
Native, frequent and widespread. Sub-oceanic temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
The reddish stem and ovate, acute, entire, opposite leaves make this creeping and trailing,yellow-flowered perennial unmistakable. While the slender, ± prostrate stems and thin herbaceous leavesmight appear delicate, the plant is completely frost hardy and evergreen, but it is not regarded aslong-lived. L. nemorum is frequent and sometimes sufficiently abundant to form dense mats orloose carpets in damp, lightly shaded ground. Typical habitats are damp to dry, poorly-drained,winter-wet or flushed, nearly bare soil patches found in woods, scrub, on lakeshores, or by tracks,hedge- or river-banks, shady ditches, or in more or less marshy grassland. It also occurs in shady spotsin upland glens and on cliffs. L. nemorum prefers quite fertile or base-rich soils of mediumreaction (ie moderately acid to neutral pH levels).
Yellow Pimpernel is absent, however, anywhere vegetation becomes densely shaded, or where the turfbecomes rank, tall or over dense, or where leaf and branchlet litter accumulates. Bare or only vernallyoccupied, open disturbed ground is essential for colonisation and establishment, allowing L.nemorum to avoid competition from more vigorous, taller, or larger-leaved species (Sinker etal. 1985; A.J. Richards, in: Preston et al. 2002). The necessary environmental conditionsfor limited growth potential are provided by a wide range of factors, but most typically they involve,cool damp shade, flushing by springs or occasional flooding, or less frequently, flushed steep slopes,damp, shady narrow cliff ledges, trampling of grassy paths, or other forms of moderate disturbance.
Fermanagh occurrence
While it is widespread throughout most of Fermanagh, having been recorded in 233 tetrads, 44.1% of thosein the VC, Yellow Pimpernel prefers fertile or base-rich soils of moderately acid to neutral pH levelsand is, therefore, only occasional in limestone areas of the county. Having said this, the tetrad mapshows it widely dispersed around most of the VC, avoiding strongly acid peat and the driest rockyground.
Reproduction
The very attractive, solitary, long-stalked, yellow, star-like flowers are often quite profusely producedand a succession of them appears between early May and mid-September. However, seed production isgenerally sparse and reproduction relies heavily on vegetative spread by nodal rooting of the prostrateshoots, which can be up to 45 cm in length (Salisbury 1942). The frequent low levels of seed productionby the species is odd, given that the flowers, while without nectar, offer pollen and attract flies,bees and other insect visitors which could pollinate them. They are also said to be hom*ogamous (ie themale and female parts mature simultaneously) and self-fertile (ie autogamous), which also ought toensure a good seed set (Vogel 1978, p. 91; Fitter 1987). The fruit is a small, globular capsule thatsplits to release the few, angular seeds (Melderis & Bangerter 1955).
Despite the observed limited seed production, a lack of any specialised dispersal mechanism andpresumably resultant poor transport, the ease with which the stems root and vegetatively produce clonalpatches, often allows L. nemorum to become locally abundant in damp to marshy grassy vegetation.
Ecological effect of mowing and grazing
Yellow Pimpernel does not survive mowing (Hansen & Stahl 1993) and, similarly, it cannot toleratemuch grazing (Sinker et al. 1985). The small scale of the plant, together with the open nature ofmany of its habitats, makes browsing depredation unlikely in any case. Gardeners have noted that unlikeother cultivated members of the Primula family, L. nemorum does not suffer the attentionsof rabbits. This suggests that, like Anagallis arvensis (Scarlet Pimpernel), it probably containssufficient saponins to make it unpalatable, or possibly even toxic to some herbivores (Cooper &Johnson 1998; Thomas 2004).
British and Irish occurrence
The New Atlas distribution map displays L. nemorum as being widespread throughout most of B& I, although less common in C Ireland and absent in Britain from much of the east coast and aconsiderable area of the E Midlands (Preston et al. 2002). Comparison of the records for thespecies in the 1962 and 2002 BSBI Atlas surveys shows L. nemorum has suffered a widespreaddecline in S England over the years, with a calculated Change Index of -0.46. Destruction of deciduouswoodland over this period and its replacement with coniferous plantation has created dense shade habitatconditions that are totally unsuitable for Yellow Pimpernel.
Fossil history
The fossil record for both L. nemorum and L. nummularia (Creeping-Jenny) is very poor in B& I. Seeds of one or other have been identified from only one site in Kent dating from therelatively recent Zone VIIb of the current interglacial period (Godwin 1975, p. 308). This does not giveany indication as to their means of transport to these islands, or the antiquity of either of theseassumed native species.
European occurrence
Beyond our shores, L. nemorum is very much confined to W & C Europe. It is absent from Icelandand from areas of Scandinavia north of 60°N. Interestingly, although the species is absent fromShetland, it has been recorded on one sandy and gravelly lake shore in the Faeroe Isles, though how itgot there is problematical (Ostenfeld & Grontved 1934). The remote population on the Azores isdistinguished as subsp. azorica (Horn.) Palh. (Hulten & Fries 1986, Map 1476).
Medicinal uses
Yellow Pimpernel had or has very few folk medicinal uses, although the yellow flowers were boiled inCounty Cavan as a cure for gallstones and there is some suggestion that it may have shared the soporificeffect of its relative Primula veris (Cowslip) (Allen & Hatfield 2004).
Threats
None.
Probably native, common. European temperate, but cultivated in gardens and widely naturalised.
1866-70; Smith, T.O.; Pollboy Bridge on the Colebrooke River.
April to December.
Growth form and preferred habitats
This evergreen, mat-forming, fast creeping perennial is common and easily recognised, but both the wildtype and a golden-leaved form (var. aurea) of it are quite commonly cultivated (usually by alpinegardeners or in hanging baskets), flowering from May to September (R.G. Woods, in: Perry & Ellis1994). They are often invasive in the deep, fertile soil of a damp garden situation, require cuttingback after flowering; they sometimes escape or are dumped with garden waste and become naturalised,confusing the native distribution to some extent (Grey-Wilson 1989). However, it would seem likely, thatfollowing the pattern of the southern portion of its British range and throughout Ireland, it isprobably native in almost all of its large number of Fermanagh stations. The typical habitats itoccupies in the VC are damp, often shady places by lakes, rivers and in woods, rather than in moredisturbed sites or close to habitation, as might be expected of a garden escape.
Fermanagh occurrence
Creeping-Jenny is quite commonly found in damp, open ground in Fermanagh. It has been recorded in 118tetrads, 22.4% of those in the VC. As the tetrad distribution map indicates, it is mainly but notexclusively found around the shores of Upper and Lower Lough Erne. The author(s) of the RevisedTypescript Flora (essentially R.D. Meikle) very definitely regarded L. nummularia as a,"characteristic plant of the Erne basin". Having said this, in some sites the status of thespecies must remain questionable and, very possibly, at least some of the Fermanagh records are gardenescapes.
Species status in Britain and Ireland
Stace (1997) describes the distribution of L. nummularia as follows (some abbreviations expandedbut otherwise quoted verbatim), "Native; damp places, often in shade; throughout most of B & Inorth to C Scotland, but a naturalised garden escape in many localities, especially in the north."The 1962 BSBI Atlas accepted the vast majority of B & I records as native, with only a tinyminority of 'recorded introductions' in the N & W of Britain and in NW Ireland (Walters &Perring 1962). That said, the main British distribution lies south of a line between Hartlepool andHeysham, thinning markedly northwards. County Floras of Durham (VC 66) and Cumbria (VCs 69, 70 and partsof 60 & 65), indicate that L. nummularia is somewhat more widespread in these areas thanpreviously thought and, in these areas, it represents a mixture of native and naturalised populations(Graham 1988; Halliday 1997).
Praeger (1934i) assembled some of the facts regarding the Irish distribution and, while acknowledgingthat L. nummularia is commonly cultivated in gardens, he argued strongly that the species shouldbe considered indigenous.
In his Botanist in Ireland, Praeger (1934i, paragraph 76) argued that the species should beconsidered indigenous in at least some of its stations. He maintained that, "the flora becomes moreand more aboriginal as the soil becomes unsuitable for agricultural operations, as on bogs, marshes andmountains".
In a paper on the standing of certain plants in Ireland (Praeger 1934b), he wrote in relation to thisspecies, "lake shores and marshes are habitats where the alien element in the flora is at aminimum", a statement with which the current author simply could not agree. However, as Webb (1985)very sensibly pointed out, a plant simply, "looking wild" and growing amongst undoubtedlynative associates, does not necessarily qualify as indigenous itself by so doing. It should be realisedthat Praeger had a strong penchant for plumping for species as indigenous which others considereddoubtful (Praeger 1934i & 1937b).
Reproduction
Creeping-Jenny produces almost no seed anywhere in these islands, yet it is a remarkably widespreadspecies. It frequently occupies waterside habitats that provide it with an obvious means of rapid seedand vegetative dispersal. Even without water transport, the plant can spread locally to form extensivepatches by means of rapid, prostrate stem growth and rooting at nodes. Taken together these methods ofincrease should ring alarm bells and hint at some form of introduction. Further investigation isrequired to clarify the status issue but, as the situation stands, RHN and the current author (RSF) feelthe Fermanagh lakeside and woodland stations of L. nummularia are best considered only 'probablynative'.
The basis of the observed high degree of seed sterility in L. nummularia was investigatedthroughout its European range by Bittrich & Kadereit (1988), examining both live and herbariummaterial. Meiotic irregularities were common (ie failure of chromosome pairing), resulting in a highpercentage of non-viable pollen and pollen grains of unequal chromosome number. Rarely, however, someindividuals were capable of seed set, so sterility is not total. In many populations, theself-incompatibility of the species and the reliance on vegetative clonal reproduction has led toincreasing levels of sterility.
Despite the above, the frequency of fertile individuals is comparatively high in SE Europe, whichprobably served as an ice-free refugium for the species during glacial periods. It seems likely thatduring the spread of L. nummularia into northern latitudes after the glaciers retreated, only afew genotypes may have been involved, perhaps migrating mainly by vegetative means, which would lead intime to a degeneration of the sexual reproduction system of the plant.
European and world occurrence
L. nummularia is present across most of middle latitude Europe, northwards into SScandinavia and stretching east to the Caucasus. The distribution also thins southwards into the Iberianand Balkan peninsulas. On account of its garden cultivation, it is widely introduced and naturalisedbeyond its native range and is quite frequent and widely scattered in eastern N America and present alsoin New Zealand (Hultén & Fries 1986, Map 1478).
Threats
None.
Native, common but rather local. Eurasian temperate, but widely naturalised.
1884; Barrington, R.M.; Lough Erne.
April to October.
Growth form and preferred habitats
Up to 150 cm tall, a rhizomatous and clump-forming perennial with opposite or 3- or 4-whorled 9 × 3.5 cm,downy-coated leaves, L. vulgaris is a vigorous, panicle-forming, yellow-flowered plant ofditches, damp to wet meadows, sheltered tall herb marshes, swampy woods and lakeshore fen-carr. L.vulgaris is easily distinguished from the only other tall, yellow-flowered member of the genusin Fermanagh, the much rarer garden escape L. punctata (Dotted Loosestrife), by that species morebranched inflorescence, calyx teeth without a darker, orange margin and leaves with at most a fewscattered hairs on their margins (Webb et al. 1996).
Fermanagh occurrence
Yellow Loosestrife is a conspicuous and characteristic perennial of wet ground around the shores of UpperLough Erne. It also occurs in similar damp to wet situations scattered throughout the county, reaching atotal of 114 tetrads, 21.6% of those in the VC. However, while it is frequent and locally abundantaround the margins of the larger lakes in sheltered, well-grown, tall herb or wet woodland conditions,it is very much more scarce or totally absent around many of the smaller lakes in the county. Furtherevidence of the local occurrence of Yellow Loosestrife is given by the fact that of the over 500 recordsin the Fermanagh Flora Database, very few derive from river margins. The species has only been found onthree – the Woodford, Finn and Swanlinbar rivers.
The fact that a vastly greater number of man hours have been spent on lake surveys in Fermanagh comparedwith those RHN and the current author (RSF) have spent walking the riverbanks, must unquestionably bepart of the reason for this apparent discrepancy. However, another factor must be the strenuous effortsthe NI Water Service and local farmers continually make to keep drainage channels open, removing tallherb communities that threaten to clog waterways and canalising ditches and streams. Harron (1986) notedvery much the same restricted pattern of occurrence of this species around Lough Neagh and itsassociated larger water bodies. He attributed this to drainage and reclamation works limiting the plantto isolated patches amid remnants of scrub and reed beds (Flora of Lough Neagh).
Irish occurrence
L. vulgaris is widespread in central and northern parts of Ireland lying northwest of the RiverShannon, but it is much scarcer or absent from both the better farming country in the SW of the islandand from the more acid, peat bog conditions of the far south and the western seaboard (Preston etal. 2002).
British occurrence
L. vulgaris is widespread throughout England and Wales, but it is much rarer in Scotland andreally is only frequent there in the central lowlands, although it does reach Aberdeen on the north-eastcoast. As it is essentially a lowland species, the high Scottish mountain ranges clearly have providedan impassable geographical barrier preventing spread further north and north-west.
The calculated Change Index between the two BSBI Atlas dates (1962, 2002) has a value of +0.22,yet despite this overall apparent increase in distribution, which could well be due to differences inrecording effort, analysis of the New Atlas survey data indicates that most of the observedhabitat losses due to drainage and clearance of watercourses, took place after 1950 (A.J. Richards, in:Preston et al. 2002).
Variation
L. vulgaris is a phenotypically very variable species, ie able to adapt and modify its growth formand characteristics depending on the particular ecological conditions under which it lives. Generallythe plant grows in small groups, rooting in organic mud, but in some wet situations, including whenstanding in permanently flooded ground, it can develop long, above-ground, horizontally spreadingstolons.
Flowering reproduction
The size, colour and, very unusually, even the shape of the flower organs can change to some extent inrelation to the level of light. Except when in deep shade, the plant bears a large number of conspicuousyellow flowers in July and August on long, branched, leafy panicles. They contain no nectar but haveplenty of pollen in their large anthers, which generally proves attractive to hoverflies, solitary bees(particularly Macropis species) and certain wasps, although the flowers can also self-pollinateand fertilise if they remain unvisited (Proctor & Yeo 1973; Simpson et al. 1983).
The fruit capsule contains only three or four discoid seed and these can float for between one and fourweeks, allowing the plant to disperse within the lake, river and stream water system. Seed has beenidentified in the crop of Woodpigeon, although this does not necessarily mean that the birds disperse it(Ridley 1930, pp. 220, 498). Yet apart from seed transport in mud, how otherwise could the species crossdry land and colonise additional, isolated waterbodies?
Horticultural use
A garden form of the species exists with slightly larger flowers with petals red at their base, var.grandiflora. Allen (1968) noted this form 'jumping over the garden wall' and actively spreadingin semi-wild boggy terrain on the Isle of Man. The species is sometimes included in wild-flower seedmixtures, widely sown in disturbed ground throughout these islands in recent years (Sell & Murrell2014).
European and world occurrence
Occasional in garden cultivation, L. vulgaris is widespread throughout most of Eurasia except theextreme north. It was introduced and became naturalised and developed into a serious invasive weed ineastern N America and to a lesser extent in New Zealand (Hultén & Fries 1986, Map 1477; Webb etal. 1988).
Names
Grigson (1987) provides two origins of the genus name; the Greek 'lusimachion' translates as'loosestrife', which means 'ending strife', but Pliny believed the name commerated Lysimachos, a King ofThrace.
Threats
None.
Introduced, neophyte, an uncommon garden escape, probably mis-named.
13 September 1974; Hackney, P.; E shore of Lower Lough Erne.
July to September.
Growth form and preferred habitats
This popular, tall evergreen garden perennial, introduced from SE Europe and SW Asia, is very vigorousand can form clumps up to 120 cm tall. Given its invasive vigour in the garden setting, the famoushorticulturalist G.S. Thomas (2004) recommends it only for broad sweeps of self-maintained colour in thewilder parts of larger gardens. On account of its thriving growth, the plant quickly outgrows itsallotted space and almost always originates in the wild from discarded material. It then persists andmay spread to a limited extent in suitable growing conditions through the possession of a short, thickrhizome.
L. punctata is readily distinguished from L. vulgaris (Yellow Loosestrife) by the flowersbeing confined to the upper leaf axils and by its entirely green sepals. In comparison, theinflorescence of L. vulgaris is a terminal panicle and its sepals have much darker margins.However, it is important to note that vigorous stems of L. punctata can produce lateralinflorescences similar to the main one (McAllister 1999).
L. punctata is sterile or strongly self-incompatible and it only very rarely sets seed (McAllister1999). Part of the reason for the lack of seed may be the close, but not usually obligate, relationshipof the flower with Macopis solitary bees as pollinator (Simpson et al. 1983).
Fermanagh occurrence
In Fermanagh, L. punctata usually occurs in disturbed wayside situations not far from houses,colonising rough grassland and hedge margins. The earliest record of L. punctata in NI appears tobe that of Paul Hackney in Fermanagh in 1974, yet there are now 16 records in the Fermanagh FloraDatabase from 13 tetrads, very thinly scattered throughout the lowlands on roadsides and in rough ordisturbed ground. This suggests that either L. punctata is spreading, or was ignored oroverlooked by earlier field workers, as often was the case with naturalised garden introductions.
An interesting observation, which probably contributes to the untended survival of the plant in the wild,is that L. punctata appears to require less soil moisture than other Lysimachia species(Ingram 1960). As a result it is not confined to damp ditches, marshy ground, or fen carr but, inaddition, is found on roadsides, waste places and old quarries, all places where people habitually dumpgarden waste (Reynolds 2002).
On the other hand, the calculated Change Index comparison of the two BSBI Atlas surveys across B& I gives L. punctata the extremely high score of +4.62. This indicates a massive increase inrecords of the species during the 40 years between the two BSBI Atlas recording efforts. Thequestion remains: is the plant rapidly increasing, or is it simply being recorded now when previously itwas ignored? L. punctata has been in garden use since 1658 and was first recorded in the wild in1853 in Angus (VC 90) (A.J. Richards, in: Preston et al. 2002). Obviously L. punctata hasbeen around for a long time and it is possible that we might be witnessing a real change in itscolonising ability.
British and Irish occurrence
Despite the lack of seed, L. punctata appears to have become more frequent in recent decadesthroughout these islands (Clement & Foster 1994; NI Vascular Plant Database). The New Atlasmap shows Dotted Loosestrife is quite frequent and widespread throughout lowland Britain, including onthe remote islands of Orkney, Shetland and the Outer Hebrides. In Ireland, it is rather betterrepresented in the north: otherwise, it is very thinly scattered southwards in the western half of theisland, with one outlier near Wexford town. However, there are grounds for believing that some recordsmade in recent years may be identification errors, due to another rather similar species, L.verticillaris Spreng., having been introduced to cultivation.
European occurrence and a possible error in B & I recording
The natural distribution of L. punctata is confined to Europe and W Turkey. A morphologically verysimilar plant with a more easterly, non-overlapping wild range is L. verticillaris Spreng.(Leblebici 1978). The latter is self-compatible and freely sets seed. In the view of the current author(RSF), it appears likely that a single, or a very few clones, of self-incompatible L. punctatahave been introduced into garden cultivation and have propagated vegetatively. More recently (perhapssince 1962), however, material of L. verticillaris must have also been introduced tohorticulture, traded, distributed and grown by gardeners as 'L. punctata' (McAllister1999). Thus the current author shares Hugh McAllister's belief that the self-seeding L.verticillaris is producing the rapid increase in records erroneously being referred to as L.punctata around these islands.
L. verticillaris differs from L. punctata in at least five characters, the most easilyobserved being the petals, which are flushed with orange near their base, and nodes and leaf bases,which are purplish, rather than green (McAllister 1999).
The self-seeding Dotted Loosestrife (L. verticillaris) was introduced to N America as a gardenplant and has likewise become a naturalised weed of disturbed ground on roadsides and in fields in theeastern states (Ingram 1960). We are probably safe in assuming the same situation applies on bothcontinents, N America and Europe.
Threats
None.
Native, frequent. Oceanic southern-temperate.
1882; Stewart, S.A.; Drumbad Scarps, Lough Navar Forest Park.
April to January.
Growth form and preferred habitats
This inconspicuous little prostrate perennial with its slender stems and numerous tiny, almost round,pale green or pinkish-buff, sub-opposite leaves is probably quite often overlooked. It creeps and rootsat the nodes on the surface of moist to wet peaty or sandy soil, or else trails over the mossy groundcarpet of bogs and heathy flushes. It grows in open, almost or fully-illuminated situations, on eitherbare mud or sandy lake shores or stream sides, on moss-covered bogs, or in short turf marshy, generallyflushed ground that frequently supports quite a wide range of flowering plant species, including many ofthe smaller sedges.
Bog Pimpernel tolerates moderate levels of grazing, trampling and flooding, eg in low-growing sedge areasof marshy grassland, acid fens and beside paths, including in more open areas along woodland tracks.These differing forms of disturbance all clearly allow it and other small plants to survive by keepingthe vegetation sufficiently open, creating bare patches, reducing competition and increasing thelikelihood of rarer or more local plants occurring.
In coastal areas of B & I, Bog Pimpernel commonly occurs on damp sand in dune slacks, a habitat typewe cannot emulate in landlocked Fermanagh.
Fermanagh occurrence
Bog Pimpernel is a quite frequent, by no means scarce species in Fermanagh, being recorded from a totalof 84 tetrads, 15.9% of those in the VC. It appears to have lost a few sites, however, since six tetradshave pre-1975 records only. As the tetrad distribution map indicates, the species is decidedly local,being concentrated in the western, wetter, more upland half of the county. That is not to say that BogPimpernel is always an upland plant, just that it is more prevalent in upland situations on theconstantly moist, nutrient-poor, moderate to strongly acidic, often peaty soils characteristic of theWestern Plateau.
At the same time, A. tenella can also be found on the shallow, somewhat drier, raw humus,ranker-type of heath soils formed directly over Carboniferous limestone. These occur, for instance,around the lowland shores of Lower Lough Erne and on the hills above Florencecourt. In such soils, itonly grows where sufficient flushing ground water keeps the plant roots moist. Very occasionally, itgrows in wet, highly calcareous situations, but it is then never abundant.
Ecological indicator species
Bog Pimpernel is a good indicator of species-rich habitats which are worthy of careful searching forother uncommon acid-tolerant plants, including Drosera anglica (= D. longifolia) (GreatSundew), Eleocharis quinqueflora (Few-flowered Spike-rush), Carex dioica (DioeciousSedge), C. viridula subsp. brachyrrhyncha (Long-stalkedYellow-sedge), C. limosa (Bog-sedge) and the slightly calcicolous Parnassia palustris(Grass-of-Parnassus).
Flowering reproduction
From June to August, Bog Pimpernel bears a mass of erect, relatively long-stalked, small, solitary, palepink flowers. The petals are finely striped with purple and the centre of the upright funnel-shapedflower is filled with fluffy white hairs on the filaments of the five stamens, making them absolutelybeautiful when viewed in close-up.
Very little appears to be known or certain about the reproductive biology of the species. The flowers arevisited by insects, but they are also capable of selfing (Fitter 1987). Seed is dispersed in mud byducks and other waterfowl (Ridley 1930, p. 547). One reference exists, dating from 1934 that claims seedis long-term persistent (ie surviving more than five years in the soil) (Thompson et al. 1997).Despite its name (see below), the plant is not tender but perfectly hardy in our western oceanic climateat least. Having said this, garden cultivated forms are described by Grey-Wilson (1989) as"short-lived and may be killed in a severe winter".
Apart from the Grey-Wilson's comment, the current author (RSF) can discover nothing in the literature onthe population biology of A. tenella.
British and Irish occurrence
The New Atlas map shows that in B & I the distribution of the species has a pronounced westernpredominance, although it does also display a complete south-north latitudinal range – although verydefinitely thinning in frequency towards the NE and the much drier SW of Britain. This trend againsuggests a possible low temperature limitation. On the other hand, the altitudinal limit of the speciesappears to be as high as 610 m in N Aberdeen (VC 93), while in Ireland it reaches 490 m in both theMourne Mountains, Co Down (H38) and in the far SW on the very highest Irish mountains, theMacgillycuddy's Reeks in S Kerry (H1), indicating the plant demonstrates considerable frost resistance(Wilson 1956; A.J. Richards, in: Preston et al. 2002).
The English distribution includes a disjunct E coast presence, clearly reflecting the occurrence of BogPimpernel in its dune slack habitat. In Ireland, A. tenella is more generally distributed incomparison with Britain, the pattern here mirroring the highest levels of precipitation and generalhydrology (Haughton et al., Atlas of Ireland, 1979, pp. 22, 32).
There undoubtedly have been losses of A. tenella in Fermanagh and elsewhere throughout B & Iin recent decades, although the New Atlas Change Index of -0.54 really is a moderately lowfigure. Losses have been consequent upon the drainage of lowland bogs and marshes, but also through ageneral nutrient enrichment associated with agricultural changes and grassland 'improvements' over thelast half century.
European occurrence
The wider, essentially milder-European region distribution of A. tenella puts it firmly into theOceanic Southern Element of Matthews (1955), which comprises 76 species disregarding those nowrecognised as introductions, a category more recently refined as the Oceanic Southern-temperate (25species) by Preston & Hill (1997). Most of the species in this grouping have their northern limit inBritain but, rather exceptionally, A. tenella is a very rare member of the coastal flora of theFaeroe Isles which lie at 62°N (Ostenfeld & Grontved 1934). The only other OceanicSouthern-temperate species to approach this northern range is Eleogiton fluitans (FloatingClub-rush) (another plant of wet, peaty habitats), which on the European mainland reaches around 57°N inS Sweden and is also present in Shetland around 60°N.
At its southern limit A. tenella has outlying localities in N Africa, W Greece and NW Crete(Hultén & Fries 1986, Map 1483; Turland et al. 1993, Map 1050).
Names
The Latin specific epithet 'tenella' is a diminutive meaning 'tender' or 'delicate', the latter morefitting when the very beautiful flower is being closely admired (Gilbert-Carter 1964). As noted above,the species is perfectly hardy in our climate, and could not overall be described as tender.
Threats
Drainage, eutrophication and changing land use.
Native, occasional. Eurosiberian southern-temperate, but widely naturalised and now circumpolar.
1882; Stewart, S.A.; Co Fermanagh.
May to November.
Growth form and preferred habitats
Rather surprisingly, this little sprawling annual is classified as one of the worst weeds in the world,having been accidentally introduced by emigrants almost everywhere around the globe from its nativerange in S Europe (Holm et al. 1977). Much of the reason for this drastic reputation rests in thefact that this small plant is a very variable and adaptable weed in a wide range of crops, includingpulses, cereals, vegetables and oil seed. Despite the small scale of the plant, the species is alsodangerously poisonous to a wide range of stock animals (Chrtek & Osbornova-Kosinova 1986). The plantcan also cause dermatitis in some individuals if the leaves or stems are handled (Holm et al.1977).
In our relatively mild, wet climate, A. arvensis is mainly a summer annual. Previously, in B &I, it commonly grew on the field margins of spring-sown crops. It can also behave as a winter annual oreven as a facultative perennial, but there is no evidence that it does so in Fermanagh (Grime etal. 1988 & 2007).
Since the 1950s, increasingly effective weed-control measures have made this once familiar agriculturalweed and widespread wildflower much scarcer. Nowadays, it is seldom found and then almost always inopen, heavily disturbed sites where competition is either absent or slight. Examples include aroundrural gateways, quarries or near habitation where bird-sown seed (some of which may originate frombird-tables) may establish.
The species can tolerate light shade, but the flower always demands full sun and will close if clouddarkens the sky, a fact that gave rise to the numerous alternative English common names relating to boththe clock and the weather, eg 'John-go-to-bed-at-noon', 'Twelve o'clocks' and 'Shepherd's watch', andwith regard to weather, 'Poorman's Weatherglass', 'Grandfather's-, Old Man's-, or Shepherd's-Weatherglass', or 'Weather-teller' (Grigson 1987).
Fermanagh occurrence
The Fermanagh Flora Database contains records from a total of 26 tetrads, 4.9% of those in the VC. As thetetrad distribution map indicates, A. arvensis is confined to the lowlands, is very thinlyscattered and has declined significantly. Only 15 Fermanagh tetrads contain post-1975 records.
The decline of arable farming together with weed-control has made A. arvensis much more occasionalor rare here. Nowadays, it appears in small, ephemeral populations confined to open, heavily disturbedsites, both urban and rural, around gateways, roadsides and quarries or waste ground near habitation.
The only semi-natural situation in which A. arvensis has been recently recorded in Fermanagh is onsandy lakeshores, but even these are grazed and trampled by cattle. The preferred soil is light andsandy, yet neither too dry nor too acidic. The plant can tolerate light shade, but the flower alwaysdemands full sun and will close if cloud darkens the sky. 'Scarlet Pimpernel' is something of amisnomer, the normal and virtually constant flower colour in the current author's experience (RSF),being a distinctive orange-red.
Evidence of decline
A pronounced decline in the occurrence of A. arvensis, similar to that in Fermanagh, was found insurveys of Danish arable fields made in 1967-70 and again in 1987-9 (Andreasen et al. 1996).These workers attributed the steep decline of the species to its sensitivity to herbicides, togetherwith low rates of seed production and the plant's lack of competitive ability against modern cropvarieties. Rich & Woodruff (1996) analysed changes in the flora of England brought to light by theBSBI Monitoring Scheme, comparing samples from 1930-60 with 1987-8. They found significant reductions in31 species of arable weeds, but did not include A. arvensis in their list of declining species.The editors of the New Atlas calculated a Change Index, comparing the results of the 1962 and2002 BSBI Atlas surveys and, for A. arvensis, the negative value is quite large (-0.73).In addition, comment is made that there have been losses in the north of the species range, the timingof which is unknown (A.J. Richards, in: Preston et al. 2002).
Flowering reproduction
Unlike many other ruderal, annual weeds, A. arvensis has quite a short flowering period (June toAugust) and, although very variable with respect to the environment in terms of the number of capsulesproduced, overall the species produces a relatively small seed crop, numbering on average just 902 ± 54seeds per plant (Salisbury 1942, p. 149).
The seed coat of A. arvensis contains a water-soluble germination inhibitor which provides aninnate dormant period and the species produces a persistent seed bank. Natural seed dormancy is at leastten years and seed longevity certainly exceeds 16 years, with other estimates ranging up to over 68years (Thompson et al. 1997).
Flower variation
The English common name 'Scarlet Pimpernel' is something of a misnomer, the normal and virtually constantflower colour in the current author's experience, being a distinctive orange-red. The five (or more)colour variants known are scarlet, pink (or orange-pink or salmon), white, purple (or lilac) and blue,all form a series, in which scarlet is simply dominant to pink, pink to white, and so on across thespectrum. The beautiful blue-flowered variant is recessive to all the other forms, except that when itis crossed with lilac there is segregation in the F1 generation (Allen 1954). Actually twoblue forms of the species exist: A. arvensis subsp. arvensisforma azurea and A. arvensis subsp. foemina, the latterhaving both narrower upper leaves and petals and with petal margins minus the numerous hairs of subsp.arvensis forma azurea (for full distinction see Stace 1997, p. 301). The formaazurea (Blue Pimpernel) is known to occur in coastal parts of counties Antrim, Dublin and Wicklow(H39, H21 & H21) at least, and the two blue forms are by a long way the most prevalent variants ofthe species in the Mediterranean region, where they behave as winter annuals. As far as the currentauthor is aware, neither form of the Blue Pimpernel has been recorded in Fermanagh.
Toxicity
The poisonous principles in A. arvensis are not yet properly understood. Aerial parts of the plantare said to contain a glycosidal saponin, while the roots supply another saponin called 'cyclamin'; anacrid volatile oil has also been isolated from the species (Cooper & Johnson 1998). When 31 cattleand buffalo died in India after been fed collected 'greens' with a high proportion of A.arvensis, Sadekar et al. (1995) considered the clinical symptoms, lesions andhistopathology pointed to oxalate toxicity. Subsequent analysis showed the plants contained 9-13% oxalicacid on a dry weight basis. In addition to the animals mentioned, in various parts of the world theplant has produced gastro-intestinal, heart, liver and kidney problems (sometimes lethal) in horses,dogs, sheep, rabbits and poultry. The seed has also affected birds (Cooper & Johnson 1998).
Medicinal uses
In the past, the plant was used in herbal medicine as a diuretic and to stimulate bile flow, butfortunately it is now considered completely obsolete in allopathic medicine, although it is stillretained in homeopathy for skin complaints such as itching and warts (Launert 1981).
European and world occurrence
Hultén & Fries (1986, Map 1484) regard the Mediterranean basin as the probable point of origin ofA. arvensis, from whence it spread as a weed, firstly to C and N Europe and then, more recently,to other parts of the world, carried around by emigrants along with their crop seed. It has now becomecircumpolar and, indeed, almost cosmopolitan everywhere Europeans became established, although there arewide gaps in the overall world distribution.
Threats
None.
Native, very rare and probably declining, although easily over-looked. European temperate, but widelynaturalised.
1884; Barrington, R.M.; Gubbaroe Point, Lower Lough Erne.
July to September.
Growth form, preferred habitats and reproductive capacity
This rare, often minute, insignificant little annual (c. 7 cm in height), of regularly disturbed orotherwise open, damp or intermittently wet habitats and sandy or gravelly, acid soils, can very easilybe overlooked on account of its unspectacular appearance. It is confined to lowland sites in B & I,on lakeshores and very often in coastal sites, including sand dune slacks, sandy sea cliffs and alongpaths and tracks on heathland and forest rides (A.J. Richards, in: Preston et al. 2002).
The axillary flowers, which are produced from June to August, are very tiny and the pink or white petalsare shorter than the sepals, making the blossom so inconspicuous as to be easily overlooked. The flowersare self-compatible and frequently pollinate themselves while still in bud. Thus cool, wet summers withpoor levels of sunshine do not preclude a normal crop of seed being set (Salisbury 1968). The plant isslightly more noticeable when in fruit, for the swollen globular capsules, white or cream in colour andoften with a red or brownish tip, are considerably larger than the flowers. As a result of this and themoss-like scale of the plant (often only about 2.5-4.0 cm tall), there is a general recognition thatA. minima is very probably under-recorded throughout B & I (Garrard & Streeter 1983; A.J.Richards, in: Preston et al. 2002).
Having said this, the Chaffweed plant can become larger and more noticeable if, as occasionally happens,it behaves as a winter annual. In this instance, which in some situations can represent around 3% of apopulation (Salisbury 1968), the seed germinates in September or October. Autumn seedlings usuallysurvive overwinter, unless they are disturbed and they recommence growth in the spring, the lowerbranches spreading and rooting at their nodes, until the plant finally reproduces in summer. The winterannual mode of growth enables A. minima to produce a small, bushy plant up to 10 × 15 cm inheight and spread. With its growth period thus extended, the winter annual plant accumulates additionalphotosynthetic reserves in comparison with spring seedlings, enabling it to flower and fruit to anextraordinary degree. While the typical April germinating spring annual produces an average of fivefruit capsules, the winter annual can develop between 56 and 144 capsules. One exceptionally largewinter annual specimen developed 443 capsules, containing on average of 16.4 seeds (Salisbury 1968).Spring germinating individuals can occasionally exhibit similar, though less extreme, exuberance ofgrowth, if they happen to occupy very favourable ground, protected from almost all competition(Salisbury 1968).
Populations of A. minima have in the past been known to fluctuate widely in abundance at somesites, sometimes making a spectacular recovery from near absence, strongly suggesting the species candevelop a long-term dormant buried seed store that in some seasons can produce a large seedlingpopulation after sufficient disturbance brings it to the soil surface (Salisbury 1968). Having saidthis, the survey of NW Europe soil seed bank literature did not feature any mention of A. minima(Thompson et al. 1997).
Fermanagh occurrence
There are a total of 16 records for A. minima from 14 tetrads in the Fermanagh Flora Database, butit has only been seen on four occasions since 1975. Thus the current status of Chaffweed in the areaneeds further investigation before it could be definitely concluded what is already suspected – that itis a declining species in this area. The local records are from damp, gravelly and sandy ground, mainlyon the shores of larger lakes in Fermanagh and, as the tetrad distribution map shows, especially fromaround Lower Lough Erne. The details of the post-1975 records are: Muckross near Kesh, 1976, N. Dawson;gravel pit N of Derrin Mountain, 6 September 2001, RHN; Drumlish, 3 km S of Derrygonnelly, August 2002,R. Birch.
Irish occurrence
The Irish Flora (Webb 1977; Webb et al. 1996; Parnell & Curtis 2012) summarises thehabitat preference and distribution of A. minima as being, "Lake shores and damp sandyplaces near the sea. Occasional in Kerry and the extreme North; very rare elsewhere." In its albeitvery limited way, the Irish Census Catalogue lists at least one record of this species in 20 ofthe 40 Irish VCs. It also indicates (by means of brackets), that Chaffweed is considered extinct, or hasnot been observed for a long period, in five of these 20 VCs (Scannell & Synnott 1987).
The New Atlas enables us to add to this overall assessment the fact that A. minima hasrecently been found in several places in W Mayo (H27) and once in W Galway (H16), as well as in a fewother scattered, almost entirely coastal sites around the country. The small number of inland recordsshown for Ireland in the New Atlas hectad map are almost all concentrated in the northerncounties and the majority of these represent pre-1970 finds.
British occurrence
The New Atlas hectad map shows A. minima is widely, but intermittently scattered along theW & S coasts of Britain, becoming more prevalent northwards into W Scotland, including on thewestern isles. It has a limited presence inland and there is clear evidence, over the last 100 years orso, of a gradual decline towards near total absence along the E coast of England, which continues tothis day (Salisbury 1968; A.J. Richards, in: Preston et al. 2002; Sell & Murrell 2014).
Likely causes of species decline in B & I
The dwindling of this usually very small, shallowly rooted species probably involves a combination offactors that have increased plant competition. These include the move away from arable farming towards'improved' pastures throughout many areas of B & I, both for grazing and for silage-making. In turn,this entails the widespread and repeated use of slurry and/or artificial fertilisers and herbicideswhich is distributed in farm runoff and enters the general drainage water. In addition, there are theeffects of a general atmospheric nitrogen enrichment of soils from pollution sources, includingtransport vehicle emissions throughout the country. These large, widespread environmental changes, thathave been taking place throughout most of B & I during the last sixty years, very definitelyencourage the growth of larger, more vigorous plant species, raising their competitive ability to levelsof aggression, with the result that small, shallowly rooted, relatively ephemeral species like A.minima are more readily ousted than ever before.
European and world occurrence
A. minima is distributed over most of Europe, SW Asia and N. Africa. It is also present inboth N & S America and regarded as native there by some, making it an amphi-Atlantic species. Thecurrent author (RSF) considers that the N American distribution mapped by Hultén & Fries (1986, Map1482) looks decidedly patchy in comparison with that displayed in Europe and does not inspire confidencethat it represents native occurrence. On the other hand, the species is small, insignificant-looking andeasily overlooked, so the American map may only reflect poor recording. A. minima is definitelyintroduced in W Australia and probably so in other parts of the world (Hultén & Fries 1986).
Threats
Increased competition due to environmental pollution and nutrient enrichment.
Native, occasional or locally frequent. Circumpolar southern-temperate.
1900; Praeger, R.Ll.; Co Fermanagh.
April to November.
Growth form and preferred habitats
In B & I, rather limited, often, but not always transient populations of this small, thinlyscattered, rosette-forming perennial most frequently occur as a salt-tolerant species of regularly orseasonally flooded or flushed, constantly wet, brackish coastal marshland, dune-slacks and cliff ledges(Flora of Co Dublin). At the same time, S. valerandi is less frequently found at inlandsites and, in Ireland, it locally occupies suitable damp ground around many of the larger limestonelakes of the lowland West and Midlands. Likewise, in Britain, S. valerandi is occasionally foundinland, mainly along the River Severn and its feeders and around the Wash. S. valerandi mighteven be considered a member of our freshwater aquatic flora, since occasionally it grows in ±waterlogged, spring-fed, fen, flush or ditch situations (Cook 1998).
S. valerandi is a pioneer colonist of bare, muddy, clay or peat, of damp to wet, but otherwisemesic or nutrient-poor, lime-rich, but typically warm, sunny growing conditions. The species has bothexacting ecological requirements and extremely limited competitive ability. Despite this, it is tolerantof some degree of trampling and of light grazing pressure (Sinker et al. 1985). A certain levelof disturbance obviously helps keep the habitat open, limiting the growth of potential competitors. Onthe negative side, however, these very specific habitat conditions must also curtail the growth ofindividual Brookweed plants, restricting their ability to reproduce and thus limiting the frequency anddistribution of the species.
Fermanagh occurrence
In Fermanagh, S. valerandi has been recorded in 67 tetrads, 12.7% of those in the VC. However,only 56 squares have post-1975 records, strongly indicating population losses. Brookweed is anoccasional or locally frequent species, found mainly on the shores of our larger lakes and especiallyaround Lower Lough Erne. Smaller lakes where it occurs include Doagh, Carrick, Bunnahone, Ross Loughnear Carr Bridge and Carran Lough, north of Boho. It was once found in quarry pools adjacent toKeenaghan Lough and it has also been recorded at Roosky, the site of important turloughs (ie vanishinglimestone lakes).
Flowering reproduction
Although S. valerandi plants are small, usually only around 30 cm tall, they can take severalyears to flower and then most of them prove short-lived. Thus, effectively, Brookweed is a monocarpicperennial (A.J. Richards, in: Preston et al. 2002). The tiny, white or pinkish, 5 mm diameterPrimula-like flowers are produced in racemes between June and August and are usuallyautomatically self-pollinated (Clapham et al. 1962). The corolla tube is extremely short and nonectar is produced, so insect visits are rare. Average seed production has been estimated at around3,000 per plant (Salisbury 1942, p. 203).
The fact that plants can occasionally occur on wet coastal cliffs strongly implies that wind is importantin dispersal of the light seed. However, transport in mud on the feet of birds and other animals is alsosignificant, allowing the species to transfer between isolated or unconnected water bodies (Ridley 1930,p. 545).
British and Irish occurrence
Brookweed occurs scattered around the whole coast of Ireland, and in Britain it stretches northwards tothe Outer Hebrides and Orkney as a rarity. On the E coast of Britain it only reaches north as far as theFirth of Forth (Preston et al. 2002). On both islands, there has been a definite decline in thepresence of the species in the last 70 years or so, mainly due to drainage.
European and world occurrence
Frequent and widespread in coastal and temperate Europe and around the Mediterranean basin, but inland itrapidly becomes scarce, scattered and increasingly disjunct, although it does stretch eastwards into S& SE Asia. Beyond Europe, S. valerandi is very widely scattered around the world in bothhemispheres, including in Ethiopia, C & S Africa and E Australia, giving it a discontinuouscircumpolar distribution.
The form in N & S America is subsp. parviflorus (Raf.) Hult. It is sometimes regarded as aseparate species, although the differences between the two subspecies do not appear to be veryfundamental and there seem to be overlapping forms on both continents (Hultén 1971, page 148; Hultén& Fries 1986, Map 1485).
Threats
Local drainage operations and eutrophication producing increased competition.
Introduction, neophyte, an uncommon garden escape. In its native range, sub-oceanic temperate, but widelynaturalised.
21 August 1986; McMullin, A.S., Corbett, P. & Phillips, J.C.L.; Inisherk Island, Crom Castle Estate.
May to September.
Growth form, status and preferred habitats
Although in parts of Britain this shrub may perhaps be native, in their catalogue of B & I alienspecies, Clement & Foster (1994) accept it as native only, "with reservations". InIreland, R. rubrum is always considered a garden escape. Red Currant does occasionallyappear in rather drier scrub, hedgerow and roadsides situations, in which cases it is often, but notalways, near habitation. Discarded plants are also more rarely found on waste ground where tippingoccurs. Nowhere is this species likely to persist long, especially in any dry ground situations.
Fermanagh occurrence
Although not recorded in Fermanagh until 1986, there are now records of Red Currant from 22 tetrads, 4.2%of those in the VC. The shrub is uncommon and local, occurring most frequently as isolated bird-sownbushes in swampy lakeshore fen-carr woods or scrubland. It is becoming quite frequently associated withFermanagh lake islands.
It is possible that some of the Fermanagh records could be mis-identifications for R. nigrum(Black Currant), but the plants at Lurganboy near Pettigo, for instance, were in fruit and aredefinitely assigned correctly.
Irish occurrence
Along with several other Ribes species, this is one of the plants which is much less frequentlyrecorded in the RoI than elsewhere in these islands. Inspection of the New Atlas hectad maphighlights the widespread coverage Red Current exhibits in NI, and the contrasting scattered scarcity ofit south of the political border (Preston et al. 2002).
Seeing this, it is difficult to avoid the conclusion that some Irish recorders must still followPraeger's example (eg in Irish Topographical Botany) and deliberately overlook this species. Thismay be an unfair, out-of-date suggestion, however, since the spread of post-1986 symbols on the NewAtlas map clearly shows that R. rubrum has been more frequently recorded throughout thewhole of Ireland in more recent years. Also, while the Irish Census Catalogue (Scannell &Synnott 1987) listed this shrub as having been recorded at least once in just 13 VCs, in her more recentAlien Catalogue, Reynolds (2002) increased that figure by a further ten, without either referenceincluding Fermanagh!
Threats
Bushes occasionally naturalise themselves in woodland, but Red Currant is insufficiently aggressive topose any invasive threat.
Introduced, neophyte, garden escape or doubtfully native. Frequent, widely and rather evenly spreadthroughout the county.
1947; MCM & D; Lough Bigwood, 1 km NNE of Rossharbour Bay, Lower Lough Erne.
March to October.
Growth form, status and preferred habitats
In the Revised Typescript Flora, Meikle et al. 1975 stated that in Fermanagh (and thus byimplication in Ireland in general), "this species is too readily assumed to be a gardenescape". Here in Fermanagh, as elsewhere in B & I, R. nigrum is a frequently found andvery characteristic shrub growing up to 2 m in damp or even swampy fen-carr scrub woodland, hedgerowsand shaded streamsides. It is very easily distinguished at all seasons by the strong perfume of its budsand leaves.
Earlier standard Floras of B & I (eg Clapham et al. 1962) and some recent local Floras in S& SE England have regarded this strongly smelling currant as being of mixed native and introducedstatus (eg Brewis et al. 1996; Beckett et al. 1999). In Cumbria, Halliday (1997) appearsto have allowed its native status in every station. At the same time, arguing against this point ofview, we know from the researched history of the garden form of the plant, that it was imported toBritain from Holland shortly after 1600 and first appeared in the wild in 1660 (Roach 1985). RecentFloras now recognise this species is a neophyte introduction and accept that plants found in the wildare of garden origin, often bird-sown (Sell & Murrell 2014).
Fermanagh occurrence
R. nigrum is over five times more frequently recorded in Fermanagh than R. rubrum(Red Currant), both fruit currant bushes occurring in exactly the same types of semi-natural lakeshorehabitats. In Fermanagh, R. nigrum is now recorded in a total of 98 tetrads, 18.6% of those in theVC and it is remarkably evenly spread throughout the whole area. Robert Northridge and the currentauthor (RSF) feel that this level of presence and its distribution suggests dual status (nativeand naturalised introductions) for Black Currant may deserve some further consideration. Thestrictures on weighing circ*mstantial evidence listed by Webb (1985) must however be remembered andinvoked in any analysis of such weighty matters (Forbes 2000).
Irish occurrence
As is the case with other representatives of this genus, the New Atlas distribution of R.nigrum in the RoI, while certainly quite frequent and widespread at the hectad level ofdistribution, at the same time remains rather sparse when compared with that north of the border in NI.This appears anomalous and requires investigation.
British occurrence
Very frequent and widespread throughout Britain in suitable damp or wet shaded sites,avoiding mainly the higher mountains and very acid peatlands (New Atlas).
European and world occurrence
Present in most of Europe, stretching northwards into Scandinavia and E to Siberia. Also, absent frommost of the Mediterranean basin, but probably native only in C & E Europe and N & C Asia to theHimalayas. Biogeographically it is described as belonging to the Eurosiberian Boreo-temperate element,but it is widely introduced, cultivated and naturalised outside its native range (Sell & Murrell2014).
Uses
Dried leaves can be used to make a herbal tea (like green tea) and is a weaker alternative to the moreusual one made from raspberry leaves used for easing labour pains in childbirth. The leaf tea was alsoused as a diuretic. Folk records of the plant's use are dominated by blackcurrant juice, made from freshor jellied fruit, which is very often prescribed for coughs, colds, chest complaints and gargles forsore throats (Grieve 1931; Roach 1985; Allen & Hatfield 2004). Nowadays, numerous named blackcurrantvarieties have been produced in government funded horticulture research facilities designed to give highvitamin C content, resist aphids, ease harvesting and provide later season harvesting. Hybrids are nowalso being bred that will adapt the crop to changes in our climate. These forms are grown commerciallyin England to make the popular cordial 'Ribena'. In the past, blackcurrant was also used to flavour andcolour wine, brandy and cheese and it is still used to make puddings, jelly and jam (Roach 1985; Grieve1931).
In addition to herbal medicine and juice production, there are several decorative horticultural varietiesof R. nigrum named, including forma apiifolium with dissected leaves; f.chlorocarpum with green fruits; f. coloratum with a white variegation; f.heterophyllum with leaves deeply cleft; f. marmoratum with leaves deeply cut and marbledcream and f. xanthocarpum with a fruit that is yellow to white in colour (Griffiths 1994).
Threats
None.
Introduction, neophyte, a rare, naturalised garden escape.
3 June 1993; EHS Habitat Survey Team; 'Wood 3' at Knockninny hill.
March to June.
Growth form and preferred habitats
There are just nine records for this now rather over-commonly planted, large (up to 4 m), decorative,pink- (or less commonly white) flowered garden shrub from the wild (or non-garden sites) in this survey,all dating from the post-1992 period. This fact tells us less than it might, since we know that Meikleand his co-workers and, indeed, originally Robert Northridge and the current writer, disdained recordingmost garden escapes. Meikle and company also ignored most agricultural and horticultural weeds duringtheir botanical outings around Fermanagh.
As listed above, the first record was made in or on the margin of the limestone woodland on Knockninny in1993. It now appears thinly scattered in nine tetrads in lowland Fermanagh, both near and remote fromhouses, in hedges, woods, roadside verges and on waste ground. In the latter two situations, it may havebeen deliberately dumped with other garden waste. It undoubtedly remains under-recorded to some extent.
Origin and introduction
Flowering Currant was introduced to gardens in B & I from NW America by the famous Scottish plantcollector David Douglas in 1826 (J.M. Croft, in: Preston et al. 2002). It was first recorded inthe wild in 1916, and since it tends to occur as isolated shrubs and produces a white-bloomed,blue-black fruit, it undoubtedly is often bird sown in the variety of habitats listed above, near towhere the vector perches!
Fermanagh occurrence
The details of the later eight Fermanagh records are as follows: near an old house, E of Gubbaroe Point,Lower Lough Erne, 18 April 1998, RHN; Clonelly, NW of Kesh, 17 March 1999, HJN & RHN; Armagh Manor,possibly planted, 9 April 1999, RSF & RHN. The last three are dated only 1987-99 and they are:Tullyweel, near Creevehill House, W of Fivemiletown, I. McNeill & D. McNeill; hedgerow at Knockroe,NE of Ederny, I. McNeill; Lurganboy, 7 km E of Kesh, I. McNeill; Roogagh Bridge, 28 July 2004, RHN &HJN; and Glen Lodge, 4 August 2007, I. McNeill.
British and Irish occurrence
R. sanguineum is widespread but patchily distributed in lowland parts of both B & I, commonin urban areas with higher concentrations of gardens, and possibly more frequent in the N & W(Clement & Foster 1994). As is the case with all Ribes species, it appears from the hectadmap in the New Atlas to be more commonly and widely recorded in NI than in the RoI, althoughReynolds (Cat Alien Pl Ir) lists stations from eight VCs south of the border with NI. Again, thisis very probably an artefact resulting from the reluctance of older or past recorders to list and givecredence to mere garden escapes.
Threats
None.
Introduction, neophyte, a local and fairly uncommon garden escape or discard. Indigenous to the Europeantemperate region, but widely cultivated and naturalised.
1 August 1986; Corbett, P. & Brain, P.J.T.; Inishroosk Td, Creaghanameelta Island shore, Upper LoughErne.
April to November.
Growth form and preferred habitats
This very familiar deciduous, much-branched, fruiting garden shrub 0.9-1.5 m tall, with its slender butvery sharp 3-spined thorns at each node and hairy berry fruit, is native to S, C & W Europe and aneophyte introduction throughout B & I. It usually occurs in damp or shaded ground fairly close tohouses – including around ruins or growing on old walls, facts strongly indicating and reflecting itsgarden origin and bird sown dispersal. At the same time it is often very well established, naturalisedand occasionally it is even locally abundant in hedges, stream sides, scrub and deciduous woodland. In awoodland setting, R. uva-crispa can deceptively, 'look, quite native'. In waste ground settings,found far too often, it very obviously is discarded garden material.
Flowering reproduction
The shrub produces its pendulous flowers in groups of 1-3 between March and May. Nectar is secreted atthe base of the bell-shaped receptacle and is protected by stiff hairs projecting from the style(Hutchinson 1972). Cross pollination is achieved by various visiting flies, bees, butterflies and moths(Clapham et al. 1962). The fleshy fruit, 10-20 mm, ripen in July and August and are probablytaken by Blackbirds and possibly other members of the thrush family, although strangely there are veryfew sightings recorded in the bird literature of this actually happening (Snow & Snow 1988).
Origin, introduction and uses
R. uva-crispa, previously known as R. grossularia L., is native in many parts of Europeincluding the Caucasus and N Africa. Forms were selected, brought into cultivation and bred in Europemuch earlier than in B & I. The earliest record of their use in Britain dates from a list of treesand shrubs supplied from France to King Edward I in 1275 for planting in the garden at the Tower ofLondon (Roach 1985). Gooseberry was not recorded in the wild until 1763, when it was listed forCambridgeshire (Perring et al. 1964).
There are said to be over 2,000 named varieties (possibly a gross over-estimate, but certainly a largenumber), grouped by fruit colour as red, green, yellow and even a 'white' (pale green) form (Vickery1995; Mabey 1996). Green varieties are less sour and are more suitable for making wine and desserts,while red berried varieties are the most acidic. Berries are capable of fulfilling a wide range of uses,including as sauces and chutneys for various meats, jam, tarts, desserts, wine and brandy making. Theyare amongst the first fruit to crop in the year, but they often are sour tasting and require a greatdeal of sweetening with sugar to make them palatable. The abolition of the English sugar tax in 1874 ledto a dramatic increase in the commercial growing of gooseberries. The public demand for jams of allkinds has greatly diminished over the last 90 years, and the use of gooseberries for pectin as a settingagent for confectionary and jam making has also disappeared as better sources for this purpose weredeveloped. Nevertheless, breeding research did go on in England to try to produce spine-free anddisease-resistant varieties, with some success in the latter case (Roach 1985).
Despite a quite marked decline in its garden cultivation since 1945, the New Atlas records asignificant increase in its presence in comparison with the earlier BSBI Atlas survey published in 1962,probably reflecting further garden escapes, seed often being transported by birds, plus a greaterinterest in recording domesticated plants in the wild than previously was the norm (J.M. Croft, in:Preston et al. 2002).
Fermanagh occurrence
Gooseberry is now known to be a local, quite widespread, if rather infrequent shrub in Fermanagh woods,scrub and hedgerows, particularly on drier soils. In Fermanagh, R. uva-crispa has been recordedin 39 tetrads, 7.4% of those in the VC. As indicated above, all our records are post-1985 in date, sinceneither Praeger and his generation of recorders, nor Meikle and his co-workers in the 1940s and 1950s,took any notice whatever of this plant during their surveys (Irish Topographical Botany;Revised Typescript Flora). It is thinly scattered throughout the VC, but more prevalent andoccasionally more abundant, in the better soils in eastern Fermanagh.
British and Irish occurrence
The New Atlas hectad map shows R. uva-crispa is common and widely distributed throughoutlowland B & I, but much more frequently recorded in B & NI than in the RoI. It is largely absentfrom the ecologically unsuitable wetter, exposed, peaty ground of N & NW Scotland, and rare orabsent also from the Scottish isles. The same edaphic reason may well explain its scarcity in W Irelandat least.
Names
The name 'Gooseberry' may well derive from the use of the berries to make a sauce to accompany goosemeat. It might also mean a berry that was eaten by geese on farms. The name is not recorded before about1532 (Grigson 1987). Another English common name, 'Feaberry', may refer to a fruit from a prickly bush,a 'thēfe' in Old English (Grigson 1987).
Threats
None.
Native, occasional, but locally frequent. Mediterranean-Atlantic.
1864; Dickie, Prof G.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Navelwort is an evergreen, fleshy perennial, generally considered native to the western parts of B &I where it is very much more frequently found. It typically grows in damp hedgerow banks and in and onold walls, often along roadsides. Perring & Walters (1976) regarded U. rupestris as anexample of a winter- and spring-growing evergreen species, which tends to become dormant during the heatof high summer. Species like this one are not necessarily frost-sensitive, but they probably arephysiologically weakened by it to some extent and subsequently fail to compete successfully after coldwinters.
Flowering reproduction
The normal flowering period of U. rupestris is from June to August, but in milder conditions thisstretches into September (Clapham et al. 1962; Webb et al. 1996). In the usually drier,sunnier parts of S & E England, plants may be burnt to a crisp by July temperatures. Plants inDerbyshire (VC 57) have been observed to flower earlier and they may complete their sexual cycle asearly as June (Hodgson 2002).
Fermanagh occurrence
In Fermanagh, U. rupestris is widely but rather thinly scattered and only occasional throughoutthe county, although it can become very locally frequent on suitable damp roadside banks and walls. Ithas been recorded in 56 tetrads, representing 10.6% of those in the VC. Although the distribution mapindicates that it has not been seen in seven Fermanagh tetrads since the MCM & D survey of 1947-53,in every case it has been found in their near vicinity and we see no decline in the species presence.Usually regarded as a calcifuge, preferring, but not strictly limited to acidic growing conditions, wefind it recorded in limestone terrain in Fermanagh, eg at the Cladagh River Glen NR (also known as theMarble Arch) and at Hanging Rock NR.
British and Irish occurrence
The distribution of U. rupestris in B & I is broadly similar to that of the much rarer speciesRubia peregrina (Wild Madder) and Orobanche hederae (Ivy Broomrape) (Lousley 1969),although the current author considers its distribution more closely overlaps that of Ceterachofficinarum (Rustyback) (Perring & Walters 1976; New Atlas). The strongly markedAtlantic distribution of U. rupestris in these isles closely follows the plot of the 1.7°C(35°F), February minimum isotherm on the overlay supplied with the BSBI Atlas. This isotherm wasbased on average means of daily minimum temperature in February from 1901-30.
Visual comparison of the hectad maps for Navelwort in the two BSBI atlases suggests there has been littlechange in the distribution of U. rupestris at this scale of resolution during the past 40 years,an impression confirmed by the calculated Change index value being as low as -0.12 (Perring &Walters 1976; Preston et al. 2002). In an interesting note, Edgington (2002) pointed out thathistorical published records in the London area stretching back to John Gerard(e) (1597), indicate howthe eastern occurrence limit of U. rupestris in SE England has undoubtedly varied over the yearswith fluctuations in winter temperatures. Thus in central London (VC 21), the species has benefittedfrom mild, almost frost-free winters (and cleaner air quality) during the last 20 years, to the extentthat it may now be observed growing, flowering and fruiting on the decaying Victorian brickwork of mossywalls above damp basem*nts around buildings, and even appearing in more exposed situations growingaround roofs.
The extension of the species eastwards in C & SE England has thus additionally been dependent uponwetter summer conditions, something which has been noted occurring in recent years.
Names
No less than 33 English common names are listed by Grigson (1987). 'Navelwort' was identified with the'Kotuledon' of Dioscorides (the father of ancient medicine), the 'Umbilicus Veneris' or'Venus's Navel' of the Romans. In English, the latter name became 'Lady's Navel' and Englishdoctors followed Dioscorides in prescribing it as a cure for stones, cuts, chilblains and inflammation(Grigson 1987). Many of the other English names involve 'penny', as in 'Penny Cake', and obviously referto the leaf shape.
Threats
None.
Introduction, neophyte, deliberately planted, rare, declining and probably locally extinct.
1902; Abraham, J.T. & McCullagh, F.; Drumclay a district of Enniskillen, walls and roofs.
Growth form and preferred habitats
A long-lived evergreen succulent perennial, very tolerant of drought, S. tectorum is a native ofthe mountains of C & S Europe and of the Greek islands introduced to B & I around the 13thcentury. It flowers sparingly in July, producing a tight cluster of blooms on the top of a leafy stem30-60 cm tall. The flowers are a red-purple colour, 15-30 mm in diameter. The flowers contain nectar andattract various insects. The fruit is a dry collection of follicles. Irrespective of seed production,vegetative offsets from the basal leaf rosettes readily spread the plant in its site.
House-leek appears to have declined greatly in popularity as a subject for cultivation on cottage orouthouse roofs, walls and in rock gardens during the second half of the 20th century in comparison withearlier centuries. Even in Victorian and Edwardian times, S. tectorum was still regarded as alucky charm, able to ward off evil spirits and prevent catastrophic events such as house fires,lightening and thunderbolts (Vickery 1995). Unlike in some other areas in B & I, in Fermanagh thissucculent perennial does not survive for long on deserted ruins. The Cat Alien Pl Ir suggestedthat only old records exist in Ireland, but the New Atlas map plots eight scattered hectads withpost-1986 Irish records.
Fermanagh occurrence
Nevertheless, despite the above traditions, evidence of the decline of this species is readily observedin Fermanagh. Between them, Abraham & McCullagh in 1902 and Meikle and his co-workers in the 1946-52period, found at least ten stations for S. tectorum in the VC. From the site names, somemight just possibly have been garden escapes actually naturalised and growing 'in the wild'. However,the current authors have never observed this distinctive plant growing outside garden confines anywherein Fermanagh, and there are no post-1952 records at all from the VC.
In the 1975 Revised Typescript Flora, Meikle et al. listed a 1902 find by Abraham &McCullagh of this species as the "Bar of Whealt" another name for a townland that includes thewestern end of the Cliffs of Magho which was published for this pair of botanical friends by Praeger(1903a). As there are no buildings or walls along the cliff top area, it is highly unlikely to be thesite meant. Site names of the Victorian era, and indeed up until map grid references came into use in VCbotanical recording around 1967, sites are often really difficult to pin down. Also, modern 1:50,000maps are quite often printed with rather different site names from older one inch and larger scale maps.It seems probable that in this instance 'Bar of Whealt' really referred to houses or outbuildings aroundthe base of the escarpment, on farms on the slopes overlooking Lower Lough Erne. In the post-warperiod, Meikle and his friends also tended to be rather vague about site names, eg listing a 1948station of their own for this species simply as, "Near Lattone Lough". Having said this, bothPraeger and Meikle describe the habitat for all twelve S. tectorum records as being "onwalls and roofs".
British and Irish occurrence
In their comprehensive book on Aliens Plants, Clement & Foster (1994) remarked that whilepreviously S. tectorum was a long-established introduction on old walls and cottage roofs, it isnow decreasing and becoming rare. The related S. montanum introduced from C & S Europe hasfared even worse. The latter reference indicates that only pre-1930 records exist for it.
The New Atlas hexad map indicates that S. tectorum, which is native to S Europe, stillremains widespread and reasonably frequent in the southern half of Britain, thinning northwards andbecoming increasingly coastal as it does so. Overall, the presence of House-leek is definitely waningthroughout these islands (Preston et al. 2002). The current author feels it rather sad that agarden plant that has been in cultivation since at least 1200 A.D. (Harvey 1981), and which was firstrecorded in the wild as early as 1629, should fade away through changing mores and a lack of faith inits life preserving and medicinal properties. The replacement of many thatch roofs by slate, which tookplace in the early 20th century has obviously also been a very significant factor, detrimental to thegrowth and survival of this species.
Medicinal uses
Apart from its supposed magical property safeguarding attributes already mentioned, S. tectorumwas also widely used in traditional herbal medicine. The fleshy leaves are astringent and diuretic andwere used to poultice corns. An application of the sap was said to ease the pain of burns and any othersore place, including lips, ears and eyes. A boiled extract was also used to treat ringworm and warts,and it was recommended for a number of women's complaints, including abortion (Grieve 1931; Vickery1995; Darwin 1996).
Threats
None.
Native, very rare. Circumpolar arctic-montane. It is also in garden cultivation and occasionally escapesin Britain, if not also in Ireland.
5 July 1904; Praeger, R.Ll.; western end of the Cliffs of Magho.
Growth form and preferred habitats
A grey-green (glaucous), leafy, tufted succulent perennial up to 30 cm tall with a thick, fleshy,branched stock. The stems are often flushed with purple. The small, dull yellow flowers borne indense, flat terminal clusters are unisexual and are produced on separate plants (ie dioecious = 'twohouseholds'). S. rosea appears to prefer inaccessible crevices and rock ledges on damp to wetcliffs, often but not always of base-rich or limestone geology.
Fermanagh occurrence
Although Robert Northridge and the current author have made several searches in the appropriate cliffsand surrounds for this 'mountain plant', all attempts to rediscover the solitary reported Fermanaghstation of this familiar and distinctive succulent species have to date been completely unsuccessful.However, the area involved is large and of very difficult access, so it is conceivable that it might yetsurvive somewhere along the cliffs where Praeger recorded it in 1904. In his wonderful account of hisvery energetic five days spent, Among the Fermanagh Hills, Praeger (1904) wrote almost casuallyof finding this species, "At one spot near the summit of the Poulaphouca western cliffs." TheCliffs of Magho and Poulaphouca are alternative names for this range of precipices, which, whenever aclear day allows, provide a fine overview of Lower Lough Erne to the north.
Irish occurrence
The next nearest known station for Roseroot is on the Carboniferous limestone cliffs of the Ben Bulbinrange in Cos Leitrim and Sligo (H29 and H28), around 12 km due west. In these latter stations, thespecies is locally abundant and the New Atlas map indicates that a number of post-1986 recordsexist. It is also well represented in W Donegal (H35) both on mountains, on sea cliffs and even in damprock crevices right down to high-tide level on west-facing Atlantic beaches.
Elsewhere in Ireland, Roseroot has previously been recorded on most of the higher mountains, nearly allof which are coastal. In NI, this includes the Mourne Mountains in SE Co Down (H38) and the NE Co Antrimcliffs, around Fair Head (H39). In several of its other Irish stations, S. rosea has becomescarce or even rare during the last four or five decades, and in others it has not been seen at all forquite some years. The latter category includes, in addition to the Fermanagh site, the Wicklow hills(H20), and several of its previous N Antrim sites.
British occurrence
The New Atlas hectad map shows S. rosea is predominantly a mountain and coastal plant of NWScotland and the Scottish Highlands & Islands, although also present on the higher mountains ofEngland and Wales. A small number of isolated stations scattered about in England and Wales arepresumably garden escapes.
Flowering reproduction
The fact that S. rosea has separate male and female plants and is thus a ± obligatory out-breeder(dioecy is rarely absolute), may become a detrimental factor seriously affecting seed production andendangering species survival whenever isolated populations decline to small numbers (Richards 1997a). Wedo not know the size of the minimum viable population necessary in the short term to protect againstinbreeding depression, but in animals, which have a great deal more mobility than plants, the equivalentfigure is often taken as 50 (Briggs & Walters 1997, p. 417). Any imbalance of the sexes wouldexacerbate this type of numerical problem. Clearly populations of both rare plants, and more isolatedpopulations of scarce species, are endangered by any degree of further decline.
Species like S. rosea, which are relict survivors from past colder climatic phases in our islandhistory, are also increasingly threatened by recent trends in global warming.
Threats
Genetic erosion in small, isolated populations, plus rapid modification of the environment associatedwith climate change.
Introduced, neophyte, a very rare, naturalised garden escape, possibly only a casual and locally extinct.Eurasian temperate, but widely naturalised, including in N America.
1939; Praeger, R.Ll.; in woods on Inisherk Island (also known as 'Garden Island'), Crom Castle Estate.
Fermanagh occurrence
There are six old records for this distinctive, cultivated, perennial succulent from wild or semi-wildsites in Fermanagh, all dating from the period 1939-52 (Revised Typescript Flora). All but thefirst appear fairly remote from local gardens and were found on the sides of tracks and roads, or on ornear lakeshores, possibly in fairly dry, shade or semi-shade of woodland or scrub margins. Apart fromthe first given above the record details are: railway track and roadside near Kesh, 1942, R. Mackechnie;Lower Lough Erne shore near Lisnarrick, 1942, R. Mackechnie; by lane on slopes of Slieve Rushen, 1949,MCM & D; and roadside N of Monea, 1952, MCM & D.
When, as at Crom, the plant occurs in deeper woodland shade, it seldom if ever flowers. The speciescannot compete with taller grasses and it is also intolerant of grazing and trampling (Sinker etal. 1985). The fact that this stonecrop has not been recorded in Fermanagh by anyone for over 50years, strongly suggests that as a garden subject this species has become unfashionable and locally itis extinct. It has been replaced in the great majority of gardens in NI by the superior decorativeattractions of S. spectabile (= Hylotelephium spectabile (Boreau) H. Ohba) (Ice Plant).The latter, introduced in 1868 from China and Japan, is currently very popular since it attractscolourful butterflies to the garden in late summer and autumn. Several rather differenthorticultural varieties of it are readily available, making it more enticing than S. telephium(Griffiths 1994; J.M. Croft, in: Preston et al. 2002).
Irish occurrence
Orpine still does occur rather rarely elsewhere on the island of Ireland from time to time, principallyin the NE, although S. telephium s.l. has been found at least once over the years in 32 Irish VCs(Reynolds 2002). A measure of frequency is provided by the fact that the survey of urban Belfast foundOrpine in just two 1-km squares, one of which was a rubbish dump (Beesley & Wilde 1997).
The most likely status of S. telephium subsp. fabaria in Irelandas a whole is as a short-term or even casual garden escape or discard, rather than a fully naturalised,established and long-persistent plant (FNEI 3; Cat Alien Pl Ir).
Threats
None.
Introduced, neophyte, a very rare garden escape, now locally extinct. Also a possible mis-identification.
1902; Abraham, J.T. & McCullagh, F.R.; Bolusty More Farm.
Fermanagh occurrence
This evergreen, succulent perennial, originally recorded by Abraham and McCullagh as S. reflexumL., was introduced to gardens from the Caucasus in the 16th century as a salad leaf crop. In the 1975Revised Typescript Flora, Meikle and his co-workers suggested this solitary old recordmight be a mistake for the rather similar S. forsterianum Sm. (Rock Stonecrop). Certainly therehave been many such identification errors in the past (Cat Alien Pl Ir), but in the absence of avoucher there is no evidence either way and Robert Northridge and the current author are provisionallyallowing this solitary record to stand.
In FNEI 3, Hackney highlighted this same identification problem, but went on to remark thatin their area S. rupestre (= S. reflexum L.) is, "certainly [the] much commoner [ofthe two], and is very frequently cultivated, often establishing itself on garden walls". [This maybe the case, but see a relevant caution made in S. forsterianum below.] In the same work, Hackneyalso observed that in NE Ireland, S. rupestre only rarely naturalises itself in truly wildsituations. This was definitely the case with the Fermanagh record, the site of which was originallydescribed as, "Bar of Whealt, well established on houses" (Praeger 1903a). 'Bar of Whealt' isan alternative name for the area around the cliffs of Poulaphouca (sometimes spelt Poulaphuca andPollaphuca) and also referred to as the Cliffs of Magho. The name encompasses the slopes below thecliffs where Bolustry More Farm lies, the ground stretching down to the southern shore of Lower LoughErne.
Nowadays, the physical characters that distinguish these two stonecrop species are much betterunderstood: S. rupestre has living leaves well spaced, not bunched at the end of the shoots.Also, dead leaves are not persistent, as is the case in S. forsterianum. Furthermore, the leaveson flowering stems are spreading or bent-back, as opposed to being held erect (An Irish Flora1996).
British and Irish occurrence
Comparison of the New Atlas maps of these two stonecrops certainly shows that in Ireland at thehectad level of discrimination, S. rupestre is slightly more frequent and widespread thanS. forsterianum. In Britain, S. rupestre is very much more common and widespread than isthe case in Ireland. South of a line between Southport and Bridlington, the current author estimates itmust be represented in comfortably over 80% of the available map hectads (Preston et al. 2002).
Introduced, neophyte, a very rare garden escape, perhaps only casual.
1949; MCM & D; on bridge over Woodford River, at Aghalane.
Growth form and British and Irish status
This succulent, grey-leaved, mat-forming stonecrop is regarded as native in parts of Wales and SWEngland. Elsewhere in B & I, it is cultivated and is widespread at least in lowland Britain as anaturalised, sometimes persistent, garden escape. It is recorded much more rarely in Ireland, however,and although sometimes plants here survive beyond garden confines for up to five years, they probablyfail to reproduce, and thus are only casual (New Atlas).
There has been and remains considerable confusion between this species and the very similar introduced,cultivated and escaped S. rupestre L. (= S. reflexum L.) (Reflexed Stonecrop), to theextent that, in FNEI 3, Hackney issued a caution with regard to the reliability of older recordsin the three VCs of NE Ireland (H38-H40). Following this, Reynolds (Cat Alien Pl Ir) re-examinedIrish herbarium material and discovered that most 19th and 20th century records of S. rupestreare not the Linnaean species, but really refer to S. forsterianum.
Fermanagh occurrence
In almost all other Irish VCs where records of S. forsterianum have been published, details ofonly one or two stations are reported (eg FNEI 3; Cat Alien Pl Ir). In Fermanagh, we areabove the Irish VC average in having a total of four records. The details of the other three recordsare: road between Tempo and Brougher Mountain, 1953, MCM & D; by main Enniskillen road, nearCoollane, July 1988, RHN; and Lisbellaw Old Quarry, 3 July 1994, RHN.
Locally, Rorbert Northridge and the current author (RSF) regard S. forsterianum as a very rare andonly casual garden escape, typically occupying dry and droughted, rocky or stony habitats. It is reputedto be most typically associated with dolerite and other basic igneous rocks producing thin, dry andmoderately acid to neutral soils of low nutrient status. In common with other small Sedumspecies, it is a poor competitor with tall grasses and herbs, but is very tolerant of drought anddisturbance (Sinker et al. 1985).
As the English common name suggests, Rock Stonecrop is quite commonly grown in old-fashioned rock oralpine gardens. In the current author's experience of this, it sometimes remains stubbornly vegetative,completely failing to produce any of its yellow, star-like flowers. In churchyards and cemeteries, S.forsterianum is also quite frequently planted on gravel-covered graves, probably on account ofits low carpet-forming, grey foliage, its drought resistance, and consequent ability to surviveconsiderable periods of neglect growing on undisturbed stony surfaces with very little soil.
Native, scarce or occasional only. European temperate, but widely naturalised including in N America andNew Zealand.
1884; Barrington, R.M.; Enniskillen Town.
June to August.
Growth form and preferred habitats
This distinctive, low-growing stonecrop with its perennial fibrous roots, creeping stems and tufts orcushions of annual flowering shoots is the most common and widespread succulent species in B & I,being found in every VC (New Atlas). Nevertheless, it is quite rare and thinly scattered inFermanagh, where it is found in a range of typical open, dry, shallow and occasionally droughted soils.It grows in sandy and rocky lowland natural or semi-natural habitats of neutral or calcareous reaction.It is also a pioneer colonist of open, man-made habitats, including crevices and the tops of walls orroadside kerbstones in both rural and urban settings. S. acre produces clusters of bright goldenyellow, star-like flowers in June and July on often dwarfed branching shoots, and is propagated by itsextremely light seeds (each weighing less than 10 millionths of a gram), or by leaves or other fragmentsof the plant being transported by wind (Salisbury 1964, p. 305; Ridley 1930, p. 29).
Fermanagh occurrence
Altogether S. acre has been recorded in 15 scattered Fermanagh tetrads, ten of them with post-1975dates. Although it is reputed to be sometimes, or perhaps frequently, deliberately cultivated in gardens(Grime et al. 1988; Grey-Wilson 1989), Robert Northridge and the current author (RSF) have neverobserved this anywhere in Fermanagh. S. acre is slightly more frequent around the limestoneshores of Lower Lough Erne, where it is sometimes found lodged on tops of large boulders.
An ecological study of plants in and around the Sheffield area concluded that Biting Stonecrop, which isslow-growing, is more or less restricted to undisturbed, unproductive, infertile, often rockysituations, which often have little or no soil (ie skeletal soils) (Grimes et al. 1988). In viewof this, these workers classified the ecological survival strategy of S. acre as astress-tolerator, meaning it avoids biological competition by surviving in a far from ideal growingenvironment. In maritime areas of B & I, S. acre is very commonly met on open,nutrient-depleted, species-rich grasslands, sand-dunes and shingle, sometimes forming quite substantialpatches. The survey of soil seed banks in NW Europe listed seed of all species of this genus astransient or indeterminate (Thompson et al. 1997).
British and Irish occurrence
The New Atlas map displays many hectads in Ireland, W Scotland and Cornwall with only pre-1970records, a fact that suggests this stonecrop species may be in decline.
Uses
The English common name refers to the very acrid, toxic sap obtained from the plant leaves, whichstrongly deters herbivores. The sap is a notable skin irritant and its pungency gave rise to alternativeEnglish names such as Wallpepper and Wall Ginger (Grieve 1931; Cooper & Johnson 1998, p. 69). Inherbal medicine, it was an ingredient of a famous worm-expelling treacle or vermifuge. It was alsorecommended for treating scurvy and useful for intermittent fever and dropsy. Great care is required interms of dosage, as it can easily cause inflammation and blisters when applied externally (Grieve1931).
Threats
None.
Introduction, neophyte, an established garden escape, occasional, locally abundant and probablyspreading. Submediterranean-subatlantic, but widely naturalised, including in N America.
1948; MCM & D; ruin on slope of Drumbad mountain.
February to November.
Growth form and preferred habitats
This creeping, evergreen, mat-forming perennial is a quite common garden escape, very widely scatteredthroughout B & I. It is mainly established on the mortar or the stone of walls, but is also presentin other open, dry, lime-rich habitats, including on rocks, gravel and stretches of concrete, especiallywhen the latter is cracked. S. album has great vegetative and flowering vigour and it roots veryreadily, enabling it to rapidly colonise open ground. In NI, it has spread very considerably during thelast 70 years and occasionally it becomes locally abundant (NI Vascular Plant Database 2005).
Fermanagh occurrence
In Fermanagh, S. album has been recorded from a total of 25 tetrads (4.7%), 22 of them withpost-1975 records. White Stonecrop is now thinly and widely scattered throughout the VC in a range ofdry artificial or disturbed wayside habitats, but locally this patch-forming succulent occurs in itsgreatest profusion on the cracked, concrete flying-boat slipways of the Second World War period on theshores of Lower Lough Erne. It has spread from these to adjacent runways and to other slipways and stonyground, where people have most access to the shores of Lough Erne.
A small minority of stations in Fermanagh are quite remote from houses, eg on tracks at Brougher Mountainand in several old quarries, which suggests that S. album is becoming thoroughly naturalised andthat it has considerable powers of dispersal. The tiny, very light, seed produced in abundance from amany-flowered, flat-topped, branched cyme inflorescence is undoubtedly readily transported by wind,evidence of which is clear from the presence of the plant high on walls and on cliffs (Ridley 1930, p.29).
Irish occurrence
In Ireland, this introduction is a certain neophyte, being first noted by Moore & More (1866) fromthe wild where it was growing on cliffs in Cork and on rocks at Fermoy. Reynolds (2002) has cataloguedits spread in Ireland. It was reported from a total of 17 Irish VCs in ITB (Praeger 1901), andlater from a further ten VCs by the early 1930s in his book The Botanist in Ireland (Praeger1934). The 1987 Irish flora Census Catalogue listed the presence of S. album in every VCexcept Monaghan and Fermanagh (H32, H33) (Scannell & Synnott 1987). With the publication of TheFlora of Co Fermanagh, Monaghan stands alone.
British occurrence
In England and Wales, S. album is a common and widespread archaeophyte, present almost throughout,while in Scotland it is very much more scattered, records having a definite eastern preponderance.However, it thins northwards noticeably beyond the Glasgow-Edinburgh conurbations, while reaching Orkney(Preston et al. 2002).
Threats
None, since it almost always occurs in artificial, man-made habitats.
Either a mis-identification or a garden escape. Oceanic temperate, but also cultivated and naturalised.
Growth form, Fermanagh absence and preferred habitats
The New Atlas hectad map for this distinctive low-growing, succulent, evergreen perennial displaystwo older period symbols in Fermanagh with records: a pre-1970 record near the SE county boundary withCavan (H30) and a 1970-86 record in the NW, close to the border with E Donegal and Tyrone (H34, H36). Asfar as RHN and RSF the current VC recorders are aware, these plots are errors and we cannot account foreither of them. Neither Meikle and his co-workers, nor we, have ever come across this species to ourknowledge during our recording outings. Furthermore, we believe the 1987 Cen Cat Fl Ir 2 iscorrect in omitting Fermanagh from the list of VCs with records, being one of eight Irish VCs whereS. anglicum has or had not been found.
Despite its specific name, S. anglicum is regarded as a native plant throughout both B & I,typically occupying dry, base-poor grassland or pockets of thin peat or organic soil with mosses onsiliceous acidic rocks, most often near the coast. It grows and competes best in damp to moist, butintermittently dry, strong to moderately acid soils, in sunny, exposed situations. Consequently, it is apioneer colonist of flat rocky surfaces and shallow turf in upland areas of high rainfall (Sinker etal. 1985), habitats that are well represented in Fermanagh. It is also said to frequent oldwalls, rocky hedgebanks and acidic substrates in quarries and mine spoil heaps (J.M. Croft, in: Prestonet al. 2002).
Garden escapes
S. anglicum is sometimes grown in gardens and very occasional escapes from cultivation are knownto occur, particularly at inland sites. The New Atlas map also displays two hectads with recordsin the extreme south of Ireland, which are designated as garden escapes, and in Britain many more suchgarden escapes or introductions are plotted, particularly in inland parts of C & SE England.
British & Irish occurrence
Apart from the far SW of Ireland where S. anglicum is more generally present, the species has avery pronounced circum-coastal distribution in the remainder of the island with just a thin scatteringof additional inland records. The latter include hectads in the northern VCs of Cavan, Monaghan andTyrone (H30, H32 & H36) adjacent to Fermanagh. In Britain, the plant is predominantly western, andit has been classified as belonging to the oceanic temperate phyto-geographical element of the flora(Preston & Hill 1997).
Native, rare. European arctic-montane, but also present in north-eastern N America and Greenland.
1902; Abraham, J.T. & McCullagh, F.R.; talus screes below Cuilcagh summit.
January to September.
Growth form and preferred habitats
This typical arctic-alpine, perennial occurs, sometimes in considerable quantity, ie in patches of a 100or more loose, leafy rosettes bearing leafless flowering stems 4-20 cm tall, in suitably wet, acidic,open sites beside streams, or in flushes where water constantly seeps through the soil, gravel, oracross ± bare rock, including occasionally on shady, near vertical, rock faces where there is someseepage of water. Leaves are a fresh green, crisp and slightly fleshy, but not leathery, spathulate,tapering to the base but without a distinct petiole. The star-like flowers are borne in a 12- ormore-flowered lax panicle, sepals reflexed and petals white, 4-5 mm, each bearing two yellow spots nearthe base, anthers coral-pink or vermillion-red, visited mainly by flies (Clapham et al. 1987;Sell & Murrell 2014).
Webb & Gornall (1989) regard S. stellaris as, "somewhat calcifuge", a definitionthat does not fit it very accurately in the Fermanagh area, since here the acidic preference isabsolutely obvious. The previously mentioned authors qualify their verdict however (p. 58), suggestingthat the species absence from limestone may be due more to lack of surface water than to chemicalfactors. It definitely appears that the species requires not just wet conditions for growth andsurvival, but also the movement of water, be it merely a steady seepage through soil or across rocksurfaces, which are often in these circ*mstances covered with a cushion or carpet of bryophytes. Theother major species requirement is for a fairly open habitat, where there is very little or no vascularplant competition, and thus where seedlings and diminutive plantlets can develop and establish. As itgrows beside or in flowing water, it inevitably can be carried far downstream from its main site.
Fermanagh occurrence
In Fermanagh, this rather variable species is very rare and is only found along the N scarp of Cuilcaghand near the summit ridge (Cuilcagh Gap), spread across just three tetrads. It can appear as small moreor less isolated tufts of a few rosettes, perhaps linked at some stage in their development bystoloniferous stems or, as already mentioned, occasionally it may form larger, crowded patches.
Irish occurrence
Starry Saxifrage is a very scarce and local plant elsewhere in NI, being completely confined to theSperrins in Cos Tyrone and Londonderry (H36, H40) and the Mourne Mountains in Co Down (H38), where someof the stations have not been seen since 1970 (New Atlas). There is a similar, moderate-altitudemountain distribution around all the major ranges in the RoI, but again it has not been seen in recentyears at some of its older stations. S. stellaris is most frequent in Cos Kerry, W Galway,Wicklow and W Donegal (H1, H2, H16, H20 & H35). A gemmiferous variant is found in Connemara, beingdescribed as 'frequent' by the Gleninagh River at around 240 m, which flows from the Twelve Pins (orBens) into the N end of Lough Inagh, Co Galway. It has leafy buds formed in the place of flowers. Theyare not released from the parent plant, but flop down onto the ground near it, and may root."Similar plants have been reported from Co Kerry, Northumberland and Scotland, but there are novouchers and no precise documentation." (Webb & Gornall 1989).
British occurrence
The British headquarters of S. stellaris is very definitely situated in the Scottish mountains,including the border Southern Uplands, but it is also well represented in Cumbria, the N Pennines and inN Wales. It shows little loss, if any, in Britain during the last 60 years (Preston et al. 2002).
European and world occurrence
In Europe, S. stellaris shows a typical arctic-alpine distribution in all the mountains ranges ofEurope. S. stellaris subsp. stellaris is absent from the high Arctic, but is found inIceland and Lapland, and is common in the mountains of Norway, W Sweden and B & I. A form recognisedas subsp. alpigena Tem is widespread throughout the Alps and is frequent in the Pyrenees,Romanian Carpathians and the mountains of Bulgaria. It is more local in the mountains of Spain,Portugal, Corsica, Auvergne and the Cévennes, N Apennines and Yugoslavia. The species reaches itssouthern limit in the Sierra Nevada of Spain and in N Greece. Beyond Europe the range of subsp.stellaris is restricted to S Greenland and a few stations in Labrador and Baffin Island, makingit one of the amphi-atlantic plants (Hultén & Fries 1986, Map 1016; Webb & Gornall 1989).
Threats
None.
Introduced, neophyte, an occasional garden escape.
May 1845; Smith, T.O.; Ardunshin Bridge, on the Colebrooke River.
Throughout the year.
The parent species of this hybrid do not overlap in the wild, although their geographic distributions arenot far apart: S. umbrosa is confined as a native to the Pyrenees, while S. spathularisranges from N Portugal, NW Spain to SW Ireland. The two species must have met and crossed in cultivationto form the sterile hybrid, sometime in the 17th or early 18th century (Stace et al. 2015). Thehybrid nature of the plant was not recognised until D.A. Webb's (1950b) study of the so-called'Robertsonian saxifrages' group.
Londonpride was once a very common and popular garden plant on account of its vigorous growth, prolificflowering and tolerance of shade, poor, dry soil, soot and occasional, prolonged neglect. It is botheasy to grow and very simple to divide and propagate, again adding to its popularity well beyond theconfines of the alpine or rock garden enthusiast. With the rapid development of commercial gardencentres from the 1960s onwards and the subsequent availability of a greatly increased range of competinggarden plant material, Londonpride has inevitably lost some of its past popularity.
Since the plant does not produce seed, stations recorded beyond the garden wall must represent survivingdiscarded garden material. Typical sites include damp, shady woods and scrub, by streams, on rocks,walls, hedge- and ditch-banks, roadside verges, disused quarries and church- and graveyards.
Fermanagh occurrence
In Fermanagh, this very familiar evergreen, rosette-forming, stoloniferous, perennial hybrid has beenrecorded in 15 tetrads (2.8%), scattered across the VC. Patches of Londonpride are occasionally found inestate woodlands, but, unfortunately, also in shady and/or damp, generally lowland places where gardenrefuse is discarded. Locally this means anywhere roadside or near streams and rivers that issufficiently remote from habitation, or concealed from public view for such 'fly tipping' to beunobserved. The plant is quite often found near bridges or other man-made structures, and it can be longpersistent. Only the pollen of this hybrid is functional (Webb & Gornall 1989, p. 69), so thatcolonies persist and spread to a very limited extent, purely by lateral growth of vegetative stolons.
Irish occurrence
In NI, this garden hybrid is quite widely scattered and established, chiefly in Cos Fermanagh, Tyrone,Antrim and Londonderry (H33, H36, H39 & H40). Elsewhere in Ireland, it appears very much rarer andmore sporadic in its distribution (Cat Alien Pl Ir; Stace et al. 2015). The map of S.× urbium published by Stace et al. (2015) plots 73 Irishhectads with records of any date, representing 7% cover on the island.
British occurrence
In Britain, S. × urbium was first noted in the wild in 1837, and isnow frequent and widely scattered throughout all latitudes, although with a definite westernpredominance in its distribution (New Atlas; Stace et al. 2015). The map in Stace etal. (2015) plots a total of 719 hectads with records of the hybrid of any date. This represents25% cover in Britain, emphasising a major difference between B & I that may represent recorderand/or gardener effort. While this hybrid is normally male fertile only and does not set seed, accordingto Webb 1950b and D.A. Webb, in: Stace (1975), there are some fertile clones in existence (eg inOrkney).
European occurrence
This hybrid in Europe is often sold and cultivated under the name S. umbrosa, and is reportedcommon in C Europe (Essl 2004). It is naturalised in France and in scattered localities in Austria,where it was first recorded in 1881 (Essl 2004).
Threats
Although it naturalises perfectly well in woods and by shady streams, locally this evergreen perennial isnot frequent enough to be a threat.
Native, very rare. European arctic-montane, also in N America and Greenland.
1902; Abraham, J.T. & McCullagh, F.; western end, Cliffs of Magho.
April to October.
Growth form, recognition and preferred habitats
A loose cushion- or thick mat-forming perennial with medium to dark green, very fleshy, linear to oblong,stalkless and nerveless, apiculate leaves that are very variable in length. The leaf margin is usually,but not always, furnished with stout, forwardly directed tooth-like hairs. One to three hydathode glands(sometimes referred to as 'chalk glands') near the leaf-tip may or may not secrete lime, so thatcalcareous incrustation is often slight or absent. From June to September, some of the stems terminatein an erect flowering portion, 7-20 cm tall, bearing a short, leafy cyme of 2-15 flowers. The petals areusually yellow (often bearing orange spots), but less commonly the flowers can range from pale yellow toorange or even brick red (Thompson 1911; Webb & Gornall 1989).
S. aizoides is most often found on the banks of mountain streams, or where water continuouslyseeps across rocky ledges, screes, or surfaces with crevices, or wet, grassy slopes. In addition to itsrequirement for a constant water supply, this fleshy, almost succulent-leaved saxifrage is also a markedcalcicole in its nutrient demands: basic rock, or base-rich flushing water is always present wherever itoccurs. Webb & Gornall (1989) describe this as a "calcicole tendency", as in their widerexperience of the species, they find it will tolerate "any except very base-poor habitats".
Irish occurrence
A rare arctic-alpine plant in Ireland, elsewhere apart from its one Fermanagh station, this perennialoccurs about Donegal Bay (mainly in Cos Sligo and Leitrim (H28, H29)), plus at one site in NE Antrim(H39) (New Atlas).
Fermanagh occurrence
In Fermanagh, S. aizoides occurs in considerable numbers over an area of cliff covered by threetetrads (or four 1-km squares), along the north-facing Cliffs of Magho (otherwise known as Poulaphouca),that overlook Lower Lough Erne. It is always found growing in very wet, unstable parts of the cliffwhere the rock face is eroded and in many, but not all, parts of this linear station, it is shaded bywet upland mixed deciduous oak-birch-ash woodland.
Lusby et al. (1996) describe the species habitat in Scotland, where the species has its B & Iheadquarters very accurately, as being, "on the banks of mountain streams, on flushed grassy slopesor on rocky banks or cliff faces where there is a constant seepage of water". Flushed or drippingrocky banks, cliff ledges and wet screes at the base of the cliff, precisely describes the Fermanaghhabitats of this plant.
Associated species
In B & I, S. aizoides is frequently associated with other interesting arctic-alpines such asSaxifraga oppositifolia (Purple Saxifrage), Silene acaulis (Moss Campion) andThalictrum alpinum (Alpine Meadow-rue), none of which have ever been found in Fermanagh (Raven& Walters 1956). While Dryas octopetala (Mountain Avens) has been recorded from the sameFermanagh cliffs over 90 years ago, the currently associated species of interest include only Listeraovata (Common Twayblade), Parnassia palustris (Grass-of-Parnassus), Pinguiculavulgaris (Common Butterwort), Primula vulgaris (Primrose) and Sesleria caerulea(Blue Moor-grass).
S. aizoides forms loose, spreading cushions by means of creeping decumbent sterile stems and it iseasy to imagine seed from the profuse flowers on separate, erect fertile stems being readily transportedto lower altitudes by the flowing water that soaks the plants (McCallum Webster 1978). Seed of thespecies is also said to be transported by birds (Lusby et al. 1996), but this is hard to imaginein the particular wet, wooded Fermanagh station occupied. In NI, S. aizoides is a protectedspecies under Schedule 8 of the Wildlife (NI) Order 1985.
British occurrence
S. aizoides is common and widespread in N & W Scotland from sand-dunes at sea level to1175 m on the mountains of Mid-Perthshire (Pearman et al. 2008). On account of its connectionwith flushing water, in Scotland plants and seeds are often washed downstream and can become establishedon river shingle and by the sides of streams (McCallum Webster 1978).
Yellow Saxifrage also features on the mountains of Cumbria and the N Yorkshire Pennines, which representthe southern limit of the species' northern Europe distribution (Webb & Gornall 1989; NewAtlas).
European and world occurrence
In mainland Europe, the distribution of S. aizoides fits the classical arctic-alpine pattern. Itstretches from Iceland and Spitsbergen to N & W Scandinavia, and locally to NW Russia. It has twostations in the French Jura and is common throughout the Alps and frequent in the Pyrenees andCarpathians. From the Alps, it extends southwards to the C Apennines and through the mountains of theBalkans to NW Macedonia (Webb & Gornall 1989, Map 39; Jalas et al. 1999, Map 3161).
Beyond Europe, it is found in Greenland and is widespread in arctic and subarctic Canada, with southwardextensions near the E & W coasts of N America. In Asia, it is found only in Novaya Zemlya and thepolar Urals (Webb & Gornall 1989, Map 38; Hultén & Fries 1986, Map 1024).
Threats
None.
Native, fairly frequent, but local. Oceanic boreal-montane.
1804; Wade, Dr; Knockninny Hill.
Throughout the year.
Growth form and preferred habitats
This fluffy, loose, evergreen, stoloniferous perennial is polymorphic, displaying considerable phenotypic(plastic) variation with changing natural environments, but it can be easily recognised at any time ofyear by its distinctive radiating, prostrate, leafy barren shoots producing a loose mossy cushion orweft growth habit. The prostrate sterile shoots often bear small, leafy bulbils in the leaf axils,called gemmae. The majority of leaves are three-lobed, but some are linear, unlobed. As is the case withmany other saxifrage species, most of the foliage of plants growing in drier situations turns bright redin summer. When growing in normal light intensities, the barren shoots are negatively phototropic, thegrowing tips bending as if to seek out darker growing conditions (Webb 1950a).
Mossy Saxifrage is a characteristic plant of shady, wet to fairly damp ledges on north-facing limestonecliffs and on damp to dripping, mossy banks overhanging streams. On account of the damp oceanic climateof NW Ireland that Fermanagh enjoys, however, it also grows in more open, sunny, rather dry, rocky,usually upland, limestone grassland, in or on eroded limestone pavement and on slightly damper areas onstabilised calcareous scree slopes. Although in Fermanagh and elsewhere S. hypnoides isfrequently closely associated with limestone or base-rich soils, Webb & Gornall (1989) point outthat over its whole distribution range, which is very much centred on B & I, it can also grow freelyon siliceous rocks, provided the soil receives some flushing by groundwater. Occasionally, elsewhere,S. hypnoides can even flourish on sand-dunes.
Genetic variation
The chromosomes of mossy saxifrage species are very small, crowded and numerous, making them difficult tocount. The existence of multiple chromosome pairing at meiosis, in some forms at least, also makesaccurate counts difficult, so that only vegetative mitoses give reliable counts. Some counts of Irishmaterial communicated by Prof Webb must be considered only tentative and approximate estimates (Webb1950a & b). However, despite these problems, S. hypnoides in B & I is known to possesstwo cytotypes with chromosome numbers 2n=26 and 2n=52, although other earlier Irish counts suggest 2n=48and 2n=64 (Webb 1950a & b; Webb & Gornall 1989). Geographically, plants of S. hypnoidesfrom W Ireland and Wales are diploid (2n=26), while those from N Ireland, N England and Scotland aretetraploid (2n=52) (R.J. Gornall, in: Preston et al. 2002). There are no consistent morphologicaldifferences between the diploid and tetraploid chromosome cytotypes (Parker 1979).
In their Hybrid Flora of the British Isles (Stace et al. 2015), the authors list fourchromosome counts for S. hypnoides, 2n=26, 48, 52 & 64, and for the closely related S.rosacea (Irish Saxifrage) which also occurs in Ireland, 2n=48, 52 & 64. Both these speciesare self-fertile and field observation shows there has been introgression and transfer of charactersbetween them in the past when their distributions overlapped (Stace et al. 2015).
A third closely related species restricted to southern continental Europe (SE France, N & C Spain andE Portugal) is S. continentalis (Engler & Irmscher) D.A. Webb (= S. fragosoi Sennen),which previously was considered a subspecies of S. hypnoides. It also has cytotypes 2n=26 &52, and differs most from S. hypnoides in having more numerous and consistently presentsummer-dormant buds that are covered with almost entirely membranous, translucent outer leaves. It hasbasal leaves that are 1-7-lobed, often 5-lobed, rather than 3-lobed as in S. hypnoides (Webb& Gornall 1989).
The basic chromosome number in this group of Saxifrages was firstly thought to be x=8 (Webb 1950a), buthas subsequently been revised to x=13, a degree of change that says much about the difficulties ofchromosome counting in these species (Webb & Gornall 1989). In any event, at least one S.hypnoides cytotype is polyploid, and must have arisen following hybridisation. Hybridisationtakes place readily between species in this section of the genus (Section Dactyloides Tausch, the'mossy' saxifrages), there being no serious sterility barrier between the members due to the degree ofpolyploidy they share (Webb 1950a & b).
The range of phenotypic modification, produced by the plasticity that S. hypnoides displays, isclosely paralleled and overlapped by the range of variation produced by its genetic diversity (Webb1950b).
Flowering reproduction
From May or late April to July a small proportion of the leafy branches develop flowers. Shading has amarked inhibitory effect on the number of flowering stems produced (Webb 1950a). The flowering stemsbranch in their upper half and bear 1-5, but occasionally up to seven pure white, shining flowers in aloose panicle. Flower buds are pendulous (nodding), when first developed, clearly differentiating theplant from the closely related S. rosacea which has erect flower buds (Webb 1950a). Otherwise thebest distinguishing feature of these two species is the aristate tip of S. hypnoides leaf lobes,ie each being drawn out into a fine, colourless hair-like point, while the leaf segment tips of S.rosacea are obtuse to mucronate (an abrupt point), but never aristate-apiculate (Parker 1979;Webb & Gornall 1989).
Flowers of S. hypnoides are regular, 10-15 mm in diameter, hermaphrodite, scentless and stronglyprotandrous (ie the anthers mature before the stigmas are ripe), meaning that cross-pollination isstrongly favoured, although self-pollination is still possible (Webb 1950b). The open nature of theunfertilised flower, with its only slightly domed central ovary (3/4 inferior, embedded in thereceptacle, ie nearly epigynous), means that the nectaries are fully exposed between the ovary and theinsertion of the petals on the flat receptacle. This makes the plentiful nectar available even toshort-tongued flies, sawflies and gall-wasps, so that a wide range of insect visitors can feed and crosspollinate them (Webb 1950a; Proctor & Yeo 1973; Fitter 1987). Individual flowers last from 8-12 daysfrom opening to petal fall. The fruit is a dry capsule and the many seeds it contains are black andcovered with numerous low, rounded papillae (Sell & Murrrell 2014).
There do not appear to be any estimates of the number of seed normally produced per capsule in wildsites. In late autumn, the dry fruit capsule partially splits to release the seeds, which are shaken outby swaying of the flowering branch in any breeze over a period of time. Seed dispersal is probablyfurther assisted by flowing water and, over small distances, by wind (Webb 1950a).
Germination does not require a chilling period, but probably most seed does not germinate until thefollowing spring (Webb 1950a). The current author (RSF) cannot find any published information on buriedseed longevity for S. hypnoides, but results for other Saxifraga species indicate that inmost of them seeds are transitory, persisting for only one year (Thompson et al. 1997).
Vegetative reproduction
Although S. hypnoides produces a spreading weft of prostrate shoots that root at intervals, itspowers of vegetative reproduction and spread by this means are probably minor compared to seed. Theshallow rocky soils, the cliff ledge and other more grassy habitats S. hypnoides usuallyoccupies, tend to be ecologically discontinuous, restricting any form of gradually spreading increase byvegetative growth (Webb 1950a).
Fermanagh occurrence
In Fermanagh, S. hypnoides has been recorded in a total of 25 tetrads, 4.7% of the total in theVC. It is almost exclusively found around the Knockmore and Marlbank limestones in the hilly W & SWof the county, but it does extend SE to an outlier on the limestone knoll of Knockninny, overlookingUpper Lough Erne where Dr Wade first recorded it in 1804.
Irish occurrence
In NI, apart from Fermanagh, S. hypnoides grows along the base-rich basaltic scarps of Cos Antrimand Londonderry (H39, H40), and elsewhere in Ireland the species is only common, widespread and locallyabundant in the Burren, Co Clare (H9) and on the Ben Bulbin mountain limestones in Cos Sligo and Leitrim(H28, H29).
British occurrence
The distribution of S. hypnoides in Britain has been reasonably stable over the last 50 years,although there may have been some decline in N Scotland (R.J. Gornall, in: Preston et al. 2002).In Britain, Mossy Saxifrage is essentially a plant of northern and western mountains, extending from thefar north of Scotland to S Wales and N Somerset (Cheddar), but there also are scattered garden escapesfurther south in England, although most of these are probably of hybrid origin (Webb 1950a; Prestonet al. 2002).
European occurrence
Beyond the shores of B & I, S. hypnoides is common and widespread in Iceland and the Faeroes,while on the continental mainland it is remarkably confined to just a few localities in W Norway, andtwo small remote outliers further south, in Belgium and in France in the Vosges, near Gérardmer. It hasnot been seen in the latter location for several decades (Hultén & Fries 1986, Map 1027; Webb &Gornall 1989, Map 55; Jalas et al. 1999, Map 3186).
Threats
Possibly sheep grazing and trampling.
Extinct. Sub-oceanic boreal-montane.
Interestingly, fossil material of S. rosacea has been found at Derryvree, near Maguiresbridge, CoFermanagh, in a Middle Midlandian full-glacial deposit, but there are no modern records this far northin Ireland (Colhoun et al. 1972).
Native. Very rare, but probably previously overlooked. European southern-temperate.
15 May 1986; Wolfe-Murphy, S.A. & Austin, L.W.; Knockmore wood.
Growth form and preferred habitats
This very variable, glandular-hairy, semi-rosette, winter annual germinates in the autumn and flowersearly in the following spring, between April and June. Depending upon available soil moisture the wholeplant can vary widely in size between tiny (c 2.5 cm tall) and single stemmed, to small but branched andtufted (10-20 cm tall). The species epithet 'tridactylites', meaning 'three-fingered', refers to thecommonest form of the leaves, although they can be 1-7 lobed (Webb & Gornall 1989). Despite theextent of the variation mentioned, most of it is phenotypic and environmental, there being littleecotypic differentiation (Webb & Gornall 1989). Being usually diminutive and fairly rare in Ireland,S. tridactylites is inconspicuous and easily over-looked unless it is abundantly present.
Elsewhere in B & I, Rue-leaved Saxifrage typically occurs in dry, open, sunny, short-turf, stony,nutrient-poor, overgrazed limestone grassland, or in droughted conditions on rock surfaces, including insolution cups in limestone pavement, in limestone or base-rich quarries and gravel pits, in crevices andon the tops of walls. S. tridactylites is classed as a stress-tolerant ruderal by Grime etal. (1988), meaning it avoids competition with larger, more vigorous, dominant plant species byfrequenting unfavourable growing conditions in drier, more shallow, exposed, rocky and less fertilesoils, often subject to summer drought and disturbance. Conditions include semi-stabilised, calcareoussand dunes in some coastal areas.
While Webb & Gornall (1989) regard it as not strictly calcicole, there is a definite tendency in thisdirection, in many instances S. tridactylites being much commoner on limestone than on any othersubstrate, and largely restricted to calcareous or base-rich soils in the pH range 6.0–8.0 (Grime etal. 1988).
Fermanagh occurrence
Data included in The Flora of Co Fermanagh (Forbes & Northridge 2012) had a cut-off date of 31December 2010. The Flora contained mention of a solitary record of this tiny saxifrage from theFermanagh western limestones, as listed above. The recorded site sounded quite atypical for the species,ie on or around the ash-wooded slope below Knockmore limestone cliffs. However, the fact that thespecies occurs further west on the Ben Bulbin limestones, of which the Fermanagh strata are a NEextension and that there is so much suitable habitat for it in Fermanagh, suggested that it justmight be present in the VC, even perhaps as a very rare colonist. Wolfe-Murphy's voucher-lessrecord was very welcome, although it remained doubtful and unconfirmed.
Shortly after the book database was finalised, S. tridactylites was discovered and confirmed froma site on the shore of Lough Melvin near a plantation, 1.2 km [originally given as 0.75 miles] NW ofGarrison found by John Faulkner and Robert and Hannah Northridge, on 11 May 2011. It was re-found hereby RHN & HN on 28 May 2012. Two additional sites have since been added by the Northridges, atInishmore Viaduct, Upper Lough Erne, 29 April 2015 and again on 15 May 2018; and at Killyhevlin, 2.3 kmSE of Enniskillen, 22 May 2015.
In view of these five new records, the current author feels Wolfe-Murphy's earlier record should probablybe accepted.
Irish occurrence
Elsewhere in NI, S. tridactylites is rare or very rare and is mainly found near the coast, onsandhills, rocks and walls, as well as on bare, base-rich gravel at one site on the shore of LoughNeagh, where it is considered a very rare, accidentally introduced colonist (Flora of LoughNeagh; FNEI 3). In the RoI, S. tridactylites is very frequent in the south and inwestern parts of the centre, but rather rare elsewhere (Parnell & Curtis 2012). It is very frequentand often locally abundant in the limestone karst area of the Burren, Co Clare (H9) and especially so onlimestone pavement, calcareous sand-dunes, and in and on lime-mortared walls near the coast (Webb &Scannell 1983).
Flowering reproduction
This little plant should be searched for early in the season, as it has more or less finished floweringby the end of May. Having said that, the white flowers, although borne either solitary or in a ratherdiffuse cymose branched inflorescence, are often so minute they do not greatly advertise the presence ofthe species. The perfect (hermaphrodite) flowers, in contrast to those of most other saxifrages, areusually somewhat protogynous (ie the female parts mature before the anthers) (Webb & Gornall 1989).Not surprisingly, however, on account of their small size, they are chiefly self-pollinated (Garrard& Streeter 1983; Webb & Gornall 1989). Seed is set from June to July, the dehiscent fruitcapsule often containing more than 100 very tiny seeds. Dispersal usually occurs close to the parentplant, but as the seeds are so minute, they are easily carried by wind or vehicle slipstream andsometimes travel further than normal, including high up on walls (Grime et al. 1988).
Being a winter annual, after a post-release ripening period, the seed germinates in the autumn and theplantlet overwinters as a tiny leaf rosette. Flowering is induced after cold vernalisation, but there isno long-day requirement for floral initiation. Flowering follows in early spring (Grime et al.1988). A short-term persistent soil seed bank is also formed, ie buried seed surviving >1 and <5years (Thompson et al. 1997).
When in full sun in a droughted, shallow, lime-rich, sandy soil, the whole plant often goes a brightscarlet red when in fruit, making it much more conspicuous for a time in June and early July. It thenquickly dries out, dies off and disappears.
British occurrence
Locally common on calcareous soils in lowland Britain, S. tridactylites becomes increasingscattered, rare and coastal northwards into Scotland, although it still reaches the northern-most pointof the mainland. The New Atlas indicates that although better recorded than in the 1962 BSBIAtlas, there has been a widespread decline of the species in S & E Britain. Analysis of theNew Atlas database reveals that the losses have occurred since 1950 (R.J. Gornall, in: Prestonet al. 2002).
However, the British partial re-survey, Local Change 1987-2004, found that S. tridactyliteswas amongst a group of lowland species of calcareous grassland and marginal or linear landscape featuresthat showed a significant increase. The calculated 'Change Factor' for the species was +15 (Braithwaiteet al. 2006).
European occurrence
S. tridactylites is the most widespread Saxifraga species in Europe, ranging over thewhole continent, including the Mediterranean isles, but absent from the extreme north and much of thenorth-east. In E Europe, it reaches only the western and southern fringes of Russia through NE Ukraine,just N of the Black Sea (Jalas et al. 1999, Map 3241). Beyond Europe, it is known only from NWAfrica and SW Asia, its eastern limit lying on the borders of NE Iran (Webb & Gornall 1989, Map 74;Hultén & Fries 1986, Map 1022).
Threats
'Improved' limestone pastures (that is, in terms of grazing productivity) through fertilizer application,leading to increased plant competition.
Native, common. Sub-oceanic temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
The prostrate stems of this low-growing, wintergreen perennial grow and spread forming adventitiousroots, thus developing ± large clonal, carpet-like mats or cushions on woodland floors, or alongpathsides in damp to fairly wet woods, and on the shaded, flushed and frequently waterlogged banks ofditches, streams and rivers. The species occupies a wide range of moderately fertile to nutrient- andbase-poor soils, from those on limestone cliffs to base-poor mud, sand and grit, but it avoids stronglyacidic or peaty soils below about pH 4.5 (Grime et al. 1988).
C. oppositifolium is able to tolerate both considerable shading by other plants and occasional temporary immersion by flood waters or ahigh water table and, while it occurs over a range of soil chemistry and reaction (pH 4.5-7.5), it mostfrequently grows on moderately acidic substrates (pH 5.5-6.0) (Grime et al. 1988).
Although it frequently occurs as a ground under-storey beneath the canopy of various herb species,Opposite-leaved Golden-saxifrage is especially common and abundant where vegetation cover is incompleteand patches of bare muddy soil occur. These sites are kept open by substrate instability, ofteninvolving occasional or periodic disturbance by cattle or other animals grazing and trampling thesurface. Rainwater scouring, temporary submergence during heavy downpours and seasonal events such assporadic winter flooding, also provide sufficient vegetation disturbance to enable colonisation by thislittle species. The plant is also commonly found in more fully lit, constantly damp, upland rock, cliffor wall situations, where lack of physical space, or a moderate degree of erosion, or other occasionaldisturbance helps keep ecological conditions suitably open for seedling colonisation and establishment.
The requirement for open ecological conditions is obvious and significant, limiting competition andpotential replacement by vigorous, larger or more ecologically demanding species.
Fermanagh occurrence
This is a very common and widespread species throughout Fermanagh, recorded in 338 tetrads, 64% of thosein the VC.
Flowering reproduction
In shaded, wet, boggy ground, Opposite-leaved Golden Saxifrage is the earliest species to flower eachyear, the blossom first appearing in March with anthesis reaching its peak in April and May. Earlyseason flower production, together with its well-maintained wintergreen leaf area, enables the speciesto coexist with much taller herbs of damp or wet ground, including Epilobium hirsutum (GreatWillowherb) and Valeriana officinalis (Common Valerian), both of which expand their leaf canopyconsiderably later in the season. The tiny, 3-4 mm, greenish, hermaphrodite, petal-less flowers havetheir parts in fours and are borne in a ± flat-topped, golden-yellow, leafy bract-subtended,sub-corymbose cyme. The flowers are protogynous (ie the female parts mature first), with 8-10 brightyellow anthers. Nectar is secreted from a disc round the two divergent styles, attracting small insectsthat act as pollinators, or else the flowers self-pollinate. Each small fruit capsule produces up to 30very small seeds, and a large clonal patch of the plant may thus generate an enormous number of seeds.
Seed and vegetative dispersal
The ripe, dry capsule splits to release the abundant small seeds which, together with plant fragmentsdetached by plant disturbance, are dispersed by wind (less likely within woodland), by floatation inrun-off water and in mud transported on both animals and man.
Buried seed survival
The published survey of NW European soil seed banks contained just two samples of this very successfulspecies: one reckoned the seed survival is transient (ie persisting less than one year), while the otherrecord was indeterminate or non-committal (Thompson et al. 1997).
British and Irish occurrence
A combination of the three seed dispersal mechanisms mentioned above, obviously confers sufficienttransport for this species, for it is certainly very common and widespread across most of B & I.However, perhaps on account of its somewhat restricted soil reaction requirement and necessity for openhabitat growing conditions, C. oppositifolium is not omnipresent throughout B & I. TheNew Atlas hectad map displays a predominantly N & W distribution in Britain, while it islocal or absent in parts of E England and the outer Scottish isles. In Ireland, Opposite-leavedGolden-saxifrage is widespread and common across most of the island, but patchy, local and less wellrepresented in parts of the W & C (Preston et al. 2002).
European occurrence
C. oppositifolium is a European endemic species and biogeographically is a member of theSub-oceanic Temperate element. Beyond B & I, it is widespread in W and parts of C Europe, extendingeastwards to W Poland and C Czechoslovakia (Hultén & Fries 1986, Map 1039; Jalas et al. 1999,Map 3249; Sell & Murrell 2014).
Threats
None.
Native, frequent but local. Circumpolar boreo-temperate.
1882; Stewart, S.A.; Callow (or Carrick) Hill.
April to December.
Growth form and preferred habitats
In B & I, this somewhat tufted, hairless perennial with a short vertical rootstock and cordate,stalked leaves, bears lovely, solitary, pure white flowers and is a plant of three completely distincthabitats: wet dune-slacks, base-rich marshes and fens, and flushed areas of otherwise nutrient-poororganic soil on limestone hillsides or below limestone cliffs (An Irish Flora 1996; Bonnin etal. 2002). In Ireland, it is mainly found either at the coast, in species-rich wet hollows infixed grey dunes, or else in damp to wet grassy ground around the upper basins of the larger limestonelakes of the W and the W Midlands. Elsewhere on the island, it is local and rather rare, including onlimestone hillsides and below cliffs (An Irish Flora 1996). In all of these situations, seepageof ground water is almost invariably present and the vegetation types supported by the preciseconditions in which P. palustris grows, possess a short, not too dense, nor very competitiveplant cover.
While in overall terms we tend to think of P. palustris as a calcicole, or a weakly-calcicolespecies, it really is unusual in frequenting soils derived either from base-rich rocks (usuallycalcareous), or of moderately base-poor peat or leached clay character (Flora of Connemara and theBurren; Sinker et al. 1985; Grime et al. 1988, p. 170). In W Ireland, these twoinfertile and unproductive soil types overlap when P. palustris grows on sufficiently damp,percolated areas of cut-over acid peat bogs, or on shallow, raw humus, whenever these types of organicsoils are formed directly overlying limestone rocks (Flora of Connemara and the Burren).
In Britain, P. palustris is generally considered to require an open, permanently moist site inmarshy, peaty grasslands (both meadows and pastures) and in short sedge and grass fen margins aroundlakes and ditches. It is also found on more upland moorland sites, in flushes or in small damp hollowsassociated with base-rich rocks and peaty soils. The form of the plant with this mainly inland ecologyis sometimes referred to as var. palustris. In coastal situations around B & I, P.palustris sometimes occurs as a genetically distinct dwarfed ecotype or ecodeme (var.condensata Travis & Wheldon) in the seasonally damp ground of dune-slacks and machair(Gornall 1988).
Ecological requirements for successful establishment
In all these various situations, seepage of ground water is almost invariably present and the vegetationthat is supported by the precise conditions in which P. palustris grows, tends to possess a shortand not too dense nor very competitive cover. A recent study has found that P. palustrispossesses an arbuscular mycorrhizal partner, which must assist it in obtaining soil nutrients incompetition with its neighbours (Eriksen et al. 2002).
A study in Scandinavia by Hultgard (1987) found that the species there is favoured by the existence ofbare ground resulting from moderate levels of either natural or human-inspired disturbance (eg periodicflooding, mowing, grazing, trampling, or in roadside conditions and in quarry pits). In many of itsFermanagh damp grassland sites, the vegetation probably appears more closed than really is the case. Theseed is tiny and very lightweight, so that bare ground and near zero competition (plus the presence ofits mycorrhizal partner) are probably all essential for germination and successful establishment.Additional ecological requirements of P. palustris were identified as a fairly high level ofoxygen supply at the roots, a need that is met by the often observed water movement in the soil itoccupies, adequate mineral nutrition (ie just enough to avoid stimulating much plant competition fornutrients, light and space), a substrate which is not too acidic and illumination generally close tofull sunlight (Hultgard 1987).
Genetic variation
P. palustris s.l. is a complex of several closely related races or cytodemes and has a widecircumpolar boreal-temperate distribution (Hultén 1971, Map 68). The species contains two principalcytodemes, a diploid (2n=18) and a tetraploid (2n=36), forms which are morphologically indistinguishablein B & I, the Netherlands and in Scandinavia (Gadella & Kliphuis 1968; Hultgard 1987; Wentworth& Gornall 1996). The tetraploid is very probably of autopolyploid origin and it occupies the morenortherly portion of the species range, while the diploid appears to be more or less confined to thearea south of the limit of the last glaciation (the Midlandian in Ireland and the Devensian in Britain)(Gornwall & Wentworth 1993).
Fermanagh occurrence
In Fermanagh, P. palustris has been recorded in 57 tetrads, 10.8% of those in the VC. It isdistributed chiefly in wet fens around the shores of Lower Lough Erne and of smaller lakes, plus influshed areas around limestone hills on the Western plateau. It also occurs more locally and lessabundantly in the SE of the county on the shore of Upper Lough Erne, at Galloon Td, near the Crom estateand along the string of marl lakes that lie between Magheraveely and Rosslea in the extreme E of thecounty.
The fact that the distribution map contains eight tetrads with pre-1975 records only indicates there havebeen definite population losses across the VC. This is readily attributed to drainage operations andother changes in land use, including conifer plantation and agricultural grassland improvements.
Ecological indicator species
In Fermanagh, P. palustris has proved to be a good high- to late-summer flowering indicatorspecies for such locally rare or scarce plants as Anagallis tenella (Bog Pimpernel),Dactylorhiza incarnata (Early Marsh-orchid), Eleocharis quinqueflora (Few-floweredSpike-rush), Eriophorum latifolium (Broad-leaved Cottongrass) and Saxifraga aizoides(Yellow Saxifrage). It is almost invariably accompanied by more common species of similar wide or'bimodal' (ie tolerating two alternative environments) ecological amplitude, such as Briza media(Quaking-grass), Carex panicea (Carnation Sedge), Pinguicula vulgaris (Common Butterwort)and Triglochin palustre (Marsh Arrowgrass).
The bimodal nature of the environmental behaviour and habitat occurrence of P. palustris isreflected in the fact that in the Great Britain-based National Vegetation Classification, the speciesfeatures in one form of mesotrophic grassland (MG2), two each of calcicole and calcifuge grasslands(CG9, CG14 & U15, U17) and in five mires (ie fens and bogs) (M8, M9, M10, M13 & M38) (Rodwellet al. 1991b, 1992).
Flowering reproduction
Grass of Parnassus blooms from July onwards but reaches its peak in August and September. The protandroussolitary, 15-35 mm diameter flower has pure white petals, decorated with conspicuous veins which maylook white or greenish. Each flower possesses five sterile stamens or staminodes alternating with fivefunctional ones. The staminodes become ± spathulate at their apex, each bearing 7-15 long, hair-likeprocesses tipped with shining yellowish glands. These staminodes act both as nectaries and aspseudo-nectaries, attracting insect pollinators with their conspicuous glistening but non-rewardingglobular tips, while each also produces nectar at its base. The main insect visitors are hoverflies andother Diptera, but the range of insect visitors can include ants, short-tongued bees, butterflies andmoths (Proctor & Yeo 1973).
Experimental studies in Norway have shown that the presence of staminodes increases both the number andthe duration of pollinator visits and thus they facilitate either cross-pollination or self-pollination,depending upon the particular population involved in the study (ie either coastal or alpine populations)(Sandvik & Totland 2003).
Selfed flowers produce a drastically reduced number of seeds, up to 95% fewer seeds per ovary thanopen-pollinated flowers. At the alpine level, the Norwegian workers found significantly moreopen-pollinated than emasculated flowers produced fruit in comparison with the same experiment atsea-level. Thus self-pollination aided by pollinators transferring pollen within the same flower,occurred more frequently than outcrossing at the alpine level, while in contrast, outcrossing was thenorm at the lower altitude and here selfing was rare. Thus, although the flowers are protandrous,releasing their pollen before the stigma ripens, the degree of outcrossing varies substantially acrossthe species distribution (Sandvik & Totland 2003).
Seed production, dispersal, survival and germination
The solitary, large, round, purple fruit capsule splits when ripe to release an approximate mean of1,600-1,900 seed for dispersal on the breeze (Ridley 1930). Salisbury (1942) estimated that with apercentage germination rate of 76%, each plant had a reproductive capacity of around 1,300 potentialoffspring per annum. According to the soil seed bank survey of NW Europe, the seed is transient toshort-term only (surviving less than one year (five studies), or 1-5 years (one study)) (Thompson etal. 1997).
Irish occurrence
The occurrence of P. palustris is typically very local throughout its Irish distribution, evenwhen the plant is relatively frequent in an area. The presence of the plant is seldom abundant, althoughit does form rather diffuse, possibly clonal patches, in old managed grassland. It is very frequent inthe W and west-centre of the island, but absent or very rare from the southern third of the country,while in the north it is scarce and mainly coastal (New Atlas).
British occurrence
P. palustris is local in much of Britain, but it is absent or very rare from S Wales and largeareas of C & S England. There has been a long-term decline in the population of the diploid form, inparticular in S England, most likely due to drainage operations over the last half century at least. Thewidespread distribution of the tetraploid form in the N & W of the British Isles seems to be verymuch more stable, although here also there is a definite westerly preponderance of the species (NewAtlas).
European and world occurrence
P. palustris, on a worldwide basis, is a polymorphic collection of subspecies or species that iscircumpolar in the boreo-temperate zone (Hultén & Fries 1986, Maps 1040 & 1041). P.palustris is widespread in Europe from Iceland and the whole of Scandinavia, to Denmark, theNetherlands and all of France except the NW. To the south, it stretches from N Spain and Moroccoeastwards to Italy, the Balkans and Greece, although it is absent from the Mediterranean islands (Jalaset al. 1999, Map 3257). In the more northerly boreal areas, Hultén & Fries (1986) map acircumpolar subarctic-montane form they distinguish as subsp. neogaea (Fern.) Hult. (= P.obtusiflora Rupr.), although they admit its occurrence is imperfectly known.
Names
The genus name 'Parnassia' is a shortened form of the Latin name given by the 16th century Flemish herbalwriter Mathias de l'Obel. He called the plant 'gramen Parnassi', a translation of the Greek 'agrostis enParnasso', mentioned by the ancient writer Dioscorides (and hence the English common name, 'Grass ofParnassus', first translated into English by Lyte in 1578). It is called after the Greek mountain holyto Apollo and the Muses, where the species does indeed occur (Gilbert-Carter 1964; Strid 1986; Stearn1992). Grigson (1987) supposes that the herbalist was inspired to make a connection to this prestigiousreligious location by the beauty of the prettily green-veined pure white flower, and to the currentwriter this does not appear too far-fetched, it being my favourite Irish wildflower.
It is, of course, somewhere between difficult and impossible to connect our P. palustris with theplant referred to by Dioscorides. According to the plant polymath David McClintock, de l'Obel includedMaianthemum bifolium (May Lily) under gramen Parnassi, and the woodblock illustrations providedfor Matthiolus in 1562 and thereafter (many editions in several European languages until 1604), or forTabernaemontanus in 1590, are shown flowerless and indeed look quite like May Lily, or even like abroad-leaved pondweed (McClintock 1966, p. 199 & Plate 9). The flower was considered by these earlybotanists to resemble (or be) a white form of Hepatica noblis (ie H. alba) and, perhapsbecause of this, Gerard (1597), in his first edition, gave the species the medicinal properties of thelatter for the treatment of liver disorders, calling it 'White Liverwort'. In the second much improvededition of 'Gerard's Herball' (1633) edited by Thomas Johnson, the latter corrected the illustrationused, and also displayed a double-flowered form of the plant (p. 840), although Johnson continued topronounce its supposed medicinal properties as a stomach remedy preventing vomiting and for helping tobreak and pass liver stones. Launert (1981) continues to illustrate and feature P. palustris as amedicinal plant to this day, although there is no mention of it in Grieve's monumental, comprehensiveherbal (Grieve 1931).
The Latin specific epithet 'palustris' is a masculine form of the two words meaning swamp or bog, 'palus,udis', and refers to the damp ground in which the plant typically grows (Gilbert-Carter 1964). OtherEnglish common names include the prosaic and inaccurate 'White Buttercup' and the somewhat better 'BogStar'. The species is grown in gardens, especially by alpine gardeners (Grey-Wilson 1989; Griffiths1994).
Threats
Drainage of marshy ground and, elsewhere, excessive disturbance from grazing or trampling stock.
ROSACEAE – Rose family
Introduction, neophyte, a rare garden escape. Native of Eurasia, from Central Europe eastwards.
6 September 2001; Northridge, R.H.; mainland shore of Lower Lough Erne, near Portinode Bridge across toBoa Island.
June to September.
This deciduous, decorative garden shrub, up to 2 m tall, has only been very recently recorded inFermanagh, as first listed above, but here as elsewhere in B & I it is probably more widespread thanis presently realised. A fluffy, cylindrical, pink-panicled, hedging species that suckers readily, S.salicifolia is a persistent relict of cultivation or a garden discard that escapes into the wildand can become thoroughly naturalised. It tends to favour colonising rough grassland, or shaded, wetground in or near hedgerows or on waste ground. So far in Fermanagh, it has appeared in lakeshore andriverside situations, all three Fermanagh records being post-2000 and discovered by RHN. In addition tothe first record above, the other details are: Kesh, 6 September 2001; and White Bridge, ColebrookeEstate, 24 June 2003. The tentative position in Fermanagh is that this plant is very thinly scattered inthe eastern half of the county in hedgerows and on waste ground on damp, rough terrain.
The true position regarding the occurrence of cultivated Spiraea species throughout B & I hasyet to become clear, since until the publication of Stace's New Flora of the BI in1991, or indeed perhaps as late as the publication of A.J. Silverside's later account in the BSBIPlant Crib (1998), there was considerable confusion in the genus. This was particularly the casebetween species, varieties and hybrids involving S. salicifolia and the more recently recognisedS. alba Du Roi (Pale Bridewort) and S. douglasii Hook. (Steeple-bush) (Silverside1990). All three species have an elongated terminal panicle of crowded, small flowers as theirinflorescence, although care is required as the leaf shape, inflorescence shape and flower colour areall variable. S. salicifolia was the first of these species to be brought into cultivation inEurope, but to some extent it has been replaced in gardens by the two more recently introduced NAmerican forms and by the subsequent hybrids between the species. Before 1991, the naturalisedSpiraea populations were usually recorded as S. salicifolia and, consequently, it isover-recorded. Having said this, most pre-1991 records can really only be ascribed to a broad aggregatecomprising all three species and their three hybrids (Stace et al. 2015).
S. salicifolia has almost hairless leaves and can be distinguished from S. × pseudosalicifolia (its hybrid with S. douglasii) which hashairy leaves and sepals bent backwards in fruit, and from S. douglasii which also has hairyleaves, but these are toothed only in the top half (Parnell & Curtis 2012). S. salicifoliahas a cylindrical inflorescence of usually bright pink flowers and leaves are widest just below halfway, while S. alba has a pyramidal inflorescence with distinct branches near its base and whiteor very pale pink flowers and its leaves are widest just above half way (Stace et al. 2015).
The native distributions of the three Spiraea species do not overlap in the wild: S.salicifolia is from Eurasia, S. alba from eastern N America and S. douglasii fromwestern N America (Stace et al. 2015). All three species are strongly suckering and the hybrids,which must have arisen in cultivation, can produce invasive thickets on damp, peaty soils. Taxonomicuncertainties developed to the extent that at one time Silverside (1990) doubted whether or not trueS. salicifolia existed in these islands at all.
This taxonomic and identification confusion is reflected in the treatment given in two books on alienspecies in B & I (Clement & Foster 1994; Reynolds 2002) and also in the maps and accountspublished in the New Atlas. All accounts of these Spiraea forms must really be regarded aspurely tentative in their statements and conclusions.
Threats
None.
Introduction, neophyte, a scarce or very occasional garden escape or relict of cultivation.
1991; McNeill, I.; Glengesh, NE of Tempo.
July to September.
This pink-flowered, suckering hedging shrub is most often found close to houses and is a relict ofplanted stock, a garden escape or a discard. Records of garden Spiraea are confused by onlyrecently recognised taxonomic splits and earlier reports of the S. salicifolia group, to whichthis hybrid belongs, still require re-determination in B & I herbaria (Reynolds 2002; Stace etal. 2015).
Identification keys are tentative (A.J. Silverside, in: Rich & Jermy 1998) and all claims made regarding the occurrence of this andother members of the group need to be treated with caution.
There are eight records of S. × rosalba in Fermanagh, equally splitbetween RHN and I. McNeill (IMcN). With one exception they are scattered in the E of the VC, the otherrecord being on farmland overlooking the SW shore of Lower Lough Erne at Ardees Lower Td.
In addition to the first record listed above, the remaining details are: roadside W of Errasallaghcrossroads, 7 km NW of Rosslea, 15 July 1997, RHN; near Moysnaght Td, NE of Clabby village, 21 July1997, IMcN; Coolcran, S of Brougher Mt, 21 July 1997, IMcN; near cottages below cliffs, Ardees Lower Td,20 September 1997, RHN & HJN; ruined house, Mullaghfad Td, E of Brookeborough, 21 September 1998,IMcN; near Garvary, E of Enniskillen, 21 July 2000, RHN; and Topped Mt Lough, 6 km ENE of Enniskillen,24 September 2000, RHN.
Introduction, neophyte, a very rare garden escape. A disjunct circumboreal species.
1947-55; MCM & D; woods at Lisgoole, Upper Lough Erne, just outside Enniskillen.
Meikle et al. (1975) recorded this in the Revised Typescript Flora as follows,"occurs as an escape at Lisgoole with Leucojum and Polygonatum multiflorum".This remains the only record of this large garden perennial occurring in the wild anywhere in Ireland.Unfortunately, we do not have a precise record date, but it falls between 1947 and 1955 at the latest –the period when Meikle and his co-workers were recording in the VC.
Although A. dioicus has been grown for decoration in gardens in B & I since at least 1633, itwas not recorded in the wild in Britain until 1950 (D.J. McCosh, in: Preston et al. 2002). TheFermanagh record is thus one of the very first discoveries of the species outside gardens anywhere inthese islands. The lateness of the first 'escape' date and the subsequent flurry of records from wildstations represented in the New Atlas occurring throughout Britain (mainly in N England and CScotland), but apparently NOT happening in Ireland, requires further study and explanation.
As a native species, A. dioicus is confined to damp or shady places in mountain districts oftemperate C & S Europe and SW Asia (from Belgium to N Albania, although absent from most of theMediterranean basin), but present also in the Pyrenees (Tutin et al. 1968). It forms part of acircumpolar species complex also known as A. sylvester Koestel, of which it may be considered aEurasian subspecies (Kurtto et al. 2004).
Growing up to 2 m tall, with very large, bi-pinnate leaves and producing very conspicuous, creamy-white,plume-like, much-branched, pyramidal panicles of small, 5 mm diameter unisexual flowers, A.dioicus is widely cultivated in gardens, planted in estate woodlands and occurs naturalised andestablished as far north as 64°N on the Atlantic coast of Norway (Kurtto et al. 2004, Map 3280).The inflorescence of the male plant is more showy than the female and thus is the much preferredcultivated form. Escapes often occur naturalised in woodland or by water and they can become verypersistent. Being ± dioecious (New Flora of the BI), the species sets seed only if plants of bothparents are present, which is an unusual circ*mstance in the wild, although known at a site in Dunbarton(Vc 99), and in another near Moulin in East Perth (Vc 89), both in Scotland (Clement & Foster 1994).Escaped populations, however, are almost always unisexual and reproduction is therefore vegetative, thestout, much-branched rhizome spreading to form clonal patches.
Native, common and abundant. Eurasian boreo-temperate, but widely naturalised, including in N America.
1881; Stewart, S.A.; Fermanagh.
Throughout the year.
Growth form and preferred habitats
The tall, leafy, wiry and furrowed flowering stems (up to 1.2 m in height) and the persistent nature ofthe inflorescence, even when it is long dead in spring, make this rhizomatous perennial easilyrecognisable in all seasons. Meadowsweet tolerates and grows in a huge range of constantly damp, but notwaterlogged, moderately acid to surprisingly dry, calcareous grassland and wayside habitats. F.ulmaria is most conspicuous and abundant in periodically wet, moderately fertile, lakeshore andriverside marshes, fens and ditches, but it is also frequent in moist or wet openings in woods, hedgerowbanks, damp meadows, calcareous upland grasslands and on the banks of streams and roadsides at alllevels.
The short rhizome and shallow roots of the typical plant make F. ulmaria a good indicator offluctuating water tables on banks, shores and hollows and it can develop dense, competitive, oftendominant patches in such situations. However, it cannot cope with prolonged waterloggedconditions and it also avoids the nutrient starvation and toxicity associated with strongly acidic peatbog soils of pH around 4.5 or below (Grime et al. 1988). Meadowsweet tolerates some shade inwoodland openings or margins and in hedgerows, but this greatly restricts its flowering and seedproduction.
Fermanagh occurrence
The wide ecological tolerances of Meadowsweet have enabled it to be the most frequently recorded plant inFermanagh. The same ecological range and flexibility permits it to be common and widespread throughoutthe whole of B & I. In the survey area, F. ulmaria probably occurs in every tetrad exceptthose entirely represented by the exposed, blanket bogland of the Cuilcagh Plateau or the open water ofLower Lough Erne. The Fermanagh Flora Database records Meadowsweet in 496 tetrads, 93.9% of those in theVC.
Flowering reproduction
Plants flower from June to August and the inflorescence, a loose, irregular panicle on an erect stem upto 120 cm tall, contains over 100 individual, bisexual flowers. The creamy-white, 5 mm diameter flowersgive off a heavy, sickly, sweet scent. They contain no nectar but still attract short-tongued insectssuch as flies, which eat and transfer some of the abundant pollen. Unvisited flowers self-fertilise.Leaf hairs on the plant also give off a refreshing fruity scent if brushed or handled, due to thepresence of oil of wintergreen (Genders 1971, pp. 86-7). The 6-10 carpels in each flower twist into ahead of achenes during ripening and the seeds are probably released only slowly from the dead stemduring the winter months. Each inflorescence produces a large number of dry seeds and after seedrelease, water transport is very likely significant since the achenes can float for up to three weeks(Ridley 1930, p. 208).
There are very different ideas regarding the ability of the seed to survive in the buried soil bank: of atotal of 34 estimates included in the survey of NW European literature, 20 regarded F. ulmaria astransient, six considered it short-term persistent (ie 1-5 years) and eight studies were indeterminate(Thompson et al. 1997).
Vegetative reproduction
The limited, but significant, vegetative extension of the creeping rhizome eventually allows theformation of large clonal colonies that can dominate other plants even at relatively low stem density(Grime et al. 1988). This is surprising considering the above ground biomass in stands of F.ulmaria tends to be less than in other commonly occurring tall-herb, stand-forming communities(Al-Mufti et al. 1977). However, the long persistence of the relatively large pinnate leavesappears to enable Meadowsweet to shade out smaller, less competitive species, especially in the morenutrient impoverished clayey soil conditions of some of the damp and shaded habitats it frequents (Grimeet al. 1988).
Ecological pressures
F. ulmaria is highly palatable and is quickly ousted by grazing, but it withstands mowing ortrampling very much better. As Rackham (1986) pointed out, on seasonally flooded or otherwise wetground, due to access problems with machinery, the farmer carries out grazing and mowing at differenttimes of year to avoid soft mud. Furthermore, in terms of selection pressure on the plant species,browsing animals carefully select the plants they prefer to graze, whereas the scythe, or nowadays themechanised blade, cuts everything!
British and Irish occurrence
F. ulmaria is common, widespread and locally abundant throughout the whole of these islands insuitable habitats up to an altitude of 880 m in the Scottish Highlands (D.J. McCosh, in: Preston etal. 2002).
Fossil record
There is a long and continuous fossil record of F. ulmaria pollen and fruits from the Cromerianonwards in B & I and it appears to be strongly persistent through both glacial and interglacialstages (Godwin 1975, pp. 182-6).
European and world occurrence
In a European context, F. ulmaria is a widespread and polymorphic species with sufficientvariation to merit the description of three subspecies in Flora Europaea 2 (Tutin etal. 1968). The three forms are: the widespread subsp. ulmaria throughout the total range;subsp. picbaueri (Podp.) Smejkal which is shorter in stature, achenes pubescent, and occurs fromE Austria and S Czechoslovakia to Bulgaria and SE Russia; and subsp. denudata (J. & C. Presl)Hayek, which has leaflets with a long, narrowly triangular apex and occurs in W, C & E Europe(Kurtto et al. 2004). The overall distribution of F. ulmaria is widespread throughout mostof temperate Europe and SE Asia, the distribution thinning out both southwards in the Mediterraneanbasin (absent from all of the Mediterranean isles) and eastwards into Russia and C Siberia (Kurtto etal. 2004, Map 3283). It has been introduced into eastern N America, but is scattered and notvery widespread there (Hultén & Fries 1986, Map 1048).
Threats
None.
Native, more frequent than occasional, especially on the western limestones, but always local. Europeanboreo-temperate, also in Greenland.
1900; Praeger, R.Ll.; Florencecourt.
May to January.
Growth form and preferred habitats
The limestones of the Burren in Co Clare (H9) and of the Ben Bulbin range in Co Sligo (H28) and theeastward extension of the latter in Fermanagh are the principal areas of Ireland where this low-growingscrambling or creeping, stoloniferous, more-herbaceous-than-woody species is most prevalent andwidespread. It tends to root in more moist, shady spots in rocky ground in open sun or semi-shade,including on knolls, scarps, screes, in ravine woods or in steeper tracts of woodland and on less acidicrocky heathland. It also occurs on many lowland limestone lakeshores, on riverside shingle and is evenrecorded around the shores of turloughs, ie the relatively small vanishing lakes in limestone districtsthat drain away through their floors.
Fermanagh occurrence
As the tetrad distribution map indicates, R. saxatilis is fairly frequent in Fermanagh, especiallyin the less agriculturally developed W of the county, having been regularly recorded in 54 tetrads,10.2% of those in the VC.
Flowering reproduction
Stone Bramble bushes flower shyly in June and July, the small, dirty-white blossom, 2-8 per compactinflorescence, being borne on somewhat stouter, more erect shoots than the normal, thin, downy,herbaceous, delicately prickled leafy stolons. The latter spreading vegetative stems, which can be up to100 cm long, are annually renewed from the deep-rooted, perennial rootstock. After pollination, eitherby visiting insects or by selfing, in late summer fruit is even more rarely produced than the flowers.
The ripe, translucent, scarlet fruit consists of just two to six large, quite separate (not coalescent)druplets (Nelson & Walsh 1991). While they are edible and indeed reasonably good to eat – althoughrather sharp in flavour – the fruit drops early, making them difficult to collect in any numbers.
It is generally assumed that birds eat the fleshy fruit and spread the seed (Lang 1987). While it hasbeen reported that Nut-crackers, Magpies and Blackco*ck take the berries in Scandinavia (Ridley 1930, pp.457, 506), Redwings in Iceland entirely ignored them in favour of the berries of Empetrum,Vaccinium and Arctostaphylos (Guitian et al. 1994). There is no direct evidence inthe literature for bird dispersal of R. saxatilis anywhere in B & I (Snow & Snow 1988).The scarcity of the fruit and its proximity to the ground suggests that mice and other small animals,perhaps even some invertebrates, might act as alternative seed vectors.
British and Irish occurrence
Stone Bramble is regarded as native in both B & I and it has a distinctly Scottish-centred northernand western distribution in these islands. The New Atlas hectad map indicates there have beencolony losses, mainly in lowland areas of England, due to site management changes and destruction,especially of woodlands. Many of these losses occurred pre-1930 and the distribution appears to havestabilised since then (D. J. McCosh, in: Preston et al. 2002).
Europe and world occurrence
From Greenland, Iceland and Arctic Russia to the Pyrenees and mountains of Italy, N Greece and theCaucasus, plus Turkey. Also stretching across temperate Asia to the Himalayas and Japan (Sell &Murrell 2014).
Threats
None.
Both native and introduced, an escape from cultivation. Very frequent. Circumpolar boreo-temperate, butwidely cultivated and naturalised.
1866-72; Stewart, S.A.; Fermanagh.
Throughout the year.
Growth form and preferred habitats
Suckering shrubs with biennial canes ranging from straggly to upright and from dwarf to tall, 100-200 cmin height. Wild strains tend to have many short, thin canes in comparison with cultivated varieties andvarying quantities of needle-like prickles. Leaves are also very variable, in shape and indentation ofthe leaflets which can number 3-5(-7), green above, chalky white and soft beneath, with dense stellatehairs (Haskell 1960; Edees & Newton 1988; Sell & Murrell 2014). The species often occurs in openwoodland or scrub, on heaths, or less often in hedgerows. In upland areas, it occurs on base-rich cliffledges and at the base of such rock faces. It also appears on waste ground and rough grassland asescapes and in quarries and other areas where garden rubbish has been tipped. The species is said toprefer sandy or well-drained soils (Edees & Newton 1988), although from the wide range of sites itoccupies in Fermanagh, this is not exactly obvious!
The origin and status of many colonies, either as native or a cultivated introduction, is sometimesdifficult or impossible to determine by inspection. However, the British authorities, Edees & Newton(1988), are definite in stating that R. idaeus is, "certainly native in many placesparticularly in hilly districts in the north." Since Raspberry is frequently bird sown, even thesite and habitat give little clue as to which we are dealing with. As might be expected, the fruits ofwild plants are generally distinctly smaller than those of cultivated varieties and, unlike the latter,they are mostly a deep purplish-red colour when ripe (Haskell 1960). After a few years in the wild,however, escaped cultivated plants inevitably suffer virus attacks and they then revert to forms bearingsmaller fruits, thus becoming inseparable from their wild cousins. However, in some apparently wildplants the fruits were not only large, but had the good flavour and the appearance of a cultivatedvariety, making it impossible to tell their origin from an inspection of the fruit alone. Plants of thetwo origins (wild and cultivated) also hybridise and genetically introgress, compounding the problem ofidentifying their status (Haskell 1960).
Male plants are readily recognised by their distinctive foliage (leaves simple or 3-foliate, leafletsimbricate, suborbicular, reniform, with broad crenate teeth) and by the rounded shape of their flowerbuds (Edees & Newton 1988)). A simple two-gene switch controls sex in the species: MF plants arehermaphrodite and Mf plants are males. The limited survey of wild R. idaeus populations byHaskell (1960) showed that the gene for male plants is widespread across Britain, stretching from WNorfolk to Inverness.
Despite the above concerns, R. idaeus is regarded as most likely or even certainly native in thehillier, more remote districts of B & I and especially so in the most northerly regions of Scotland(New Atlas). In addition to seed production, dispersal and establishment, Raspberry plantscommonly sucker from the roots and spread locally, particularly in more open or rocky, base-rich sites,forming dense prickly clumps or thickets (Edees & Newton 1988, pp. 18-9).
Fermanagh occurrence
In Fermanagh, R. idaeus is frequent and widespread, present in 266 tetrads, 50.4% of those in theVC. It is found on the margins of woods, on cliffs, in hedges, on heathland and the drier edges of bogs.Raspberry also occurs quite frequently as a garden escape or discard on waste or rough ground nearhabitation, when its origin is unambiguous.
Irish occurrence
R. idaeus is very widely distributed in Ireland, but the New Atlas hectad map indicates,even at this coarse level of resolution, that it is less prevalent down the west of the island (Prestonet al. 2002). Presumably this is due to the wetter, very acidic, peaty soils that predominatethere.
British occurrence
In Britain, R. idaeus is very frequent and widespread almost throughout, the main areas where itis less prevalent being fenland around The Wash, the far north of Scotland and a limited number ofscattered coastal sites (New Atlas). Again, since it is so widely cultivated, it is oftenimpossible to be certain of the origin of the plants recorded.
European and world occurrence
R. idaeus s.l. is a polymorphic species with a wide Euroasiatic and N American distribution.Subsp. idaeus is widespread throughout most of Europe, but only on mountains in the south. BeyondE Europe it also stretches across from the Caucasus and Turkey to W, N & C Asia where subsp.melanolasius Focke continues eastwards through most of E Asia. It also occurs over large areas of NAmerica. R. idaeus s.l. belongs to the circumpolar plants and has been introduced into eastern NAmerica, Greenland and New Zealand (Hultén & Fries 1986, Map 1053; Sell & Murrell 2014).
Threats
None.
Native, very rare.
1948; MCM & D; Scottsborough, near Magheraveely.
Fermanagh occurrence
This hybrid between two of the more commonly identified brambles in B & I, the Raspberry (R.idaeus) and the Dewberry (R. caesius), has only been recorded once in Fermanagh as listedabove. While R. idaeus (both wild and cultivated) has been recorded frequently and widely in Fermanagh,R. caesius is rare, having only been found on eleven occasions in the VC, although it is veryprobably under-recorded (see the R. caesius species account).
Hybrid identification
The parent species are very different in many respects, but their hybrid resembles R. caesius inhabit and stem characters. The leaves are closer to those of R. idaeus, but mostly with 3-5leaflets and the inflorescence is short and corymbose (Newton, A., in: Stace 1975). Hybrid plants withternate leaves can be distinguished from R. caesius by having pubescent carpels, curled hairs onthe abaxial leaf surface (ie the under-surface) and reddish-black fruits (if formed). Hybrid plants withfive or seven leaflets per leaf can be distinguished from R. idaeus by having leaves only looselyhairy on the under-surface, the stipules are broader, flowers have larger petals and, again, the fruitcolour is reddish-black. The flowers of hybrid plants can be either white or pink (Stace et al.2015).
British and Irish occurrence
Edees & Newton (1988) commented that hybrids of R. caesius with brambles of SectionCorylifolii are probably frequent and, indeed, this whole bramble Section possibly arose as aresult of crossing between R. caesius and Section Rubus. This could mean that up to 200European bramble taxa originate from pairings with R. caesius. Nevertheless, Newton (in: Stace1975), regarded R. caesius × idaeus as fairly rarely recorded in B& I and in his own subsequent book he listed a total ofjust nine English VCs from which records of the hybrid were known (Edees & Newton 1988).
The hectad map in The Hybrid Flora of the British Isles plots just two hectads with records ofthis hybrid in Ireland, one in Fermanagh and the other in N Tipperary (H10). However, for Britain, thesame map plots 35 widely scattered hectads where the hybrid has now been recorded (Stace et al.2015). With just one exception, where R. × pseudoidaeus wasaccompanied by R. idaeus only, the hybrid is always recorded in the presence of both parentspecies. The habitats where the parents coexist are listed as woods, hedgerows, wooded railway banks,commons and rough grasslands, roadside banks and the sides of ditches (Stace et al. 2015).
Cytology
R. idaeus is a sexual diploid species (2n=14 (mostly)), while R. caesius is either atetraploid (2n=28), or a pentaploid apomict (2n=35) (A. Newton, in: Stace 1975). The F1hybrid is, therefore, largely infertile (2n=35 (mostly) but the figure can range as follows: 21, 28, 35,42) (Stace et al. 2015). As the F1 generation is not completely sterile, latergenerations do exist and backcrosses with the parent species are said to occur (A. Newton, in: Stace1975).
Introduction, neophyte, cultivated, occasionally escaping and becoming naturalised.
24 July 1976; Dawson, Miss N.; Muckross, near Kesh.
March to November.
Growth form and introduction
A deciduous perennial shrub with an extensive branching rhizome and annual canes up to 4 m tall, nativeof western N America, R. spectabilis was introduced to B & I in 1827 by the famous Scottishplant hunter David Douglas. It was quickly taken up for its decorative flowers, for breeding ofautumn-fruiting and disease-resistant raspberries, as an almost prickle-free cane-forming shrub for gamecover in estate woodlands and to stabilize steep banks along roads and streams (Roach 1985; Oleskevichet al. 1996; Mitchell & House 1999). R. spectabilis avoids cold temperatures and isrestricted to mild, maritime climates. It has a relatively high shade tolerance and shows a strongpreference for open, moderately disturbed, moist, water-receiving sites with well-aerated soils, ofwhich it is a vigorous pioneer colonist. In a mild winter, with mean temperature of 6°C or above, R.spectabilis can continue minimum shoot extension throughout (Oleskevich et al. 1996).
It is not known exactly when it escaped from cultivation into the wild, but it had certainly done so andbecome thoroughly naturalized by the end of the 19th century in many areas of B & I and it nowappears to be increasing (D.J. McCosh, in: Preston et al. 2002).
Flowering reproduction
This very vigorous, tall, deciduous, thicket-forming shrub declares its presence early in the spring,producing solitary, conspicuous, 20-30 mm diameter, bright purple flowers on short, leafy lateral shootson mature canes from the middle of March through into June, followed by large, orange, edible fruits(Sell & Murrell 2014). The flowers are self-sterile and cross-pollination by insect visitors isobligatory. While the fruit looks like a large unripe raspberry (bright red, orange or yellow incolour), it is disappointingly rather tasteless. Thrushes do not seem to mind, however, and they consumethem and may spread the seed over considerable distances. Seed can survive burial in soil for at least100 years, making the species very persistent and extremely difficult to eradicate (Oleskevich etal. 1996).
Vegetative reproduction
Once established in a site, the plant consolidates itself by suckering vigorously from its spreadingunderground rhizome, rapidly forming dense, multi-layered, clonal stands. These can outgrow andeffectively exclude local competitors, eventually establishing dense single-species stands (Oleskevichet al. 1996).
Northern Ireland occurrence
Salmonberry is actively spreading throughout NI and the hectad map in the New Atlas shows thatUlster (ie the nine-county northern portion of Ireland) has the greatest continuous hectad distributionof the species anywhere in B & I (Preston et al. 2002). R. spectabilis was firstobserved in the wild in E Donegal (H35), Tyrone (H36) and Down (H38) in 1931, Londonderry (H40) around1945, Antrim (H39) in 1956, Fermanagh (H33) in 1976 and Armagh (H37) in 1981 (FNEI 3; CatAlien Pl Ir).
Fermanagh occurrence
R. spectabilis is now known in Fermanagh from 29 tetrads, 5.5% of those in the VC, butinexplicably it is confined to the N & W of the county. It occupies disturbed ground along linearlandscape features (often the first ground to be colonised by alien species) in moist, semi-shadedconditions in a wide variety of soils along river and stream banks, lakeshores, hedgerows and roadsides.The shrub is very persistent in gardens around long-derelict properties where it was obviously planted.
Irish Republic and British occurrence
R. spectabilis appears to be rather scarce and widely scattered in the RoI, although Reynolds(2002) does catalogue its presence in ten VCs there. Many of the sites listed in the RoI are fromroadside hedges, in estates or near gardens or churchyards where it has been planted, although some areclearly 'escapes' or garden discards in damp lakeshore or riverside woods and in thickets. In Britain,it is likewise very thinly but widely spread, except in the Scottish Midlands where it is concentratedaround the two major cities and their hinterlands. In some Scottish sites, the species has persistedmore than a century (Clement & Foster 1994; Preston et al. 2002).
Threats
Naturalised, actively spreading, persistent, potentially invasive alien that requires monitoring.
Native, common, widespread and locally dominant. European southern-temperate.
1900; Praeger, R.Ll.; Fermanagh.
Throughout the year.
Growth form and preferred habitats
Brambles or Blackberries are all-too-familiar, strongly invasive, dominant, stand-forming, woodypolycarpic perennials 3-7 m tall, with viciously sharp, spiny, arching, usually biennial branches and3-5 foliate, semi-evergreen leaves. The Revised Typescript Flora of MCM & D had many Rubusfruticosus segregates identified by the bramble referee W.C.R. Watson and these are listedbelow. Very little recent work has been done on Fermanagh's brambles, most recorders simply listingthese clothes-clinging plants as this species aggregate.
The name R. fruticosus agg. covers all the multiple segregates, the most common habitats of whichare hedgerows and wayside banks. Any disturbed, open, rocky, or waste ground is rapidly colonised bythem when neglected and undisturbed for any length of time greater than a few months. This includeslowland pastures and woodland, the latter especially after felling, fire or wind-throw opens up thecanopy. Brambles are most abundant on damp to wet acid soils and comparatively few microspecies grow onchalk or limestone. However, given a moderate level of vegetation disturbance and ample moisture, R.fruticosus agg. microspecies can grow on most soils, although they prefer more fertileconditions and do poorly on sandy or skeletal soils (Amor & Richardson 1980).
Variation, apomixis and taxonomic difficulties
Members of the R. fruticosus agg. belong to the subgenus Rubus and are extremely plastic ingrowth form and morphology to a wide range of environmental factors including light levels, whichtogether with the high level of genetic variation inherent in the apomictic (non-standard) seedproduction they exhibit, that makes identification very complex and difficult and, in B & I, largelya matter for a small number of bramble specialists. Thus the R. fruticosus agg. comprises acomplex group of more than 330 microspecies or agamospecies in B & I and over 2,000 worldwide (Amor& Richardson 1980; Edees & Newton 1988). The most recent count is now 353 microspecies for B& I (Sell & Murrell 2014), of which 100 have been found in Ireland mainly by two specialists,D.E. Allen and A. Newton (Newton & Randall 2004). The genus Rubus is arranged into fivesubgenera and very many sections and subsections, making it a little less confusing and bewildering forthe uninitiated.
Apart from one diploid species that reproduces in a normal sexual manner, R. ulmifolius (CommonBramble), all other Rubus microspecies are polyploids having multiple sets of chromosomes. Themajority (c 91%) are tetraploids (2n=28) and the rest range in a series from triploids to octoploids (ie2n=21 to 56). The complications only start there, as in order to set fertile seed most Rubusflowers require pollination with viable pollen (some cross- others self-pollination), to stimulate seedproduction, which is an asexual process, ie agamospermy is involved and the development of the embryo isautonomous, arising always and entirely from maternally derived tissues.
Seed reproduction, where there is a requirement for pollination of the stigma, but no fertilization of afemale gamete takes place, is called 'pseudogamy' (ie false marriage). Seed-formation in this situationis a form of asexual reproduction, ie it gives rise to seeds without sex and it is described asapomictic or agamospermous. Essentially, this is clonal reproduction by seed, giving the plant some ofthe advantages of seed formation (eg a 'clean egg' without any viral or genetic load, plus ease ofdispersal and dormancy), but doing so by the genetic equivalent of vegetative reproduction (Richards1997). Self-pollinating (autogamous) pseudogamous Rubus apomicts will be most successful inpollen-limited environments (eg arctic and alpine sites), as they are independent of pollinators(Richards 1997).
A consequence of apomixis and agamospermy is that the progeny are genetically identical to the femaleparent plant. However, in some Rubus microspecies a proportion of pollen is viable, so thatagamospermy remains facultative, rather than obligate. Partial or facultative apomixis is much morecommon than obligatory as we can never be certain that sexual reproduction is totally ruled out (Briggs& Walters 1997). Thus, from time-to-time sexual reproduction involving fertile pollen and ovulesdoes occur in microspecies within R. fruticosus agg., creating new hybrid microspecies genotypesthat may then persist and reproduce agamospermously. In this way, a large number (over 2,000) of closelyrelated microspecies have evolved over thousands of years, some dating back into the last ice-ages (Amor& Richardson 1980).
The large number of named microspecies in Rubus and the lack of standard sampling procedures usedin the past to delimit them has led to a great deal of long-term confusion in their taxonomy andnomenclature (Amor & Richardson 1980). However, more recent research and field work in B & I hasundoubtedly resolved some of the previously intractable identification and naming difficulties (Edees& Newton 1988; Sell & Murrell 2014).
Fermanagh occurrence
This aggregate has been recorded in 501 tetrads throughout Fermanagh, representing 94.9% of the squaresin the VC. The aggregate is the 17th most frequently recorded taxon in the Fermanagh Flora Database,lying between Mentha aquatica (Water Mint) and Hedera helix (Common Ivy). It ranks thirdin the VC in terms of number of tetrads in which it has been found, immediately after Juncuseffusus (Soft-rush) and Ranunculus repens (Creeping Buttercup). Like these, it too has avery wide ecological tolerance and it is most frequent to almost ubiquitous in lowland areas below 300m, apart from purely aquatic or wet peat situations. Nevertheless, R. fruticosus agg. ismainly found on neglected, derelict or ill-managed roadside banks, hedgerows, margins of woods, heathsand moors, scrub thickets and in rough grassland on rocky, more or less ungrazed or unmown ground.
Almost all brambles of this aggregate are long-lived, clonal entities and they are produced by thestrongly predominant vegetative reproduction that characterises this group. In our mild, damp climate,the leaves remain alive and functional over the entire winter period, presumably adding to the alreadygreat vegetative vigour of the plants. Strongly rooted crowns produce two to six arching woody shoots,well protected from browsers by large numbers of stout thorns and prickles. Despite this heavy armour,horses appear to relish soft, young, bramble shoots and they will sometimes tackle older foliage too.
Vegetative reproduction
Towards the middle or end of the growing season, arching branches root and form an overwintering restingbud when their tips touch the ground one or more metres distant from the original crown. Undergroundspreading adventitious shoots (suckers) may also be produced occasionally by roots in better, deepersoils (Amor & Richardson 1980). The individual woody stems usually persist for only two or threeyears but the plant has meantime produced sufficient younger stems to continue its invasive strategy ordominance of the site (Grime et al. 1988). Bramble can produce root suckers from a depth of 45cm, which makes it difficult or almost impossible to eradicate the species from ground of any real scaleby physical cutting, mowing and grubbing (Amor & Richardson 1980).
Flowering reproduction
Flowering branches are formed on canes more than one year old, usually second year canes. Theinflorescence is a leafy elongated rounded or pyramidal cyme, usually containing around 20 flowers. Thewhite- or pink-petalled flowers are hermaphrodite (perfect), actinomorphic (regular), 5-merous and 2-3cm in diameter. Numerous stamens with slender filaments and versatile anthers surround the many freecarpels, each of which contains two ovules, only one of which develops. Pollination is essential to bothfruit and seed production but, as mentioned above, there is no sexual fusion (ie pseudogamy occurs).Having said this, it is reckoned that some of the tetraploid microspecies may be capable of occasionaltrue sexual reproduction (Sell & Murrell 2014).
The flowers provide both nectar (partly concealed) and pollen as insect food and attract an abundance ofvisitors, including flies, bees, wasps, beetles, butterflies and moths (Proctor & Yeo 1973).
The seed becomes enclosed in the familiar shiny black edible drupe that in multiples form the blackberryfruit-cluster on a conical receptacle or torus. In contrast to raspberries, the blackberry fruit-clusterdoes not separate from the receptacle.
The variation, vigour and ubiquity of brambles highlights the success partial agamospermy is for them asa breeding system, perhaps particularly well adapted and flexible enough for the peculiar, occasionallong-term, repeating kind of unstable habitats they most frequent, such as gaps in forest canopy openedup by the death of old trees or by wind-throw (Proctor & Yeo 1973).
A small proportion of the bramble edible fruit clusters are eaten by birds, mammals (including foxes) andhumans, providing internal transport for the bramble seed to fresh sites. Fortunately, in terms oflimiting its invasiveness, regeneration from seed is infrequent, much of it apparently being non-viable;thus seedling establishment is poor. Despite the infrequency of seedling establishment, seed transportis important, enabling the plant to colonise newly available sites offering suitable habitat (Amor &Richardson 1980).
The flowering shoots have indeterminate growth and canes frequently form daughter plants in autumn bytheir tips when they touch the ground and root, providing an additional source of vegetativereproduction and giving rise to new plants in the following spring.
Weed control
Bramble plants are very difficult to physically eradicate, being difficult to uproot and capable ofregenerating from small root fragments left in the soil. However, a very specific systemic herbicide forwoody plants is available and works well against it, although it requires at least two applications peryear for several years to be effective. Spraying top growth in summer, followed by burning dead canes inwinter and respraying more thoroughly the following spring, offers a good measure of control (Amor &Richardson 1980).
British and Irish occurrence
Bramble is very common and widespread throughout both islands, except on the wettest, driest and highestground. It has probably lost some populations due to the removal of hedgerows when farmers increase thesize of arable and pasture fields, but it will have benefited from site disturbance around motorways andforestry operations. Together with other invasive weeds like Ivy (Hedera helix) and StingingNettle (Urtica dioica) the nutrition of the species has benefitted from widespread environmentalpollution with ammonia and nitrous oxides from agricultural slurry spreading and from general trafficexhausts (D.J. McCosh, in: Preston et al. 2002).
European and world occurrence
R. fruticosus agg. is common and widespread throughout Europe, the Middle East and NW Africa.Cultivated varieties have been introduced to S Africa, the E & W of N America, SW, SE & WAustralia, Tasmania and New Zealand for fruit and other purposes including hedging and soilstabilization. In many instances, the introduced material escaped into native vegetation and has becomea serious invasive weed (Amor & Richardson 1980).
Native, apparently very rare, but probably under-recorded.
July 1946; MCM & D; Lough-a-Hache, near Mount Darby.
All three existing records of this suberect bramble date from 1946-53 and were determined by Dr Watsonfrom specimens collected by Meikle and his co-workers. No recent records have been made due to a lack oflocal Irish expertise in this critical plant group. The remaining site details of the other two stationsare: Clareview Road, near Lisnarrick, 1947; and Imeroo Crossroads, NW of Tonyglaskan Bridge, 1953. Thesethree sites lie on the shores of Lough Erne or to the east of it and the Revised TypescriptFlora described the habitats as, "woods and hedgerows".
The 2004 Bramble Atlas hectad map shows that R. nessensis is the third most frequent andwidespread microspecies of Subsection Rubus in Britain behind R. plicatus and R.scissus. In Ireland, it is poorly recorded, however, appearing in only ten widely scatteredhectads. In Britain, R. nessensis is chiefly a plant of damp wooded riversides, where it formsrobust thickets. The few Fermanagh records fit this habitat pattern reasonably well, including both damplakeshore woodland and roadside hedges. If we generalise the habitat range of all the suberect bramblesoccurring in Fermanagh, they are inclined towards lime-rich, damp or wet, shady situations on or nearlakeshores, or along roadside hedges, rather than in the cut-over peat workings proposed by Hackneyet al. in FNEI 3.
Native, rare, but under-recorded.
1881-2; Barrington, R.M.; on shore opposite White Island, Lower Lough Erne.
Ten old records exist, all but one from the MCM & D survey of 1947-55, mainly scattered around thelarger lakes in the county. There are no recent records due to a lack of local identification expertise.All the MCM & D records had vouchers in K and seven were determined by W.C.R. Watson. Thelast two listed below were more recently determined by D.E. Allen. The remaining record details are:amongst stones (ie large rocks), Rosskit Island, Lough Melvin, 1948; lough shore at Crom Td, Upper LoughErne, 1948; Drumcose Lough, Ely Lodge Forest, 1949; Inishroosk shore, Upper Lough Erne, 1950; CorraghLough, near Upper Lough Erne, 1950; Florencecourt, 1950; E end of Boa Island, Lower Lough Erne, 1945-55;laneside near Keenaghan Lough, 1945-55; laneside near Gadalough, N of Keenaghan Lough, 1945-55.
With the exception of the Florencecourt estate record, for which RHN and the current author (RSF) have nofurther details, all the other stations are from sites near lakeshores which are clothed with bogs (orfens), wet woodland, or scrub thickets. There appears to be some doubt or disagreement surrounding thetypical habitat of this taxon. Edees & Newton (1988) describe this as, "heaths and moors,preferring dry sandy soils". However, in NI at least, R. plicatus appears to favour wetwoods and thickets, cut-over bogs and open rocky places, the latter probably being the driest of theseenvironments (NI Vascular Plant Database). The Fermanagh vouchers may therefore require furtherexamination.
R. plicatus belongs to the so-called suberect brambles, which originated as ancient hybridsbetween the true brambles or blackberries of the R. fruticosus species aggregate (Bramble) andR. idaeus (Raspberry). The Bramble Atlas hectad map suggests that this microspecies iswidely scattered in Ireland, mainly in northern counties, but with a further slight accumulation offinds in western areas from Cos Mayo (H26, H27) and Galway (H15-H17) and stretching S to Cork (H3-H5).In Britain, it is very much more commonly recorded than in Ireland and it is the most widespreadsuberect bramble throughout the larger island, matched in frequency only by R. scissus(Bramble Atlas).
Native, very rare but under-recorded.
1946; MCM & D, K, det A. Newton, 2005; damp woods by Lough Yoan, Castle Coole.
There are four records of this bramble in the Fermanagh Flora Database, two made by Meikle and co-workersand determined by Dr W.C.R. Watson and two by Mrs Y. Heslop-Harrison. All of her Fermanagh records aregiven with very vague site details only and, as they have not been verified by anyone, RHN and thecurrent author (RSF) are unsure as to their real status. In addition, her record dates are also vague,being given as 1940-60. Her two 'sites' are simply described as, "Roadside between Correl Glen andGarrison" and, "Near the head of the Correl Glen." The other MCM & D record isInishroosk, Upper Lough Erne, 1950. Originally Dr Watson identified a voucher specimen at K ofthe first record listed above as being R. nessensis Hall, but it was redetermined by Alan Newtonin 2005.
On the albeit very flimsy basis of just four records one might describe the likely Fermanagh habitats ofR. scissus as peaty ground in woodland, lakeshore and roadside situations.
This suberect bramble of Subsection Rubus is shown by the Bramble Atlas to be the secondmost frequently recorded and widespread member of that grouping in Britain. In Ireland, however, thesame map source shows this microspecies as being very much rarer and widely scattered. There are justthree hectads on the map of this taxon in the NI Vascular Plant Database (2005), two in Fermanagh (H33)and one in Co Armagh (H37), the latter dating from the 1970-86 period. Very probably, the apparentdifference in occurrence of this bramble on the two islands simply reflects the lack of local Irishinterest and expertise in this critical group of plants.
R. scissus is a very prickly, seven-leaflet bramble of peaty ground on the margins of heaths,moors and woods and especially of birch woods (Edees & Newton 1988) so, in theory at least, itshould find a multitude of suitable sites throughout Ireland.
Native, endemic, very rare.
1947; MCM & D; damp woods by Lough Yoan, Castle Coole estate.
Until 2004 there was a solitary Fermanagh record of this microspecies as detailed above, but D.E. Allen'sAugust visit to the VC in that year added a further four sites. The site details of his records are:Crom Visitor Centre, det A. Newton; Moher River, 2 km SE of Kinawley, det A. Newton; old railwaytrackbed at Lough Bresk, det Allen; Castle Archdale, Lower Lough Erne, det Allen.
The B & I distribution of this endemic taxon is quite inaccurately described in Edees & Newton's(1988) bramble monograph as, "widespread, but particularly frequent in the Pennine foothills".While the latter suggestion contained in this phrase is undoubtedly true, the subsequently publishedBramble Atlas of 2004 shows that in Britain this shrub is completely confined to the southernhalf of the island and it is concentrated in the W Midlands. Elsewhere in Britain, it is scatteredmainly along the S coast of England.
R. calvatus has also been found in two hectads on the Isle of Man, while in Ireland, all but oneof the 17 hectads plotted with records lie to the north of the distribution in England.
With respect to the Irish records, FNEI 3 simply states that R. calvatus is frequent in itsarea of remit, but it lists just three hectads in Cos Down and Londonderry (H38 & H40), but none inCo Antrim (H39). This is odd, since six years earlier Harron's Flora of Lough Neagh (p. 69)regarded this plant as being, "generally distributed" and listed all five Irish countiessurrounding the Lough, including Co Antrim. In their Rubus monograph, Edees & Newton (1988,p. 308), mapped the plant in nine hectads up along the River Bann and around Lough Neagh, confirmingtheir belief in this situation and the Bramble Atlas confirms this interpretation.
Native, very rare.
1948; MCM & D; Rosskit Island, Lough Melvin.
There are two Fermanagh records, both found by Meikle and co-workers, at Rosskit Island, Lough Melvin in1948 and in Ely Lodge forest on the shores of Lower Lough Erne in 1953. The habitats are given aswoodland margins and clearings and Meikle clarified this by adding, "on the shores of largerlakes" (Revised Typescript Flora).
Flora of Lough Neagh lists Newton & Hackney's find of this microspecies in a birch and alderwood at Killycolpy, on the Co Tyrone (H36) shore of Lough Neagh in 1985. It is described as the onlyIrish station, but clearly Newton & Hackney were not aware of the published Rosskit record (Meikleet al. 1949).
Nobody appears to know of the Ely Lodge record since it does not appear in the maps in Edees & Newton(1988), nor in the subsequent Bramble Atlas, although the Rosskit record listed above isincluded in both of these references. Newton (1986) also appears to have ignored or overlooked R.gratus in his interesting résumé and catalogue of Irish brambles of that date.
Native, very rare.
1949; MCM & D; near Newtownbutler.
This bramble is common and widespread in hedgerows and less so on woodland margins in most of lowlandBritain. It tends to need shelter, avoids exposed hills and coastal situations and becomes increasinglywestern in its distribution as one goes northwards into Scotland (Bramble Atlas). The BrambleAtlas hectad map shows that in Ireland R. lindleianus is very much more scarce orunder-recorded than is the case in Britain and especially so in the Republic of Ireland.
Until David Allen's August 2004 visit to Fermanagh there were just five scattered records: four found byMeikle and co-workers dating from 1949-53 (all verified by W.C.R. Watson), plus one made by Alan Newtonin 1984. D.E. Allen (DEA) added a further nine records to the VC, so this bramble is now known from atotal of 13 tetrads. In addition to the first record listed above, the details of the others are: scrubat Lattone Bridge near Garrsion, 1950; Wattle Bridge, 13 July 1950, R.D. Meikle, K, det A. Newton2005; Drumacrittin Lough near Rosslea, 15 July 1953, R.D. Meikle, K, det A. Newton; InishmoreBridge, ie E of the Inishmore viaduct, Upper Lough Erne, 1984, A. Newton; Moher River, 2 km SE ofKinawley, DEA, det A. Newton; near Pettigo, DEA, det A. Newton; Swanlinbar River S of Kinawley, DEA, detA. Newton; hedgerow on limestone, Roosky, DEA; field and plantation on clay, Roosky, DEA; woodlandaround eastern end of Glencreawan Lough, near Lough Navar Forest Park, DEA; Castle Archdale, Lower LoughErne, DEA; and Monea Castle, DEA.
In comparison with Fermanagh, the FNEI 3 lists one record for Co Down (H38) and two for CoLondonderry (H40) (one of which dates from 1909), plus a few further old unconfirmed records from CosDown and Antrim (H38 & H39).
Native, endemic, very rare.
1949; MCM & D; hedgerow near Gortaree gravel pits, Slieve Rushen.
This endemic bramble microspecies is widespread in the southern half of England and in coastal Wales. Itis also scattered throughout Ireland to the extent that Newton & Randall in the Bramble Atlasregard it as one of the most common brambles on the island. After examining the BEL herbariumspecimens, Alan Newton commented in the FNEI 3, that R. amplificatus was, "verycommon in all areas, as in Ireland generally".
Until David Allen's Fermanagh visit in August 2004 there were just two Fermanagh records. The first wasfound by Meikle and his co-workers as detailed above and Newton's own 1984 find at "Inishmorebridge, H23", which RHN and the current author (RSF) take to mean the E end of the InishmoreViaduct, Upper Lough Erne, as opposed to the definitely named 'Carry Bridge' which lies at the far endof Inishmore island. Allen added a further twelve records and R. amplificatus is now known tohave occurred in 14 tetrads scattered in lowland Fermanagh, 13 of them with post-1975 dates.
Newton (1986) listed the habitats of this microspecies as hedges, hedgebanks and the margins of heath,bog and woodland, the plant occurring on both acidic and calcareous soils. The Fermanagh habitats fitthis range well but also include a riverbank, a long-derelict railway track-bed and lakeshores.
Native, very rare, probably a mis-identification.
1948; MCM & D; drain at Sand Lough, Killyclowny Td, Upper Lough Erne.
While this microspecies is thinly scattered in C Ireland, it has in fact been recorded in a total oftwelve Irish VCs, including Co Cavan, W Donegal and Co Londonderry (H30, H35 & H40), adjacent oralmost so to Fermanagh (Bramble Atlas). The British stronghold of this bramble is in N Waleswhere it is very common, but it is also rather widespread but much less frequent in other parts of SEngland and Wales. Meikle and his co-workers found the solitary known Fermanagh station in 1948, on thedamp bank of a ditch on the NE shore of Upper Lough Erne. The more usual habitats of this microspeciesare in open woods or on woodland margins, heaths, hedges and banks (Edees & Newton 1988). In 2005,both A. Newton and D.E. Allen examined the voucher in K. Newton commented that it was not R.incurvatus, while Allen suggested it could be a stunted form of R. nemoralis.
The plant has one locality in Denmark, so it is not endemic in B & I (Newton 1986).
Native, very rare.
1950; MCM & D; a scrub thicket in the Correl Glen woodland.
This bramble was collected in 1950 and identified as R. broensis W.C.R. Watson by the author ofthe name who was the BSBI Rubus microspecies referee at that time. This is one of severalsynonyms of this plant, subsequently renamed by him R. septentrionalis W.C.R. Watson.
The solitary Fermanagh record is the earliest Irish record of this microspecies, although it has beenoverlooked and does not feature in any published form until now. The Bramble Atlas hectad mapcontains two post-1988 Irish records from E Mayo and Sligo (H26 & H28). The same map shows that thismicrospecies, which is widespread in N Europe, has its British stronghold very firmly located inScotland where it is really very common, including the Western Isles and Orkney (Bramble Atlas).
The Fermanagh thicket habitat is atypical, the plant generally being found in open woods, the margins ofmoors, on the banks of rivers and streams, or on gravel spreads (Edees & Newton 1988).
Native, endemic, rare but very probably under-recorded.
August 2004; Allen, D.E.; Monea Castle.
This microspecies in Series Rhamnifolia is a B & I regional endemic with scattered populationswhich the Bramble Atlas plots across southern England and up the Welsh coast, along the Kintyrepeninsula in SW Scotland, plus twelve hectads dispersed from W to E from coast to coast in middle Irishlatitudes (VCs H21-H23, H27,H30 & H31). English records are most concentrated in East Anglia centredon Norwich and also along the Suffolk coast, so that there appears to be a definite, but not exclusivelylittoral distribution. The four Fermanagh records, all made by D.E. Allen on a visit in the summer of2004, are the most northerly Irish occurrences known.
The site details of Allen’s other three records are: old railway track at Lough Bresk; Ederney square,hectad grid reference H26; Ross Lough near Carr Lough, hectad grid reference H14. The last two siteshave been given surprisingly vague locations for modern records (Allen 2005b).
Native, rare and certainly under-recorded.
1948; MCM & D; laneway near Garrison.
This is another microspecies in Series Rhamnifolia which is mapped in the Bramble Atlas ina manner that indicates it is one of eight brambles which, relatively speaking, are the most common andwidespread in both B & I. There is nothing remarkable about its range of typical habitats, whichinclude heaths, rough moorland, woodland margins and other open marginal situations including roadsidesand quarries (Edees & Newton 1988). R. nemoralis is quite certainly under-recorded inIreland, yet even with this serious data shortcoming, it definitely is more frequently recorded innorthern parts of the island than elsewhere. Despite this situation there are very occasional, sporadicor rare records of R. nemoralis from 29 of the 40 Irish VCs.
In Britain, R. nemoralis is described as very common in N England and Scotland, but it is alsowidespread elsewhere in suitable habitats (Edees & Newton 1988; Bramble Atlas). As with allstatements regarding the occurrence of brambles in an Irish context, it is necessary to qualify anyobservation that is made by recognising the overall poor coverage of the recording in comparison withBritain. In Fermanagh, there are a total of eleven records of this bramble from separate tetrads. Sixdate from the previous survey by Meikle and his co-workers (1948-53) and five were located and collectedby D.E. Allen (DEA) in 2004 and subsequently determined by him or by Alan Newton (AN). Apart from DavidAllen's visit in August 2004, RHN and the current author (RSF) recognise that almost certainly no onehas spent more than a few hours researching brambles in Fermanagh since 1953.
As usual, five of the six MCM & D R. nemoralis vouchers from Fermanagh were determined by DrW.C.R. Watson. The sixth record is the earliest, as listed above, and was a voucher at Kre-determined to be this species by AN in 2005. The habitats were described by Meikle et al. inthe Revised Typescript Flora as roadsides, thickets and lakeshores. Apart from the firstrecord listed above, the details of the other five are: Tempo River near Shanco Td, 1949; Arney village,1952; E end of Boa Island, Lower Lough Erne, 1953; Killyvilly Lough, NE of Rosslea, 1953; CoolyermerLough, 5 km ESE of Enniskillen, 1953. David Allen's August 2004 five sites are listed as: Ports Tdshore, Upper Lough Erne, det AN; Crom Visitor Centre, det AN; near Pettigo, det DEA; field andplantation margin on clay, Roosky, det DEA; on clay, old railway track-bed near Lough Bresk, det DEA.
Perusal of the Bramble Atlas shows that R. polyanthemus, which is in the same brambleSection, is the microspecies which shares the most similar B & I distribution pattern with R.nemoralis (Bramble Atlas, p. 29). Both of these bramble microspecies cover almost theentire latitudinal range of Britain and, while they are scattered throughout Ireland, they areprincipally recorded in the northern half of the island. However, this Irish distribution may do no morethan reflect the itineraries of the several Rubus experts who have visited the island for briefperiods, especially in recent years (Newton 1986; Allen 2005a & b).
Native, rare but undoubtedly under-recorded.
1904; Praeger, R.Ll.; Bunnahone Lough, Lenaghan Td.
There are eleven Fermanagh records of this bramble in ten tetrads and their dates stretch across 100years from Praeger's find listed above to the six records made by D.E. Allen on his visit in August2004. Interspersed with these are Meikle and co-worker's "Glens at Slieve Rushen", 1949 andtheir "Ely Lodge forest", 1953, Mrs Y. Heslop-Harrison's "Between Belleek andGarrison", 1949 (a K voucher re-determined by D.E. Allen), plus Alan Newton's record made atInishmore viaduct in 1984. The site details of D.E. Allen's records (some of them rather vague) are:Crom Visitor Centre; Moher River 2 km SE of Kinawley; near Pettigo; Ely Lodge Forest; roadside hedge onlimestone, Roosky; on clay on old railway track-bed near Lough Bresk. The first three of these recordswere determined by Alan Newton.
The local habitats of R. polyanthemus include thickets in lakeshore woods, upland glens and alongold railway tracks, plus in lowland hedgerows. The seven tetrads are widely spaced and all but one lieto the south of Lough Erne, the more northerly outlier being on the long disused railway track nearLough Bresk.
Even when recognising the extent of bramble under-recording in this area, the dearth of records of thisparticular microspecies is rather surprising in view of how widespread and often abundant it isthroughout Britain and even in fact in relative terms in other parts of Ireland (Bramble Atlas).
In comparison with Fermanagh, the FNEI 3 lists records from a total of 28 hectads in the threenorth-eastern Irish VCs and Hackney et al. described this form of bramble as, "verycommon" in their survey area. In other parts of B & I, apart from the habitat types representedin the Fermanagh, R. polyanthemus is commonly found on heaths and in sand and gravel quarries(Edees & Newton 1988).
Native, occasional or rare, but certainly under-recorded.
1882; Stewart, S.A.; Co Fermanagh.
In comparison with other bramble microspecies of the R. fruticosus agg., R. ulmifolius is asexual diploid of rather distinctive appearance (Grime et al. 1988). The chalky-white feltedunderside of the leaves, together with the deep pink or lilac colour of the petals, filaments andstyles, plus the well-defined habitat preferences, make this a relatively easy form of bramble to get toknow and recognise. Typically, R. ulmifolius occupies south-facing, sunny, sheltered situationsin limestone or coastal ground, with warm, dry, calcium-rich, often sandy soils.
Despite the fact that the Revised Typescript Flora authors only knew of seven sites for thisbramble microspecies in Fermanagh, they still regarded it as being the prevailing bramble in thelimestone areas of the county (Revised Typescript Flora). Undoubtedly, the distinctive appearanceof R. ulmifolius helps explain why they made this statement, which really appears to express anexpectation as if it were an actual finding. The distinctive nature of this bramble alsojustifies the widespread, indeed almost ubiquitous occurrence of this microspecies in Britain south ofthe Lake District and Yorkshire, as well as its relative commonness in lowland Ireland – although it isstill under-recorded in the last (Flora of Co Dublin; Bramble Atlas).
This bramble is represented in a total of 14 Fermanagh tetrads, although only seven tetrads containpost-1975 records. Apart from Stewart's first record with its unspecific site listed above, there are 14additional records, seven of which were discovered by Meikle and his co-workers between 1949-53. Asolitary record was made by Alan Newton (AN) during a visit in 1984 and a further six sites were listedby D.E. Allen (DEA) in August 2004. The site details of these are as follows: Ballagh crossroads, 1949;Hanging Rock NR, 1949; cliff to S of Doagh Lough, NW of Knockmore, 1950; Bunnahone Lough, 1950; nearKesh, 1953; Belleek, 1953; Lough Scolban, 1953; E of Inishmore Viaduct, Upper Lough Erne, 1984, AN; CromVisitor Centre, det AN; near Pettigo, det AN; Swanlinbar River, S of Kinawley, det AN; woodland N ofMagho viewpoint, overlooking Lower Lough Erne, det DEA; on clay on old railway track-bed at Lough Bresk,det DEA; Monea Castle, det DEA.
The habitats of the seven Fermanagh sites discovered by MCM & D were summarised by them in theRevised Typescript Flora as, "roadsides, banks, woods and screes", and theirrecords were thinly scattered in the limestones to the W of Lough Erne. The seven Newton and Allenrecords add this bramble to just four more tetrads, but it is now more definitely scattered across theVC and it is less obviously associated with lime-rich soils.
The English Rubus expert Alan Newton revising the records for the FNEI 3 regarded R.ulmifolius as, "very common and widespread and especially abundant near the coasts of allthree counties". However, R. ulmiflorus is a highly variable, genetically polymorphicbramble that changes its form in response to environmental conditions, ie phenotypically it is veryplastic. In addition, this bramble being sexually active, it hybridises with R. caesius and withother stable forms of bramble microspecies. It crosses particularly frequently with R. vestitusand, when it does so, it forms either fertile or sterile offspring (Edees & Newton 1988).Clearly even when dealing with this relatively distinctive bramble, identification is not always plainsailing!
Seemingly native, apparently rare, but probably overlooked and under-recorded. An Anglo-Frenchtransmarine species with an Irish connection.
August 2004; Allen, D.E.; birch-willow woodland, Lough Achork.
This bramble is a member of the woolly and thinly glandular series, Vestiti. It was described by D.E.Allen (1994) from material first discovered abundant and widespread in Jersey (the alleged Roman name ofwhich it bears). Apart from a group of roadsides in S Hants (VC 11) between Portsmouth and Southampton,this was only known elsewhere from old herbarium specimens from single locations – a dock-side inGloucestershire (VC 33, 34) and an unspecified site in Co Cork (H3-H5) until David Allen visitedFermanagh in late August 2004 (Allen 2005a). Here, Allen was quite shocked and amazed to discoverspecimens, unquestionably belonging to this same bramble species, in three sites within a few days ofone another. The first patch at the site listed above was discovered looking perfectly native, in openbirch-willow woodland 100 m from the E shore of Lough Achork, Lough Navar Forest Park. This station onthe Western Plateau is quite remote from human habitation.
A second patch was discovered a few days later in more artificial circ*mstances around the edge of theroadside car park below the viewpoint on the Cliffs of Magho overlooking Lower Lough Erne. This secondlakeshore site is approximately 3 km NE of Lough Achork, but at an altitude 150 m lower. A day later athird even larger patch of R. caesarius was recorded in a clay pasture, close to the ruins ofMonea castle, approximately 13 km SE of the second site (Allen 2005b).
The Bramble Atlas map, which takes account of notified finds or re-determinations up until 2000(Allen 2005c), plotted only the Channel Island and S Hants records of R. caesarius. Theassociated caption for the map described the taxon as, "a recently described Anglo-Frenchtransmarine species". Clearly, however, this bramble has a newly revealed Irish connection and, atleast in the case of the Fermanagh plants, this is not associated with the coast, nor with ports (Allen2005a, b).
Native, endemic, very rare.
1948; MCM & D; hedgerow by Lough Melvin, near Garrison.
There are a total of five Fermanagh records: two from the extreme west of the county date from the late1940s or early 1950s were recorded as R. hebecaulis Sudre, a frequent synonym for thismicrospecies, plus three records made by D.E. Allen in August 2004. The first station detailed abovefound by Meikle and co-workers had as usual a voucher determined for them by the Rubus referee DrWatson. The second record was made by Mrs Y. Heslop-Harrison who apparently identified the plantherself. All her Fermanagh plant records have only vague site and date details available, in this case,"roadside between Belleek and Garrison, 1945-55". The site details of Allen's three finds are:Swanlinbar River S of Kinawley; area of cleared conifer plantation, Lough Navar Forest Park; and E endof Lough Achork.
On the evidence of the Bramble Atlas, this endemic bramble is now quite widespread in Ireland,although still scattered both north and south of the border with the Republic of Ireland. The hectad mapshows that a considerable number of post-1988 records have been made in B & I including on the Isleof Man. For comparison with the Fermanagh situation, elsewhere in NI, the FNEI 3 lists only tworecords and there are now three additional post-1988 hectads in Co Down (H38) and one station plus anadditional hectad in Co Londonderry (H40) (FNEI 3; Bramble Atlas).
While in Fermanagh this plant has been found in roadside, riverbank and lakeshore hedgerows, the moreusual habitat in B & I is on wood margins. The recently cleared softwood plantation mightapproximate to the latter. In his Irish bramble summary paper, Newton (1986) regarded it as,"occasionally [found in] hedges and banks".
Native, a regional endemic, very rare.
1949; MCM & D; wooded glen at Slieve Rushen.
The first record listed above was collected by Meikle and his co-workers in 1949 and subsequentlydetermined by Dr W.C.R. Watson from vouchers in K as usual. Another voucher in K collectedby MCM & D was originally somewhat tentatively identified as R. rhobophyllus Muell. &Lefèv. by Watson. It was re-determined as this microspecies by D.E. Allen in 2005. The site of thesecond record is, "road between Tempo and Brougher Mountain". The addition of these twopreviously unpublished Fermanagh records widens the range of this regional endemic that is named afterthe 19th century Co Down clergyman and botanist, Canon H.W. Lett (FNEI 3).
R. lettii appears confined to the north of Ireland (ie the province of Ulster), and the Isle ofMan (VC 71) and the addition of Fermanagh to its range, brings to eight the total number of VCs in whichit appears. The other Irish VCs that have records are Co Cavan (H30), E Donegal (H34), Tyrone (H36), CoArmagh (H37), Co Down (H38) and Co Antrim (H39) (Bramble Atlas).
A number of records of R. lettii that were published by Allen (1990, 1993) in Co Dublin (at Howth)and in Meath (H21 & H22) appear to have been discounted as errors or renamed since they do notfeature in the Bramble Atlas. Clearly even bramble experts like David E. Allen and Alan Newton(acknowledged for their assistance with identification and for reading drafts of the two aforementionedAllen papers), can and do make mistakes with this bramble group. This is comforting knowledge for thoseof us who find bramble variation completely baffling! It therefore appears as if Co Cavan (H30)represents the present, true southern distribution limit of this microspecies and there are records fromthere by G.C. Druce in 1912, R.Ll. Praeger in 1929 and A. Newton in 1992 (Reilly 2001).
Native, locally frequent.
1947; MCM & D; Mullylusty Td, referred to as, "the Lurgan River Glen".
Fourteen Fermanagh records of this bramble exist, eight dating from the 1947-53 period discovered byMeikle and co-workers and determined by W.C.R. Watson, plus six recorded around Lower Lough Erne by D.E.Allen in August 2004. This is sufficient to make R. vestitus one of the most common Rubiin the Fermanagh Flora Database: it occurs in 13 tetrads, although only six of these contain post-1975records. The authors of the Revised Typescript Flora suggested that this particularmicrospecies, which belongs to Series Vestiti, is locally common in Fermanagh, especially off thelimestone (but see comment on this below). The locally recorded habitats were woods, lakeshores androadsides. The records are quite widely scattered across the county, but are slightly more frequent in SFermanagh.
The Bramble Atlas map indicates that with the exception of Scotland where it is rare and coastalthis is one of the most frequent and widespread identified forms of bramble elsewhere in B & I andespecially so in S England, the Midlands and in Wales. In common with other bramble microspecies, inIreland R. vestitus appears very much more rare and scattered than in Britain. As usual, thismust be attribute to a lack of recorder effort with this critical group. Having said that, with recordsymbols in the Bramble Atlas plotted in 62 Irish hectads (RSF's visual count), this has to beranked in terms of frequency as one of the top ten most widespread bramble microspecies in Ireland. InNI, however, and particularly in the three counties of NE Ireland, Hackney et al. (FNEI 3)regarded this plant as, "apparently scarce", listing a total of just nine sites.
R. vestitus is a plant of woods, wood margins and hedge banks, described by Edees & Newton(1988) as being often abundant on calcareous or clay soils and rarer on acid soils on heaths and moors.As such, it has calcicole habitats, but not absolutely so, which helps account for its occurrence in themore acidic Fermanagh situations. Taylor (2005) found R. vestitus could tolerate conditions inbrown-earth soils in the very wide range pH 3.8-7.6 in terraced scarp woodland in Cumbria, where patchyglacial drift overlay Carboniferous limestone. He showed that efficient nutrient-cycling by themycorrhizal roots was of vital importance. The brown-earth woodland soils involved were highlyphosphate-fixing, this element being the main factor limiting plant growth in three local soil variantsstudied.
Red-flowered and white-flowered forms of R. vestitus occur, sometimes side-by-side, although whiteforms are usually much more common off the limestone. The red-flowered form (var. roseiflorus N.Boul.) was recorded by Meikle and his co-workers at Lower Lough Macnean and also in the Correl Glen.
Native, very rare.
1949; Meikle, R.D.; hedge in lane near shore of Bunnahone Lough, Lenaghan Td.
The solitary Fermanagh record of this bramble was originally identified as R. drejeri by Dr W.C.R.Watson, but in 1993 David Allen revised the determination from a herbarium voucher in BEL (Allen1993). In 2005, Allen likewise re-determined another voucher of this same record held at K.
R. wirralensis is recognised to be a hedgerow and hedge bank plant, which is widely scattered inIreland, most frequently in the north. The FNEI 3 lists two stations in Co Down (H38) plus 14other hectad grid squares in Cos Antrim and Londonderry (H39 & H40).
This microspecies is also widespread in Britain from Islay to Devon, but it is most frequent in the westand especially so in Lancashire. Newton & Randall (Bramble Atlas) regarded it as, "a keymember of the Irish Sea florula".
Nobody has rediscovered this bramble microspecies in Fermanagh for almost 60 years, but RHN and thecurrent author (RSF) do not doubt but that an expert visiting batologist might do so. This highlightsthe problem with critical groups of species of any sort. It is obvious that the distribution mapsproduced in such cases more than anything else reflect the travel itineraries and the time spent in thefield by sufficiently expert observers, rather than representing a valid scientific sampling of theactual geographical distribution of the organism of interest. This is not a recipe for doing nothing,nor a council of despair; rather it ought to be a spur to do better!
Native, very rare, but possibly a mis-identification.
1949; MCM & D; glens at Slieve Rushen.
The Bramble Atlas hectad map shows this microspecies has a cluster of recent (ie post-1988)records in nearby Cos Cavan and Monaghan (H30 & H32),plus a more remote one on the coast of Co Antrim (H39). Newton & Randall (Bramble Atlas),apparently unaware of the above listed Fermanagh record by MCM & D, incorrectly considered them asbeing the first Irish records. D.E. Allen examined the voucher for this record at K in 2005 andcommented that it is possibly this species since it is present in quantity in nearby Cavan, but in hisview, it is more probably R. anisacanthos G. Braun (see below). A second MCM & D recordcollected at Bunnahone Lough, Lenaghan Td in 1949, identified by Dr Watson from the K voucher asR. drejeri, has similarly but more definitely been re-determined by D.E. Allen in 2005 as R.wirralensis (see above).
R. drejeri is a northern bramble of unexceptional habitats, ie wood and heath margins andhedge banks (Edees & Newton 1988). It is a widespread bramble in NW Europe. In Britain, thismicrospecies is thinly scattered from Inverness south to Northamptonshire, with a strong frequencyaround the Firth of Forth reaching coastal Northumberland (Bramble Atlas).
Native, very rare.
1949; MCM & D; glens at Slieve Rushen.
The voucher of this bramble was originally identified by Dr Watson as R. drejeri but D.E. Allenexamined and re-determined it in 2005 to be this microspecies. A second MCM & D voucher in Klabelled, "Side of a lane between Kesh and Muckross" dating from the 1945-55 period, wasoriginally identified as R. lasiostachys but has also been re-determined as this microspecies byAllen.
R. anisacanthos is described in the Bramble Atlas (p. 53) as a "North SeaFlorula" member having a majority of its sites on the E coast of Britain, although it is also quitewell represented in the English NW and Midlands, and also north of the Scottish border in Galloway. InIreland, the representation in the Bramble Atlas map shows that it has a widely scatteredoccurrence, being slightly more prevalent in the northern half of the island. However, 17 of the 26Irish hexads with records belong to the pre-1988 date class, so as usual it is very obviouslyunder-recorded. Its typical habitats are described by Edees & Newton (1988) as hedgebanks and themargins of woods and moors.
Native, endemic, very rare.
1948; MCM & D; hedge lane near Garrison.
This bramble, of which there are just three Fermanagh records, was first noticed and collected in theperiod 1893-1901 by two clergymen botanists in western and southern parts of Co Down (H38) (ie atAghaderg, Annaclone, Scarva and near Newry, Banbridge and Saintfield towns). The two gentlemen regardedthe plant as locally common in most of these areas. They sent their unidentified specimens to thebramble referee of the time, the Rev W. Moyle Rogers, who identified and named it R. drejeri var.hibernicus. Subsequently he changed the name to the present one in 1899 (Edees & Newton1988). Rogers wrote up this and many other Rubi he received from NE Ireland in a paper hepublished in the Irish Naturalist in 1901 (10: 213-20).
Nowadays this microspecies is regarded as a B & I endemic form and like most brambles it is a plantof hedgerows and the margins of woods, heaths and moors (Edees & Newton 1988). This ecological andhabitat range allows it to crop up in other forms of disturbed ground, eg on roadsides and in wasteplaces. The FNEI 3 account adds very little to the early Co Down records, except to comment thatthis bramble is closely allied to R. dunensis W.M. Rogers, another local endemic microspecies. Itdoes report, however, that in 1985 Alan Newton and Paul Hackney refound R. hibernicus at Aghaderg(GR J1142).
Newton has also examined the DBN herbarium Rubus specimens and in Cen Cat Fl Ir 2,R. hibernicus is listed as occurring in Cos Down and Armagh (H38 & H37). Presumably thismeans that the early, "near Newry" record was not in Co Down, but actually across thecounty boundary in Co Armagh.
The first two Fermanagh records of R. hibernicus were made by Meikle and his co-workers in 1948,the specimens as usual being determined by Dr W.C.R. Watson. They were published in INJ 9:225 (Meikle et al. 1949) and the given site details were, "On a ditch on the slopes ofSlieve Rushen" and, "In a hedgerow near Garrison." D.E. Allen has recently discovered thetwo vouchers in K (Allen 2005c). A third MCM & D record was listed in the 1975 RevisedTypescript Flora as E end of Boa Island, Lower Lough Erne.
RHN and the current author (RSF) have no definite record of the whereabouts of most Fermanagh Rubivouchers but we surmise that they are scattered among the very many British and Irish Rubusfolders at K since R.D. Meikle worked there for many years (Allen 2005b). However, it appears asif these three Fermanagh records have been long overlooked by subsequent Rubus experts, sincethey do not appear in the Cen Cat Fl Ir 2, nor in Edees & Newton's (1988) monograph.
The authors of the Bramble Atlas also overlooked these Fermanagh R. hibernicus records, butthe hectad map does display a post-1988 record of this bramble in W Galway (H16), which is ratherremote from all other stations of the plant. Elsewhere there is a crop of records in the Isle of Man (VC71) and in S Wales – which is very much the major centre of the known distribution, plus a number acrossthe Bristol Channel in Somerset (VC 5 & 6), Wiltshire (VC 7 & 8), S Devon (VC 3) and Dorset (VC9). This distribution makes R. hibernicus a member of the 'Severn Bay Florula', in terms ofbramble phytogeography, although for some unexplained reason the published map associated with thisgrouping does not include any of the Irish territories of this microspecies (Bramble Atlas, seemap on p. xv).
Native, endemic, very rare.
15 July 1953; MCM & D; laneside near Rosslea.
Two bramble vouchers in K separately collected, first by Meikle and his co-workers and,subsequently, by Mrs Heslop-Harrison between 1940 and 1960, were re-determined as this microspecies in2005 by David Allen. The first record listed above was previously labelled R. adenenthus by DrWatson. Mrs Heslop-Harrison's record from a roadside between Belleek and Garrison was identified by heras R. wedgwoodiae. A third record was made by Alan Newton on a brief excursion into Fermanagh in1984 and David Allen added a further 14 records in August 2004.
R. adenanthoides is now known from 16 Fermanagh tetrads, 15 of them with post-1975 records. This B& I endemic is a plant of open woods, margins of heaths and moors and of roadsides, particularlyfound at higher altitudes. Ecologically it does not shun limestone soils (Newton 1986). This brambleappears to be widely scattered throughout Ireland, although it is much more frequent to abundant in thenorthern half of the island.
R. adenanthoides is common and widespread in N & C England, Wales, and the Isle of Man (VC 71)and there are outliers in both N Scotland and in Oxfordshire (VC 23) (Bramble Atlas).
Native, very rare.
1904; Praeger, R.Ll.; Monawilkin Lough.
July.
In his fascinating Irish Naturalist paper entitled, 'Among the Fermanagh Hills', Praeger (1904)recorded this plant as R. longithyrsiger Bab. var. botryeros, Foche. He found it near theedge of Monawilkin Lough and described it as a rather small but strongly armed form. After making acomparison with two specimens collected by botanists elsewhere in B & I that he regarded as similar,Praeger went on to comment, "This bramble is hitherto unrecorded for Ireland, either type orvariety." RHN and the current writer (RSF) now regard the Monawilkin specimen as R.longithyrsiger Lees ex Focke, since R. botryeros (Focke ex W.M. Rogers) W.M. Rogers is infact confined to SW Britain and NW France. In contrast, R. longithyrsiger is found in both B& I.
In Ireland, as far as we can tell, given the usual under-recorded bramble situation, R.longithyrsiger is widely scattered, but is chiefly found somewhat inland in counties borderingthe south coast. The pattern in Britain is very much clearer and more certain, the centre ofdistribution being in SW England, S. Wales and the Welsh Borders, with outliers in N Wales, Berks (VC22) and N Essex (VC 19).
The legend of the Bramble Atlas hectad map of R. longithyrsiger describes this microspeciesas, "a major element of the general Lusitanian bramble flora". In our view this appears odd ormistaken, since in the 'Phytogeographical analysis' chapter of their book these authors list it as amember of the 'Severn Bay Florula'. In addition, R. longithyrsiger does not appear to have anygeographical affinity with the 14 species that these batologists group into the west of Ireland'Lusitanian Florula' (Bramble Atlas).
Native, a rare endemic.
1953; MCM & D; waste ground at E end of Boa Island, Lower Lough Erne.
A voucher for this record at K originally identified by Watson as R. hibernicus wasexamined and re-determined as this microspecies by D.E. Allen in 2005. The Bramble Atlas mapshows a very heavy concentration of English records in Shropshire (VC 40) and very few elsewhere on theisland. In Ireland, the same map shows a scatter of new finds dating from 1988-2000 among a very fewolder records. These more recent Irish records are predominantly associated with the Dublin area and theE coast northwards to Dundalk, further emphasising the manner in which known bramble distributionreflects the travel routes of a few expert visiting batologists.
Native, a local endemic and here a definite mis-identification.
1881; Stewart, S.A.; near Derrygonnelly.
This 19th century record was not checked by W.C.R. Watson, but apparently a voucher wastentatively identified by Prof Babington from rather inadequate material as belonging to thismicrospecies (or rather to R. emersistylus Mull., the synonym of the time) and it was publishedby Stewart (1882) with this uncertainty hanging around it. The Bramble Atlas map shows that thisis a very local regional endemic microspecies, found only in S Devon (VC 3), so that it can now besafely discounted as a definite error.
Native, endemic, very rare.
1949; MCM & D; hedgerow on the Tempo River, near Shanco Td.
As the Fermanagh tetrad map shows, there are five Fermanagh records in scattered tetrads of this quitewidespread endemic B & I bramble. The first was collected on the bank of the Tempo River by Meikleand his co-workers and determined by Dr W.C.R. Watson. On his August 2004 visit to the VC, David Allenrecorded a further four sites around the VC. The sites and determiners of the latter records are: CromVisitor Centre, det A. Newton; Swanlinbar River S of Kinawley, det A. Newton; hillside plantation andhedge of carpark, Magho Td, det D.E. Allen; on clay, old railway track-bed, near Lough Bresk, det D.E.Allen.
The Fermanagh plants of R. echinatoides are associated with hedges, scrub in derelict railwaytrackways and river banks (Allen 2005b, c), but elsewhere it is additionally found on the margins ofwoods, heaths and moors (Newton 1986; Edees & Newton 1988). The Bramble Atlas map shows thatR. echinatoides is relatively frequently recorded across the northern half of Ireland, but it isvery much rarer further south.
In Britain, it is very much more widespread, with its heartland in N & C England, ie Lancashire (VC59 & 60), Yorkshire (VC 61-65) and the Midlands, but with outliers beyond Inverness and also towardsthe English south coast in S Hampshire (VC 11)(Bramble Atlas).
Native, a regional endemic, definitely a mis-identification.
1949; MCM & D; roadside, Ballagh Crossroads.
The solitary record, as usual identified by W.C.R. Watson as this microspecies, is certainly incorrect.The Bramble Atlas map displays R. fuscus as a regional endemic of S & C England, whichhas no Irish, Welsh or Scottish records. RHN and the current author (RSF) feel that the most probablesubstitute would be R. fuscicaulis Edees, of which there are a few Irish records in nearby CoRoscommon (H25), but we await with interest to see if a voucher appears in K and, if it does,what the current Rubus referee will make of it.
Native, a regional endemic, a mis-identification.
1940-60; Heslop-Harrison, Mrs Y.; roadside between Belleek and Garrison.
This is another of the brambles that Mrs Heslop-Harrison collected somewhere between Belleek and Garrisonat some time in the period between 1940 and 1960! In this case, her identification is definitelyincorrect since R. rotundifolius is an extremely local and rare endemic bramble with adistribution lying entirely within one hectad in the English Midlands (Bramble Atlas). It isconsidered probably extinct there (Edees & Newton 1988).
Again we do not know if a voucher specimen exists. There is none for this plant in BEL. Since Kewwas Meikle's workplace for many years, it is the most likely depository herbarium. If a voucher doesexist for this record, we may hope for an eventual re-determination.
Native, rare.
1946; MCM & D; lane near Carry Bridge, Inishmore Island, Upper Lough Erne.
July and August.
It is indisputable that brambles are a neglected area of study in Ireland and have been so since theclose of Praeger's time, ie in the late 1940s. However, A. Newton (1984) and D.E. Allen (1990) andsubsequent papers by both of these gentlemen, have tried to stimulate more interest in Rubi onthe island. Of the 23 records of this bramble in the Fermanagh Flora Database (previously referred to asR. balfourianus A. Bloxam ex Bab.), 13 date from the 1946-53 survey by Meikle and co-workers andall have vouchers determined by Dr W.C.R. Watson. A further MCM & D voucher in K from the Eend of Boa Island, Lower Lough Erne previously labelled R. badius has been re-determined as thismicrospecies by D.E. Allen in 2005. Alan Newton added another record from E of Inishmore Viaduct, UpperLough Erne in 1984, and Allen a further eight in August 2004. As the distribution map indicates therecords are widely scattered across 22 tetrads, the great majority lying around the Lough Erne lowlands.Nine of the tetrads have post-1975 records.
Unquestionably, brambles are a difficult group to identify and we have no Ireland-based expert. Apartfrom the two visitors already mentioned, no one has really spent any time looking at local Rubiin the last 50 years, a position that generally applies throughout the whole island. The majority ofIrish bramble records of the last 20 or more years have been made on a few visiting tours by Britishspecialists invited (and funded) to undertake the task and publish the results (eg Newton (1986) andAllen (2005a, b)).
The Revised Typescript Flora authors noted that this 'species' (ie microspecies in SectionCorylifolii) is the predominant bramble in the Lisnaskea to Newtownbutler area of Fermanagh.Similarly, Newton (1986) commented that on his Irish foray in 1984 he found it was, "locallyfrequent ... along the upper reaches of the River Shannon and Lough Erne ... on banks, hedges andthickets, mostly on calcareous soils".
In their book, Brambles of the British Isles, Edees & Newton (1986) listed twelve Irish VCsfrom which records of R. nemorosus exist and they described the typical habitat and theoverall distribution as being, "hedges, banks and damp low-lying woods on sand or clay. Widespreadin NW & C Europe." In Britain, R. nemorosus is rather thinly scattered south of a linebetween Liverpool and Grimsby, but is most frequently found in coastal regions from Norfolk to Dorset(Bramble Atlas).
Native, rare, but under-recorded.
1891; Focke, W.O.; Lough Erne (unspecified site).
June to September.
Dewberry has been recorded eleven times in Fermanagh in a total of nine tetrads, but very probably itremains under-recorded in lowland limestone districts here as elsewhere in B & I, due to perfectlyunderstandable identification confusion with forms of the apomictic R. fruticosus speciesaggregate (Bramble). The identification difficulty notably, but not exclusively, involves microspeciesof Section Corylifolii (D.J. McCosh, in: Preston et al. 2002). This latter RubusSection is undoubtedly related to the Dewberry and it may well be derived from one or more hybridsbetween R. caesius and other forms of the R. fruticosus species aggregate. Backcrosses ofR. caesius with forms of Section Corylifolii are probably of quite frequent occurrence,although again there is no record of this being the case in Fermanagh (Edees & Newton 1988, p. 272).
In truth, due to a local and general lack of expertise in Ireland in recognising and identifyingRubi, the only bramble of Section Corylifolii recorded in Fermanagh to date isR. nemorosus (see RSF's brief account above). However, two other micro-species in this Section,namely R. hebridensis and R. pruinosus, have been recorded a few times in adjoiningcounties (Bramble Atlas). Thus, these two microspecies might well be also present in Fermanagh ifthe area were actively searched by someone with the necessary knowledge of this critical group.
R. caesius grows in a variety of semi-shaded habitats, generally on basic or lime-rich soils,including rather dry marginal grassland and hedgerows. It also is found in very much damper woodedlakeshore and fen-carr scrub situations (D.J. McCosh, in: Preston et al. 2002). Dewberry isregarded as a native species and, as far as knowledge appears to go, in Ireland it is thinly and widelyscattered, with a greater concentration of records in the southern and western Irish Midlands. However,this may well be an artefact created by the routes taken by visiting bramble experts on their occasionalIrish tours.
In Britain, by comparison, R. caesius has a widespread, somewhat disjointed English distribution,but it becomes very much rarer and coastal, or absent in most of Scotland, Wales and SW England. In bothislands, there is some suggestion of a decline in the presence of this species (D.J. McCosh, in: Prestonet al. 2002), although it is hard to ascertain why this should be the case, since the typicalhabitats are numerous and remain relatively unthreatened.
Native, common. Circumpolar boreo-temperate.
1881-2; Barrington, R.M.; Lough Erne.
Throughout the year.
Growth form and preferred habitats
This perennial has attractive, star-like flowers, plus interesting grey-green, palmate leaves and weakstems which grow either erect or sprawling, rooting like stolons. These stems and a far-creeping woodyrhizome sometimes help form floating multi-species mats of vegetation in shallow, slow-moving waters.
P. palustris is found in both very acid and very obviously calcareous mainly lowland habitats, onpermanently, intermittently or seasonally flooded ground.
The species has wide ecological tolerances and it ranges from very acid, nutrient-starved, peaty sites(eg in drains and wet hollows on bogs, or in sheltered, shallow pools on lowland pastures or in uplandgrassy heaths) to productive, base-rich conditions, in muddy or silty soils on marshy grasslands, or inshallow, moderately acidic, but lime-rich waters around lakeshores and along the banks of slow-movingrivers. It can also tolerate semi-shade under trees in wet woodland at all levels and in willow andalder fen-carr around lakeshores.
P. palustris can be locally dominant and is very commonly associated with other rhizomatousspecies, including Carex rostrata (Bottle Sedge), Equisetum fluviatile (Water Horsetail)and Menyanthes trifoliata (Bogbean), in wet or flooded, poor- to medium-nutrient, acidicpeaty ground. In these, and in many other situations, Marsh Cinquefoil can clearly survive competitionfrom taller, shading plants over a wide range of organic and mineral soils.
Flowering reproduction
Since its rhizome and stolons form clonal stands, local survival of P. palustris does not requiremuch seed production in stable habitats.
Seed is vital, however, both in terms of long-term maintenance of variation and vitality and dispersalability to fresh sites. The dark crimson flowers, with petals half the size and much less conspicuousthan the sepals, appear from May to July and are visually conspicuous. They also give off a terriblesmell and their partially concealed nectar attracts insects such as craneflies, mosquitoes and stinglesswasps (Proctor & Yeo 1973; Proctor et al. 1996). Unvisited flowers are self-compatible, butselfed individuals set a reduced amount of seed (Olesen & Warncke 1992). As in P. anserina(Silverweed), seeds can float for a long period, but nothing is known about their survival, transportand establishment. A clear case of further work required.
Fermanagh occurrence
Marsh Cinquefoil is very frequent, widespread and locally abundant in Fermanagh, being represented in 265tetrads, 50.2% of those in the VC. It is most frequent in the area south of Lough Erne. It also occursin semi-shade in wet woodland and in fen-carr around Lough Erne and, more rarely, in damp, uplandwoodlands on the Western Plateau.
British and Irish occurrence
Marsh Cinquefoil is widespread in most of B & I, the distribution showing a N & W preponderance.Considerable local extinction has occurred over the last 120 years, especially in the drier SE of bothislands and the English Midlands (Preston & Croft 1997). Loss of habitat through drainage andagricultural improvement is largely responsible for the losses of the past 60 years (D.J. McCosh, in:Preston et al. 2002).
European and world occurrence
P. palustris is a member of the circumpolar boreo-temperate group and is common throughout arcticand boreal regions around the northern hemisphere and extends south to C Spain, N Italy and S Bulgaria,but is absent from the Mediterranean shores and islands (Kurtto et al. 2004, Map 3390).
Uses
The attractive, dark crimson, star-like flower and the interesting grey-green, palmate leaves makes thisa garden-worthy plant, or almost so, were it not for its far-creeping, invasive woody rhizome. Havingsaid this, it is obviously suitable for larger gardens with aquatic features and a white-flowered formalso exists in the horticultural trade (Griffiths 1994).
Threats
In Fermanagh, increasing nutrient levels from agricultural runoff and atmospheric nitrogen pollutionmight oust this species from previously mesotrophic habitats, especially around our larger lowlandlakes.
Native, common and very widespread. Circumpolar boreo-temperate, but widely naturalised.
June 1862; Smith, T.O.; Ardunshin.
April to November.
Growth form and preferred habitats
This very variable, low-growing, carpet-forming, stoloniferous, rosette perennial is a common pioneerweedy colonist of open bare ground and of gaps in pastures. Silverweed is shade-intolerant but is quitefrequent along moderately disturbed roadside verges, waste ground, heavily trampled soil around gatewaysand in compacted, otherwise more or less bare ground beside tracks and paths, on damp to perfectly dry,sometimes very stony soils. However, it definitely prefers and is most common and abundant in almostpermanently damp, moderate to richly fertile, near neutral, base-rich or calcareous soils, whereseasonal flooding inundates and nutrient-enriches grassland on lakeshores and riverbanks. The vigorousvegetative reproduction of the species allows it to invade vegetation gaps in overgrazed or moderatelydisturbed pastures and, once established, it can become very enduring.
While it is certainly able to perform well as a pioneer coloniser of bare, disturbed soil and vegetationgaps, in Sweden on a grazed Baltic seashore pasture, P. anserina was found to be competitivelyinferior to three common grass species, Agrostis stolonifera (Creeping Bent), Festucarubra (Red Fescue) and Poa pratensis subsp. irrigata(Lindm.) H.Lindb. (= P. humilis) (Spreading Meadow-grass). Silverweed could only out performthese grasses on patches of cattle dung (Eriksson 1986b).
P. anserina is also common where persistent rain regularly produces temporary pools in hollows inwet pastures (Sinker et al. 1985; Hill et al. 1999). As it also features on sea coasts onsalt marshes and sand dune grasslands, this is very definitely a species (or species aggregate) of wideecological versatility and, consequently, it is widespread all around the northern hemisphere.
Fermanagh occurrence
Apart from very acid peaty uplands and regularly managed agricultural ground, P. anserina iscommon and widespread in Fermanagh and has been recorded in 376 tetrads, 71.2% of those in the VC, themain exceptions being the upland acid ground of the Cuilcagh and Lough Navar areas. It is less frequentor absent in the better agricultural ground east of Lough Erne.
In Fermanagh, Silverweed is most constantly present and abundant around the winter high water level onmany local lakeshores, and by far the greatest local concentration of P. anserina is in the UpperLough Erne basin. In part, this is certainly an artefact produced by the enormous amount of recordingtime taken by the government conservation team that surveyed this area in the mid-1980s. However, theeminently suitable terrain and the traditional water-meadow management regime employed by the localfarmers, provides an ideal habitat for this stress-tolerant, moderately competitive, ruderal,vegetatively far-creeping, clonal species (Grime et al. 1988).
Flowering reproduction and genetic variation
While P. anserina flowers from May to August, it tends to do so relatively sparingly; eachsolitary bright yellow blossom, 2-2.5 cm in diameter, is formed either axillary on the mother rosette,or terminally on stolon internodes (Eriksson 1986b). Flower production is inversely related toplant vigour (Ockendon & Walters 1970). The dish-like, unspecialised flowers stay open for 2-3 days,are self-incompatible and are pollinated by various short-tongued insects. While this is the case, theystill frequently fail to set seed and fruiting plants are often difficult to find (Ockendon &Walters 1970; Fitter 1987; Miyanishi et al. 1991).
The production of seeds in a Swedish study was reduced by limited pollen availability in the Balticcoastal population studied, mean seed set being less than ten per flower. Pollen availability variesfrom year-to-year and with prevailing weather conditions and it is particularly sensitive to lowtemperatures and precipitation, although the species does occur north of the Arctic Circle (Miyanishiet al. 1991). Pollen limitation will be most significant in populations dominated by few clonesor incompatibility types (Eriksson 1987).
Another reason for fruiting failure, although probably only an extremely minor element, is that P.anserina exists in both tetraploid (2n=28) and hexaploid forms (2n=42), which aremorphologically inseparable (Rousi 1965; Ockendon & Walters 1970). No diploid representatives havebeen found so far and Rousi (1965) concluded that they were unlikely in Europe or N America, but theymight still survive in unexplored Asia. Hexaploids are probably autoalloploids arising in a single stepfrom the fusion of reduced and non-reduced tetraploid gametes (Cobon & Matfield 1976). A cytologicalsurvey of P. anserina in Britain found hexaploids scattered throughout otherwise tetraploidpopulations (Ockendon & Walters 1970).
The tetraploid is the widespread, fertile and most probably ancient form of the plant in both B & Iand around the world, while the hexaploid shows no discernible geographical pattern in these islands. Itonly rarely sets seed and thus is something of an evolutionary blind alley (Rousi 1965; Ockendon &Walters 1970). The fact that Erlandsson (1942) found two pentaploid Silverweed plants, however, provesthat the hexaploids are not totally sterile. Having said this, no natural hybrids of P. anserinawith any other Potentilla species have been reported, although it can cross with the Fragaria× ananassa Duch. (2n=56) the Cultivated Strawberry (Miyanishi etal. 1991).
Morphological variation, hybridization and gene flow
There is considerable variation in form within Silverweed populations throughout the species wide totalrange, and especially so further north. Taxonomists and plant geographers have named up to eightsubspecies and/or varieties, treating P. anserina as a species aggregate (Rousi 1965; Hultén1968). Hybrids have been recorded in Finland between some of the subspecies, particularly between subsp.anserina and subsp. egedii Wormsk. (a more arctic form found today in N & C Europethat is sometimes given full species status (Rousi 1965)). A number of other botanists are not convincedabout the reality of some of these northern variants, eg Porsild & Cody (1980), who could find nodistinctions between specimens identified as P. anserina and P. egedii subsp. yukonensis [in Arctic Canada] on the basis of their acheneor bractlet characteristics.
On the basis of mainly glasshouse hybridization experiments between populations and observations ofnatural hybrids in populations on the coast of Finland, Rousi (1965) has argued that the two forms mayhave coexisted during the Late-glacial period, hybridized then, and transferred some subsp.egedii characters to subsp. anserina which still persist until today. Subsp. egediiis usually distinguished by its smaller size, its glabrous stolons and the lack of silvery pubescence onits leaflets (Miyanishi et al. 1991).
The fact that these two taxa (subsp. anserina and subsp. egedii) have managed tokeep distinct from one another in the N Hemisphere for such a long period raises questions as to howthey achieved isolation and avoided intermingling? Presumably this is due to differing ecologicaltolerances, but the fact that subsp. egedii is self-compatible must also help keep the twoseparate, their flowering periods only have a brief overlap and the geographical ranges have also partedcompany nowadays to a large extent (Rousi 1965).
The latest critical Flora of B & I distinguishes four varieties on the basis of the hairiness of theupper leaf surface and the size of various plant parts, including leaves, flowers and petiole length:these forms are given as Var. nuda Gray; Var. sericea Hayne; Var. anserina and Var.grandis Lehm. (Sell & Murrrell 2014). These taxa do not appear to coincide with the formsrecognised by Rousi (1965).
Fossil history in B & I
The achenes of P. anserina are sufficiently distinguishable that the species has been recorded inBritain back as far as two stages in the Hoxnian interglacial 400,000 BP which lasted about 53,000years. It has appeared in all glacial and interglacial periods since then, although there was a hiatusbetween Flandrian zone IV and zones VIIb & VIII, the latter falling within Roman and Bronze Ageperiods. This suggests vegetation or woodland clearances going on that created open, disturbedconditions more favourable for increase of the species. Periglacial outwash from retreating glaciers andbare river terrace deposits would favour the pioneering behaviour of P. anserina and, in morerecent times, the species has spread with human culture thanks to the environmental habitat disturbancewe create (Godwin 1975).
Vegetative reproduction
P. anserina spreads efficiently by its annual stolons, 30-100 cm long, with internodes 10-15 cmlong at which daughter ramets are formed. The often reddish stolons wither each autumn and thereafterthe rooted ramets live independently (Eriksson 1987). Dense mats of ramets can develop with up to 2,000ramets per m² (meaning in this case, each ramet may occupy as little as 5 sq cm, although there could besome degree of overlap)(Eriksson 1986a). The vigorous vegetative reproduction shown by P.anserina is the major means of ramet recruitment to the population, far outstripping recruitmentfrom seed and ramet mobility enables plants to find space and avoid competition (Eriksson 1986b). Eachramet consists of a short vertical rhizome bearing swollen adventitious roots containing starch. Theramets are perennial, all above ground tissues withering in the autumn, the plants overwintering simplyas the short rhizome with winter buds developing on their uppermost part (Miyanishi et al. 1991).In the Swedish Baltic shore study, the half-life of an established ramet was 3.1 years (Eriksson 1986a).
Vegetative reproduction is regularly associated with polyploidy. Other examples of this property includeAgrostis stolonifera (Creeping Bent), Potentilla reptans (Creeping Cinquefoil),Ranunculus repens (Creeping Buttercup) and Trifolium repens (White Clover)(Salisbury 1964, p. 335). Plants that grow very vigorously and reproduce asexually often reducetheir investment in flower production and sexual reproduction. This appears to happen in many Silverweedpopulations, which may produce several stolons or 'runners'up to a metre in length in a single season, but which flower only sparingly and sporadically.These vegetatively produced clonal patches are potentially long-lived, perhaps sometimes ancient(Eriksson 1987).
Fruit production and dispersal
P. anserina plants that do manage to fruit successfully produce a head of between 6-50 (maximum60) lightweight, dry achenes (single-seeded fruits). These are dispersed either by water flotation (theachene wall has a layer of air-containing cells enabling them to remain buoyant for up to 15 months),rain wash, adhesion in mud to other organisms, or by being eaten along with foliage and passing throughbirds or other animals (Ridley 1930; Salisbury 1964, p. 276; Miyanishi et al. 1991). Achenesremain viable for up to a year buried in soil (four reports), but there is only one report of long-termsurvival (ie longer than five years) so it is unlikely that there is a permanent seed bank (Thompsonet al. 1997).
British and Irish occurrence
P. anserina is common and widespread throughout the whole of B & I except for inland parts ofN Scotland. This pattern reflects the species avoidance of extreme acid soils and colder, high altitudesites. Around the coasts of B & I, P. anserina is also an almost universal component of thesalt-tolerant maritime vegetation of the upper part of shingle or coarse sand beaches, where it forms adistinct pioneer zone below the turf of the true dry land vegetation (Preston et al. 2002).
Specimens from coastal Scottish populations were found to approach the more northern or arctic form P.anserina subsp. egedii, which is characterised by entireor shallowly toothed epicalyx segments and a low number of teeth on the terminal leaflet of leaves,seeming to have a mixture of anserina and egedii characters, similar to the situationdescribed by Rousi in Baltic Sweden (Ockendon & Walters 1970).
Species origin
It is believed by some botanists that P. anserina originated on sea shores and other saline soilsand that it subsequently developed a wider ecological tolerance of trampling, disturbance and nutrientstatus to become a common and widespread stress-tolerant weed of disturbed ground in cool temperateregions. Other evidence from the extent of variation points to the Himalaya, or somewhere in SE Asia asthe centre of origin of the species or species aggregate (Rousi 1965, p. 106). At some unknown stage inhistory, tetraploid P. anserina spread around the globe in both hemispheres. As with other weedyruderal species, man undoubtedly assisted its transportation, but despite the excellent review by Rousi(1965, pp. 106-9), it is difficult to be convinced that anyone knows the true native range of P.anserina (Rousi 1965; Hultén & Fries 1986, Map 1097; Miyanishi et al. 1991).
European & world occurrence
P. anserina is common and widespread throughout N, W & C Europe, extending south into thenorthern half of the Iberian peninsula and extending eastwards north of the Alps to N Greece and theCrimea (Kurtto et al. 2004, Map 3400). P. anserina s.s. is worldwide in distribution,although it is not yet possible to separate geographically the varieties that have been distinguished inthis taxon. It is certainly most widespread and frequent in temperate parts of the N Hemisphere where itis a member of the circumpolar boreo-temperate element, but it has also been introduced and has becomean invasive, ruderal weed in the S Hemisphere in Chile, SE Asia, New Guinea, S Australia and New Zealand(Hultén & Fries 1986, Map 1097).
Threats
None.
Native, common and abundant. Eurosiberian boreo-temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
A rosette-forming, shallow-rooted perennial, P. erecta has a stout, woody, almost tuberousrootstock from which slender annual aerial stems with sessile or subsessile, mostly trifoliate, compoundleaves with large, divided, leaf-like stipules at their base and erect flowering stems arise. The basalrosette leaves die back each summer before flowering or else in the autumn, although, in a few cases,these leaves survive and remain wintergreen. While the species is phenotypically very flexible, theplant usually has two types of leaves; basal leaves are long-stalked, palmately compound with five ovalleaflets that are toothed at the apex, while the very much more numerous stem leaves are unstalked, oralmost so, and have three unevenly toothed leaflets, borne opposite a pair of large, lobed, toothedstipules that look like two extra leaflets. Leaf size decreases noticeably as the plant stem elongatesduring the season, with the smallest leaves being formed towards the terminal flowering region of thestem (Harold 1994).
With its small, usually 4-petalled yellow flowers, the petals slightly notched, P. erecta is oneof the most widespread and familiar flowering plants in the British Isles, yet despite its botanicalname, the long flowering stems of P. erecta are really more trailing and clambering than trulyerect in habit. However, unlike the stems of P. reptans (Creeping Cinquefoil), or P.anserina (Silverweed), they do not root as they spread outwards from the established leafrosette, so there is no vegetative reproduction in this species.
Tormentil grows commonly on just about every heath, bog (raised, blanket and valley), pasture, roadsidebank and quarry at all altitudes in the VC. Undoubtedly, it is most abundant in upland areas oninfertile acid soils and in moderately disturbed habitats. It is intolerant of shade and is only rarelyfound in more open areas of woods or on their margins. P. erecta tolerates a wide range ofsubstrate pH (from 3.7 to 7.5), but in terms of frequency, it tends towards acidity and is uncommon insoils above pH 6.0. In limestone grasslands, Tormentil is confined by its shallow root system to wetterareas of ground and it is generally much less vigorous under these conditions (Grime et al.1988).
Vegetative growth strategy
The complete winter dieback and annual spring regrowth of the aerial parts of P. erecta is anunusual survival strategy for a herbaceous perennial of more or less infertile, mainly acid conditionsthat are often subject to grazing pressure. This is only made viable by the fact that all parts of theplant taste astringent to browsers and, therefore, can survive and maintain active growth when lessprotected competitors suffer grazing pressure (Grieve 1931; Milton 1933). In terms of competitiveability and established strategy, P. erecta was classified as being intermediate between astress-tolerator and C-S-R (meaning it displays a balance of ability between competitor,stress-tolerator and ruderal) by Grime et al. (1988). The substantial, thick rootstock isparticularly unpalatable since it contains 18-30% tannin and, in the past, it was used for tanning bothleather and fishing nets and as a source of dye and medications (see below) (Grigson 1987).
Leaf ecology
Leaf life spans of herbaceous plants are less investigated than those of shrubs and trees, but Diemeret al. (1992) showed that mean leaf longevity in a range of 29 herbaceous plants, split almostevenly between lowland meadow flora and high alpine species, had leaf longevities that ranged (with noclear altitudinal trend), from 41-95 days. Mean leaf longevity in P. erecta at the two altitudesdid not differ significantly, being 71 ± 5 days in lowland areas and 68 ± 4 days at high altitude (ie2,600 m) in the Austrian Alps near Innsbruck. Low-altitude herbs and grasses studied produced severalleaf cohorts during the growing period, while apart from Geum and Potentilla the plants ofmore alpine vegetation studied produced just one set of leaves. This Austrian study also found that leafduration of herbaceous plants is associated with properties other than the efficiency of lightutilization, eg mechanical strength, the avoidance of herbivory and resistance to pathogens (Diemeret al. 1992).
Flowering reproduction
Flowering can be delayed in P. erecta individuals subject to heavy competition in meadows, perhapsfor up to three or more years (Salisbury 1942). The bright yellow, 7-11 mm diameter, four-petalledflowers of P. erecta (an exceptional number in the Rose family) are almost omnipresent from Mayor June to September, or even into October in sheltered ground in Fermanagh, when the stems die.While the great majority of flowers have four lightly notched petals, some with three, five orsix can occur (Sell & Murrell 2014).
The many-flowered inflorescence is a loose terminal cyme and the flowers are self-incompatible makingoutbreeding obligatory. Nectar is produced and is partly concealed by the filaments of the 14-20stamens. Pollinators include flies, parasitic wasps, short-tongued wasps, beetles, butterflies and mothsfeeding on pollen and nectar (Proctor & Yeo 1973). Seed is set from July onwards. The achenes(single-seeded dry fruits) are slow to ripen and are released late in summer, often in a green condition(Grime et al. 1988). Seed does not germinate until the following spring and a long-termpersistent soil seed bank develops (>5 years) (Thompson et al. 1997).
Seed dispersal
The seed is relatively heavy (0.58 mg), usually with 4-8 seed in each fruit head (Grime et al.1988). The typical plant produces between 100-1,000 seeds and they tend to fall very close to the motherplant. This makes it difficult to imagine exactly how the species has managed to disperse so widely andacross a range of different habitats in B & I and elsewhere. Without the plant being browsed, theseed swallowed and transported internally by animals such as cattle, sheep, goats, deer and rabbits, itis very difficult to explain the observed current distribution of this and many other low-growingherbaceous species that produce small, dry fruits and seeds and have no obvious adaptations to enableefficient dispersal (Ridley 1930, p. 361). Yet, as mentioned above, all parts of P. erecta plantsare reputed to contain astringent substances such as tannins that make them unpalatable and, therefore,apparently the species is avoided by browsing animals (Grieve 1931; Grime et al. 1988). Obviouslythe avoidance does not include pollen and nectar.
The possibility remains that human interest in the plant for its herbal medicinal properties (see below)leads to its collection (especially the rootstock), transport and, even possibly its cultivation. All ofthis could, over many years enable wider occurrence of the species to arise, at least near habitation.Also, hay has been made for winter fodder from meadow vegetation containing P. erecta forthousands of years, and although stock might still avoid eating the dried plant tissue if it remainedunpalatable, its seed would be carried within bales. There has always been a commercial trade in hay,and bales exported to other areas would inadvertently help distribute the species. Nevertheless, evengiven a possible dispersal role for man, there does not appear to be any convincing evidence of adispersal mechanism capable of creating and sustaining the common and widespread plant distribution ofP. erecta that we see today.
Variation
P. erecta is a tetraploid species (2n=28) like P. reptans (Creeping Cinquefoil), from whichit is morphologically very distinct. Both these species are highly fertile and self-incompatible andthere are incompatibility barriers between them which make hybrids very difficult, but not impossible,to obtain (Matfield & Ellis 1972). A highly sterile wild hybrid of the two species has been foundjust once in Wales in 1983 and is named P. × italica Lehm. (Staceet al. 2015).
A robust upland race of P. erecta exists with larger flowers occurring in dwarf-shrub heaths andhas been described as subsp. strictissima (Zimmeter) A.J. Richards (Richards 1973). A review of B& I herbarium material showed that the latter has occurred in 16 Irish VCs, but it has yet to befound in Fermanagh (Rich & Scannell 1990). RHN and the current author (RSF) expect that it will befound in suitable ground, eg on the upper slopes of Cuilcagh mountain.
Hybrids
P. erecta can form hybrids with P. anglica (Trailing Tormentil) and P. reptans, bothof which occur thinly and widely scattered in Fermanagh. The hybrid with P. anglica was recordedas long ago as 1856, but has not been seen since about 1950 and it is clearly being overlooked.
There are no obvious isolating barriers between the many varied habitats that P. erecta occupiesand, theoretically, there are unlimited opportunities for genetic mixing and gene flow betweenpopulations, although even a small amount of maternal effect could mask the true situation. Experimentalstudies with Scottish populations showed that the difference between a Molina-dominated and aFestuca-dominated habitat was roughly as effective as a north-to-south difference of 200 miles(325 km) in terms of selective forces on Tormentil (Watson 1969, 1975).
Fermanagh occurrence
P. erecta is widespread, abundant and has been very commonly recorded in 410 of Fermanagh tetrads,77.7% of those in the VC. It ranks as the 12th most frequently recorded species in the Fermanagh FloraDatabase, after Ranunculus acris (Meadow Buttercup) and before Succisa pratensis(Devil's-bit Scabious). Tormentil is particularly frequent in lowland grassland around Upper Lough Erneand on acid heaths, bogs and grassy moorland in the south and more upland west of Fermanagh.
Fossil record
P. erecta has been found in England at the Hoxnian and Ipswichian interglacials and in the morerecent Flandrian, from zone VI onward, it becomes far more frequent, including presence in Roman andMediaeval sites. The fossil records show a preference for more open, northern habitats, including onblanket bog (Godwin 1975).
British and Irish occurrence
Common and widespread throughout B & I, although it is ± absent from the fens around the Wash andsparse in SE England, where it has declined from the 1950s onwards (Preston et al. 2002).
European and world occurrence
P. erecta is widespread and common throughout N, W & C Europe, thinning southwards tothe Mediterranean and eastwards into W Asia. It is present to the southern tip of both Spain &Italy, although it is absent from all the Mediterranean islands except Corsica (Kurtto et al.2004, Map 3521). It is also listed as occurring in N Africa, including Algeria, Morocco and the Azores(Sell & Murrell 2014). Apart from B & I, P. erecta is very much a mainland Europeanspecies and its presence in Iceland, the Faeroes and the Azores is doubtfully indigenous. It has a verysmall presence in eastern N America where it is almost certainly introduced (Hultén & Fries 1986,Map 1120).
Uses
Apart from tanning, P. erecta is also a source of a dye, called 'Tormentil Red', previously usedfor colouring leather, plus the production of resins and complex acids. In modern herbal medicine,Tormentil is regarded as one of the safest and most powerful astringents for use in treating diarrhoeaand other discharges, ulcers and long-standing sores. Grieve (1931) lists numerous other medicinal usesfor P. erecta, commenting that many of the 150 species of Potentilla have been similarlyused in medicine. Apart from a long list of medicinal uses in humans, P. erecta has also beenwidely used for a range of ailments in veterinary practices (Allen & Hatfield 2004).
Name
The name 'Tormentil' is considered derived from the Latin 'tormentum', referring to the torture of colic('Tormina' in Latin) (Grigson 1987).
Threats
None.
July 1977; Nelson, Dr E.C.; Correl Glen (woodland).
Occasional.
According to P. Hackney, this subspecies is very common in the north-east of Ireland, while the otherBritish Isles form, subsp. strictissima is apparently very rare. The latter subspecies occurs inupland areas (Hackney et al. 1992).
Threats
None.
Rare, certainly under-recorded and probably occasional with its parents.
1849; Mathew, Dr; Newtownbutler.
There are only three records of this partially fertile hybrid in the Fermanagh Flora
Database and they are dated 101 years apart. The hybrid has not been found in the VC by anyone since thesurvey by Meikle and co-workers recorded it twice in 1950. The reason P. × suberecta is rarely recognised is that it is intermediate incharacter between its parents to a very remarkable extent – sometimes even with respect to the number ofcarpels formed – and, when it is found, it is almost always amongst mixed populations of its parents. Inaddition, there is great seasonal and environmentally induced variation in both P. anglica(Trailing Tormentil) and P. × suberecta, so that these two taxa areoften extremely difficult to distinguish. The fact that the F1 hybrid, P. × suberecta, is only partially sterile means that some degreeof backcrossing is feasible. The extent of this is unknown due to the amount of variation within theparent species and also in the hybrid itself (B. Matfield & S.M. Walters, in: Stace 1975).
The habitats where the parent species overlap include low-growing vegetation on heathy pastures(including those near the coast), inland heaths, dry roadside-, riverside- and railway-banks,tracksides, hummocks in damp fields, woodland rides, paths and margins, field edges, quarries andshingly waste ground near the sea (Stace et al. 2015). Most of these sites feature well-drained,acidic soils. The occurrence of P. × suberecta is probably mainlylowland, but the upper altitudinal limit and the true frequency level and the distribution remainunknown (D.J. McCosh, in: Preston et al. 2002).
The possible presence in these habitats of another pair of very similar Potentilla hybrids, thosebetween P. anglica and P. reptans (Creeping Cinquefoil) (P. × mixta Nolte exRchb.), and the extremely rare hybrid between P. erecta (Tormentil) and P. reptans (P.× italica Lehm.), compound an already difficult identificationproblem (B. Harold (née Matfield), in: Rich & Jermy 1998). These latter two hybrids are so alikethat effectively it is impossible to distinguish them in the field. Secure determination requiresmicroscopic examination of their pollen (D.J. McCosh, in: Preston et al. 2002). P. × italica is so elusive, it has only been recognised once inthe wild in Britain (Stace et al. 2015).
The only certain feature enabling field identification in hybrids involving P. anglica is in thedegree of fertility displayed. P. anglica is seed- and pollen-fertile, but the hybrids it makesare sterile, or almost so. However, it is necessary to recognise that the level of seed set cannot beassessed early in the flowering season. A very useful indication of maturity is the presence of at leastthree withered flowers on the same stem distal to an open flower or a flower bud. The oldest of thesewithered flower heads should contain several conspicuously swollen carpels if the plant is fertile.P. × suberecta may sometimes manage to set a few seed, butundeveloped carpels will always be in the majority. In comparison, P. ×mixta rarely sets any seed, or only around one per flower at most (B. Harold (née Matfield), in:Rich & Jermy 1998).
In addition to this, regarding fertility, it is important to realise that P. reptans is fullyfertile but self-incompatible. Since the species can form large clonal patches by vegetativereproduction, some P. reptans colonies may consist of just one genome and, therefore, the flowerswill fail to set seed unless a suitable cross-pollinating partner is available nearby.
Apart from sterility or near-sterility, it is essential for the supposed hybrid to also display a mixtureof 3-, 4- and 5-nate, petiolate stem leaves and the mixture of 4- or 5-merous flowers typical of the twomost probable hybrids (P. × suberecta and P. × mixta) that distinguish them from P. erecta and P.reptans (Harold 1994; B. Harold (née Matfield), in: Rich & Jermy 1998). As its namesuggests, P. × suberecta is much more like P. erecta,with a predominance of ternate stem leaves on short petioles, generally only one leaf per node,and the leaf size decreases from plant base to the tip of the stem. Also, like P. erecta, thishybrid does not spread vegetatively and P. × suberecta usually setsjust a few seeds per flower head. On account of these properties, P. ×suberecta is only found in close proximity to both its parents. On the other hand, if P.erecta is not growing nearby, the plant in question could be either P. anglica or P.× suberecta, and a closer look for P. erecta should be made(Harold 1994).
Apart from the first record of P. × suberecta given above, the detailsof the other two Fermanagh MCM & D sites are: roadside at Drumbad House and near Inishmore Viaduct,Upper Lough Erne, both dated 1950.
P. × suberecta is said to be fairly frequent, or even common in someparts of B & I in comparison with other natural hybrids in these islands (New Atlas). RobertNorthridge and the current author believe it is certainly under-recorded in Fermanagh, as indeed isprobably often the case elsewhere. The hectad maps in the New Atlas and in The Hybrid Flora ofthe British Isles (Stace et al. 2015) display a remarkably patchy distribution for P.× suberecta, which suggests that relatively few recorders candistinguish this hybrid and also that they work their local areas thoroughly for it.
For a more local NI comparison with respect to this hybrid, the FNEI 3 lists just six records forthe three most north-eastern Irish VCs (Down (H38), Antrim (H39) & Londonderry (H40)), all but oneof them dating from the 1849-1935 period. The recording situation has greatly improved in the last 20years since a careful examination of the BSBI Database, made by the current author on 30 November 2020,now provides a list of 40 records (plus a few duplicates) for the same three Irish VCs. No less than 27of these records were made by the late John Harron between 1991 and 2003 and a further seven by thecurrent BSBI VC Recorder (H38), Graham Day, between 2001 and 2018.
The status of this and other Potentilla hybrids clearly merits further detailed investigationwhenever time permits.
Native, frequent but under-recorded. European temperate, but with a few naturalised outliers in eastern NAmerica.
1900; Praeger, R.Ll.; Belcoo, Lough Macnean.
April to December.
Growth form and preferred habitats
As its English common name indicates, P. anglica is a trailing, procumbent, mat-forming perennial,possessing a thick, branched rootstock bearing a varying mixture of 3-, 4- and 5-nate leaves and withleaf stipules entire or 3-lobed, not deeply divided or lobed as in P. erecta (Tormentil). Small,simple, undivided leaves also occur. The spreading aerial stems root at their tips later in the growingseason and the four or five petalled flowers regularly contain up to 20 carpels and set good seed, incomparison with the 4-12 carpels that are usual in P. erecta (Webb et al. 1996).
Although the two closely related taxa overlap in their ecology, P. anglica tends to be a morelowland species than P. erecta and it also frequents less acid, better drained, light or sandysoils than the latter: it especially avoids damp, cold, iron podsols with their impeded drainage. Thehabitats where these two species overlap include low-growing vegetation on heathy pastures (includingthose near the coast), inland heaths, dry roadside-, riverside- and railway-banks, tracksides, hummocksin damp fields, woodland rides, paths and margins, field edges, quarries and shingly waste ground (Staceet al. 2015).
Hybrid origin of the species
P. anglica is an example of an allopolyploid species of sudden origin that arose by hybridizationbetween two tetraploids (2n=28), P. erecta and P. reptans (Creeping Cinquefoil). Boththese species are highly fertile and self-incompatible, and there are incompatibility barriers betweenthe two which make hybrids between them very difficult but not impossible to obtain (Matfield &Ellis 1972). At some stage, a hybrid involving both species was formed, possibly involving an unreducedgamete from P. reptans, creating a hexaploid hybrid (2n=42). This hexaploid must then haveback-crossed with P. erecta in a second stage of the process, again without reduction of itsgamete, combining to form a new, fully fertile octoploid hybrid (2n=56) that is P. anglicaLaichard. (Matfield & Ellis 1972; Harold 1994).
As one would expect of a species of hybrid origin, its morphology is intermediate between its twoparents, but P. anglica also has some features that are directly attributable to its higherchromosome number. Each stem node bears one to several leaves with petioles of variable length. Theleaflet number, leaf size and petiole length all decrease through the growing season so that a plant maylook more like P. reptans in early summer, but becomes more like P. erecta later on. P.anglia roots at its stem nodes, but does so less readily than P. reptans. Directly due tothe higher chromosome count, the leaflets tend to have a lower length:breadth ratio than those of theparent species and the pollen grains are larger. Most significantly, the additional sets of chromosomesin P. anglica lead to a breakdown in the self-incompatibility mechanism that is so characteristicof the two parent species. This means a single, isolated specimen of P. anglica is able toself-pollinate and set good seed, making seed fertility a useful diagnostic character for this species(Harold 1994).
However, matters are complicated by the relatively frequent occurrence of another hybrid, P. × suberecta, a cross between P. erecta and P. anglica.P. × suberecta is morphologically very similar to P. anglicaand always occurs in the presence of both its parents. Although P. ×suberecta is not completely sterile and may set a few seed, it does not reproduce vegetativelyand, therefore, it tends to die out. Nevertheless, its existence confuses the identification of P.anglica to a very considerable extent (see my account of this hybrid for more details).
Fermanagh occurrence
Trailing Tormentil is widespread and locally frequent in Fermanagh in short grass on well drained banks,slopes and roadsides. Although recorded from 117 Fermanagh tetrads, representing 22.2% of the totalnumber, P. anglica remains almost certainly under-recorded in the current Flora survey. Itis probably quite often confused with the more common and widespread P. erecta (Tormentil) withwhich it ecologically overlaps and forms the partially fertile hybrid, P. × suberecta. RHN and the current author (RSF) are sorry to report wehave never found this hybrid in Fermanagh.
There are nine tetrads in Fermanagh with pre-1975 records only for P. anglica, but this is too fewto suggest there has been any real decline in presence.
British and Irish occurrence
The hectad map in the New Atlas (accompanied with a warning regarding confusion with the hybrid)displays P. anglica as widespread throughout Ireland, but more local and patchy in its occurrenceand with many old records in C Ireland.
The same map indicates a widespread presence in W England to the south of Lancaster and also in Wales.The distribution becomes more scattered and local further N & E and also in the E & SE ofEngland, where evidence suggests that it is declining. In Scotland, P. anglica is frequent onlyin Dumfries and Galloway and in the Clyde and Forth conurbations, although isolated outliers exists inthe western isles and in NE Aberdeen (D.J. McCosh in: Preston et al. 2002).
European and world occurrence
Beyond these isles, the native range of P. anglica is restricted to a small region of cooltemperate W & C Europe, with a few introductions beyond. The SW limit is in the Pyrenees and itstretches north to reach the S Baltic region of Sweden and Finland. It is absent from the Iberian andItalian peninsulas but stretches east to the western border of the former USSR (Sell & Murell 2014).P. anglica is probably endemic to Europe. The plant mapped on Corsica (Kurtto et al. 2004,Map 3522) is tetraploid and should be recognised as a separate species. A claim by Fernald (1933) thatthe population in Newfoundland is native appears unsupportable, particularly in view of other widelyscattered outliers of this species in N America that are acknowledged naturalised introductions (Hultén& Fries 1986, Map 1121). The same applies to outliers in Madeira, the Azores and Iran, where thespecies is almost certainly introduced (Kurtto et al. 2004). See also P. sterilis below.
Threats
None.
Native, frequent and widely scattered. Eurosiberian southern-temperate, but widely naturalised.
1881-2; Barrington, R.M.; shores of Lough Erne.
Throughout the year.
Growth form and preferred habitats
In common with related Potentilla species, P. reptans possesses a deep, branching rootstockas its perennating structure and it bears a persistent terminal rosette of 5(-7)-nate palmate leaves onlong petioles. Although very variable in habit with growing conditions, the long, trailing, floweringstems and four or five (-15) red, prostrate, surface-running stolons rooting at the nodes, with palmatestem leaves mostly of five leaflets and stipules entire or rarely toothed, together make this an easyperennial to recognise (Harold 1994; Sell & Murrell 2014).
A vegetatively vigorous pioneer colonist of disturbed, well-grazed, trampled or bare ground, CreepingCinquefoil often forms clonal patches or carpets at the edge of roads or concrete tracks. However, italso grows, competes well and persists amongst other vigorous colonising species in sunny, open,somewhat disturbed or compacted soil conditions in short turf lowland grasslands, or on dry bare soil orstony ground. It is also one of the most troublesome garden weeds due to its rapid spread and the depthand vigour of its roots (Salisbury 1964). It is sometimes an abundant competitive ruderal on gravellylakeshores, on the margins and in openings in woods, hedge-banks, tracksides, or in old quarries, onroadside verges and waste ground.
P. reptans prefers a well-lit, damp but free draining, moderately fertile, mildly acid to basiccalcareous soil, although to some extent it can tolerate semi-shade and dry or more moderately acidicconditions (Sinker et al. 1985; Hill et al. 1999).
Fermanagh occurrence
This perennial is frequent and widely scattered in lowland Fermanagh and elsewhere in the county off thepeat. It has been recorded in 103 tetrads, 19.5% of the total in the VC. As the tetrad distribution mapindicates, seven of the tetrads contain only pre-1975 records, but this is not believed to represent anyreal reduction in the presence of this otherwise frequent species.
Flowering and vegetative reproduction
In sunny situations, Creeping Cinquefoil flowers from June to September, producing solitary, brightyellow, 17-25 mm diameter flowers on long slender pedicels from prostrate, quickly rooting shoots. Each5-petalled flower contains around 20 stamens and 60-120 ovules, with nectar secreted by the receptaclearound the base of the stamens (Hutchinson 1972). The pollen and nectar attracts bees and flies.Although the flowers are fertile, seed set often fails because of a genetic self-incompatibilitymechanism, making cross-pollination essential (Harold 1994). The fruit is a head of single-seeded dryachenes varying in number from 25-200, usually about 90 (Salisbury 1964). Although some reports suggestseed is transient, other studies suggest it persists for five or more years buried in soil (Thompsonet al. 1997).
Thus P. reptans can reproduce by seed and even more effectively vegetatively by the production ofnumerous surface stolons or runners that spread rapidly and radially from the parent rootstock rosette.An individual stolon can travel 2 m or more in a season and develop rooted plantlets at c 15 cmintervals along its length. Removal of the aerial shoots and upper parts of the roots in the gardensetting is ineffective as a weeding measure, since the deeper root remnants regenerate adventitiousshoot buds and quickly re-establish the plant. Vegetative growth is especially active in damper groundand in wet seasons, and also in more shaded habitats where the ability to flower is reduced. Therelatively light seed allows it to wind-disperse, although some is taken by birds or is carried in mudand soil (Salisbury 1964). In any event, the seed manages to rapidly colonise new sites, where it canagain spread vegetatively to form clonal patches or larger carpets.
Hybrids
P. reptans sometimes coexists in disturbed ground alongside
P. erecta (Tormentil) and P. anglica (Trailing Tormentil) and several hybrid forms,intermediate in many respects, are known to occur. The two hybrids involved, P. × mixta (P. anglica × P. reptans)and P. × italica (P. erecta × P. reptans), are bothsterile and are impossible to separate in the field. P. × italica isthought to be exceedingly rare, having only once been cytologically confirmed in Britain (Stace etal. 2015). However, some forms of the parent species can be very similar to their hybrids and,since the hybrids can display both morphological seasonal variation and apomixis in some circ*mstances,they really are only distinguishable on the basis of their very low or zero fertility (Czapik 1975;Stace 1975).
It is not surprising that these hybrids are only extremely rarely recorded in Ireland and RHN ad thecurrent author (RSF) doubt if anyone has even looked for them in Fermanagh for the last half century. Amap for P. × mixta s.l. plotted in the New Atlas isaccompanied by a warning that these hybrids are under-recorded to an unknown and very obviously unevenextent throughout B & I (D.J. McCosh, in: Preston et al. 2002).
British and Irish occurrence
P. reptans is frequent and widespread in most of S, E & C Ireland, becoming less prevalent inthe more acidic terrain of the W & NW, which is often peaty and infertile. In Britain, P.reptans is common throughout lowland England and Wales, but further N in Scotland, it becomesrarer and more coastal, and beyond the Clyde and Forth conurbations it is only sporadically present andthen chiefly accidently introduced (New Atlas).
European and world occurrence
P. reptans has a widespread, almost continuous native distribution in warm temperate Europestretching from the Mediterranean basin to the southern Baltic region of Sweden and Finland (Kurttoet al. 2004). It is also native in W Asia and N Africa, but it is believed to have spread withsettlement both within and beyond its indigenous range. It has been introduced in Ethiopia, N & SAmerica and New Zealand (Hultén & Fries 1986, Map 1122).
Threats
None.
Native, common. Suboceanic temperate, but also naturalised in SE Newfoundland.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
The smallish, grey-green, ternate leaves with the terminal tooth shorter than those on each side, makethis low-growing, tufted, stoloniferous, wintergreen, semi-rosette, white flowered perennial easy todistinguish at all times of year from the otherwise morphologically very similar Fragaria vesca(Wild Strawberry). This species also flowers much earlier, from late February onwards to about May orearly June and the fruit is a collection of dry, hairy, achenes, unlike the red, fleshy and deliciousWild Strawberry.
Compared with our native yellow-flowered Potentilla species, P. sterilis is shallow rootedand it therefore prefers moist to dry but definitely never droughted, base-rich, generally Carboniferouslimestone soils. It also prefers open, sunny or only partially shaded, mainly lowland sites. Under suchconditions, competition from taller and more vigorous neighbours is limited by a number of additionalstress factors such as infertile, unproductive, fairly shallow or rocky soils, or moderate levels ofgrazing.
P. sterilis is chiefly found on dry banks, open woods, scrub and hedgerows or their margins,crevices on limestone cliffs, screes and quarries and in less fertile pastures and grassy roadsideverges.
Although Grime and his co-workers classified P. sterilis in terms of its ecological strategy as astress-tolerant ruderal or an intermediate balance of competitor/stress-tolerator/ruderal (C-S-R), thetolerances only stretch to cover some habitat situations. The species, for example, completelyavoids peaty or more strongly acidic soils of pH below 4.5 and it is equally intolerant of any form ofmoisture extreme, deep shade or heavily disturbed ground at any altitude (Grime et al. 1988).
Despite the above, population regeneration of this relatively uncompetitive plant is favoured by moderateor occasional levels of disturbance sufficient to create vegetation gaps that keep growing conditionsopen to colonising species. Examples of suitable forms of disturbance comprise trampling on waysides,grazing in pastures and soil creep on steep slopes or cliff ledges.
Fermanagh occurrence
Barren Strawberry is common and widespread throughout Fermanagh, but especially so in the Carboniferouslimestone areas. It has been recorded in 315 tetrads, 59.7% of those in the VC.
Flowering and vegetative reproduction
P. sterilis flowers and fruits early in the season (February to late May), 1-3 flowers beingcarried on each slender, procumbent, 5-15 cm axillary stem, produced from the central leaf rosette. Theflower is 10-15 mm in diameter, with many stamens and nectar secreted by a circular disk between thestamen bases and the numerous carpels. Pollen and nectar attract insect visitors, flies, butterflies,moths and bees (Fitter 1987). If insect pollination fails to occur, the stamens bend inwards andself-pollinate the stigmas. The dry fruit consists of numerous small achenes on a flat hypanthium with acentral domed, hairy receptacle which does not enlarge and become fleshy like the Wild Strawberry doesin fruit (Hutchinson 1972). The fact that the flower appears to wither away, "leaving behind abarren or chaffe head, in shape like a Strawberrie, but of no woorth or value", gave rise to theEnglish common name (Gerard 1633), and hence the erroneous notion that this Potentilla is in someway sterile, as indicated also by its scientific name.
The seeds are capable of prolonged survival in the soil, at least under woodland conditions (Warr etal. 1994). Other estimates regard seed survival as transient (Thompson et al. 1997).
In common with Wild Strawberry, the possession of several stolons per plant allows P. sterilis tospread locally to a limited extent, forming more or less diffuse clumps or larger carpet-forming clones(Grime et al. 1988).
British and Irish occurrence
Barren Strawberry is common and widespread in most of B & I except for the wetter and moreacid areas along the W coast of Ireland, around the English Wash and in N Scotland and the W & NIsles. The low figure of Atlas Change Index (-0.30), indicates a stable distribution throughout theseislands (Preston et al. 2002).
European and World occurrence
Beyond the shores of B & I, P. sterilis is mainly restricted to W & C Europe, reachingnorthward only to E Denmark. There are two outliers further east, in W Turkey and the Caucasus (Hultén& Fries 1986, Map 1123). A solitary station on Avalon Peninsula, SE Newfoundland dating from 1927was considered indigenous by some N American botanists (eg Fernald (1950)), although the site was almostas disjunct as a plant species could possibly be. This notion has now been has been refuted by laterAmerican Flora editors, Scoggan (1978) and Kartesz (1999), both of whom regard the solitary New Worldstation as alien.
Threats
None.
Native, common and widespread. Eurosiberian temperate, but also in N America and widely naturalised.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This very familiar, low-growing, wintergreen, rosette-forming, weedy perennial colonises dry to damp butwell-drained, shallow, rather stony soils in woods, scrub, hedgerow and roadside banks, open wasteground, old walls, gravel and in crevices on upland limestone cliffs, rock outcrops, pavement andscrees. F. vesca prefers a soil reaction near neutral and it seldom tolerates conditions below pH5.0 (Grime et al. 1988).
F. vesca typically occupies a wide variety of less-than-ideal, stressful habitats wherecompetition with potential dominants is reduced or absent due to the physical or chemical severity ofthe growing conditions. Factors that inhibit competition include shade, dryness, low-nutrient levels, orabsolute physical space limits imposed by crevices in bare rocks, pavements or in walls. In theSheffield area, the established ecological strategy of F. vesca was assessed as lyingintermediate between stress-tolerator and competitor-stress-tolerator-ruderal (C-S-R) by Grime etal. (1988).
Fermanagh occurrence
F. vesca is common and widespread in Fermanagh except in very wet or very acid peaty soils. It hasbeen recorded in 379 tetrads, 71.8% of those in the VC.
Vegetative reproduction
The thick, woody rootstock of the plant helps it survive drought situations remarkably well and itscharacteristic vigorous stoloniferous spread forms substantial clones in open sites. The arching stolonsare produced after flowering finishes. In common with related clonal species, individual colonies maywell be very long-lived in stable, albeit often severe and limiting environments.
Flowering reproduction
Hermaphrodite (perfect) flowers appear from April to July in few-flowered cymes on axillary branches. Theflower parts are 5-merous, the obtuse white petals nearly touching or overlapping, unlike those of P.sterilis (Barren Strawberry) which are definitely separated. As in the latter, numerous stamenssurround the 40 or so central carpels and nectar is secreted from a disc on the hypanthium inside thestamen ring. Insect visitors including flies, bees, butterflies and moths are attracted by pollen andnectar food and they cross-pollinate the flowers. In the absence of insects, selfing occurs (Grime etal. 1988). Small, red, fleshy strawberries, 10-20 mm in diameter, begin appearing in June. Theedible 'fruit' of the strawberry is a textbook example of a false fruit. It is not a berry at all, butis formed from the greatly swollen, fleshy, hairless receptacle which grows up through the thin, ± flathypanthium and encloses the true fruits, which are the dry, black achenes that appear embedded in theglossy red surface of the strawberry, the so-called 'seeds' (Sell & Murrell 2014). Achenes aresingle-seeded dry fruits, not actual seeds, although we often refer to them as such.
Fruit and seed dispersal
Despite the obviously edible, red, sweet, fleshy fruit, obtaining definite direct evidence that birdscollect and disperse them is not easily achieved. In common with most botanists, Snow & Snow (1988,p. 100) believe that birds definitely ingest the fruits and disperse the seed, although their eight anda half hours of observation proved fruitless, if the current author may be allowed the pun! Theseobservers suggest that even when wild strawberries are locally abundant, their very small size meansthey still offer only a sparse food reward for the seed-transporting animal.
German literature records members of the Thrush family, Chaffinches and Blackcaps as consumers of wildstrawberries. In view of their obvious scent and the fact that the fruit is produced at ground level,they are poorly presented to birds searching from above. This led Snow & Snow (1988) to proposethat, in this case, mammals and slugs may be more significant than birds in their frequency in acceptingthe food reward and enabling internal F. vesca seed dispersal. Ridley (1930) reports manyinstances from around the world of birds transporting F. vesca seed and he also mentions a caseof snails doing the same with F. × ananassa duch*esne (GardenStrawberry).
Again as a consequence of scale, it is not surprising that this rather dwarf plant is reputed to betolerant of light grazing (Sinker et al. 1985). Browsing animals of any real size are unlikely towaste much time foraging on very small plants.
While vegetative reproduction is probably responsible for most regeneration in existing F. vescacolonies, seed is very important in allowing dissemination to fresh sites. Seed is also capable ofpersisting buried in the soil for up to five years, or perhaps longer (Thompson et al. 1997).
In appearance, Wild Strawberry fruit is like a garden strawberry in miniature and, nowadays, it issometimes far more tasty than imported unseasonal representatives many times their size! F. vescais a relative of the now cultivated "Alpine Strawberry" (Garrard & Streeter 1983).
British and Irish occurrence
F. vesca is common and widespread throughout B & I. It becomes less universal in the morestrongly acid, peaty and wetter soils along the western seaboards of both B & I and this isparticularly the case on the N & W Scottish isles. There is some evidence of a decline of thespecies since the 1962 Atlas, particularly in S England (D.J. McCosh, in: Preston et al.2002).
European and world occurrence
The species, in the broad taxonomic sense, is common and widespread in most of temperate Eurasiaincluding some of the Mediterranean isles (Sicily, Sardinia and Corsica) as well as Iceland, but not theAzores, the Faeroes or the Macronesian islands (Kurtto et al. 2004, Map 3550). It is also presentin N America, although there it is known as var. americana. F. vesca is also introduced inmany countries around the world (Hultén & Fries 1986, Map 1125).
Threats
None.
Native, frequent. Eurosiberian boreo-temperate, but also native in N America.
1866-72; More, A.G.; Co Fermanagh.
March to January.
Growth form and preferred habitats
Like the closely related, but considerably more common and widespread G. urbanum (Wood Avens),this is a perennial species of mildly acid to calcareous, moderately fertile soils. In comparison withthe latter, it prefers, or rather it much better tolerates, wetter, slower draining, ± permanently dampgrowing conditions. In the damp, cool oceanic climate of Fermanagh, the environmental requirements ofG. rivale are readily met in a wide variety of habitats, including the damper flushed parts ofdeciduous woodlands, in marshy grasslands or fen-carr scrub by lakes, turloughs (so-called"vanishing lakes" with no proper inflow and outflow) and rivers, or on shady, ledges onnorth-facing cliffs where water seeps down them.
G. rivale persists in much wetter soils and in more upland sites than G. urbanum, includingground that floods and becomes waterlogged for periods in winter or after heavy rainfall. This islargely thanks to its well-developed, much branched, horizontally spreading rhizome-like caudex, afeature represented only by a very short thickened organ in G. urbanum (Taylor 1997b). Inmorphological terms, the caudex is an aggregation of leaf bases with their associated dormant buds andit is seen as intermediate between a true rhizome and an over-ground stem or stolon. The caudex of G.rivale is very well developed and it can grow and spread over a considerable horizontaldistance.
Survival of G. rivale in waterlogged conditions is due to the numerous vigorous adventitious rootsthe caudex produces (Waldren et al. 1987). Since it simultaneously develops annual floweringshoots, the horizontal stem can rot away at one end, extend at the other, produce a dense clone andunder favourable growing conditions persist for many years (Taylor 1997b).
Fermanagh occurrence
G. rivale is frequent in Fermanagh, having records in 148 tetrads, 28% of those in the VC. As thetetrad distribution map shows, it is widely scattered but much more frequent in the damp uplands to theSW of Lough Erne.
Considering the amount of wet ground that exists in Fermanagh, it is rather surprising that Water Avensis less than half as frequent and widespread in the VC in comparison with Wood Avens. Of the two,G. rivale is somewhat more competitive in grassland situations, slightly more tolerant ofoccasional mowing (for example, on verges or waterside banks) and, likewise, with respect to lightgrazing of pastures (Sinker et al. 1985). On the other hand, Water Avens does not colonise thenumerous weedy or dry soil situations exploited by G. urbanum.
Another aspect of competition affecting G. rivale is that when aggression is sufficiently strongto limit the species' growth and resources, maturity of the plant measured in terms of first ability toflower can be delayed for some years. The species is polycarpic and individual plants continue to flowerand fruit for a number of years after attaining maturity, so that the loss of two or more years' seedoutput may be but a small percentage of the total seed production throughout life. Nevertheless, theprolongation of the juvenile phase greatly increases the risk of mortality prior to reproduction andthis will inevitably affect the survival capacity of the population in a particular community (Salisbury1942, p. 54).
Vegetative reproduction
Reproductive strategy in G. rivale is a combination of vegetative and sexual, weighted towards theformer strategy. Consequently, G. rivale is able to compete directly with neighbours, or whenthey are more vigorous, it can avoid competition for ecological space through the mobility conferredupon the species by its vegetative dispersal, a property also displayed by the related stoloniferousspecies Potentilla anserina (Sliverweed) and P. reptans (Creeping Cinquefoil) (Erikkson1986).
Flowering reproduction
Pendulous, bell-like (campanulate) pink to dark crimson, 5-merous flowers are produced in a 2-5 flowerednarrow cyme from late May to September. The flowers are protogynous, the stigmas ripen first, protrudingbeyond the immature anthers, inviting cross-pollination by bees, flies or beetles attracted bysemi-concealed nectar, secreted by the saucer-shaped hypanthium at the base of the stamen filaments.Secondarily, after the anthers have matured and their filaments elongated, automatic self-pollinationbecomes a possibility (Taylor 1997b).
Either way, after pollen-transfer and fertilisation, around 100-150 crowded achenes are produced perflower, each one equipped with a strong hooked awn developed from the original stigma, enabling it toadhere to the coat of a passing animal or human (Ridley 1930, p. 142). A Swedish study found that G.rivale achenes attached easily to the fur of both fallow deer and domestic cattle and had thepotential to be transported from tens of metres up to a kilometre or more, depending upon the range ofthe animal species and the amount of grooming it performs (Kiviniemi 1996). The seed appears to betransient in the soil, persisting for less than one year (Thompson et al. 1997). Presumably theseed germinates in spring after overwintering, although nothing appears to be known about fieldgermination and seedling establishment (Taylor 1997b).
Variation
A taller variant of the plant with stems up to 30(-50) cm, with more coarsely toothed leaflets andflowers with initially white petals that later turn somewhat pinkish, occurs in northern regionsincluding Iceland, the Faeroes, N Scandinavia, N America and Britain. It has been variously named f.subalpinum, var. subalpinum or subsp. subalpinum (Neumann) Selander. In Britain, ithas been recognised as the latter by Sell & Murrell (2014).
British and Irish occurrence
Water Avens is widespread in both B & I, but has a definite northern bias, especially noticeable inIreland. There is also a much lesser western leaning in its distribution in England, and it iswidespread throughout Wales (Preston et al. 2002). Being a species of wetter ground, G.rivale has suffered a decline in its presence in both islands due to drainage associated withthe intensification of agriculture during the last 60 years or so. This change in land management andspecies loss is particularly marked in C & S England. The plant is also grown in gardens andoccasionally escapes (D.J. McCosh, in: Preston et al. 2002).
The Fermanagh data are insufficiently detailed historically to allow any estimate the local extent ofthis general trend in losses.
European and world occurrence
G. rivale occurs throughout most of W, N & C Europe but thins markedly southwards inFrance, Spain, Italy and the Balkans. It is entirely absent from the Mediterranean isles and Macronesiaand becomes scattered, and perhaps less well recorded, eastwards into Russia, the Caucasus, Turkey andSiberia (Kurtto et al. 2004, Map 3374). Although it is not circumpolar, the native range of G.rivale is amphi-atlantic extending to N America, from Newfoundland to British Columbia, Coloradoand New Jersey (Sell & Murrell 2014; Hultén & Fries 1986, Map 1091).
Threats
None.
Occasional and very local.
1884; Barrington, R.M.; N end of White Island, Eastern Lower Lough Erne.
March to June.
The Fermanagh distribution of this relatively frequent fertile intermediate hybrid closely reflects thatof its less widespread parent G. rivale (Water Avens), or rather the overlap in distribution andhabitats of the two parent species. It is claimed that the frequency of the hybrid in the area of theparent species overlap is positively influenced by disturbance (Taylor 1997a), which would tend to bringthe more weedy G. urbanum (Wood Avens) into greater contact with the natural wet ground species,G. rivale. While this may be generally true, only three of the 18 or more distinct sites knownfor this hybrid in a total of 17 Fermanagh tetrads might conceivably be considered public enough to fitthis pattern (eg the Crom Castle and Florencecourt National Trust estates and the Cladagh River GlenNR).
The ecological separation of the two parent species is not the whole story, since in the very mild, dampclimate of W Ireland, the flowering periods of these Geum species certainly overlap more thanthey do in drier, more continental climatic conditions (Briggs & Walters 1997, p. 282). Thesecirc*mstances also pertain at the much larger scale of B & I and the New Atlas map shows thatthis hybrid is much more frequent in NI than elsewhere on the island.
The Irish Flora Census Catalogue indicates that G. × intermedium hasbeen recorded at least once in 19 of the 40 Irish VCs (Scannell & Synnott 1987), a figure which wecan revise upwards to 24 thanks to the New Atlas map (Preston et al. 2002).
G. × intermedium is fully fertile and it can therefore backcross withboth parents, sometimes displaying an array of forms intermediate in character between the parents whichare typically both present in the near vicinity, ie it forms a hybrid swarm with a continuous linkingrange of intermediates between the two parent species.
While it is relatively frequent in some VCs, although still always very local, eg in Fermanagh and CoAntrim (H33, H39)(Hackney et al. 1992), in the Flora of County Dublin (H21), it wasdescribed as extremely rare, there being just one record dating from 1983 (Doogue et al. 1998).
As with G. rivale, the New Altas survey suggests that there have been some losses,particularly in C & S England, presumably due to drainage and other factors responsible fordestruction of damp, shady habitats (D.J. McCosh, in: Preston et al. 2002).
Threats
None.
Native, common and widespread. Eurosiberian temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This variable, wintergreen, rosette-forming, pubescent, herbaceous perennial possesses a short, thick,overwintering rhizome that seldom branches. A new rosette of basal leaves is produced by the rhizomeapex each year, but the lifespan of individual rhizomes is only a few years (Taylor 1997a). G.urbanum relies on seed to disperse and colonise suitable new ground and therefore is usuallysparsely distributed in most sites, since it originates from widely scattered propagules and possesseslittle capacity for vegetative spread (Grime et al. 1988). Wood Avens occurs in full- orsemi-shade and shelter of almost every small or large patch of deciduous woodland, scrub and substantialhedge in Co Fermanagh. Individual plants are also frequent as sporadic weeds of fully-lit, fairlydisturbed, open ground in gardens, urban waste ground and in neglected corners in quarries and similarunmanaged rural situations.
The species shows a definite preference for mildly acid to calcareous, sloping, free draining, moistsoils in the pH range 5.4-7.7 that are average in terms of plant nutrients and fertility. It possiblybenefits in habitats with a modicum of additional nitrogen, but it is not a strong plant competitor assuch, being more of an environmental stress tolerator (Ellenberg 1988; Grime et al. 1988). Assuch it cannot compete in grassland communities and is virtually absent in grazed habitats (Taylor1997a). G. urbanum can tolerate summer drought conditions rather well in some calcareous habitatsin S England, although similar conditions would be most unusual in NI with its frequent damp periods andstrongly expressed Western Atlantic climate. Having said this, Wood Avens can be considered amoist site indicator, being absent both from places that dry out and from wet ground (Taylor1997a).
Fermanagh occurrence
Since its ecological requirements are readily met in Fermanagh, Wood Avens is regularly recorded in 332tetrads, 62.9% of those in the VC. It is absent mainly from strongly acid soils, wetlands, closed turfgrasslands, more exposed uplands and regularly grazed ground (Taylor 1997a). While generally a lowlandspecies, the plant is hardy and easily survives the mild winters in our oceanic area, the wintergreenrosette probably being physiologically active right throughout the year. Perhaps on account of itsability to persist in some types of disturbed site, G. urbanum is more than twice as frequent andwidespread as G. rivale (Water Avens) in Fermanagh.
It is not easy to establish the upper altitudinal limit from the local Fermanagh data, but it is probablysomewhere around 350 m on the Belmore limestones, while the highest recorded site in B & I is 512 min the English Lake District (Wilson 1949).
Flowering reproduction
Flowering takes place from June to August, the inflorescence consisting of only 2-5 flowers borne on a 90cm tall stem that waves its relatively small (10-15 mm) yellow blossom to attract a range of insects,mainly flies, offering food rewards of little nectar but plentiful pollen. Despite this effort, mostflowers are probably unvisited and, in this event, they are automatically selfed (Taylor 1997a). Eachflower contains around 20 carpels.
In addition to the normal summer flowering, very tiny flowers are produced on side branches in September,when the fruits of the topmost heads are already well on the way to producing seed (actually achenes aresingle-seeded dry fruits).
Fruiting heads consist of brown, burr-like clusters of achenes, each with a 6 mm long, hooked awn whichreadily attaches to the coats and fur of passing animals and disperses the seed (Ridley 1930, p. 142;Hutchinson 1972).
Seed survival in soil is, at best, short-term persistent (1-5 years), although other studies suggest itis transient (less than one year) (Thompson et al. 1997). In one English outdoor, pot-basedstudy, after a cold spell of weather germination began in December, reached a peak in March and ceasedat the end of May. A few additional seedlings appeared in the following spring (Roberts 1986).
British and Irish occurrence
The New Atlas hectad map indicates that the distribution of G. urbanum is widespread andstable throughout B & I, but it becomes rare or absent in NW & C Scotland including Orkney,Shetland and the Outer Hebrides. It is also rare or absent on the ± permanently wet, strongly acidicpeats of W Ireland. However, the distribution does indicate that G. urbanum has a very wideclimatic tolerance (Taylor 1997a).
European and world occurrence
G. urbanum does not occur in Iceland and the extreme N of Europe, but it is widelydistributed southwards from S Scandinavia and Russia (65°N) to the Mediterranean, where it thins out insouthern parts of Spain, Portugal and Greece. It is well represented in Corsica, Sardinia and Sicily,but absent from all the other Mediterranean isles and from the Macaronesia islands (Kurtto et al. 2004, Map 3379).
Beyond Europe, G. urbanum extends eastwards to W Asia, W Siberia and the Himalaya and it is alsocommon on the NW African coast from W Algiers to the Tunisian border (Hultén & Fries 1986, Map 1092;Taylor 1997a). It has been introduced, presumably accidentally and to a minor degree, in eastern NAmerica and New Zealand (Hultén & Fries 1986). In New Zealand, it is easily confused with thesimilar and more common introduction G. alepicum Jacq., so that G. urbanum may beoverlooked to some extent and perhaps is more common than imagined (Webb et al. 1988, p. 1082).
Threats
None.
Native, rare and local. Circumpolar arctic-montane.
1804; Wade, Dr W.; eastern range of Cliffs of Magho.
May to July.
Growth form and preferred habitats
This variable, prostrate or low-growing, large creamy-white flowered, dwarf shrub, 5-20 cm in height, hasa stout woody rootstock that penetrates deep into rock crevices and sends out lateral roots. At groundlevel it produces several spreading shoots from the rootstock, each of which branches laterally androots adventitiously in suitable base- or lime-rich soil, forming dense or loose vegetative clones thatoften together carpet the rocks, cliff ledges and limestone pavement over which it grows.
The alternate, simple, leathery, evergreen, 2.5-4.0 cm leaves are very variable in size and shape andthey are crowded along the prostrate stems overlapping one another to give an almost total cover of thesoil. The rolled-down (revolute) margins of the leaves are crenately lobed or bluntly toothed, givingthem a scalloped look reminiscent of a miniature oak leaf. This appearance is reflected in the genusname, since 'dryas' is thought by some botanists to be from 'drus', the Greek for 'oak' (Johnson &Smith 1946). The upper surfaces of the small leaves are dark glossy green, ± hairless and rough-textured(rugose) with deeply impressed veins, while their under-surfaces are densely felted with a grey-whitetomentum of hairs.
While described as evergreen above, this may merely reflect the relatively mild oceanic climate of WIreland where D. octopetala evergreen leaf longevity is the norm. Elsewhere, in habitatssuffering more severe, extreme or variable temperature regimes, the plant is more likely deciduous,losing its mature leaves gradually through the winter, although retaining juvenile ones around the shootapices that will expand in spring. It might be better, therefore, to regard D. octopetala plantsas semi-evergreen (Elkington 1971). Nelson goes even further, suggesting that while the rootstock isdefinitely woody and the oldest stems can be well over 50 years old, the stems are less woody and theplants might be better considered an evergreen, perennial herb (Nelson & Walsh 1991, p. 153). Instating this opinion, the current author feels Nelson is being controversial and is out on a limb on hisown.
Factors limiting distribution
The distribution of D. octopetala is confined to the occurrence of basic or calcareous rocks andsoils. In Ireland, this means scattered Carboniferous limestones and northern basic basalts, while inBritain it also occurs on metamorphosed sugar limestone in the Cairngorms and Teesdale, mica-schists inthe central Grampians, lavas in the Lake District and Wales and on calcareous shell sands on the N &W coasts of Scotland (Lusby et al. 1996).
In addition to being restricted to typically calcicole or base-rich soils in B & I, D.octopetala is climatically limited to cool, humid regions with annual rainfall above 1000 mm (40inches) (Elkington 1971). On European continental mountains, it is usually well protected from winterfrost by a deep blanket of snow cover, while in B & I it is more exposed to freezing lowtemperatures due to the frequent absence of snow. However, observation shows the species is completelyresistant to frost and there is no evidence of drought affecting growth or damaging plants either(Elkington 1971).
A number of studies suggest the modern distribution of D. octopetala is probably primarily limitedby climatic factors and only secondarily affected by its edaphic (soil) requirements. It is well knownthat alpine and northern plants grown in lowland gardens suffer during the hottest spells of summer. Astudy by Conolly & Dahl (1970) found that the distribution of many arctic-montane species in B &I can be correlated with maximum summer temperatures and, in particular, with isotherms of the estimatedmean annual maximum temperature of the highest points they reach, both in terms of altitude andpole-ward proximity. Their map of the distribution of D. octopetala shows that in England, Walesand Ireland localities of the species are all included within the 25°C isotherm of this parameter, whilein Scotland all localities are bounded by the 23°C isotherm. Of course, correlation with a single factordoes not necessarily indicate or prove a causal relationship and a climatic factor that truly affectsthe distribution or growth of a species must act in some way on the physiological processes ofindividual plants of that species. It is not known if high temperatures have a direct effect on theplant or act in a more complex manner involving temperature. Since D. octopetala is closelyassociated with free-draining substrates, a very likely effect of increased summer temperature isincreasing moisture stress on the plant, affecting its growth, reproduction and, eventually, itssurvival (Elkington 1971).
Interestingly, in the horticultural setting D. octopetala is a very easy subject and can besuccessfully grown, "in any reasonable garden soil, and in any sunny place, requiring nothing butto be well planted and then left alone (with a top dressing at times) to get larger and wider forever". (Farrer 1930).
Lowland occurrences
Mountain Avens occurs in coastal areas down close to sea-level in parts of both W Ireland and NW Scotlandand also on relatively lowland hill slopes inland, including on Knockmore in Fermanagh. At these lowersites, it typically grows on free-draining slopes of sand and gravel or on steep cliff faces, bothsituations where erosion helps keep the habitat open and provides soil nutrient renewal by rockdisintegration. Thus habitat instability, together with the inherent low soil nutrient levels ofunproductive vegetation, limit any overly aggressive plant competition, while grazing pressure, to whichD. octopetala is rather sensitive, is much reduced by the inaccessibility of steep ground tosheep and other herbivores (Ingrouille 1995; Lusby et al. 1996).
In the Burren, Co Clare (H9), D. octopetala grows with a spectacular profusion over a wide areathat is unequalled anywhere else in NW Europe. Here it occurs at or near the coast, carpeting barelimestone rock on slopes and valleys leading to the shore, sometimes present in enormous abundance. Itavoids deep, clayey limestone drift but is perfectly at home and able to dominate vegetation over porousdrift and solid rock on N- or S-facing slopes from sea-level to around 300 m (Webb & Scannell 1983).In view of this, it is most surprising that there are large areas of ground in and around the Burrendistrict of NW Clare (H9) and SE Galway (H17), apparently sharing the very same geology, geography andhistory, where the species is totally absent. It is especially mysterious that D. octopetala iscompletely absent from the adjacent Aran Islands, which represent a slightly offshore section of themainland Burren and share much of its geology and characteristic mixed-geographic regional flora.
Fermanagh occurrence
Dryas octopetala is one of Fermanagh's most notable Arctic-Alpine Late-glacial survivors. It hasbeen found as a macrofossil in a full-glacial freshwater deposit of Middle Midlandian age, radio-carbondated to 30,500 BP, located at Derryvree, near Maguiresbridge, Co Fermanagh (Colhoun et al.1972). The flora and fauna of the deposit indicated that open tundra and a periglacial climate prevailedwhen the sediment was laid down.
Outside the Burren region of Cos Clare (H9) and Galway (H16), where it is still common and locallydominant on the extensive karst limestones, this is a very rare plant in Ireland. In NI, it occurs onlyon the base-rich basalt cliffs of Knock Dhu in Co Antrim (H39) and of Benevenagh in Co Londonderry(H40), plus this surviving Fermanagh station on the Carboniferous limestone summit of Knockmore hill,where S.A. Stewart discovered it in 1881. Here it still grows in abundance on a steep slope immediatelysouth of the trigonometrical point and in lesser quantity around the immediate vicinity of thetrigonometrical point plinth.
In July 2020, there was considerable delight and surprise when Hannah Northridge found a small patch ofD. octopetala, 1 m × 2 m, in a new site on a south facing slope on limestone at Monawilkin to theeast of the lake at an altitude of 190 m. Associated species included: Antennaria dioica(Mountain Everlasting), Anthyllis vulneraria (Kidney Vetch), Briza media (Quaking-grass),Campanula rotundifolia (Harebell), Carex flacca (Glaucous Sedge), Euphrasia spp.(Eyebright), Galium verum (Lady's Bedstraw), Juniperus communis (Juniper), Koeleriamacrantha (Crested Hair-grass), Leucanthemum vulgare (Oxeye Daisy), Linumcatharticum (Fairy Flax), Lotus corniculatus (Common Brid's-foot-trefoil), Pilosellaofficinarum (Mouse-ear-hawkweed), Sesleria caerulea (Blue Moor-grass) and Thymuspolytrichus (Wild Thyme).
At Dr Wade's other original 1804 Fermanagh site for the species, namely the north-facing Cliffs of Maghooverlooking Lower Lough Erne, D. octopetala has not been seen since the 1945-53 survey of Meikleand his co-workers. When Praeger refound Dr Wade's original station here exactly a century later, hedescribed the site as follows, "At the base of the east end of the eastern cliffs of Poulaphuca [analternative name for the site – appearing on maps and in the literature with several differentspellings], where the cliffs are low." Robert Northridge and the present author have searched allover this site several times without any success. It is a large area of cliff, some of it difficult orimpossible to access, so that a very small patch of D. octopetala might simply elude us, but wehave not seen the plant anywhere from along the cliff base.
Flowering reproduction
The main flowering period in lowland sites in western Ireland such as The Burren, Co Clare and Knockmorein Co Fermanagh is in late April and May. In more mountainous situations, anthesis is delayed until Juneand July, with much smaller numbers of flowers being produced throughout the summer and into autumn andwith a lesser late flush occurring in August (Elkington 1971; Nelson & Walsh 1991). Thehermaphrodite and male, large, 25-40 mm diameter flowers are borne solitary on erect, dark-red or green,white-tomentose peduncles 2-8 cm long with purplish-black glandular hairs above.
Unusually for a member of the Rosaceae, most flowers have eight white to cream petals, although thenumber can vary from 7-16. Stamens and carpels are both numerous, the latter bearing long, featherystyles that in fruit elongate further to 20-30 mm and become hairier. The receptacle is domed or conicaland the hypanthium saucer-like. A circular nectary on the mouth of the hypanthium sits between thestamens with their golden anthers and the clustered carpels. The flowers in B & I are usuallyhom*ogygous (anthers and stigmas maturing simultaneously) although protogyny and protandry have beenrecorded elsewhere in Europe. While most flowers are bisexual (perfect, hermaphrodite), a smallproportion are male only, containing undeveloped carpels. The flowers attract a range of insectsincluding flies, bees, butterflies, moths and beetles, although flies are considered the majorpollinators in B & I, Scandinavia and the Arctic. If a flower remains unvisited it mayself-pollinate and produce at least some viable seed, although less than if cross-pollination wasachieved. This suggests there is some trace of an incompatibility mechanism in the species (Elkington1971).
An interesting flowering adaptation known as sun-tracking (or heliotropism), allows the flowers of D.octopetala to constantly glow on a sunny day. The dish-shaped flowers constantly turn to facethe sun, thanks to a rotating physiological movement of their peduncles. It is believed that in colderarctic and mountain environments, the heating provided within the near-parabolic dish of the blossomhelps focus the rays of the sun onto the anthers and carpels. This feature warms the reproductiveorgans, presumably increasing the rate of their development; it will also warm the nectar, making theflowers more attractive to the relatively few flying insects that remain active in these cold habitats.Insects are also known to sun-bask within some types of arctic and alpine flowers such as D.octopetala, their prolonged flower visits allow them to warm themselves, increasing theefficiency of their flight muscles and assisting their subsequent flight (Hocking 1968).
Seed production and dispersal
Achene counts vary across B & I from 48 per fruiting head in Co Clare to 61 per head in Perthshire.The number of flowers per plant probably varies enormously, perhaps as many as 100 on a large plant,although very often it is impossible to recognise an individual as the species is so very gregarious andmat-forming. The achenes are each c 3 mm in size and have a persistent style 2-3 cm long in fruit. Theachene and its style are both covered in long silver-white lateral hairs at this stage, making thefruiting head look like "beautiful silver fluff-whirls", as the rock gardener Reginald Farrer(1930) described them. In the Burren, Co Clare where the plant is locally dominant and abundant, thefeathery fruiting heads are described by Webb & Scannell (1983) as, "almost as conspicuous asthe flowers". The peduncle that bears the fruiting head elongates to about twice its originallength as the achenes ripen. The achenes thus appear very well equipped for wind transport by theirfeathery styles and by their elevation above the creeping vegetative mat of the plant.
An unknown proportion of achenes disperse on the breeze, distances again unmeasured, but in gardenexperience many achenes appear to end up around the base of the parent, or close to it. There is atendency for the long, hairy styles to twist around one another, either when immature or in wet weather,which at least temporarily, must seriously reduce the efficiency of their dispersal (Elkington 1971).The fact that so many populations of D. octopetala appear small, isolated, relict and confined tolong-known sites, and in other areas like the Burren, Co Clare, super-abundance rubs shoulders withtotal absence in very similar habitats, suggests to the current author that under present environmentalconditions (including climatic warming), dispersal has become a major problem limiting the distributionand affecting the longer term survival of this once extremely widespread arctic-alpine species.
Seed germination and survival
Seed normally germinates in the subsequent spring after overwintering and buried seed is transient in thesoil, surviving for one year or less (Thompson et al. 1997).
Effective reproduction appears to be normally by seed, although in modern times at least, seedlings andyoung plants seem to be only rarely recorded under natural conditions in the field. This cannot alwayshave been the case since, from the plentiful fossil record, D. octopetala was capable of rapidinvasive colonisation of recently de-glaciated, unoccupied soils (silts, sands, gravels) in the LateGlacial and early Post-glacial periods. Indeed, it became so widespread and abundant in the exposedtundra as northern European glacial ice-sheets retreated, that earlier palaeontologists labelled twoperiods in the Late-glacial 'the Older Dryas'(12,500-12,000 BP) and 'the Younger Dryas'(12,300-11,500 BP), with the brief much colder 'Allerød' glacial snap separating them. This labellingarose because fossil leaves and pollen of the subshrub were so plentiful during these two climaticallywarmer periods when Devensian (or in Ireland 'Midlandian') glaciers began to melt. The abundance of thespecies in the Late Devensian and early Post-glacial is evidence of the low intensity of grazing atthese times. D. octopetala and other members of the Dryas heathland vegetation communityare characterised by low growth rates, so that as the climate continued to warm, they could not competeagainst a broad range of colonising species. Only in the cooler N & W of Ireland & Scotland didopen Dryas-heath continue to cover large areas before the advent of sheep grazing and beforeleaching made soils more acidic (Ingrouille 1995).
While today, evidence of Dryas seed reproduction and colonisation appears surprisingly scant, anexception was the report of seedlings growing among mature D. octopetala plants and on barepatches in Sesleria caerulea (Blue Moor-grass) dominated grassland in the Burren, Co Clare,recorded in 1959 and 1960 by Prof C.D. Pigott (quoted in: Elkington 1971). Further study is needed toclarify the reproductive capability and establishment of D. octopetala in today's warming climateand changing soil conditions.
Vegetative reproduction and individual longevity
A very limited amount of vegetative reproduction may be achieved by the decay of old stems in existingclones, but this is unlikely to be significant in terms of species increase or spread. Shoot growth onolder stems is often asymmetrical, with the pith being formed to one side, often the upper, and withincomplete growth rings. Counts of growth rings on the oldest woody stems in the Kola peninsula inLappland (68°N) showed the individuals were between 50 and 108 years old (Kihlmann (1890), quoted inSchroeter (1908, p. 185)). Measures like this indicate the plant has a definite, rather surprisinglylimited lifespan and its local survival in stations that today are geographically isolated must involvemaintenance recruitment from seed or genets (offset plantlets) of the existing stock (Elkington 1971).
Fossil history
The leaves of D. octopetala can be readily identified as macrofossils in peat and silt deposits,even in fragmentary condition on account of their strongly recurved margins, densely hairy under-surfaceand characteristic venation. There are Middle and Late Weichselian macrofossil records from most of thepresent day Mountain Avens sites in northern Britain and N & W Ireland, plus a few sites furthersouth in Wales and SE England where the species no longer occurs (Godwin 1975, Map of B & I in Fig.62). There are also pollen records from the Late Weichselian. The Older and Younger 'Dryas clays'widespread in N & W Europe are deposits now recognised as corresponding with zones I & III ofthe Late-glacial period (see also above).
A second European map (Fig. 63) reproduced in Godwin (1975), shows how the modern Arctic and alpineEuropean distribution of D. octopetala has become fragmented compared with its fossil record bythe loss of intermediate lowland stations from the Netherlands, Denmark, NW Germany, southern parts ofNorway and Sweden, plus isolated areas of C Europe towards the Black Sea. The post-glacial Flandrian(Littletonian) decline across the whole of its European range has been very considerable and must beattributed to habitat changes affecting plant competition involving factors such as rising summertemperatures, soil chemistry acidification as it ages, plus disturbance by man and his animals.
Variation
All chromosome counts made to date regard D. octopetala as diploid with 2n=18. However,being a very widespread species, D. octopetala contains considerable variation, both phenotypicwith respect to different environments and genotypic across its wide geographic range, so it isunsurprising that people have named forms, varieties and subspecies from time-to-time. The FloraEuropaea account (T.T. Elkington, in: Tutin et al. 1968) mentions two forms, plants ofthe first lacking branched hairs on the leaves named as D. babingtoniana A.E. Porsild, and asecond form named D. punctata Juz. from Arctic Russia, which has large glands on the uppersurface of the leaves. However, the differences are slight and herbarium studies prove there areintermediate forms, so neither of these taxa are considered worthy of more than varietal rank, and it isprobably best to regard D. octopetala as one polymorphic species (T.T. Elkington, in: Tutin etal. 1968; Lusby et al. 1996).
Hybrids
D. octopetala and the closely related northern boreal and Arctic form D. integrifoliaoverlap and crossbreed in Greenland (Elkington 1965) and Alaska (Hultén 1968), with subsequentwidespread introgression taking place (Elkington 1971).
British and Irish occurrence
In Britain, for the most part, D. octopetala is thinly scattered across a few mountain sites in NWEngland and NW Wales, while in Scotland it is much better represented across a wider range of habitats,altitudes and stations, stretching from sea-level in the Inner Hebrides to 1035 m in the Cairngorms. InIreland, Mountain Avens is common and dominant in a few areas in the Burren, Co Clare from sea level to300 m on the hills. Elsewhere in Ireland, it is thinly scattered (although locally abundant in a fewsites in SE, NE & W Galway (H15, H16 & H17)), with a few isolated, mainly relict mountain cliffsites in northern and western VCs from Sligo (H28) and Leitrim (H29), Fermanagh (H33), W Donegal (H35)to Londonderry (H40) and Antrim (H39). (D.J. McCosh, in: Preston et al. 2002).
European and world occurrence
In Europe, at least, D. octopetala is considered a single species and it has a very definiteArctic-montane distribution stretching in the north from Greenland, Iceland, Spitzbergen, the Faeroes, WNorway, N Sweden and Finland to N Russia and then SW to B & I. In S Europe, it stretches acrossmountain ranges from the Pyrenees, Alps and Apennines, to the mountains of the Balkans and Macedonia(Kurtto et al. 2004, Map 3367). Beyond Europe it ranges from the Caucasus to Turkey and fromSiberia, Korea to the Yenisei region of Japan. In N America, taking it in a wider s.l. sense to includesuch sometimes disputed forms as subsp. alaskensis (Porsild) Hult. and subsp. integrifolia(Vahl) Hult. (sometimes recognised as separate species), it is widely distributed from Alaska and theYukon east to the Mackenzie River and south along the Rocky Mountains to Colorado (Elkington 1971;Hultén 1974, Map 45; Hultén & Fries 1986, Map 1190). In the broad sense, D. octopetala isconsidered a circumpolar Arctic-montane species.
Threats
Fertiliser 'improvement' of the pastures on Knockmore, or excessive sheep grazing, would thus threatenits slender hold on its solitary surviving Fermanagh site.
Native or possibly an archaeophyte, occasional. Eurosiberian southern-temperate, but widely naturalised.
1882; Stewart, S.A.; Carrick Td.
May to October.
Growth form and preferred habitats
This variable, 30-50(-150) cm tall, erect, yellow- flowered, rhizomatous perennial with 3-6 pairs oflarge pinnate leaves and small leaflets between them, occurs as scattered individuals on dry,well-drained, rough grassy places in lowland areas. It prefers calcareous or basic soils that arenear-neutral in reaction. A. eupatoria typically grows on drier hedgebanks, sloping waysideverges, lakeshore meadows and on the margins and in openings in woodlands and scrub in conditions offull sun to half-shade. The types of habitat suggest that it is well able to compete with tall grassesand other herbs and it can also tolerate occasional mowing or light grazing pressure (Sinker etal. 1985). Sheep and goats will browse on the plant, but horses, cattle and pigs leave it alone(Grieve 1931).
As a result of its dispersed individual nature, ± unbranched form and occasional occurrence, Agrimony isreally only conspicuous when it produces its long, slender, tapering, spike-like racemes of yellowflowers. The vegetative differences between A. eupatoria and the closely related, usually taller,larger and coarser A. procera Wallr. (Fragrant Agrimony) are rather slight and, despite theEnglish common name of the latter, leaves of both species give off a fragrant balsamic scent whencrushed. For this reason, ripe fruits really are the only certain character to distinguish the twospecies (Stace 2019).
Flowering reproduction
In common with many members of the Rosaceae, plants of A. eupatoria possess a rather woody,perennating rhizome, but there is very little vegetative spread and for its reproductive increase anddispersal the species relies entirely on seed (Butcher 1961). Plants flower from June to August,developing long, terminal, rod- or spike-like racemes of numerous, closely clustered, small, 5-8 mmdiameter, bright yellow, 5-merous flowers each containing just two carpels in a deeply concave, cup- ortop-shaped ribbed hypanthium or receptacle. While it is purely a pollen blossom, containing no nectar,the spike gives off a spicy odour like apricots and manages to attract bees, flies and other smallinsects which pollinate it. Failing this, the flowers frequently self-pollinate (Clapham et al.1964; Garrard & Streeter 1983; Fitter 1987).
The pendant, now rather woody fruits contain only one or two seeds and are bell- or top-shaped, armedaround their apex rim with numerous stiff bristles hooked at the tip. The spike or raceme elongatesconsiderably after fertilization has taken place. The fruits thus function as burrs, attaching firmly tothe rough coats of passing animals or human clothing in a very efficient form of seed dispersal alongroadsides or in pastures (Ridley 1930, p. 590).
Uses and names
A. eupatoria has a long history as a medicinal herb with magical properties dating back to ancientGreek and Roman times, Pliny and Dioscorides. The ancient Greeks knew it as 'Argemone' and prescribed italong with other similar herbs in the treatment of eye complaints (Le Strange 1977). Dioscoridesrecommended Agrimony for the treatment of snake bites, dysentery and liver complaints. In Anglo-Saxon B& I, it was regarded as 'a simple' and was well known to all country-folk as one of the best herbsfor external salves, healing all kinds of wounds and sores, including bites, battle-wounds and warts.
Agrimony (both A. eupatoria and the less common A. procera), was one of 57 herbs in theAnglo-Saxon magical 'Holy Salve', said to provide active protection against goblins, evil and poisons.It was also valued in medieval times for its astringent, diuretic properties, which kept it popular fora long period in herbal medicine for sprains and bruises (Grieve 1931). In addition to collection of theherb in the field, A. eupatoria was brought into garden cultivation for medicinal use and as adye plant (see below). It is very easy to grow (Miller 1741; Grieve 1931; Grigson 1987).
By the end of the 16th century, the herb was being taken internally, mainly used to make a tea as a mildtonic and stimulant. It was considered a general prophylactic and purifier of the system, as well as atreatment for sore throats and the worst sort of colds and coughs (Le Strange 1977). However, it wasalso recommended for more serious ailments such as rheumatism, backache and ailments of the liver,kidneys and bladder including jaundice (Grieve 1931; Darwin 1996; Allen & Hatfield 2004). Whenintroduced to N America, A. eupatoria was said to have been used to treat fevers with greatsuccess by the Indians and Canadians (Grieve 1931). There is a widespread N American species equivalentin the related A. striata Michx. (Hultén & Fries 1986, Map 1086).
In the autumn, when it is gathered, the whole plant yields a pale yellow dye, but later in the season thecolour becomes stronger (Grieve 1931).
Numerous English common names allude to these folk medicinal and magical supernatural uses, such as 'Teaplant', 'Aaron's Rod' and 'Fairy's Rod', but other names refer more directly to the clinging burrfruits, including 'Clot-bur', 'co*ckle-burr', 'Stickwort', Stickle-wort', 'Sweethearts' and'Harvest-lice' (Grigson 1987).
The question of status
A. eupatoria is usually regarded as native in both B & I and there have been a couple offossil fruits found in two earlier interglacial periods (Hoxnian and Ipswichian), but not in the currentFlandrian (Littletonian) warm period. However, there is a pollen record from Zone VIIb of the Flandrian,although it is only regarded as tentative (Godwin 1975, p. 194). Later finds date from the Neolithic andBronze Age (Sub-boreal), Roman and Medieval periods, suggesting that the species could have beenintroduced to B & I by Neolithic or later farmers and traders along with numerous other weed andruderal species, making it an archaeophyte (an ancient introduction) (Godwin 1975, Table 43).
In view of its ancient and longstanding herbal reputation and use, A. eupatoria could well be anintrusive introduction, brought into cultivation and spread from gardens into wayside habitats, assuggested or hinted at by Allen & Hatfield (2004).
Fermanagh occurrence
In Fermanagh, A. eupatoria is occasionally recorded in 39 tetrads, 7.4% of those in the VC. It hasnot been found in seven of these tetrads during the post-1975 period, however, which suggests somethingof a species decline that is also noticeable across much of B & I at the hectad level in the NewAtlas. Locally, Agrimony is chiefly associated with hedgebanks and lakeshore meadows, especiallyalong the Lough Erne basin, being very much more occasional on limestone elsewhere in the VC.
The association of many of the Fermanagh records with lakeshores suggests fruit dispersal could not onlyengage the coats of animals and man, but might also involve flotation in water. Ridley (1930, p. 208)quotes work by Praeger (1913) on buoyancy of fruits and seeds, mentioning that A. eupatoria isunusual amongst Rosaceae in that its fruits remain afloat for up to a week, while most other herbaceousmembers of the family manage only 2.5 days.
British and Irish occurrence
Although certainly an uncommon species in Fermanagh and N Ireland, it is fairly frequent and widespreadthroughout Ireland as a whole, although probably with something of a southern and eastern bias in itsoccurrence. In Britain, it is widespread: more common in the S & E, but increasingly scarcenorthwards and absent in a good part of C Wales and in N Scotland (New Atlas).
European and world occurrence
In Flora Europaea 2: 32 (Tutin et al. 1968) three subspecies are listed, but theyare rather ill-defined and completely intergrade in parts of S & E Europe and it is probably betterto ignore them when considering the species range (Kurrto et al. 2004). As a native, A.eupatoria stretches from 64°N on the W coast of Norway, throughout all of temperate Europe andadjacent parts of N Africa and Asia (Hultén & Fries 1986, Map 1086; Kurrto et al. 2004, Map3340). Agrimony is also widely introduced and naturalized from the Azores to S Africa, New Zealand and afew eastern states of N America.
Threats
None.
Native, or possibly an archaeophyte, rare, but possibly over-looked and under-recorded. Europeantemperate.
1939; Faris, R.C. & Cole, J.M.; Legakelly, SW of Clones.
September to October.
There are only three recent records in two sites for this erect perennial in Fermanagh, compared withseven pre-1975 records. As the tetrad distribution map indicates, apart from the first record listedabove, the species is confined to the shores of Lower Lough Erne and Lough Melvin. In common with thesmaller, more frequent and widespread A. eupatoria, this species has a history involving herbalmedicine, and has burr fruits that float in water, enabling dispersal by attachment to animals and alsoby flotation. Thus, like A. eupatoria, it may be an ancient introduction, grown for itswide-ranging medicinal properties, since both species were more or less interchangeable in this respect.See the A. eupatoria account for more detail.
There is no obvious reason why this species should have so markedly declined and perhaps it is beingover-looked to some extent by modern botanists. In Ireland generally, however, A. procera appearsmuch less common than A. eupatoria (Agrimony). The two species are alike in their tolerances andusually occupy a very similar range of marginal habitats in woods, grassland, lakeshores and waysides.A. procera shows a greater preference for acidic conditions than A. eupatoria, however,and essentially it is a species of moderate soils, neither too calcareous nor too acidic (D.J. McCosh,in: Preston et al. 2002).
The two Agrimony species also share a similar overall distribution range in B & I, although A.procera is much more scattered and a lot less frequent. While A. procera (previouslynamed A. odorata auct. non (L.) Mill.) is called 'Fragrant Agrimony' on account of its plentifularomatic glandular hairs, both species are fragrant; the glandular difference is not absolute,but quantitative, which again may cause confusion. As mentioned in the A. eupatoria speciesaccount, the only really certain character for distinguishing the two taxa is the ripe fruit: in A.procera the bell-shaped receptacle or hypanthium has short, shallow grooves, not extending toits base and often not reaching the apex, and the outermost bristles of the burr are strongly reflexed;in A. eupatoria the pendulous, bell-shaped fruiting hypanthium is deeply grooved almost all theway from its base to the rim, and the bristles of the burr are spreading, patent or erecto-patent, butdefinitely not reflexed (Stace 2019).
The record details additional to the one above are: Castle Archdale, Lower Lough Erne, 1939, R.Mackechnie; five records by MCM & D – White Island, off Castle Archdale, 1946; Rosskit Island, LoughMelvin, 1948; Garrison, near Lough Melvin, 1948; Rossmore Point, Castle Archdale, 1948; Ely LodgeForest, Lower Lough Erne, 1953; two records by RHN, both at Old Castle Archdale, 1978 & 1990; HareIsland, Lower Lough Erne, 1989, M. Tickner.
Introduction, casual, probably locally extinct. Eurosiberian southern-temperate.
August 2001; Rippey, I. & Northridge, H.J.; Long Island, Lower Lough Erne.
August and September.
This semi-rosette, wintergreen, tap-rooted perennial is accepted as native by botanists in S Ireland(Cen Cat Fl Ir 2; An Irish Flora 7th edition; Flora of Co Dublin), but in NI it haslong been considered a casual introduction (Stewart & Praeger 1895; More et al. 1898). It hasbeen discovered at just two sites in Fermanagh, both in 2001.
The first find listed above was growing with Daucus carota (Wild Carrot) at the edge of an areapreviously cultivated as an island garden. Then, in September 2001, RHN & RSF found several plantsscattered among tall grass on an overgrown roadside bank just outside the town of Lisnaskea. The plantswere growing with numerous Cichorium intybus (Chicory) plants below a recently planted beech andhawthorn hedge along with tree saplings, including poplar and oak. The fact that woody material,probably of imported origin, had recently been planted, suggested to us that Salad Burnet seed arrivedhere as a soil contaminant.
Throughout B & I, Salad Burnet is almost entirely confined to dry, infertile, Carboniferous limestoneor chalk grasslands, or occasionally on base-rich boulder clay. The chief habitats are species-rich,relatively infertile pastures and waste ground, but it can also grow in other situations wherecompetition from more vigorous plants is either completely absent or severely limited, eg in rockcrevices, on limestone pavement, screes, quarries and steep, fairly unstable roadside banks (Grime etal. 1988).
As the English common name indicates, previously Salad Burnet was commonly grown for its salad leaves,which have a pleasant cucumber-like scent and flavour on the palate or in summer drinks including wine.Like S. officinalis L., from Dioscorides onward it was regarded as a vulnerary herb, useful fortreating bruises and other wounds and for staunching blood. 'Burnet' is from Old French 'burnette' or'brunette' meaning 'dark brown' and refers to the colour of the flower heads. S. minor is not sofrequently grown nowadays (Grigson 1987).
Threats
None.
Introduction, neophyte, a rare, naturalised, garden escape.
1952; Mackechnie, R.; Killadeas Td, Lower Lough Erne.
There are three records in just two Fermanagh stations (Killadeas and Tempo Manor) for this low-growing,mat-forming, perennial garden escape. In the grounds of the Manor House Hotel at Killadeas on the shoreof Lower Lough Erne it has persisted for at least 25 years, while the second location is a recentdiscovery along a path-side in estate woodland.
Mention of only one record of this persistent naturalised alien appears in the Revised TypescriptFlora. A form of 'Pirri-pirri-burr', as members of this mainly New Zealand genus are oftencalled, was found at Killadeas on the shore of the lough by the very reliable Scottish botanist, R.Mackechnie who was working locally for a time at Lisnarrick near Kesh. Almost all of his Fermanagh plantrecords appear with very limited site information. Although there is so little detail associated withthis record, Mackechnie's reputation as a field botanist is so well-founded there is no reason whateverto doubt his find.
However, McClintock (1979) has pointed out that prior to the early 1970s, and the publication on thisgroup of plants by Yeo (1973), all published records of this genus were made under the name A.anserinifolia (as used in this instance by Mackechnie), or its synonym A. sanguisorbae.Without voucher specimens, the true identity of these records cannot be known. Many of Mackechnie'srecords, including garden escapes, have herbarium vouchers chiefly located in the Royal Botanic Garden,Edinburgh (E) and the Kelvingrove Museum in Glasgow (GL), but a few also exist inBEL. McClintock listed the Mackechnie record and another by Paul Hackney at the Manor HouseHotel, Killadeas dated 11 September 1975. Subsequently RHN discovered the plant in November 1997,naturalised by a path-side in plantation woodland at Tempo Manor, Upper Lough Erne.
Acaena is a curious genus, native primarily in New Zealand, but with a few species scattered in SAmerica and Polynesia. Species such as A. anserinifolia (J.R. Forst. & G. Forst.) Druce(Bronze Pirri-pirri-bur), A. novae-zelandiae (Pirri-pirri-bur) and A. ovalifolia (which isS American) are commonly grown in gardens as low, ground-cover mats, suitable for planting along withsmaller bulbous plants, on rockeries with alpines, or along pathways and between paving slabs(Grey-Wilson 1989, p. 13). The burr fruits attach themselves to animal coats or human clothing andrecords of them seem to be on the increase in B & I. Beyond the garden wall, the plants tend tooccupy sparsely vegetated, somewhat disturbed ground, either in sun along wayside paths (the plants seemvery well adapted to occasional droughts), or in partial shade on the margins of woods or scrub.
Threats
The plant can be invasive and forms of Acaena have become troublesome pathside weeds in a numberof forest parks in NI.
Native.
1882; Stewart, S.A.; Co Fermanagh.
Most of the Fermanagh Alchemilla records have been ascribed to one of the three species thatfollow, and this species aggregate is simply a 'bin' for the 15 unassigned records which remain, threeof which are very old 'end-of-the' 19th century records. Thehabitats include roadsides, banks and a scree; the scattered stations cover the area of Lower LoughErne and the ground to the west of it, across a total of twelve tetrads.
Native, frequent. European temperate.
1900; Praeger, R.Ll.; Lower Lough Macnean.
Throughout the year.
This apomictic perennial has been frequently recorded in Fermanagh across a total of 115 tetrads, 21.8%of those in the VC. In terms of local frequency, it only just ranks second to A. glabra (SmoothLady's-mantle), but it is considerably more widespread than the latter in tetrad terms. At the sametime, both species are much more frequently found in the western half of the county, in ground lying SWof Lough Erne and, as the tetrad distribution map shows, A. xanthochlora is only thinly scatteredelsewhere in the VC.
A. xanthochlora is recognised by the degree of hairiness of its stems and leaf stalks(petioles), which are more or less clothed with spreading hairs; it is distinguished from the very muchhairier (and less frequent) A. filicaulis subsp. vestita (a HairyLady’s-mantle), by the fact that its leaf blades are hairy only on their underside and the inflorescencestalks are hairless, or almost so (Webb et al. 1996).
A. xanthochlora and A. glabra both prefer damp to constantly moist grassland habitats in awide variety of situations, semi-natural to definitely man-managed. However, A. xanthochlora alsoshows a definite affinity with base-rich or calcareous soils, chiefly in lowland sites, while A.glabra generally favours more upland conditions. The local habitats of A. xanthochlorainclude grassland in open areas within damp woods, wayside verges, stream and ditch banks, lakeshores,cliffs, screes and quarries. It is certainly most frequently found on neutral or lime-rich soils, or onheavier clay.
The New Atlas hectad map indicates that A. xanthochlora is widespread in northernand western Britain, although absent from the more acidic, peaty conditions of much of N & WScotland, including all the more offshore isles. In S & E England, it is very much more rare andscattered, while in Ireland this species again displays a northern and central island distribution,becoming rarer and much more scattered both southwards and westwards in the Republic. When compared withthe earlier BSBI Atlas (1976), the distribution pattern appears quite stable with few losses.
Threats
None.
Native, occasional but possibly under-recorded. European boreal-montane, but also in N America.
1947; MCM & D; laneway near Fardrum Lough.
March to December.
This is the scarcest and most locally occurring of the three Alchemilla species in Fermanagh, witharound half the frequency of records and tetrads of A. glabra (Smooth Lady's-mantle) and A.xanthochlora (Intermediate Lady's-mantle). All three of these apomictic perennials occupy rathersimilar damp, rough grassland habitats, differing ecologically in only minor ways and showing a degreeof overlap. It is possible that all three are under-recorded, since many less experienced observersstill find distinguishing them quite tricky and, therefore, they tend to ignore them or lump themtogether.
A. filicaulis contains two subspecies, but only subsp. vestita occurs in Ireland (Garrard& Streeter 1983; An Irish Flora 7th edition). One old record of subsp.filicaulis does exist for Ben Bulbin mountain in Co Sligo (H28), the current status of which isunknown to RHN and the current author (RSF). The New Atlas hectad map shows that subsp.vestita is the most widespread form of any Lady's-mantle in both B & I occupying suitableshort, rough grassland throughout the whole latitudinal range of both islands, but with a definitewestern tendency in the distribution.
In Fermanagh, this subspecies has been recorded well over a hundred times in a total of 63 tetrads, over11.9% of those in the VC. As the distribution map indicates it is widely scattered across the county,but is most frequent in the west. Eight tetrads have pre-1975 records only, suggesting either a declinein suitable habitats or maybe the identification confusion mentioned above. Local habitats of subsp.vestita include rocky limestone grassland, wayside verges and banks, gravelly watersides, cliffledges, screes and quarries.
Threats
None.
Native, frequent but local. European boreo-temperate, but widely naturalised.
1900; Praeger, R.Ll.; shore of Lough Melvin.
Throughout the year.
This apomictic perennial can sometimes form quite large, robust clonal patches and it is the mostfrequent (just!) of the three widespread Alchemilla species or subspecies that are found in CoFermanagh. As with the other apomictic members of the genus, the pollen is defective so that the seedproduced is vegetative, asexual, arising without any fertilisation taking place. Before the 1960s theseasexual clones were lumped together as A. vulgaris agg., but thanks to of Drs Margaret Bradshawand Max Walters in England, a workable subdivision has been achieved which also appears to besatisfactory in Ireland (Walters 1949b & 1952; Bradshaw 1963 a & b).
As the Latin binomial implies, A. glabra is the least hairy of the three most commonly met membersof the genus Alchemilla. In this plant, only the lowermost couple of stem internodes have a fewscattered hairs and they are appressed, not spreading: in addition the leaves are almost hairless,except towards the ends of the veins on the under-surface, near the leaf margin.
Although in B & I A. glabra is generally found in quite a wide range of damp grassy habitatsat all levels, in Fermanagh, when compared with other members of the genus, it is chiefly a plant ofwetter, upland grassy habitats, including moorland pastures, lakeshores, cliff ledges, in rocky gorges,stabilised screes and especially near waterfalls (where it is often luxuriant). Elsewhere in its B &I range, this species is most commonly found in lowland, ± constantly moist, occasionally flooded grassyplaces, including damper pastures and tall-growing hay meadows, as well as in rough grassland on banksbeside streams and along roadside verges.
Fermanagh occurrence
While the three commoner species of Alchemilla occupy very similar habitats, A. glabra issomewhat less widespread in the VC than A. xanthochlora (IntermediateLady's-mantle), being represented in 97 tetrads, 18.4% of those in the VC, compared with 21.8% for A.xanthochlora. In six of these tetrads there are only pre-1975 records, indicating a slight lossof suitable habitat that probably is associated with drainage and other agricultural grassland'improvement' measures. Both these Alchemilla species are much more common and widespread thanA. filicaulis subsp. vestita (a Hairy Lady's-mantle), which isapproximately half as frequent.
In Fermanagh, A. glabra is widely scattered but, as the tetrad distribution map shows, locally itis more frequent in the W & SW of the county.
British and Irish occurrence
In both B & I, A. glabra has a distinct northern and western distribution, the overall patternbeing very similar to that of A. xanthochlora, although the current species is better representedin N & W Scotland than the latter. In Ireland, A. xanthochlora extends further south in theMidlands than A. glabra (Preston et al. 2002).
Threats
None.
Native, occasional. European temperate, widely naturalised in both hemispheres.
10 July 1985; Corbett, P. & Weyl, R.S.; Carrick Td.
January to November.
The tetrad maps of the two segregates of this species aggregate probably illustrate the distributionalseparation of the two quite accurately, with A. arvensis occurring on the limestones of the westand A. australis (Slender Parsley-piert) more frequent on the sandy soils around Tempo in theeast of the VC.
Fruiting specimens are required to distinguish these two apomictic annual species and in their absence,or when an inexperienced observer or occasional intermediate plants are involved, the plant may berecorded as this aggregate. The small fruit is generally present from about May onwards, although itwill appear somewhat later in the season if the plants originated from spring germinating seed.
In the Fermanagh Flora Database, there are 39 records of this species aggregate, spread across 29 thinlyscattered tetrads, 5.5% of those in the VC. Parsley-piert is typically found on dry, shallow soil onrocky or sandy, often disturbed ground, offering a high proportion of bare surfaces available forcolonisation. Alternatively, it also appears in hollows and cracks in bare rock, on screes and the topsof walls. These ecological conditions are less favourable for perennial species and provide openingswhere small annual species may rapidly colonise in the near-absence of competition.
Native, occasional, but very probably under-recorded. European temperate, but widely naturalised in bothhemispheres.
1881; Stewart, S.A.; Co Fermanagh.
April to December.
Growth form and preferred habitats
This little annual is found on dry or droughted, shallow, warm, open, bare or patchy lowland vegetationover more-or-less disturbed, lime-rich, rocky or sandy soils, or on bare rocks or walls, or gravel orunsurfaced tracks (eg in forestry plantations). In other parts of Ireland, this is a good indicatorspecies for other rarer annuals of similar stress-tolerant, perennial-avoiding, non-competitive habits,such as Erophila glabrescens (Glabrous Whitlowgrass) and Saxifraga tridactylites(Rue-leaved Saxifrage), both of which are extremely rare in Fermanagh.
The curious English common name 'Parsley-piert' alludes to the often rocky nature of the terrain theplant frequently occupies. The name appears to be a garbled version of the French 'perce-pierre', aplant which grows in rocky ground, piercing the rock, from the verb 'percer' meaning 'to pierce' or 'tobore' (Grigson 1974).
Although some observers regard A. arvensis s.s. as being indifferent to soil acidity (FNEI3), other work indicates that it rarely colonises substrates below pH 5.0 and appears to seekout pockets of deeper soil to occupy (Grime et al. 1988).
Flowering reproduction
As with other well-adapted small, therophyte, weedy species, germination can occur either in autumn orspring, which results in a prolonged flowering period stretching from April to October. In parallel withmost members of the genus Alchemilla (to which grouping this species previously belonged), thetiny green flowers are apomictic, setting seed without sexual fusion taking place. Fruiting occurs fromMay onwards and some seed persists in the soil seed bank for five or more years (Thompson et al.1997).
Uses
Previously, A. arvensis was used both in herbal medicine (against bowel inflammation and stones)and as a minor salad vegetable (Grigson 1987).
Fermanagh occurrence
In Fermanagh, A. arvensis has been recorded in 22 tetrads, 4.2% of the total. RHN and the currentauthor (RSF) regard it as occasional, the majority of sites occurring on limestones in the west of theVC. As the tetrad distribution map highlights, eight of the tetrads have only pre-1975 records, which inthis case suggests that it is an under-recorded species.
While characteristically occurring as a small, low-growing annual, A. arvensis can form quitesubstantial patches or even low tussocks in short turf in more fertile, less challenging soils or innear-bare ground conditions, especially if regular mowing or grazing minimises competition from morevigorous species. However, being fairly insignificant in appearance, in Fermanagh Parsley-piert oftenneeds to be actively searched for in suitable habitats, or it can be very easily overlooked.
British and Irish occurrence
In Ireland, A. arvensis s.s. is thinly widespread or local, but rather better represented in the N& S of the country and along the E coast. The distribution in Britain is also widespread, but theplant is more consistently recorded in the SE and thins gradually northwards, becoming more easterly inScotland (Preston et al. 2002).
European and world occurrence
Found in W, S & C Europe reaching S Scandinavia, Latvia and NE Poland and stretching east to Turkeyand Iran. Also present in Ethiopia, the NW African coast and the Macaronesian Isles. Introduced in NAmerica, Chile, S Australia and New Zealand (Hultén & Fries 1986, Map 1155).
Threats
None.
Native, local and apparently uncommon, although very probably under-recorded. European temperate, butwidely naturalised.
1946; MCM & D; bank by Lower Lough Erne, near Gubbaroe Point.
April to December.
Growth form and preferred habitats
A small, much branched, usually pale greyish-green, summer annual therophyte up to 10 cm tall, withdigitately or palmately lobed leaves, this sexually reproducing form of Aphanes is very much morelocal than the apomictic A. arvensis (Parsley-piert). It is greener and more slender than thelatter, flowers from April to October and has even smaller fruits than A. arvensis s.s., showingno constriction between the upper and lower parts, and the sepals are convergent (New Flora of the BI2019, p. 277, Figs 1, 2). It is less common than A. arvensis s.s. and appears to beconfined to short turf, mossy areas on acidic sandy or gravelly soils, or dry rocky ground, eg onroadsides, along tracks and in quarries and sand-pits. It is not as confined to well-drained soils asA. arvensis, but is more definitely a plant of acidic conditions (Garrard & Streeter 1983).In Fermanagh, these conditions are also found locally on or near lakeshores and on river banks.
Fermanagh occurrence
The limited number of records that have accumulated in the Fermanagh Flora Database (18 finds in 14tetrads) are mainly the work of RHN, facts that strongly suggest this rather insignificant-lookinglittle species is under-recorded. Otherwise, as the tetrad map shows, it appears to be very local aroundthe Tempo area, with very few (six or seven) records elsewhere in the county.
Additional to the first record are the following: Poll Beg District, NW of Boho, 11 June 1978, M.J.P.Scannell, DBN; all the remaining records involve RHN – Knockennis, 3 km NE of BrougherMountain, 7 July 1988; Pubble Bridge, Tempo River, 1 October 1988; fen at Feddan Bog, 8 June 1992;fields at Largy Lough, 13 August 1992; sand pit at Pubble Bridge, 11 September 1994 & 20 August1999, with RSF; Drumcreen, Ballinamallard River, 16 April 1995; roadside Ballyreagh, 5 km NW of Tempo,31 December 1995, with HJN; roadside at Tempo, 13 April 1996; Tully, W of Edenmore, 21 June 1997, withRSF; N of Many Burns Bridge, Many Burns River, 3 May 1999; Pubble Forest, 1 December 2001; sandpit atPubble Bridge, 28 August 2004, with RSF; Agnaglack, 20 April 2009, with HJN; Gublusk Bay, Lower LoughErne, 28 February 2010; Killyreagh House near Tamlagh, 10 June 2010.
British and Irish occurrence
The New Atlas map shows the species widespread throughout both islands, but much more thinlyscattered in Ireland, yet with a slightly greater presence in the south and the sunny south-east corner,both areas which attract more visitors and where the local recorders are more energetic than the norm(Preston et al. 2002). Thus the mapped distribution of A. australis across the whole ofIreland suggests probable under-recording in comparison with the situation in Britain.
European and world occurrence
It stretches northwards from a scattered presence in Spain and Portugal through W Europe to S Sweden andeastwards to NE Poland, the Carpathians and the Adriatic. Very local in the Balkans and present only onthe W Mediterranean Isles (Minorca, Mallorca, Sardinia and Sicily), but also recorded on Madeira (Press& Short 1994; Sell & Murrell 2014). Beyond Europe, it occurs in Morocco and Algeria and isintroduced in E & S parts of N America (Hultén & Fries 1986, Map 1156).
Threats
None.
The Fermanagh recording experience has taught RHN and the current author (RSF) that roses are difficultplants to identify for at least two reasons: their breeding system, which involves polyploidy, and theextent of their hybridisation, which regularly produces partially fertile plants and introgression of genesfrom crossing taxa producing intermediate morphological characteristics. Old Floras interpreted thevariation and deviations that these two factors produced by naming very large numbers of separate varieties,but recognising very few hybrids. Little or no serious work has been done on Fermanagh's roses untilrecently, although Meikle and his co-workers collected carefully during their 1945-53 survey and thereferee, Mr N.Y. Sandwith, checked many of their records. The poor treatment of the genus Rosa in the6th edition of Webb's An Irish Flora (1977) is possibly a reflection of the confusion in rosetaxonomy that was partially created by the Klastersky treatment of it in Flora Europaea 2(Tutin et al. 1968). Prof Webb was heavily involved in this important taxonomic exercise, as one ofthe editors and as a major contributor to Flora Europaea. Melville's paper of 1967, which drewattention to rose hybrids and the problems of species classification, together with his treatment of thehybrids published in Stace's Hybridization (1975), may have also played a part, helping to underlinethe difficulties recorders face making correct identifications in this plant group.
Whatever the historical and scientific reasons were, the Irish field guide Flora in use over the major periodwhen the vascular plants of Co Fermanagh was being surveyed was An Irish Flora (1977) and it provideda particularly poor, indecisive treatment of the genus Rosa. This was especially the case in thehandling of Rosa canina s.l. The outcome of this is that most botanical recorders working inFermanagh simply did not bother to try to identify the majority of the roses they encountered.
The publication in 1991 of Stace's New Flora of the BI and of the BSBI Roses Handbook in 1993,appeared at first to offer a better treatment of all our roses. However, for the extremely variable SectionCaninae in particular and, more generally, on account of the very brief and inadequate descriptionsthat these two new standard works provide, recorders have unfortunately been left with many rose specimensthat fall into no clear cut taxon – hybrid or otherwise.
The fact that ripe or late season fruit characters are sometimes required to reach a definitiveidentification also means that field identification of some specimens may be impossible on a single visit,which is usually all that most visiting field workers can manage to a particular site and rose bush.Regrettably, for these reasons, a very large collection of undetermined Fermanagh rose records haveaccumulated, of little value to anyone. Much work remains to be done to sort out the roses in Fermanagh.
Four rose species in B & I reproduce sexually in the normal way: Rosa arvensis, R.multiflora, R. rugosa and R. spinosissima. The remaining species in B & I areoften referred to as 'unbalanced polyploids', most often pentaploids (2n=35) but also tetraploids andhexaploids (2n=28, 42) (Stace et al. 2015). The hybrid progeny of these species inherit more sets ofchromosomes from the female (seed) parent, than from the pollen parent and thus are described as'matroclinal' (or sometimes, 'matriclinal'). Because of this, it is often possible to judge with reasonablecertainty in which direction a cross has occurred, ie which parent was the female. In roses, by convention,the female parent is listed first (R. Melville, in Stace 1975, p. 213). The recognition of directionalhybrids is regularly feasible in first generation crosses, but subsequent secondary crossing and resultantintrogression of genes and associated recognisable characters complicates the picture, giving rise to rosecomplexes of indeterminate ancestry (Roses Handbook, p. 13). Directional and non-directional recordsof rose hybrids are never of this complexity, but in the latter case, the field recorders have beenunable to decide which parent was the female.
Native, common, widespread and locally abundant. European southern-temperate.
1968; unspecified recorder; Marble Arch/Cladagh River Glen NR.
Throughout the year.
The indecisive treatment of roses in An Irish Flora (1977) was especially poor in its handling ofRosa canina s.l. The outcome of this has been that most botanical recorders working in Fermanaghduring the duration of the Flora survey, simply did not bother to try to identify the majority ofthe roses they encountered, resulting in an almost pointless accumulation of over 715 records in thisunspecifically identified grouping (Rosa sp.) scattered across 345 Fermanagh tetrads, 65% ofthose in the VC.
Native, occasional. European temperate.
1882; Stewart, S.A.; Florencecourt.
June to September.
This is a weak-stemmed, sprawling, trailing and clambering rose which tolerates only a little shade andwhich therefore often climbs over other woody species for support and to reach the light. While it growsand can form quite dense sprawling patches on a wide range of soils, it appears to prefer heavyconditions and avoids more acidic substrates (R. Maskew, in: Preston et al. 2002). This rosespecies has stems that display the very strongest colour contrast between their sunlit and shade sides:wine-red on the sunny side and greenish-glaucous on the shaded side. This type of pigmentation is commonenough in other rose species, but the contrast is never as strong as in R. arvensis. The leavesare glabrous (hairless) or with a few sparse hairs on the midrib below and consist of five leaflets. Thestyles are also glabrous and are fused into a long column that sticks out of the flat disc on top of thedeveloping hip like a little peg. Other useful recognition characters are the long, glandular flowerstalks and the almost entire, simple sepals that are unique in B & I in being purplish, notnormally green in colour (Graham & Primavesi 1993, p. 65; Primavesi & Graham, in: Rich &Jermy 1998).
In Fermanagh, R. arvensis has been recorded in 41 tetrads (7.8%), but only 27 of them havepost-1975 records, a statistic which suggests to the current author (RSF) that it is under-recorded. Thelocal habitats it occupies are perfectly typical for the species, ie hedges and the margins ofwoods and scrub, on roadsides, lakeshores and riverbanks. As the tetrad map indicates, Field-rose iswidespread but very unevenly scattered in the VC, with most of the sites being around Upper Lough Erne.
In the rest of Northern Ireland, it is also unevenly scattered, with many sites around Lough Neagh and inCo Down (H38), but comparatively few elsewhere. Field-rose is widespread in the Republic of Ireland, buthere it has a decidedly southern and eastern trend in its occurrence, the distribution apparentlybecoming much more fragmented N of Dublin.
The New Atlas map shows that except in Scotland, where it is rare and very probably introduced, inBritain R. arvensis is widespread – yet with a very pronounced southern distribution. This mainlyEuropean rose very much reaches the northern limit of its distribution in B & I, while on thecontinental mainland it does not extend further N than the Netherlands (Hultén & Fries 1986, Map1072).
R. arvensis is rather variable and in their critical Flora of Great Britain and Ireland2, Sell & Murrell (2014) recognise four varieties of it, distinguished by leaf serration(uni- or biserrate), pedicels (glandular or smooth) and hip shape (globose or ovoid). Of these fourvarieties, var. arvensis occurs throughout the whole range of the species, while the other threevarieties occur occasionally.
No less than ten hybrids are listed for B & I by Sell & Murrell (2014) and nine by Stace etal. (2015). Only one of these is at all widespread, R. ×irregularis Déségl. & Guillon (= R. × verticillacanthaMérat), the other parent being R. canina L. (Sell & Murrell 2014; Stace et al. 2015).Four of the nine or ten hybrids have been found at least once in Ireland (Stace et al. 2015).
Native, occasional. Eurasian temperate, very disjunct in E Europe and Asia and widely naturalised,including in N America.
1884; Barrington, R.M.; Inish Dacharne peninsula, Lower Lough Erne.
May to November.
More commonly, this is a plant of coastal dunes, sandy heaths and sea cliffs around most of B & I. Itis an often low-growing, erect, densely prickled deciduous shrub and in land-locked Fermanagh it occursonly locally, occasional and thinly scattered in rocky limestone terrain. Typical habitats are rockylimestone hills, cliffs, screes and quarries, and in scrub on calcareous lakeshores, including aroundturloughs (vanishing lakes that are occasional or rare in limestone terrain that have no fixed inflowand outflow streams, but fill with rainwater and drain through their base).
Apart from the Carrickreagh area on the lowland shore of Lower Lough Erne, Burnet Rose is never plentifulin Fermanagh. As the distribution map indicates it has been recorded in 22 widely scattered tetrads(4.2%), mainly lying to the west of Lower Lough Erne.
The plant has a long rhizome and it frequently suckers from this, forming clonal clumps of slender aerialstems. In more sheltered sites, including hedgerows and amongst other taller shrubs, it can grow talland densely bushy, sometimes helping to form impenetrable thickets (Roses Handbook). Elsewhere,and more typically in open, exposed, rocky or stony ground, usually with shallower, drier soils, theshrub is invariably low-growing, little more than knee-high (50 cm) and it may then be only sparselybranched. In the latter situation, it is easily overlooked except when in flower. The solitary flowers,up to 4-5 cm in diameter, are produced from late-May to June. They are a beautiful creamy-white,sometimes more or less flushed with pink. Larger flowered double forms of R. spinosissima aresometimes horticulturally planted, but none of the Fermanagh plants display any trace of garden origin.
R. spinosissima suffered major losses at inland sites throughout B & I prior to 1930.Unfortunately the reasons for this were not documented at the time, and observations made duringsubsequent years have cast no additional light on the matter. The main inland sites surviving in Irelandare on outcrops of Carboniferous limestone in the Burren, Co Clare (H9), in Connemara (H16), along theRiver Shannon and in the wider range of Ben Bulbin limestones from Cos Sligo (H28) to Leitrim (H29) andFermanagh (H33) (New Atlas).
R. spinosissima is distinctive but variable and six varieties of it are listed and keyed out bySell & Murrell (2014). Burnet Rose also regularly forms hybrids with six, or possibly seven other B& I wild roses whenever they meet (Sell & Murrell 2014; Stace et al. 2015). Three ofthese crosses have been rarely recorded in Fermanagh.
Native, very rare.
1882; Barrington, R.M.; Muckinish or White Island, Western Lower Lough Erne.
This appears to be a very rare hybrid between a very common and a rather common rose. Barrington's recordwas only tentatively identified by his expert referee, Mr J.G. Baker at K. Barringtonwrote of it, "An imperfect specimen of a rose gathered on the shore of White Island [, which] Mr.Baker was unable to identify.", but he added, "See if R. hibernica, which speciesshould be looked for." (Barrington 1884). The Fermanagh Flora Database listing of this plantsuggests that a voucher exists for this record in DBN, a suggestion RHN and the current author(RSF) greatly doubt.
However, there seems no reason to doubt the other (second) 1947 Fermanagh record determined for Meikleand his co-workers by Mr N.Y. Sandwith, which was described as being, "plentifully, [in a thicket]by the shore of Fardrum Lough, near Drumcose" (Carrothers et al. 1949). The distribution mapin Stace et al. (2015) plots Irish records of R. × hibernicafrom Cos Down (H38), Antrim (H39) and Kerry (H2), but for some reason does not include Fermanagh.
The existence of this rose hybrid was first discovered in 1795 by the Belfast naturalist John Templetonand in the FNEI 3 a handful of old records are listed from Cos Down (H38), Antrim (H39) andLondonderry (H40). In Fermanagh, R. × hibernica should still besought in the Carrickreagh and Drumcose areas, for it is known to be long persistent elsewhere in NEIreland, eg near Holywood, Co Down, where it was first recorded by Templeton in 1795 and persisted thereuntil sometime around 1954 when John Harron saw it, although the site was subsequently destroyed(Kertland & Lambert 1972).
The suckering branches of R. × hibernica bear a mixture of straightnarrow-based prickles and acicles (from R. spinosissima) and curved prickles (from R.canina). The hips are ovoid, somewhat urn-like, or small and poorly developed (Sell &Murrell 2014). In Ireland, the plant is confined to Cos Down (H38), Antrim (H39) and Fermanagh (H33) andalthough scattered records exist in England, N Wales and Scotland, they are very few in number andalmost all pre-date 1950 (Roses Handbook, Map 6; Stace et al. 2015).
Native, very rare.
July 1949; MCM & D; shore of Lough Melvin.
This hybrid was determined by the rose referee Mr N.Y. Sandwith after the plant was collected by Meikleand his co-workers, "on sandy ground by lake shore at Garrison" (Carrothers et al.1950). It is an erect, freely suckering deciduous shrub that is usually accompanied by its parentspecies in hedges and in rocky grassland. Otherwise, and indeed more typically, in coastal parts of B& I it is found in scrub on sand-dunes, a substrate resonating with the sandy, but stony Fermanaghupper lakeshore habitat (T.D. Dines, in: Preston et al. 2002).
R. mollis (Soft Downy-rose) is the most similar of the downy roses to R. spinosissima inits general habit and appearance, and of the five native roses with which R. mollis crosses, thefrequency of R. × sabinii occurrence in Britain is second only tothat of the hybrid with R. canina (R. × molletorumHesl.-Harr.), the latter hybrid being one that is not recorded anywhere in Ireland (Stace et al.2015).
In addition to this solitary Fermanagh record of R. × sabinii, thereare a few historical records from NE Ireland, including one found by R.D. Meikle in 1946 at Lough Cowey,Co Down (H38) (Carrothers et al. 1949). Most of the Co Antrim (H39) records are from the 19thcentury onwards to the 1920s, while for Co Londonderry (H40) four coastal stations are given in theFNEI 3. The editors of the FNEI 3 state that R. ×sabinii is frequently recorded in their area, but they go on to admit that in their view,"probably many of the hybrids found were actually R. spinosissima ×R. sherardii (= R. × involuta Sm.)". The BSBI RosesHandbook lists only two Irish VCs with "non-directional" records of R. × sabinii (in which the maternal parent was not determined),originating from Cos Clare (H9) and Antrim (H39), and it maps neither of these (RosesHandbook, Map 8). Stace et al. (2015) do map these two Irish VCs and also include recordsfrom Connemara (H16 & H26).
Rosa spinosissima × R. rubiginosa (R.× biturigensis Boreau (= R. × cantiana (Wolley-Dod) Wolley-Dod))
Native, very rare.
22 July 1900; Praeger, R.Ll.; shore of Lower Lough Macnean.
Praeger made the solitary Fermanagh record of this vigorous hybrid on a brief visit to Fermanagh inconnection with the fieldwork for his book, Irish Topographical Botany that appeared in print thefollowing year. He described this rose as the only notable plant he found that morning on the lakeshore,which he also described as being, "singularly unproductive" (Praeger 1901a).
The name he gave it was Rosa involuta var. Nicholsonii Crépin, now recognised as the hybridbetween R. spinosissima and R. rubiginosa and previously called R. × nicholsonii (Crépin) Wolley-Dod (R.D. Meikle in the 1975 RevisedTypescript Flora). This hybrid has the very strong, erect branches, suckering habit and mixedarmature with large curving prickles standing out among the general clothing of slender nearly straightprickles and acicles. It also displays the general leaf appearance of R. spinosissima, but likeits other parent, the hybrid can grow much taller, reaching heights of up to 250 cm (Sell & Murrell2014). The leaflets are small and more rounded than in Burnet Rose (R. spinosissima), and whilethey are only sparsely pubescent beneath, they are like other parts of the plant, dotted with thesticky, apple-scented glands of R. rubiginosa (R. Melville, in: Stace 1975; RosesHandbook; Stace et al. 2015).
The Roses Handbook lists non-directional records only of this hybrid from a mere three Irish VCs,W Galway (H16), E Mayo (H26) and Co Down (H38), making no mention of the Fermanagh occurrence. Likewise,the Hybrid Flora of the British Isles (Stace et al. 2015), maps only the records from thesame three Irish VCs, neglecting the solitary old Fermanagh occurrence.
All of the remaining native roses of B & I belong to the Section Caninae of the genus. Theyare mostly pentaploid and three-fifths of their inheritable characters are passed to offspring unchangedin the female line (R. Melville, in: Stace 1975, p. 212). Hybrids are commonly formed; they are oftenpartially fertile and thus are capable of further hybridisation of increasing complexity with each newgeneration. Add to this individual bush longevity and an ability of the flowers to self-pollinate ifcrossing fails and field roses can display a quite bewildering array of characters in a very wide rangeof combinations. It is not at all surprising that with the limited botanical manpower and expertiseavailable, Irish roses remain a neglected area of work.
Native, very rare, or possibly a mis-identification. European temperate.
24 July 1986; Northridge, R.H. & Forbes, R.S.; Gubbaroe Point, Lower Lough Erne.
July to August.
There are just two records of this climbing rose in the Fermanagh Flora Database, both made by RHN andthe current author (RSF). The second record was made by RHN on 17 August 1992 at a rath (a prehistoriccircular fort earthwork) 0.5 km N of Cullen Hill, which itself lies 1 km N of Monea village.Unfortunately, there are no voucher specimens for either of these records, so their presence and statusis very insecure and it requires further investigation.
In B & I, R. stylosa is a rare species of semi-shaded wood margins and hedgerows onwell-drained, lime-rich soils. In Ireland, the New Atlas hectad map indicates that it is confinedto the southern half of the country. Of the eleven VCs listed in the Cen Cat Fl Ir 2, the nearestto Fermanagh is Co Offaly (H18). At the same time, in FNEI 3, Paul Hackney recorded plants withprominently conical discs atop the hip, collected by him near the Carnlough River. The BritishRosa experts Graham and Primavesi saw the vouchers and considered that these were probablyderived from hybridization with R. stylosa. The southerly lie of R. stylosa is alsoreflected in its British distribution; Stace in the New Flora of the BI (1997) describedit as, "B & I south of a line from Co Dublin to E Suffolk, but now rare." Having saidthat, the previously very confused rose taxonomy, stemming from the reproductive biology of the plants,has made the whole genus under-recorded and this is especially the case in Ireland.
R. stylosa was considered an unremarkable Dog-rose of no garden interest by Praeger (1934 f), whor*-established the native status of the species in Ireland, a situation first doubted by Isaac Carrollin Cybele Hibernica (1866), where he suggested records of it on rocks at Myrtle-hill, near Corkcity were perhaps planted. Praeger (1934 f) quoted one of his rose experts working in Ireland, a MrPhillips, who wrote in correspondence to him, "R. stylosa seems to always occur in smallquantity under circ*mstances and in habitats similar to those in which native species such as R.agrestis [= R. sepium] and R. rubiginosa occur in their outlying stations."Praeger (1934 f) also reckoned that its British range, which he described as, "England south of theWash-Severn line", and its Continental range (Germany and France), suggests that this westernspecies should be considered native in Ireland, a position that is now very widely accepted.
An indication of the extent of variation within R. stylosa is given by the fact that Sell &Murrell (2014) recognise and name eight varieties within the species.
Native, common and widespread.
1882; Stewart, S.A.; Co Fermanagh.
May to November.
Recorded 388 times across 231 Fermanagh tetrads (43.8%), this species aggregate appears to be thecommonest rose in Fermanagh, occurring in a wide variety of habitats. However, RHN and the currentauthor (RSF) believe that in reality, like 'Rosa sp.' which has 715 Fermanagh records, this namehas more of the character of a 'rag bag' or 'dust bin' into which all unidentified roses have beencrammed or deposited for simple convenience. We do not want to waste either our time or the reader'swith such a piece of business.
Native, occasional. European temperate, but widely naturalised.
21 June 1985; EHS Habitat Survey Team; marsh to E of Inishroosk Td.
April to October.
Problems and confusion regarding the true identity of early records has meant that pre-1985 finds withoutvouchers have had to be discarded. The remaining Fermanagh records are located in just 19 tetradsscattered across the western half of the VC, as shown in the distribution map. The habitats representedinclude the margins of woods (often by water), scrub, hedges, limestone cliffs and turloughs. Atpresent, R. canina s.s. can only be described as occasional, although RHN and the current author(RSF) are confident that bushes are frequently over-looked, or more often for convenience they arelumped into the Dog-rose species aggregate. On the other hand, in the Fermanagh Flora Database there arealso at least 25 records that can be apportioned to the four subgroups recognised by the RosesHandbook (1993) and New Flora of the BI (1991 & 1997), and for whatever they areworth (a matter RHN and the current author (RSF) do not feel strongly about either way) and few as theyare, they appear in the frequency ranking Pubescentes, Lutetianae, Dumales and Transitoriae. In view ofall this, RHN and RSF believe that a little more work directed at specific rose identification wouldquickly confirm R. canina s.s. to be a widespread and common species in Fermanagh. Having saidthis, the recently published critical account of R. canina published in the Flora of GreatBritain and Ireland by Sell & Murrell (2014) recognises and names no fewer than 38 varietiesof this species, indicating just how much work on roses is required in Fermanagh to meet such achallenging level of variation.
The New Atlas map suggests that R. canina is common and very widespread in both B & I,although even at the hectad scale it is not a ubiquitous species. The hectad map supports Graham &Primavesi's (1993) view that while R. canina s.s. is the commonest rose in most of B &I, it tends to be replaced by R. caesia further north, especially in N Scotland. Since R.canina s.s. prefers well-drained, calcareous soils and tolerates only moderately acidconditions, it is largely excluded from the wet, peaty or silica-rich substrates heavily predominant inN & W Scotland and W & C Ireland.
1947; MCM & D; Fardrum Lough.
Thirteen records can be fitted into this informal group, all or most of which were previously reckoned asbelonging to forms or varieties of R. dumetorum Thuill., which then became R. corymbiferaBorkh. (Revised Typescript Flora). The taxonomy is so confused that RHN and the current author(RSF) are weary of the group, but still want to include the record details for the sake of completenessand since vouchers possibly or probably might exist in K and BEL.
The first eleven records were made by Meikle and co-workers between 1947-50 and were as usual determinedby N.Y. Sandwith. Specimens of the last two records were collected by Paul Hackney in 1974-5 and weredetermined by R. Melville around 1980-2. Apart from the first given above, the sites and dates are asfollows: Crom Castle Estate, 1948; limestone pavement near Crevinish Castle, Gubbaroe Point, Lower LoughErne, 1948; Ballagh crossroads, 1949; Ardunshin Bridge, Colebrooke River, 1949; Glens at Slieve Rushen,1949; Lisgoole, Upper Lough Erne, 1949; Hanging Rock NR, 1949; Muckross, near Kesh, 1950; between LoughDigh and Corrard Peninsula, Upper Lough Erne, 1950; Corrard Peninsula, 1950; old railway track, BigwoodTd, near Boa Island, 18 September 1974, BEL; Killadeas Church, 11 September 1975, BEL.
1948; MCM & D; lane by Lough Melvin near Garrison.
There are just three records of this rose category in the Fermanagh Flora Database and RHN and thecurrent author (RSF) quote them purely for the sake of completeness. The details of the other two are:Inisherk Island, Crom Castle Estate, Upper Lough Erne, 1949, MCM & D (both Meikle records weredetermined by N.Y. Sandwith); wood on shore of Lower Lough Erne at Muckross, near Kesh, 13 September1974, P. Hackney, BEL (originally determined by R. Melville as R. canina f.oxyphylla W.-Dod.).
13 September 1974; Hackney, P.; wood on shore of Lower Lough Erne at Muckross, near Kesh.
Again, three records exist in the Fermanagh Flora Database all of them made in September 1974 by P.Hackney, with vouchers in BEL determined by R. Melville in 1982. The site details are given forthe sake of completeness and they are: hedgerow west of Rushin Point, Upper Lough Macnean; and disusedrailway track, Bigwood Td, near Boa Island, Lower Lough Erne.
1948; MCM & D; scarps around Tullynasrahan Td, in Big Dog Forest.
Two records of this rose group exist, both made by Meikle and co-workers during their survey anddetermined by N.Y. Sandwith. The remaining details of the second record are: Lower Lough Macnean, nearBelcoo, 1952.
Under-recorded, probably quite frequent.
1929; Carrothers, E.N.; Farnaght, SE of Tamlaght.
There are five or possibly six Fermanagh records for this hybrid. The earliest record, detailed above,was originally identified by the Kew expert of the time, Mr Wolley-Dod, as R. canina var.dumalis, which in modern terms can be either this hybrid, or R. caesia subsp. glauca, which is the more common of the two R.caesia subspecies. There may well be a voucher for this record in either BEL or possiblyDBN since it was Praeger (1946) who published the record.
R. canina and R. caesia grow in similar habitats and these are also where the hybrid mostlyoccurs, ie all three are mainly found growing in hedgerows. The hybrid has well-formed fully fertilehips and it also grows on woodland edges and along rides, in scrub, both inland and coastal, on roadsidebanks and on the banks of lakes, rivers and streams (Sell & Murrell 2014). Its fertility allows thehybrid to also colonise numerous kinds of newly available artificial habitats, including abandonedquarries, spoil heaps, dismantled railways and roadside cuttings. This colonising ability also meansthat the occurrence of R. × dumalis extends well beyond thedistribution of the less widespread parent, R. caesia, to a much greater extent than occurs inany other rose hybrid in B & I. This hybrid actually behaves more like as if it was a species and,indeed in mainland Europe, many authors regards it as such, referring to it as R. subcollina(Stace et al. 2015).
Both of these parent roses and their hybrid are attractive to grazing animals and in more heavily grazedlandscapes of the N & W of Britain they are restricted to cliffs and other inaccessible sites thatferal goats and deer cannot reach (Stace et al. 2015).
The details of the other Fermanagh records of this hybrid are: Rosskit Island, Lough Melvin, 1948, MCM& D; lakeshore hedgerow at Muckross, near Kesh, 13 September 1974, P. Hackney, BEL; bay androadside by peninsula W of Rushin, Upper Lough Macnean, 14 September 1974, P. Hackney, BEL; hedgeat Killadeas Church, 11 September 1975, P. Hackney, BEL; Lisrace, 1 km NW of Lacky Bridge, 6September 1997, RHN & RSF.
So little time has been devoted to the discrimination of our local Dog-roses that RHN and the currentauthor (RSF) suspect that, as is the case elsewhere in NI, R. ×dumalis is seriously under-recorded. It may be more frequent than R. caesia and it mighteven be more common than R. canina s.s. (P. Hackney, in: Flora of Northern Ireland website 2005). Having said that, there are only six modern R. × dumalisrecords in the Flora of Co Dublin, despite the primary editor having completed his doctoralresearch on Irish roses (Doogue et al. 1998).
The thinly scattered but widespread distribution of R. × dumalismapped for Ireland in the New Atlas and in the Hybrid Flora of the British Isles(Stace et al. 2015) is at variance with this view, however, and the isolated, comprehensivehectad representation of the plant in Co Kildare (H19), only serves to highlight the very incompletenature of rose recording on the whole island. The British distribution of this hybrid is also patchy andunfortunately, in such cases, it is the occurrence of competent interested recorders that the map reallydisplays.
Native. Mapping error or possible mis-identification.
There is a Fermanagh hectad of date class 1970-86 shown in the New Atlas map for this hybrid(square H05), but RHN and the current author (RSF) do not have any record of it in the Fermanagh FloraDatabase. Probably it is a misplaced hectad error, although this rose hybrid does have a considerablepresence in the eastern Irish Midlands and it is thinly and widely scattered throughout, most frequentat or near coasts elsewhere on the island. On the basis of the published hectad map in the NewAtlas, it appears to be only very rarely reported in NI. The later map in the HybridFlora of the British Isles (Stace et al. 2015) appears to plot a total of elevenhectads for NI, six of them concentrated in Co Down (H38).
Native, possibly under-recorded or an error. European temperate.
1950; MCM & D; Rosskit Island, Lough Melvin.
There are three records from 1950 that seem to best fit into this taxon. They were all made by Meikle andco-workers and as usual they were determined by N.Y. Sandwith. Originally, they were assigned to R.canina var. dumalis and it is conceivable that they might better fit the hybrid R.× dumalis. Without vouchers RHN and the current author (RSF) cannotre-determine them, but they are included in case voucher specimens appear later. The location detailsare: Castle Archdale; Farnaght, SE of Tamlaght; plus the Rosskit Island record above.
There are hybrid Dog-rose records of R. × dumalis from the latter twoFermanagh stations, each with different dates from these finds.
Native, very rare, possibly under-recorded or mis-identified.
1949; MCM & D; glen to the west of Lough Fadd.
This subspecies of Dog-rose is apparently extremely rare throughout Ireland. It might well be so, but incommon with all Irish roses, it is very possibly under-recorded too. The FNEI 3 suggests thatsubsp. caesia is or was recorded, albeit rarely, in all three VCs in NE Ireland (H38-H40). Someof the records listed in FNEI 3 date from 1899 and the most recent date listed is 1955.
The New Atlas map, on the other hand, features just four hectads in all Ireland (no Fermanaghrecords are included). Two of the four Irish stations that the hectad map features are in Co Down (H38)and one in Co Londonderry (H40); all three are pre-1970 in date. The remaining Irish station is locatedfurther SW in Co Sligo (H28) and it dates from the 1987-99 period.
The two Fermanagh records in the Fermanagh Flora Database were made by Meikle and co-workers over 70years ago. Geographically they lie between the extremely few stations for this taxon mapped in theNew Atlas. The Fermanagh sites are remote from one another in the west of the VC. The dates andlocations are: a glen west of Lough Fadd, 1949, determined by N.Y. Sandwith; and Garrison, near LoughMelvin, 1950. Mr N.Y. Sandwith, as ever, followed Wolley-Dod's approach to rose identification andnaming. He identified the plant as R. coriifolia Fr. var. typica W.-Dod. This taxon andname translates as the rarer of the two R. caesia subspecies, subsp. caesia, but ideallyRHN and the current author (RSF) would like to see bushes of the plant in Fermanagh today to prove thatthese earlier botanists were correct in their findings. The most likely alternative would be that theplant in question is another R. caesia cross with R. canina, since the hybrid R. × dumalis appears to be much more common than either or indeed bothforms of R. caesia. Another possibility is a pubescent form of R. canina.
The most recent treatment of R. caesia by Sell & Murrell (2014) recognises the extensivevariation within the species by naming no less than 19 varieties of it.
Rosa caesia subsp. vosagiaca (N.H.F. Desp.) D.H.Kent (R. caesia subsp. glauca (Nyman) G.G.Graham & Primavesi, R. afzeliana Fries) Glaucous Dog-rose
Native, very rare but probably under-recorded.
1948; MCM & D; limestone pavement near Crevinish Castle, Gubbaroe Point, Lower Lough Erne.
July to September.
There are just three records for this rose subspecies in the Fermanagh Flora Database, but it is verypossibly frequent, probably being ignored, overlooked and lumped into R. canina agg. or perhapsmistaken for R. × dumalis, although again there only are a handful ofrecords of the latter in any case. Two of the probable R. caesia subsp. vosagiaca records are by Meikle and his co-workers, thefirst being listed above. The difficulty here is that this castle is several kms NE of Gubbaroe Pointwhere the limestone pavement occurs. However, there is a second record for 'Crevinish Castle', dated1950, so it appears safe to assume that these details refer to two separate stations. The first recordis listed as having been determined by N.Y. Sandwith and a voucher very probably exists somewhere, mostlikely at K where Desmond Meikle worked.
Paul Hackney found the only other Fermanagh record of this taxon in September 1974, the station beinggiven as, "below bridge on abandoned railway embankment near Lough Bresk, near Kesh". Thereare vouchers in BEL (Accession Nos H5546 (two sheets) and H5555). This plant was determined asR. afzeliana Fr. by R. Melville in 1980.
Glaucous Dog-rose can be distinguished from other Dog-roses by its caesious leaflets (ie of a lavendergrey-green colour), being almost hairless and blue-green beneath – certainly far from tomentose. Theleaflets are also often folded along the length of their midrib, which exposes the arching stems morefully to the sun, resulting in an even stronger wine-red stem coloration in comparison with subsp.caesia (Roses Handbook).
There are only five other Irish stations for R. caesia subsp.vosagiaca shown in the map of this subspecies in the BSBI Roses Handbook, two of whichare pre-1950. The stations stretch from N Tyrone (H36) and Co Antrim (H39) in Ulster, southwards to CoWicklow (H20). The New Atlas hectad map improves very slightly on this representation, withfurther stations in coastal Cos Sligo (H28) and Antrim (H39).
Native, occasional, possibly under-recorded or mis-identified to some extent. European temperate.
1881; Stewart, S.A.; Co Fermanagh.
May to October.
In terms of record numbers, this is the third most frequently found rose in Fermanagh with 58 recordsfrom 46 tetrads, identified by many recorders over a hundred year period. Almost all of these records,however, were identified using earlier taxonomy and specifically Prof David Webb's 1977 6th edition ofAn Irish Flora, which on account of the then perceived great difficulty with their separation,lumped R. mollis and R. sherardii along with R. tomentosa into a species aggregatetermed the 'R. tomentosa group'.
Where roses are taken beyond 'Rosa sp.', most of the common roses, certainly from 1974 on, arelisted as either R. canina agg. or as R. tomentosa, meaning the aggregate group. Some ofthe earlier published records give us a little more detail. In the paper detailing his finds around theshores of Lough Erne, R.M. Barrington (1884) wrote, "many forms of this species occur; some comingnear R. mollissima (Willd.). White- and red-flowered varieties seem pretty equallydistributed." Then he goes on to say, "Mr. Baker [the Kew referee, J.G. Baker] considers oneof my specimens var. scabriuscula; the other varieties were not sufficiently perfect with fruit,etc., to identify positively." It is very likely that this represents the hybrid with R.canina.
In the Revised Typescript Flora of Fermanagh R.D. Meikle (1975) explains that most of the roserecords made by him and his friends in the 1946-50 period were checked by the referee Mr N.Y. Sandwith.The seven records made by Meikle and his co-workers for the R. tomentosa agg. are given asfollows: limestone pavement near Crevinish Castle; near Ardunshin; upper part of Donaghmore Glen; nearHanging Rock; Inishrook; by Lough Melvin near Garrison; and Knockmore.
There are 35 further records in the Fermanagh Flora Database dating from 1974-98. Any of these withspecimens would need to be reassessed since, with hindsight provided by the New Atlas, RHN andthe current author (RSF) believe they might now be mainly assigned to R. sherardii or possiblysome might well be the hybrid between R. tomentosa and R. canina (R. × scabriuscula Sm.), as in the case of Barrington's record mentionedabove. All in all, a very unsatisfactory situation exists.
In their very detailed rose treatment, Sell & Murrell (2014) recognise eight varieties within R.tomentosa Sm.
Native, occasional. European temperate.
June 1949; Brenan, J.P.M. & Simpson, N.D.; Belcoo, Lough Macnean.
May to October.
In common with R. tomentosa, the taxonomy of this downy-leaved rose species has been complicatedand confused, with numerous variety and form names being appended, particularly in the Praeger andMeikle eras up until 1950. The Roses Handbook suggests that there appears to be more regionalvariation in R. sherardii than in most British wild roses and Sell & Murrell (2014) supportthis idea. The latter reference lists no less than eight separate varieties of R. sherardii.
The rose treatment in the FNEI 3, the Roses Handbook and the New Atlas all stronglylead RHN and the current author (RSF) to believe that R. sherardii is a much more common rose inB & I than the Fermanagh Flora Database would suggest. Allowing for the suggestion made by Hackneyin the FNEI 3 (p. 194), repeated in the Flora of Co Dublin (p. 232), that a largeproportion of the records listed in the past as the R. tomentosa group may well belong to R.sherardii, this would still leave the latter trailing far behind R. canina in terms offrequency in Fermanagh. Until concerted systematic work on roses is undertaken in the VC using theimproved modern identification texts that are now available, it is not possible to comment further onthis possibility.
As R. sherardii is currently represented in the Fermanagh Flora Database, it is the third mostwidespread hedgerow rose, lying behind R. canina and R. tomentosa with records in 25tetrads (4.7%). Fifteen tetrads contain post-1975 records. R. spinosissima with 36 records ismore frequent than R. sherardii, but it occurs in a mere 22 tetrads! The New Atlas mapshows R. sherardii to be widespread in both B & I, but while there is a shared northernpredominance observable on both islands, in the RoI this rose is more frequently recorded in the eastand the far south, while in Britain the species has a definite western tendency throughout. The apparentIrish distribution of this rose may simply reflect the inescapable facts that on account ofhybridisation, roses are not easy to identify, field botanists are rare and in the Republic most of themreside in and around Dublin. The secondary RoI concentration of rose hectads around Cork city reflectsthe presence of one very active and competent recorder, Mr T. O'Mahony.
Native, very rare.
1938; Brenan, Rev S.A. & Simpson, N.D.; Lower Lough Macnean near Belcoo.
Only three records of this hybrid exist in the Fermanagh Flora Database – one of them post-1975. Allthree are non-directional hybrid records, ie there is no indication given as to which rose wasthe dominant female parent in the cross. Additional to the first record shown above, the other two are:Corrard peninsula, Upper Lough Erne, MCM & D, 1946-54; and, scrubby woodland on limestone soil, neara farm, Finlane, Florencecourt, P. Hackney, 27 August 1976 (vouchers for the latter in BEL andDBN). The Meikle record was originally listed as R. sherardii var. suberecta (Ley)W.-Dod. in the Revised Typescript Flora.
Hybrids between these two roses are to be expected where they overlap in distribution and since R.sherardii is probably the second most common rose species in NI and R. rubiginosa tendsto grow on limestone, it is not peculiar to find the hybrid confined to lime-rich areas in Fermanagh(Roses Handbook). R. × suberecta has well-developed fullyfertile hips, usually with a few acicles present on them (Sell & Murrell 2014; Stace et al.2015). The list of habitats in which this hybrid is found in B & I include open woodland, woodlandmargins, scrub, hedges, roadsides, cliffs, rocky hillsides, disused quarries and coastal dunes (Staceet al. 2015). The seed fertility of the hybrid helps explain its considerable colonising abilityand occupation of a wide range of semi-natural and artificial (man-made) habitats.
The FNEI 3 lists seven mostly old re-determined records from Cos Down (H38), Antrim (H39) andLondonderry (H40), all of them with R. sherardii as the female parent, plus one 1990 record fromGlenarm, Co Antrim which has R. rubiginosa as the female parent. The Flora of Co Dublincontains just one non-directional record from near Balbriggan. The specialist authors of the RosesHandbook list six Irish VCs with non-directional hybrids, these being Mid-Cork (H4), NE Galway(H17) and Cos Leitrim (H29), Louth (H31), Fermanagh (H33) and Antrim (H39).
Native, rare, but very probably under-recorded. European boreo-temperate.
1881-2; Barrington, R.M.; Devenish Island, Lower Lough Erne.
June to October.
R. mollis has ashy-grey leaves, softly hairy (tomentose) on both sides, but the hairs are notglandular. It occurs in woodland, hedges, scrub and rough grassland, on rocky streamsides and rockoutcrops, screes, cliffs, sand dunes, waste ground and by paths and roadsides on a variety ofwell-drained soils (Sell & Murrell 2014). Despite the wide range of habitats and soils that thissuckering, thicket-forming species generally occupies elsewhere in B & I, there are just twelverecords for it in Fermanagh, equally split between pre-1954 and post-1975. It has been found in a totalof ten tetrads, thinly scattered, mainly but not exclusively on limestone. In view of this and the factthat it is described in the FNEI 3 as being, "frequent, especially on the basalt", itis very likely that a more concerted approach to roses would be certain to show that it is seriouslyunder-recorded in Fermanagh. The New Atlas hectad map appears to show R. mollis isdecidedly rare and scattered in Ireland and the current author (RSF) believes it very probably isunder-represented throughout, especially in comparison with Britain.
RHN and the current author believe the very poor taxonomic treatment provided for the Downy-roses in thewidely used An Irish Flora (1977) is the major factor that created this situation. R.mollis is fairly readily distinguished from the other common Downy-roses, R. tomentosaand R. sherardii, and especially so at the very end of the season since the erectsepals persist on the fruit hip until it decays on the bush. Real difficulty with identification doesarise, however, since R. mollis hybridises with five other common roses, including the two justnamed. It crosses most frequently with R. spinosissima, however, and when this occurs thehips of the hybrid, R. × sabinii are red – not black, and they mostresemble smaller versions of R. mollis hips (Roses Handbook).
There is only one local record of this particular hybrid (see separate species account above), butundoubtedly it too is under-recorded, since Fermanagh has plenty of suitable habitats for both of theparent species.
Sell & Murrell (2014) now recognise six varieties of R. mollis, the distribution of themremaining unknown.
Native, occasional. European temperate, but widely naturalised.
1899; West, W.; Florencecourt.
June to October.
All but one of the 21 records for this bright pink-flowered rose are scattered from the shores of LoughErne through the limestones to the W and SW of the two loughs. The typical habitats are roadside verges,hedges, scrub and stony margins of lakes, but it also crops up as an early colonist of under-grazedcalcareous grassland, in the crevices (ie the grykes) in limestone pavement and in neglected areas orwaste ground in quarries and around rock outcrops. In coastal counties of B & I, R.rubiginosa regularly occurs on shingle and other open scrub situations. The densely glandular,sticky, brown hairs on the under-surface of the leaflets give off a distinctive ripe apple scent whenrubbed, allowing most people to easily recognise the 'Sweet-briar', also called 'Eglantine' a name thatis derived from the medieval Latin meaning 'prickly' (Grigson 1974).
RHN and the current author (RSF) regard R. rubiginosa as occasional in Fermanagh, it beingrecorded in 16 tetrads, 3% of those in the VC. There are post-1975 records in ten tetrads. In Irelandoverall, it appears an uncommon, rather thinly scattered species. The New Atlas map shows it ismore frequent in the northern half and along the eastern coast of Ireland, but only very rare or absentelsewhere. The FNEI 3 account of this species aggregate strongly suggests it has declined sincethe 19th century in Cos Down (H38), Antrim (H39) and Londonderry (H40), there being comparatively fewmodern records for it: a total of just twelve stations with post-1970 records are listed. On the otherhand, around the Lough Neagh area, Harron (Flora of Lough Neagh) described this rose as,"frequent but sparingly distributed". A more careful reading of his list of stations, however,shows that he is heavily reliant on observations made at the end of the 19th century by botanists suchas Stewart and Praeger!
In Co Cavan (H30), Reilly (2001) regards R. rubiginosa as "rare", his local Floramentioning only two old records from 1901 and 1938. Eighty kms further south the also recent Flora ofCo Dublin records R. rubiginosa as "occasional" in hedgerows and scrub, theeditors listing a mere eight stations, at two of them the shrub persisting for at least 60 years.
Sell & Murrell (2014) now recognise five varieties within R. rubiginosa. Very little is knownregarding the occurrence and distribution of these recently described varieties, some of which may turnout to be very local.
Introduction, neophyte, a rare garden escape, probably extinct.
1957; MCM & D; Crummer.
An escape from gardens, the two records of this deciduous, suckering shrub with solitary flowers weremade by Meikle and his co-workers at unknown dates between 1946-56. The stations were listed as,"Crummer, near Drumcard crossroads" and, "near Derrybrick, W. of Trasna Island, UpperLough Erne". There are no record cards for these and they first appeared under the name R.cinnamomea L. in the Revised Typescript Flora produced by R.D. Meikle in 1975.This name was used twice by Linnaeus for two different plants in works published in 1753 and 1759. Asthere is no mention of vouchers (although they might possibly exist in K where Meikle worked formany years), RHN and the current author (RSF) do not know whether the garden escape was R.pendulina L. (R. cinnamomea L. 1753 non 1759) (Alpine Rose), or R. majalisHerrm. (R. cinnamomea L. 1759 non 1753) (Cinnamon Rose) (Griffiths 1994).
Reynolds (Cat Alien Pl Ir) lists records of seven alien roses but she does not mention either ofthe species relevant to Fermanagh. Both of these roses are, however, included by Clement & Foster(1994) in their account of alien plants in B & I, and both roses appear to be capable of persistencein hedgerows. This is particularly the case for R. majalis, which originated in N & C Europe.R. pendulina, on the other hand, is a mountain species, native of rocky ground in C & SEurope.
Native, very common and locally abundant. European temperate, but widely planted and naturalised,including in N America.
1882; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
There can only be a few field botanists who have not drawn blood on this very common, rigid branched,sharply thorned deciduous shrub or small tree; indeed, the woody thorns have a reputation for causingseptic wounds. Blackthorn or Sloe is one of the most abundant native shrubs and is particularlyconspicuous in hedges and scrub when in bloom in early spring. In hedges, it is very commonly planted(in recent years sometimes with quicks of non-native, European origin), but in more mixed hedgerows itmay be feral or bird-sown. The shrub grows rapidly to around 4 m and branches and suckers very strongly,forming wickedly dense thickets of dark blackish-brown branches, impenetrable to all grazing mammalslarger than rabbits on account of its long, very hard, woody thorns and its dense, irregular branching(Edlin 1964; Rackham 1980, pp. 351-2). Such thickets can develop in a matter of a few years if ground isneglected and the shrub remains unchecked. Widespread in scrub woodland and hedgerows, it also colonisesrocky ground, fens and sea cliffs, dunes and shingle.
P. spinosa reproduces by bird-dispersed seed and even more commonly by rapidly spreadingvegetative suckers. P. spinosa colonises under-grazed pasture and waste ground, especially in theFermanagh area on limestone. Having said that, it grows on a very wide variety of soils, avoiding onlythe most acidic, wet peat. On undisturbed rocky ground, it forms wickedly dense thickets impenetrable toall mammals larger than rabbits on account of its long, very hard woody thorns which strongly detergrazing, and its dense, irregular branching pattern (Edlin 1964; Rackham 1980, pp. 351-2).
In many areas, native stands are augmented by planted stock in hedges, or as anti-browser nurse plants intree copse plantations, although Crataegus monogyna (Hawthorn) is generally the preferred andmore readily available shrub for these purposes. While useful in such tree plantations, P.spinosa is not long persistent in secondary woodland, eventually being shaded out by tallertrees. Larger Blackthorn trees cannot stand coppicing either, as cutting down to ground level generally,or very often, kills the plant (Rackham 1980).
Fossil history
Blackthorn is an insect pollinated tree or shrub and therefore does not form sufficient pollen for it toappear in the fossil record. However, macrofossil remains do appear as wood or fruit stones.Interestingly, although P. spinosa has been recorded in this way from early interglacials fromthe Cromer Forest Bed series to some stages of the Hoxnian and Ipswichian, it does not feature in thecurrent Flandrian (Littletonian in Ireland) until the Mesolithic period when it frequently appears inarchaeological sites, often as charcoal. It can be deduced from this that P. spinosa was used byearly man as firewood, charcoal and for its edible fruit (eg a large quantity of fruit stones was foundin a Glastonbury Lake Village deposit). In view of the earlier interglacial records, it is probablysensible to regard the absence of P. spinosa fossils from the early and middle Flandrian as moreapparent than real (Godwin 1975, p. 196).
Flowering reproduction
P. spinosa flowers from March to late May, with a peak in April, the blossom opening wellbefore the leaves appear. The 10-15 mm diameter axillary flowers are borne solitary or in twos or smallclusters on short smooth pedicels. They have five pure white petals, around 20 stamens and a singlegreen style and stigma projecting above the stamens. Nectar secreted by a disc on the saucer-likehypanthium is partially concealed at the base of the stamens. The flowers attract a wide range of insectvisitors including bees, overwintering hoverflies, butterflies and early flies, but the very earlyblossom is rather often destroyed by late frosts in March and early April (Proctor & Yeo 1973;Pollard et al. 1974).
Fruit dispersal
The fruit is a fleshy drupe, a 3-layered structure with a central stone containing the single seed of theflower. They ripen in September, some shrivelling and soon dropping, others surviving in good conditionwell into the winter. Song Thrushes take more fruit than Blackbirds; the size of the sloe is close tothe upper swallowing limit of the smaller members of the family. Robins and Starlings merely peck at theflesh rather than ingest and transport the fruit (Snow & Snow 1988, pp. 58-9).
Uses
Although difficult to collect on account of the ferocious thorns, the plum-like sloe fruit is stillsometimes used to make jam, wine, or especially to flavour gin. Rackham (1980) mentions that P.spinosa charcoal and sloe stones have been found fossilised in archaeological sites ofMesolithic and later ages
Sloe juice and the inner bark are very strong astringents and the plant was used by herbalists for,"cooling, binding and drying" bleeding wounds, especially internally, including bowelailments, piles and diarrhoea (Gent 1681). Sloe jelly is recommended for coughs and sore throats, sloewine for colic and sloe gin for kidney problems (Lang 1987; Allen & Hatfield 2004).
"The lower branches of Blackthorn make hard and finely coloured walking or riding sticks with a veryaristocratic knobbliness.", according to Mabey (1977). The knobbliness is thanks to the verynumerous, thickly set, dwarf lateral branches that the thorns represent. The Irish 'shillelagh' is aprime example of the Blackthorn stick, its large knobby head being the base of the shoot where it comesoff the root. As a formidable club weapon, it is euphemistically referred to as, 'an ancient Hiberniantranquilliser'. Up until the 18th century, tannin-rich Blackthorn bark was also used to make ink forwriting. The tannin fixed the ink on the paper and prevented fading (Mabey 1977).
There are also a multitude of superstitions about Blackthorn, including bad luck associated with bringingit (white flowers) into a house when in blossom; it was considered (like Hawthorn) a premonition ofdeath (Vickery 1995).
Fermanagh occurrence
Blackthorn, or Sloe, is the fifth most frequently recorded tree or shrub in Fermanagh, found throughoutin 435 tetrads, over 82.4% of those in the VC.
British and Irish occurrence
Blackthorn is a very widespread and common shrub throughout B & I, absent only from the extreme W ofIreland and NW & Highland Scotland on account of the unsuitable sodden, strongly acidic peaty soilsprevalent in those regions. It is also absent from the outer W & N Scottish isles, except forscattered introductions (New Atlas).
European and world occurrence
P. spinosa is native and widespread throughout Europe except the NE and beyond 60°N in S Sweden.It stretches across the continent from S Spain to Iran and SE Siberia and is present on all or most ofthe islands of the W Mediterranean. It is also introduced in eastern N America and New Zealand (Godwin1975; Hultén & Fries 1986, Map 1178; Sell & Murrell 2014).
Threats
None.
Introduced, archaeophyte, occasional.
13 September 1974; Hackney, P.; Rossigh Bay, Lower Lough Erne.
May to October.
Wild Plum is a medium-sized deciduous tree, 6-10 m tall in good growing conditions, with a rounded crownand a trunk up to 30 cm in diameter. The 'wild' trees occasionally found in hedgerows, on watersides,roadsides and waste ground are usually smaller than this and sucker very readily from the base. Most ormany of the records made of this taxon should probably be transferred to subsp. domestica (WildPlum), but definitively separating this subspecies from subsp. insititia (cultivated Damson andthe wild Bullace) can be difficult. For this reason, RHN and the current author (RSF) have concludedthat plants identified in the Fermanagh field work as P. domestica should remain in thisundifferentiated plum taxon.
P. domestica is an archaeophyte (ie an ancient introduction), a relict of cultivation, previouslyplanted in orchards, gardens or in hedgerows around cottage farms. It often persists long afterhabitation has been abandoned. P. domestica is distinguished from subsp. insititiaby the almost total absence of thorns or spines and by the hairs on the twigs being sparse, rather thandensely pubescent (Webb et al. 1996; Stace 1997). For other comparisons see P.domestica subsp. insititia below.
Fermanagh occurrence
There are Fermanagh records of P. domestica s.l. from a total of 26 tetrads, 4.9% of those in theVC. It is widely but thinly distributed in the lowlands, mainly E of Lough Erne in hedges, roadsidescrub, riverbanks, lakeshores and waste ground.
British and Irish occurrence
The New Atlas hectad map shows P. domestica s.l. is more consistently recorded in NI thanin the RoI and although this shrub is widespread in lowland areas of Ireland, it is much more scarce, orabsent, in many western counties. The distribution in Britain is widespread in the lowlands, becomingvery much rarer and more scattered further northwards into Scotland, with the exception of theGlasgow-Edinburgh conurbations (New Atlas).
Threats
None.
Introduction, archaeophyte, derived from planted material, occasional.
1900; West, W.; Drumskew Td, to the west of Enniskillen.
May to October.
Usually planted, sometimes in quantity, though occasionally perhaps self-sown and apparently wellestablished and long-persistent in situations more remote from habitation. In the Revised TypescriptFermanagh Flora, the eight pre-1970s records of P. domestica were all without exceptionassigned by Meikle to subsp. insititia. There are now records of this subspecies in the FermanaghFlora Database from 38 tetrads, 7.2% of those in the VC. Bearing this in mind, Damson, or rather itswild form, Bullace, is very slightly more frequent than what RHN and the current author (RSF) wouldregard as Wild Plum, P. domestica s.l., but both really are only occasional in Fermanagh,appearing on wood margins and hedgerows along roadsides, riverbanks and lakeshores. They display a verysimilar thinly scattered distribution mainly in the lowland east of the VC.
All forms of P. domestica, including the Plum (subsp. domestica), Bullace (or itscultivated form, Damson) and Greengage (subsp. italica (Borkh.) Gams ex Hegi) are hexaploids ofhybrid origin, the triploid progeny of self-compatible, tetraploid P. spinosa (Blackthorn)(2n=32), crossed with self-incompatible diploid P. cerasifera Ehrh. (Cherry Plum) (2n=16),followed by chromosome doubling to arrive at hexaploid status (Stace et al. 2015). Although thishybrid probably occurred in other places besides the Caucasus region where it is common, the Cherry Plumis not a native species in B & I (Roach 1985, p. 144). Thus, all forms of this hybrid in these islesare undoubtedly ancient cultivated introductions, or subsequently derived forms.
Subsp. insititia is somewhat spiny and it has densely hairy twigs, whereas subsp. domesticahas almost spineless, hairless twigs and a more flattened stone than subsp. insititia, thefruit of which is longer than broad and smaller than a plum stone (Hadfield 1957). However, lifeis rarely or really never this easy when hybrids are involved, since intermediate forms, developed fromfurther generations of crosses abound. The extent of hybridisation between the two or three subspeciesof P. domestica s.l. is so great in places that, in the New Flora of the BI (1997, 2019),Stace reckons character-correlation has partly broken down, often making the subspecies, "hardlydiscernible" in his view.
Threats
None.
Native and planted, locally frequent. European temperate, but widely naturalised.
1882; Barrington, R.M.; roadside between Blaney and Poulaphouca.
January to November.
Growth form and preferred habitats
Native in woodland and occasionally planted and bird-sown in hedges and elsewhere, this lovely whenflowering, medium- or small-sized tree grows on a wide range of reasonably fertile soils, although it isprobably more common in limestone and base-rich clay areas. Well grown specimens can reach 25 m inheight. While found individually, sparingly or occasionally frequent, the species is always local,growing in more open areas, often in linear habitats, such as the margins of woods, in hedges and alongrivers and lakeshores. Apart from these generally somewhat shaded situations, P. avium alsooccurs sporadically as the result of bird sown seed in more unusual colonised situations where it faceslittle competition, including drained cutover bogs, less exposed areas on cliffs and in disused parts ofquarries.
Reproduction
Apart from reproducing freely by seed after obligatory cross-pollination involving bees and otherinsects, P. avium can also spread by suckers. This only occurs to a limited extent, however, andnot with anything like the same vigour displayed by P. spinosa (Blackthorn) and P. cerasus(Dwarf Cherry) (Clapham et al. 1962; An Irish Flora 1996).
The fruits, which ripen early, in late June or July, have a large woody stone contained in a thin butsweet pulp which birds love so much they quickly disappear off the tree! The larger Thrushes,Blackbirds, Crows and Woodpigeons are probably the most significant seed vectors for the species, assmaller birds cannot swallow the relatively large fruit whole, although they may peck at them (Snow& Snow 1988, pp. 55-6).
P. avium is one of the parents of cultivated cherry varieties and seedlings and suckers have beenused as rootstocks for budding or grafts of the latter for many hundreds of years (Roach 1985).
Fermanagh occurrence
On account of the fact that it is regularly transported and introduced by birds, Gean or Wild Cherry,which is also known as Sweet Cherry on account of it being sweeter-than-some related fruits, iswidespread in lowland Fermanagh, being represented in 140 tetrads, 26.5% of those in the VC. However, asin other areas of these islands, it is often difficult or impossible to distinguish planted fromnaturally dispersed trees (D.J. McCosh, in: Preston et al. 2002).
British and Irish occurrence
Ignoring these differences in origin, as with both P. domestica and P. padus, this diploidspecies is more widespread in N, E & SE Ireland than elsewhere on the island. In Britain, it iswidespread throughout, except in NW Scotland and the Highlands (Preston et al. 2002).
Threats
None.
Introduction, archaeophyte, occasional.
1899; Praeger, R.Ll.; Castle Coole estate.
May to November.
The geographical origin of this tetraploid, very acid-fruited (previously very aptly known as SourCherry) shrub or small tree is unknown, but it probably came from somewhere in SW Asia (Edlin 1964).P. cerasus has been known from ancient times and the Morello cherries and numerous cultivatedhybrids are derived from it. Nowadays, it is found wild in B & I, in C & S Europe and intemperate Asia, extending northwards into Scandinavia (Roach 1985).
P. cerasus most typically grows naturalised or planted in hedges, the shrub often suckering freelyalong many metres. Lacking the vigour of P. avium (Wild Cherry), in hedgerows, however, it seldomexceeds about 3 m in height, although it is said to be capable of reaching up to 7-8 m or more in opengrowing conditions (Clapham et al. 1962). The branches tend to be rather blackish in colour, verymuch darker than those of P. avium.
Reproduction and dispersal
Dwarf Cherry reproduces both by seed and vegetatively by suckering from the roots. It is very oftendifficult or impossible, therefore, to discern if it has been planted as hedging, or is of bird-sownorigin, subsequently establishing and spreading itself horizontally. Although isolated bird-sown P.cerasus plants are commonly reported, eg on walls, tall buildings, or in enclosures (Ridley1930), there seems to be very little published information on which bird species are involved. CarrionCrows and the larger thrushes are the most likely transporters (Snow & Snow 1988, p. 166). P.cerasus stones have also been found in the droppings of badgers and foxes (Ridley 1930, pp.352-3).
Fermanagh occurrence
As the tetrad distribution map illustrates, Dwarf Cherry is widely but thinly scattered throughoutlowland Fermanagh, being represented in 80 tetrads, 15.2% of those in the VC. Due to this level ofoccurrence, RHN and the current author (RSF) believe most populations of this cherry in Fermanagh mustrepresent persistent, naturalised hedgerow plantations on the margins of woods, along roadsides, riversand lakeshores. Only the very occasional or fairly rare, more remote lakeshore thicket or solitary cliffshrub outliers can most probably be attributed to bird dispersal (Cat Alien Pl Ir).
British and Irish occurrence
In Ireland, the plant is widespread but most prevalent in the north. In Britain, it is more evenlyscattered throughout England and Wales, becoming much more occasional or rare, eastern and coastal inScotland (Preston et al. 2002). Having said this, confusion with P. avium both past andpresent makes the true distribution, and any changes occurring in it, uncertain to say the least (D.J.McCosh, in: Preston et al. 2002).
Native, local and declining. Eurasian boreo-temperate, but extensively planted and naturalised.
1896; Browne, Rev W.J.; Cooneen, 5 km NNE of Brookeborough.
May to October.
Growth form and preferred habitats
This is a very conspicuous and easily distinguished tree when in flower in May and June due to its long,drooping racemes of 10-40 scented white flowers. When not in flower, P. padus can still bereadily distinguished from P. avium by the possession of tufts of white hairs along the mid-ribof many of its leaves. Bird Cherry occurs as a shrub or small tree, suckering and spreading clonally andseeding itself in moist woods in glens and in hedgerows by streams and rivers. It prefers fairly moistto wet, calcareous or base-rich soils, often associating with Fraxinus excelsior (Ash) andAlnus glutinosa (Alder) (Leather 1996).
As a tree, P. padus is decorative and garden worthy and is planted occasionally in public amenityparkland as well as private gardens. Like other cherry species, it sometimes escapes into the wildthrough birds consuming and transporting the small, black, bitter berries. Noone has attempted todistinguish escaped populations of cultivated forms or varieties in this survey, although it would be aworthwhile exercise for the future.
Fermanagh occurrence
Bird Cherry has been recorded in 46 tetrads in Fermanagh (8.7%), but in nine such squares the onlyrecords are pre-1975, indicating a definite local decline in recent years. It is widely scattered acrossthe county, but most frequent and prominent in the N & E.
British and Irish occurrence
In Britain, as one might expect, P. padus is the most northerly of the three native cherryspecies, although none of them make it to Orkney and Shetland. South of a line between Cardiff and Hull,the New Atlas displays all records except those in E Anglia as introductions; there are also anumber of this designation further north, especially around the Firth of Forth. There are clearlyproblems distinguishing native from introduced populations, a fact that needs to be remembered (Prestonet al. 2002). In Ireland, Bird Cherry is described as being, "frequent in the NW and rareand scattered elsewhere" (An Irish Flora 1996), a view strongly reinforced by the NewAtlas hectad map. In the FNEI 3, Hackney and his co-workers regarded P. padus asscarce in Co Down (H38) and, indeed, they listed no post-1975 records for the county. However, thespecies is more frequent, yet still decidedly local, in the other two VCs covered by the Flora,Cos Antrim (H39) and Londonderry (H40).
European and world occurrence
P. padus is the hardiest Prunus species of them all, having both the most northerly and themost widespread Eurasian distribution. Allowing for some variation within the species, its distributionstretches continuously from the shores of the Arctic to the Japanese Pacific and south to the Pyrenees,Alps, Carpathians and the Himalaya (Hultén & Fries 1986, Map 1179).
Toxicity
All parts of P. padus are poisonous, especially the leaves and berries, which contain cyanogenicglycosides that break down when eaten and produce symptoms of cyanide poisoning (Lang 1987). Humanpoisoning is highly unlikely as the individual berries contain only small quantities of the toxicglycosides and they taste so foul and bitter nobody would consider eating them. One case of animalpoisoning by P. padus has been recorded in Britain in 1996; three cows in Scotland died aftereating the foliage and flowering shoots which had grown through a netting wire fence (Cooper &Johnson 1998). Prunus padus should be removed, therefore, from any hedge used to contain stock.It is referred to in very derogatory terms by Grigson in his book, The Englishman's Flora (1955,1987), describing it as, "a useless little tree, with black and wry fruit".
Folk lore
In NE Scotland the wood of the 'Hackberry', as P. padus is called there (derived from the OldScandinavian 'heggr', giving Hag(berry), Hack(berry) and other similar alternative English common names(see Grigson 1955, 1987) is not used for any purpose, as it is considered a witch's tree (Vickery 1995).
Threats
None.
Introduction, neophyte, cultivated; occasional, but probably somewhat under-recorded.
1983; Kelly, D.L.; Florencecourt estate.
March to December.
Growth form and preferred habitats
In many areas of B & I, this densely branched, laurel-like, evergreen shrub or small tree grows up to10 m in height with 15 cm long, glossy-leaves and small white flowers in erect racemes up to 12 cm inlength, followed by 1 cm cherry-like, glossy red fruits that turn black as they ripen.
Since the middle of the 17th century, when the species was introduced from SE Asia and the Balkans,Cherry Laurel has been commonly planted in demesnes and large gardens, around building for shelter andas cover for game. It has slowly spread beyond these plantations and has become naturalised on woodlandmargins, along paths or in clearings, but occasionally it also appears on lakeshores and islands. In allthese situations, it can form dense thickets and it has the potential to become as invasive and dominantover native species as Rhododendron ponticum (Rhododendron), particularly if the predicted risein global temperatures results in its more frequent seed production.
Toxicity
As with P. padus (Bird Cherry), the leaves and the seeds within the fruits of P.laurocerasus contain cyanogenic glycosides (mainly prunasin and amygdalin). The fleshy part ofthe ripe fruit contains only very small amounts of these glycosides. In cut or crushed material of thespecies, the toxin is hydrolysed to form hydrocyanic acid that can starve animal tissues of oxygen,affecting the central nervous system and even causing death. The glycoside concentration is higher inyoung plants and during the summer (Cooper & Johnson 1998). Even hedge cuttings of the planttransported in a car to the local recycling centre can cause headache, nausea and mental confusion dueto cyanide production by the cut material. This danger remains relatively unknown and needs to be mademore public.
Fermanagh occurrence
Currently, in Fermanagh, there are records of P. laurocerasus from 31 lowland tetrads (5.9%) andin only a meagre handful of cases are the established plants outside estate boundaries or 'over thegarden wall' and thus possibly self-sown and naturalised. Thus, the current author (RSF) suspects thatmany Fermanagh recorders have ignored or dismissed it as being 'a mere introduction', unworthy of noteand, therefore, it is probably under-recorded.
Apart from a very few remote situations in Fermanagh in woods along the base of cliffs, this shrub hasalso been recorded on some wooded islands on Lough Erne, eg Crevinishaughy Island, just offshore of theCastle Archdale estate. Although, very possibly, it might be planted on these islands, escape fromplantation sites may possibly involve not just vegetative propagation by means of the lower brancheslayering, a method of increase and dispersal that this species regularly employs, but transport ofviable seed by birds. Webb (1982) has previously observed this circ*mstance on lake islands inConnemara. Nairn & Crowley (1998) reckoned that this shrub was so invasive and dominant in CoWicklow oakwoods (H20) that it constituted a major threat to native species. In Co Waterford (H6), ithas spread rapidly since first recorded in the 1950s and the majority of records are of self-sown trees(Green 2008). As yet this is certainly not the situation in Fermanagh.
British and Irish occurrence
The New Atlas map shows P. laurocerasus is much more frequent and widely scatteredthroughout B & I than was previously recorded, the calculated change index for the four decadeperiod between the two BSBI flora atlases being as high as +4.7 (Perring & Walters 1976; Prestonet al. 2002). Although widespread in Ireland, the records of P. laurocerasus show adefinite southern and eastern bias in distribution, while in Britain the prevalence is very much moresouthern and western. Having said this, Cherry Laurel is also quite well represented, at least at thehexad level of discrimination, in lowland Scotland as far north as Inverness.
Threats
While in Fermanagh this invasive species is still mainly reproducing vegetatively by layering,occasionally Cherry Laurel sets seed. Were seed to become more frequent due to increasing summertemperatures, this species could quickly become as big a problem in local woodlands and other nativevegetation as Rhododendron ponticum.
Both native and introduced, occasional.
1976; Dawson, Miss N.; Muckross near Kesh.
March to November.
Since the Cultivated Apple, M. domestica and the native Crab Apple, M. sylvestris s.s. areoften difficult or impossible to distinguish on an isolated field visit, there are a total of 44 recordsin the Fermanagh Flora Database of this non-commital taxon. When mapped, they show that almost all 30tetrads in which they lie are on the shores of Upper and Lower Lough Erne. This is easily enoughexplained in that the records were frequently made by the members of the EHS Habitat Survey team duringtheir survey looking for suitable conservation sites on the shores of both parts of Lough Erne in the1986-9 period. There is also a definite habitat bias towards woods and hedgerows near water, includingboth riverbanks and lakeshores, again reflecting the lake survey.
Native, occasional. Eurosiberian southern-temperate, but very widely naturalised.
1739; Henry, Rev W.; Knockninny Hill.
March to November.
Variation and identification problem
Stace (1997) believes that the native Crab Apple is much over-recorded for M. domestica, theCultivated Apple. At first sight the two taxa should not be hard to distinguish, M. sylvestriss.s. having glabrous mature leaves, pedicels and outer calyx surface, while the corresponding parts ofM. domestica are pubescently hairy. Of course Real Life and Nature are never as simple as wecontinually like to believe. Plants of M. domestica are regularly found with leaves hairy only onthe veins beneath, clearly intermediate between the two taxa in this respect (Webb et al. 1996).
Large numbers of cultivars of both these apples exist, representing a vast amount of variation but, ofcourse, particularly so in the Cultivated Apple. The intermediate forms regularly found might behybrids, or as Stace (1997) hints, these two apple species may never have been all thatspecifically distinct. Extensive genetic analysis of old apple cultivars by Robinson et al.(2001) concluded that, "hybridisation, lineage sorting, recent evolution and the spread of'cultigens' and associated weeds by humans have disrupted formerly distinct [apple] taxa andhabitats". The same authors also said, "Morphological characters used to delimit species andsubspecies in series Malus are continuous and overlapping."
Fermanagh occurrence
Crab Apple is one of the very first plants ever recorded in Fermanagh, being written about by Rev WilliamHenry in an early descriptive travel account of Upper Lough Erne published in 1739. This work refers tothe plant as, "crabbes" (in facsimile: Henry et al. 1987). Today, this small tree isuncommon and occasional in lakeshore woods, hedgerows and scrub, especially around the larger lakes, egboth parts of Lough Erne, plus Lough Melvin. It is very scarce and scattered elsewhere in the VC,although there appears to be some linkage with large estates, eg Castle Caldwell, Crom Castle andColebrooke Park, which might suggest a certain degree of planted introduction. The larger estates andthe wooded islands belonging to them are the most likely places for scraps of ancient woodland tosurvive, and it is believed that real, wild Crab Apples are confined to these old shades (Brewis etal. 1996). In Fermanagh, there are records of Crab Apple from 25 tetrads (4.7% of those in theVC), although only 17 of them have post-75 dates.
British and Irish occurrence
Recognising the identification problem and this latter possibility, the New Atlas editors combinedall the apple records into one hectad map as Malus sylvestris s.l. The distribution of thisentity thins northwards in Britain and, rather more patchily, westwards in Ireland. It is a littlereassuring that in Fermanagh there are 44 indeterminate apple records which are also listed as thistaxon.
Threats
None.
Introduction, archaeophyte, occasional.
1902; Praeger, R.Ll.; Co Fermanagh.
February to September.
Found in woods, including those on lakeshores, hedgerows and waysides. Sometimes this small tree iseither deliberately planted in hedges, bird-sown or probably extremely rarely established from seeds indiscarded apple cores. The balance between these three sources is, as usual in such circ*mstances,absolutely impossible to unravel. The cultivated apple is very widely and apparently randomly scatteredthroughout the whole of Fermanagh. As the tetrad map shows, it has been recorded in 66 very widelyscattered tetrads (12.5%), making it more than twice as frequent as the native Malus sylvestris(Crab Apple) in the county although, as noted under that taxon, the two are often very difficult orimpossible to accurately distinguish.
It is important to remember that apples in the 'wild', even if they were derived from seed of perfectly(or nowadays reasonably) edible cultivated forms, often revert and bear small, yellow, sour fruit(New Flora of the BI). For this reason it is suspected that at least some of the records of CrabApple probably belong here.
Threats
None.
Native, common, sometimes planted. Eurasian boreo-temperate.
1882; Stewart, S.A.; Fermanagh.
Throughout the year.
Growth form and preferred habitats
Rowan or Mountain Ash, Sorbus aucuparia is the commonest member of a large genus, occurring notonly native and wild, but also as an ornamental garden subject. While decidedly variable, fivesubspecies being distinguished in Europe (Tutin et al. 1968, 2), thetypical form, subsp. aucuparia, which occurs throughout B & I, grows to a slendermedium-sized tree around 20 m in height, with a narrow crown and erect or somewhat spreading branchesand pinnate leaves. As the English common name suggests, Mountain Ash is a tree particularly associatedwith higher ground in areas of greater rainfall and it is much more local in drier, lowlands areas andin the agricultural midlands of B & I (Rackham 1980; Rich et al. 2010). Typical habitatsinclude deciduous woods in upland areas or in river gorges, open heaths, bracken-covered hillsides,cliffs and rocky hillsides. The small, bright scarlet fruits, although sour and bitter tasting, attractmany bird species so that seeds can be bird-sown anywhere, including on waste ground, along hedges andon roadside and railway banks (Rich et al. 2010). The specific epithet 'aucuparia' is from theLatin 'aucupatorius', meaning 'used for catching birds' ('avis', bird and 'capio', catch), indicatingthat fowlers used the fruit as lures or bait (Gilbert-Carter 1964).
In woodland, Rowan prefers open illumination or semi-shade situations that allow it to freely flower andfruit successfully. It reproduces entirely by seed and cannot survive in the longer term beneath a dark,heavy, deciduous tree canopy like oak. Seed transported into woods by birds and other berry-eatinganimals may allow some seedlings and saplings to develop in shade and they may persist there for longperiods but fail to flower and fruit. This is why S. aucuparia does best on steeper, uplandslopes and glens, where shallow, rocky, leached soils prevent taller dominant tree species forming acomplete or heavy canopy. The more open, partial canopy on slopes allows shorter lived, rapidlydeveloping trees like Rowan and the even more light-demanding Birch to thrive and form a secondarycanopy layer, or even to dominate canopy gaps in deciduous broadleaved woods for short periods. Thespecies is described as a stress-tolerant competitor and it is not long-lived, having a maximum lifespanof about 150 years (Grime et al. 1988).
S. aucuparia tends to avoid calcareous and heavy soils, but it can occur on calcareous groundwhere there is sufficient depth of overlying peat or litter to produce the dry to damp but well-drained,low-nutrient, at least slightly acidic soil conditions that this small to medium-sized tree requires tocompete with its neighbours. It is probably most frequent on soils below pH 5.5, the mean acidity ofmost agricultural soils (Grime et al. 1988).
Rowan can tolerate remarkably exposed conditions on cliffs and steep moorland slopes where it can avoidgrazing and the encroachment of blanket peat. Often, in such situations, it becomes reduced in scale toa small tree, or dwarfed further to shrub size. In other areas of Britain and Ireland, with highermountains than Fermanagh, it grows as isolated shrubby trees on cliffs inaccessible to grazing animalsup to around 650 m in Britain. Occasionally it occurs above this altitude, up to as much as 900 m, asstunted saplings, a feature that makes it the highest growing tree species in these islands (Grime etal. 1988; Raspé et al. 2000).
Flowering reproduction
S. aucuparia is a sexual diploid. The ± flat-topped inflorescence is a densely packed compoundcorymb of between 120-250 small, 8-10 mm diameter, 5-merous flowers each with c 20 stamens. Theinflorescence is heavily scented (Sell & Murrell (2014) describe it as "sweet and sickly")and the nectar is partially concealed, being secreted by the ring of the hypanthium between the stamensand the carpel bases. In warm weather, the stamens spread apart to expose the nectar, but in dullconditions they converge, conceal and protect it (Thomas 2000).
The extremely numerous flowers are self-incompatible, protogynous, outbreeders, pollinated in May andearly June by flies, bees, wasps, moths and a wide variety of other insects (Raspe et al. 2000).Not all flowers set fruit: the proportion doing so in W Scotland can be as low as 16% (Doar 1989, quotedby Rich et al. 2010). The fruit is a 2- to 5-celled, berry-like pome, 6-9 mm in diameter, eachcell containing one or two small, brown seeds. Seed bearing does not begin until the tree is about 15years old (Raspé et al. 2000).
The species is well adapted to the short growing seasons that prevail at both high altitudes and highlatitudes and, in Fermanagh, all the fruits are fully ripe and ready for bird- or mammal-dispersal bymid August or earlier. A good fruit crop is produced virtually every year, but in our mild, dampclimate, uncollected fruits deteriorate fairly rapidly. Seed requires chilling to break deep dormancyand only one of seven publications suggested that seed is more than transient in the soil (Thompsonet al. 1997).
Fermanagh occurrence
Mountain Ash or Rowan forms a significant part of Fermanagh's upland woody flora, being found in a widevariety of habitats throughout the VC at all levels. It is very widespread throughout the countyand has been recorded in 327 tetrads, 61.9% of those in the VC. Although in Fermanagh it is mosttypically found in damp upland acidic woods, rocky glens and scrubby hillsides, whenever it is protectedfrom heavy grazing pressure, Rowan can also feature in areas of calcareous terrain, including lowlandlakeshores and on upland cliffs, eg at Knockmore Hill and at Hanging Rock NR.
British and Irish occurrence
S. aucuparia is common and widespread throughout most of B & I, but becomes more scarce inlowland S & E England and in the Irish Midland plain, probably due to the predominance ofnear-neutral, base-rich soils of these areas, in which the tree is a poor competitor (Grime etal. 1988; Raspé et al. 2000; Preston et al. 2002). In some of its lowland E &C England stations it is probably not native, but derived from planted material (Raspé et al.2000).
European and world occurrence
In Europe, the distribution of S. aucuparia appears limited by hightemperature induced water stress, although another view is that a combination of poor drought tolerance,adaptation to a short early growing season and a cold requirement for bud burst, may be the maindetermining factors (Raspé et al. 2000). S. aucuparia subsp.aucuparia occurs throughout most of Europe from S Scandinavia and S Russia (although not intheir Arctic regions), south to the mountains of C Spain and Portugal, Corsica, Italy, Macedonia and theCaucasus. In the northern regions of Iceland, Scandinavia, Russia and in the mountains of C Europe it isreplaced by subsp. glabrata (Wimmer & Grab.) Cajander.
S. aucuparia is absent only from the Azores, the Faeroes, Spitsbergen, Balearic Isles, Sardinia,Crete and Turkey (Raspé et al. 2000; Hultén & Fries 1986, Map 1158).
Uses
All parts of the Rowan tree are astringent and have been used in leather tanning and for dying black. Thetimber is tough and hard wearing and has been used for all sorts of implements such as staves, poles andhoops for barrels. The berries yield a delicious jelly which is excellent with cold game or wildfowl anda wholesome kind of perry or cider can also be made from them.
The bark, leaves and fruit were all said to have herbal medicinal properties. A decoction of the bark hasbeen given for diarrhoea and coughs and the ripe berries provide an astringent gargle for sore throats,inflamed tonsils and (somehow) as a remedy for haemorrhoids! The leaves were used as a poultice for soreeyes. The anti-scorbutic properties of the berries have also been used as a treatment for scurvy (Grieve1931; Allen & Hatfield 2004).
Folklore
In Irish folklore (and in Scotland and Scandinavia also to some extent), the Rowan has always beenconsidered a tree of formidable magical and protective powers against evil forces, due to its brightflame red berries. An alternative name 'Quicken' refers to its 'quickening' or life-giving powers, whilethe Irish Gaelic name 'Caorthann' derives from the word 'caor', which means both a berry and a blazingflame (Mac Coitir 2003). A Rowan was planted near cottages or a branch was hung inside to preventfire-charming (ie avoiding danger from fire) and used to keep the dead from rising. It was also tied ona hound's collar to increase its speed in the hunt.
A Rowan walking stick offered good protection from the fairies, as did a cross made of the wood or asprig of the plant on the hat. Above all, it was used to protect milk and its products from supernaturalharm, was kept in the byre to protect the cows, and put in the pail and around the churn to ensure thatthe 'profit' in the milk was not stolen. Branches were put over the door lintels of houses, barns,stables and any other farm buildings to ward off witchcraft. In Scotland, flail rods were made ofMountain Ash to keep witches from threshing the corn and stealing the grain. Rowan magic appears innumerous Irish legends and myths also, including The pursuit of Diarmaid and Gráinne (Vickery1995; Mac Coitir 2003).
Threats
None.
Introduced, planted, very rare and widely scattered. European temperate, but widely planted.
4 June 1988; Northridge, R.H.; Brockagh, 8 km E of Belcoo.
May to September.
S. aria is a very variable tree at least 20-25 m tall, with unlobed, ovate leaves with 20-26veins, uniserrate to biserrate marginal teeth, densely white tomentose beneath and with fruits longerthan wide (Rich et al. 2010).
Locally, this is mainly a planted decorative garden species, although the Brockagh record above is of atree growing amongst native vegetation on a dried out, cut-over bog. Two trees that were subsequentlyfound growing on an old wall at Castle Caldwell on 23 August 2003 must have been bird sown. There are atotal of 21 records for this aggregate species in the Fermanagh Flora Database, spread across 15tetrads. They were all found by RHN, sometimes assisted by HJN.
The species prefers well-drained calcareous soils, but it can grow in a wide variety of other more acidicsituations when planted and tended. S. aria is one of three sexually reproducing whitebeams in B& I that fruit and seed themselves around, chiefly using birds as vectors despite the insipid tasteof the berries – at least to the typical human palate. The tree is most conspicuous at the end of Apriland into early May, when the recently opened, white-backed leaves flutter and draw attention to it.S. aria agg. is quite widely planted in gardens and demesnes and more rarely in hedges and alongrivers. It occasionally or rarely naturalises itself, sometimes at considerable distance from thenearest planted tree, again strongly indicative of avian transport and good, competitive, colonisingability.
The species is introduced in Ireland and while the tree is occasional in the wild in NI and apparentlyquite widespread, it is very much more rarely reported in the RoI (NI Vascular Plant Database 2005).
In Britain, S. aria is very much more frequent and widespread than in Ireland, although thedistribution is increasingly patchy further north. It is possibly native in some limestone and chalkareas of the south of England and is certainly well naturalised, especially on calcareous soils. Manynon-native trees probably originate from bird-sown fruits carried from nearby planted specimens, and thetrue extent of native English distribution may never be known (New Atlas; Rich et al.2010).
Native, an Irish endemic, rare.
1984; Northridge, R.H.; scattered on the S shore of Rossergole Peninsula, Castle Caldwell estate, LowerLough Erne.
Identification
S. hibernica is a shrub or small, deciduous tree up to 10 m tall with a broad, open crown and atrunk up to 30 cm in diameter. Leaves are un-lobed, elliptic to obovate and are clearly toothed alongthe whole of their margin except at the extreme base. The leaf teeth are straight, symmetrical andcrowded in the upper half, acuminate, even and almost like a fine comb. The blade usually has 18-20veins and the petiole is about 1.4 cm long. In Ireland, the leaves of S. hibernica most resemblethose of S. aria (Common Whitebeam), although the leaves of the latter are slightly larger andtheir under-surfaces are densely white tomentose, rather than pale greyish-white with silky hairsbeneath as in S. hibernica (Rich et al. 2010; Parnell & Curtis 2012).
Irish and British occurrence
The distribution in Ireland of S. hibernica is widely scattered, trees generally occurring insmall numbers or as solitary individuals, in a wide variety of soils in both semi-natural and artificialhabitat types, chiefly in Midland Ireland (Rich et al. 2005, 2010). Unlike many other species inthe Subgenus Aria in Britain, S. hibernica occurs in a range of habitats on mountains andon lowland rocks and cliffs, rocky lake shores and islets, river gorges, open rock pastures androadsides, hedges, open woods and copses (Rich et al. 2010).
Fermanagh occurrence
Since RHN first found specimens of this endemic apomictic Irish whitebeam on the shoreline and the scrubmargins of plantation woodlands on the Castle Caldwell estate at the W end of Lower Lough Erne, he hassince discovered it in a total of five Fermanagh tetrads, four of them around the same small area. Theoriginal discovery was made quite early on in Robert's recording career when he was naturally unsure asto exactly what he had found. Fortunately, he had the good sense to keep herbarium vouchers ofunrecognised plants. During a fleeting visit to Fermanagh in August 2007, Dr Tim Rich, the BSBISorbus referee, confirmed the identification of the S. hibernica vouchers.
Spurred by the knowledge that this interesting, rare Whitebeam was indeed present in the county, in theautumn of 2007 RHN and HJN searched the scrub shoreline and sections of two peninsulas on the CastleCaldwell estate where the tree had first been encountered. Sixteen trees were discovered in two tetradson Rossergole Peninsula, 14 specimens on the northern shore and two on the southern shore. A solitarytree was located near the eastern end of the adjacent larger Rossmore peninsula on the same estate. Bothpeninsulas were planted up with conifers sometime in the 1930s and the 17 specimens of S.hibernica are growing rather randomly scattered on the strip of shoreline that became exposedwhen the water level in Lower Lough Erne was lowered by drainage works in the 1870s. This strip wasenlarged in the 1950s by further lake drainage. The latter adjustment was a desirable consequence of thedevelopment of the River Erne hydroelectric generation scheme, situated between Belleek, Co Fermanaghand Ballyshannon, Co Donegal and constructed between 1942-57.
The lake level reduction helped drain farmland around both parts of Lough Erne, ground which previouslyhad been subject to unmanageable flooding despite the 19th century drainage efforts. The rocky limestoneground around the new shoreline has since been colonised by dense, almost impenetrable scrub, which madethe task of surveying for S. hibernica extremely difficult. The survey is incomplete for thisreason and perhaps dozens more trees might stand undetected, particularly on the long Rossmore peninsulawhere access is the most difficult. The trees discovered on the Castle Caldwell estate were 4-7 m inheight and some were fruiting sparingly.
Certainly, the main area where this endemic species occurs is in the Irish Midlands, but the 17individuals now known to occur at Castle Caldwell in Co Fermanagh make this the most important site forthis endemic rarity in the whole northern province of Ulster.
Additional Irish Whitebeam specimens were subsequently discovered in Fermanagh at a second site lying onthe S shore of Lower Lough Erne, 1 km E of Hill's Island in 2004. Three specimens were found at the endof a path from a small car park. They appear to have been deliberately planted in a line behind a wirefence. A fourth tree stands isolated further along the path. Apart from accompanying native willows,another definitely introduced tree at this site was Acer campestre (Field Maple). This secondaryS. hibernica site is approximately 2 km south of the Rossmore peninsula, so the possibility ofthe specimens being of bird-sown origin should not be completely ruled out.
A third local station was discovered in 1989 lying about 6 km SSW of Enniskillen. Here, a solitary treeof S. hibernica is growing amongst birch on a cut-over bog at Gransagh, not far from the shore ofUpper Lough Erne. Very probably this tree is another bird-sown individual.
It is very clear from this account that S. hibernica has been previously overlooked in Fermanagh,almost certainly because it has only recently been properly described (Warburg 1957), identification isnot easy and relatively few Irish field workers can confidently recognise it from S. aria (CommonWhitebeam). Thus, although there are many more plants and sites of this endemic recorded than previouslywas the case and there now are records from 31 of the 40 Irish VCs (the main exceptions being in the farSW), at the moment S. hibernica still remains under-recorded throughout Ireland (Rich etal. 2010).
2003; Northridge, R.H. & Northridge, H.J.; lakeshore scrub, Gubdarragh Point, Corrard Peninsula,Upper Lough Erne.
Three shrubs of this endemic apomictic species were found in 2003 growing up to c 180 cm tall and about30 cm apart, approximately 5 m from the edge of the Upper Lough Erne at Gubdarragh Point. The associatedvegetation contained Euonymus europaeus (Spindle), Rhamnus cathartica (Buckthorn) andPrunus spinosa (Blackthorn). On revisiting the site in August 2007, the S. devoniensisindividuals had grown to over 2 m in height and the plants were fruiting. Having taken some samplefruits and leaves for identification RHN and Hannah Northridge met and fell into conversation with theowner of the land, Mr C. Brown of Corrard House. Interest was expressed in the Sorbus growing onGubdarragh Point and Mr Brown indicated he was not surprised that it grew there. He knew there were anumber of similar shrubs growing in hedges on his farm and he believed all were bird-sown from an oldtree that he showed us growing in the field in front of his house (GR H299345). The bole of the tree,about 40 cm in diameter, had been broken off about 2 m from the ground and had produced numerous newsprouts from this point; all the shoots were covered in fruits and leaves.
During a brief visit to Fermanagh in August 2007, Dr Tim Rich, the BSBI Sorbus referee, confirmed theidentification of the vouchers as S. devoniensis. We then visited a bird-sown S.devoniensis growing in a hedge about 100 m to the north-west of the house. This tree was about 6m tall and was covered in fruit. Mr Brown stated that more Sorbus trees grew in the hedges justto the north, on Inishbeg Hill (GR H295348), which is also in Corrard Td, and these were visited by RHNand HJN in August 2007.
He proceeded to tell the history of Corrard House. The King family, who he said originated in Devon, cameto Corrard some time prior to 1739 and stayed until the Brown family bought the property during theSecond World War. The current house, which replaced an earlier one, was built about 1825. The Kingfamily planted many exotic trees on the estate and it was probable that the original S.devoniensis tree in front of the house had been planted some time before 1900.
This is key evidence that S. devoniensis has been planted in the north of Ireland and has beenspread by birds into numerous natural situations.
Introduction, neophyte, very rare and probably extinct or possibly a mis-identification.
1939; Faris, R.C.; roadside, Magheraveely, ENE of Newtownbutler.
S. torminalis is a small to medium-sized, rare or local, uncommon tree, usually with a short,stout trunk and ascending branches forming an oval crown. The large, oval (not pointed), greenish budsare prominent in winter and the leaves are maple-like, but are alternately borne and not palmatelyveined like true maples (Hadfield 1957).
There is just the one old record for Fermanagh, detailed above, which was communicated by Praeger (1939).Charles Faris was an important Co Cavan contributor of records of both plants and insects (Praeger1949). His is the only known Irish record for this medium-sized tree in the wild anywhere in the wholeisland. As was often the case with older records, the site is extremely vague. We would dearly likethere to be a voucher somewhere to support it, but otherwise we can only very tentatively accept it as aFirst County Record.
Elsewhere in Ireland, a large S. torminalis tree has been recorded from demesne land in Co Sligo(H28) (Praeger 1939). These two records published by Praeger (1939) do not feature in the CenCat Fl Ir 2, nor in Cat Alien Pl Ir, which suggests they have not met the necessarycriteria for acceptance. Arboretum specimens of S. torminalis are listed for Glenveagh Gardens,West Co Donegal (H35) and the J.F. Kennedy Park, New Ross, Co Wexford (H12) (Forrest 1985).
S. torminalis is another of the three sexually reproducing Sorbus species that occurs inthese islands, but while it is native at the great majority of its sites in S England and Wales, it isplanted, or spread from introduced populations further north beyond its native range and, of course, itis an extremely rare alien in Ireland (New Atlas). Even in its native area, WildService-tree is nowadays often local and scarce. Previously, the tree was much more common in ancientwoods and old hedges over a wide area of lowland England and Wales as far north as the Lake District.
In these areas, and beyond them, it might also have been cultivated for its "pleasantlyacid-tasting" (Vickery 1995, p. 399) small fruit, which is orange at first and a distinctiveolive-brown colour when frosted and ripe in late October and November (Rackham 1980, p. 358-9; Roper1993; Mabey 1996, pp. 204-5). Although elsewhere described as "rather gritty to taste" (Milner1992), it made prune-flavoured jams, preserves and liqueurs and was certainly also used medicinally.Another account describes the fruit as being, "hard, with rather scant dry flesh, and preciouslittle flavour" (Lang 1987), which suggests the sample collected was not yet ripe and 'bletted' byfrost and subsequent partial decay (Mabey 1996).
In common with Whitebeams, S. torminalis is very variable in its flowering and fruit production.The species is considered to probably have evolved in dry, open woodland and its reproductiveperformance is strongly controlled by local climate (Termena 1972). Most trees come into flower towardsthe end of May, but some are earlier, consistently flowering from the end of April. In the warmer, drierparts of Britain, many trees only fruit every other year and, in more marginal habitats and at the edgeof its geographical range, it fruits even more irregularly than this. Towards the northern edge of itsEnglish distribution in Derbyshire (VC 57), only 5% of S. torminalis seeds proved fertile(Wilmott 1977).
Although seedlings are very seldom recorded, this is considered due to seed and seedling predation bybirds and a range of small mammals and invertebrates, rather than sterility. Birds avidly consume thefruit, but the tree appears only rarely bird-sown, probably on account of the noted levels ofinfertility and seed predation (Roper 1993).
Vegetative reproduction by suckering from the roots is common, especially if the roots in question areonly shallowly buried or have been subjected to any form of damage. Suckers can arise at considerabledistance from the base of the parent tree, even at 110 m from it (Rich et al. 2010).
Being very often solitary, Wild Service-tree is easily over-looked and it may well be under-recorded andespecially so in Ireland where recorders are not on the lookout for it.
Introduction, neophyte, an occasional and locally frequent garden escape.
1947-53; MCM & D; field by railway at Belcoo.
March to October.
This is the most common and widespread member of a species aggregate of twelve difficult to distinguishforms which are all native of N India, the Himalaya and China (species 21-32 in the New Flora of theBI 1997). Webb's An Irish Flora 1977 mentioned only two Cotoneaster species, C.microphyllus and C. simonsii (no naming authorities are given in this field Flora).However, the most recent edition, An Irish Flora 1996, now lists seven Cotoneasters and theCat Alien Pl Ir mentions no less than 19. Apparently, almost all specimens previously identifiedas C. microphyllus Wall. ex Lindl. in B & I actually belong to C. integrifolius and inthe Fermanagh Flora Survey they are treated as being virtually synonymous. There is only one station forC. microphyllus listed in the Cat Alien Pl Ir, from Co Dublin (H21), dating from 1932. RHNand the current author (RSF) are unsure which of the other small-leaved cotoneaster species arerepresented among the Fermanagh records and further work is required if this level of splitting isnecessary.
In Fermanagh, this frequently cultivated, evergreen garden species now has stations in a total of 20tetrads, 3.8% of those in the VC. On Knockninny Hill, where it has become bird-sown and appears to beactively spreading, this very locally occurring sub-shrub is now very well established and persistent.The plant can cover quite large areas of rock outcrop with its low, carpet-like growth. When it is wellestablished and has developed a largish local population, its seedlings may then begin to invadeadjacent stony pastures from the original stand or stronghold, as it is doing for example at RahallanTd, S of Belmore Mountain.
C. integrifolius is more strictly calcicole than other members of the genus and it requires orprefers well-drained soils on sloping ground. Typical local habitats are limestone cliffs, screes,woodland margins, pastures and quite often it appears in quarries. In Fermanagh, it occurs sporadically,principally around the western limestone upland plateau and on widely scattered outlying rock outcrops,including a number of rock quarries. Elsewhere in NI, it is thinly and widely scattered in Cos Antrim(H39) and Londonderry (H40), but it appears much scarcer in the other three VCs in NI.
The New Atlas hectad map shows C. microphyllus agg. is very widely scattered throughoutIreland, with a greater concentration of stations in VCs along the River Shannon and in Connemara. InBritain, it is also spread over a wide range of latitude and shows a definite western tendency.
Threats
Potentially this neophyte could become a nuisance if it invaded a nature reserve, but generally, it istoo infrequent to pose any real threat to native species.
Introduction, neophyte, a very rare garden escape, but certainly under-recorded.
1998; Northridge, R.H.; on wall near Model School, Enniskillen Town.
Bird-sown plants of this commonly grown, sub-shrub, with its distinctive arching, near-horizontal,herring-bone branches and numerous red berries undoubtedly occur in a number of other scattered dry soillocations in both urban and rural Fermanagh habitats. However, the Fermanagh Flora Database containsonly two definite records: the one listed above plus a second found at Tedd Crossroads, Raw Td, 4 km NEof Irvinestown, on 9 September 2010, by RHN and HJN.
Until recently, RHN and the current author (RSF) have always tended to ignore these Cotoneasters,considering them rather uninteresting or even insignificant! We are not alone in this respect. TheCen Cat Fl Ir 2 lists just three VCs in the far SW & S with known records, N Kerry (H2), MidCork (H4) and E Cork (H5).
The Irish picture all changed in 2002 when Reynolds published her Cat Alien Pl Ir for sheenumerated records from a further nine VCs scattered across the island, though not including Fermanagh.The species seeds freely and, apart from self-sown and bird-sown individuals, well-naturalised coloniesmay readily establish from garden waste or from throw-outs. In Co Waterford (H6), C. horizontaliswas first recorded in 1983, and in his county Flora, Green (2008) has mapped around 35 tetradswhere it now has been found. The majority of his sightings (all bird-sown) are on walls, but it alsooccurs on roadside banks, along disused railways and on rock faces.
Introduction, neophyte, an occasional garden escape.
March 1989; Northridge, R.H.; Conagher Upper Td.
March to December.
This Himalayan shrub was not recorded in Fermanagh until 1989. Although it may simply have beenpreviously overlooked, more probably this now quite common bird-sown garden escape really is increasingin the survey area. To date, it has only occurred as single plants in Fermanagh, but it is regularlyfound in natural vegetation and some sites are quite remote from habitation. The Fermanagh FloraDatabase now contains 17 records from ten tetrads, habitats ranging from scrub on cliffs and inquarries, to roadside hedges, a ruined castle and a bridge.
Elsewhere, C. simonsii has generally become rather common and widespread in B & I, at least inthe more gardened lowlands with higher population densities, but also to some extent in their immediaterural surrounds (D.J. McCosh, in: Preston et al. 2002).
First recorded in the wild in Britain in 1910, C. simonsii is now very common and widespread in SWEngland and in W Wales, plus the more populous areas of Scotland. In Ireland, it is rather more thinlyalthough widely scattered, except around the Belfast and south Co Down conurbation, where it has becomelocally very common. It has also been reported as widespread and sometimes becoming well-established inCo Waterford where it was first recorded in 1987. Here again, it occurs in a range of open habitatsincluding along disused railways, rides in conifer plantations, in open broad-leaved woods, on fieldbanks and walls, in hedges, on rock faces, waste ground and in rocky areas (Green 2008).
Garden escape, rare.
4 October 1998; Northridge, R.H.; Derrygore, NW Enniskillen town.
This garden shrub, native of W China, was found on the bank of the River Erne Narrows just as it entersLower Lough Erne – a site where it undoubtedly was bird-sown. C. rehderi is a medium tolarge-sized, deciduous shrub that is very similar to C. bullatus Bois (Hollyberry Cotoneaster)and, without a doubt, it is sometimes confused with it.
It reproduces readily by seed and, in lowland Britain at least, is regularly bird-sown and frequentlybecomes naturalised. The only other Irish record of C. rehderi known to the current author (RSF)is in Cahergal, Co Waterford (H6), where Paul Green discovered it in 2000 (Cat Alien Pl Ir;Green 2008).
Garden escape, very rare.
December 2001; Northridge, R.H.; roadside, Carrickreagh Bay, Lower Lough Erne.
The solitary Fermanagh find detailed above occurred, bird-sown in a roadside situation. From the evidenceof other Irish records, this evergreen 3 m tall shrub tends to grow on rock faces, rubble, walls orroadsides, all generally dry soil situations. It has been recorded in seven other Irish VCs widelyscattered across the island, all finds dating post-1968 (Cat Alien Pl Ir). C. franchetiiis a native of SW China, Tibet and Burma. It is much more frequently and widely recorded in Britain thanin Ireland.
Native, common, widespread and abundant. European temperate, but also widely naturalised.
1864; Dickie, Dr G.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
For centuries, ever since the old Medieval open-field system was abandoned for permanent enclosures, thisextremely familiar and very variable deciduous species with its wickedly sharp, dark crimson, 1-2.5 cmthorns has been by far the most commonly planted woody plant in hedgerows throughout B & I. Youngsaplings of C. monogyna well deserve the English common name 'quicks' for their ability torapidly produce stock-proof enclosures. Apart from linear hedgerows, where it is obviously planted,Hawthorn appears in an extensive range of habitats including as an understorey in more open areas orcanopy gaps in woods, on woodland margins and in scrub on waste or neglected ground.
Seedlings and small saplings also crop up in many additional grazed and ungrazed open situations,especially where competition from vigorous perennials is in some way limited. In terms of its'established strategy' (ie the ecological functional type of the plant once it is established in itssite), C. monogyna is described as a 'stress-tolerant competitor' by Grime et al. (1988,2007) in their C-S-R model. Hawthorn avoids permanent wetlands and aquatic sites, heavily or regularlydisturbed ground and any very acidic, peaty soils of pH below about 4.0 (Grime et al. 1988,2007).
Studies in England suggest C. monogyna, which relies on seed for reproduction, does not regeneratewell in dense woodland shade due to a lack of flower initiation in dark conditions (Pigott 1969; Rackham1980). It is a common colonist of neglected open ground, rapidly forming dense scrub thickets and it canalso persist in relatively low numbers thinly scattered in wood pastures. However, Hawthorn is uncommonin all types of woodland except under the light canopy of Fraxinus excelsior (Ash). SuitableAshwoods are typically found in limestone districts and especially on infertile, north-facing, rockyslopes.
When Hawthorn produces a scrub thicket it develops a very dense, dark canopy of its own which effectivelyexcludes all other woody species. Impenetrable thorny scrub 2-6 m high (occasionally up to 10 m) maythen persist as a permanent ecological landscape feature, perhaps for a century or longer, rather thanforming a temporary seral succession stage in the development of secondary woodland (Rackham 1980, p.353).
Flowering reproduction
Flowers are normally produced from late April or early May to June and seed is set from July toSeptember, although other variant forms flower in two seasons, both spring and either autumn or winter.The timing is reflected in the English common names 'May', 'May-flower' and 'May-bush', which emphasisethe month of peak anthesis (flower opening and functionality) (Grieve 1930; Grigson 1955, 1987). The10-15 mm diameter, 5-merous flowers vary from the usual white to a more local strong pink. They arehermaphrodite, self-incompatible irrespective of colour and are carried in loose corymbose clusters of c15 on pedicels 4-33 mm long. The flowers give off a heavy sugary scent (sometimes tinged with a hint ofurine or stale dung) and partially concealed nectar is secreted by a ring on the cup-shaped hypanthiumjust inside the 15-20 stamens (Proctor & Yeo 1973). The style and stigma are solitary and the fleshyhaw fruit is a single-seeded drupe (hence the specific epithet monogyna).
The flowers attract short tongued insects including predatory flies that feed on pollen and nectar, plusother unspecialised insects that visit and together carry out pollination (Proctor et al. 1996).The deep red, smaller 'haw' fruits of var. nordica are normally plentiful and the fleshy coveringof the stony pip containing the seed provides a very important winter foodstuff for many bird species(Lang 1987), but especially for sedentary species such as Song Thrush and Blackbird. Mistle Thrushes andother migratory members of the thrush family such as Fieldfares and Redwings are more attracted to thelarger drupes of var. splendens (Snow & Snow 1988, pp. 45-8; Sell & Murrell 2014).Unsurprisingly, as a result of this, apart from obvious linear hedgerows laid out by man, it is oftenimpossible to distinguish native bird-sown Hawthorn from planted populations.
The seed inside its woody endocarp cover takes around 18 months to weather, chill and enable germinationto occur. There is no evidence of any persistent soil seed bank (Thompson et al. 1997).
Variation
Evidence of variability exists in every hedge, including flower colour (white or pink), leaf shape andgeneral phenology, to the extent that Tutin et al. (1968) in Flora Europaea 2: pp.75-6 recognised no less than six geographically distinct subspecies in Europe (1968). In Britain, Stace(2019) mentions two subspecies, the common 'wild' form, subsp. nordica Franco as being the normin B & I ("our plants") with fruits 6-9 mm, bright orange-red to purplish-red, and thegarden form subsp. azarella (Griseb.) Franco, from S Europe, that has much more hairy twigs andleaves and often escapes and becomes bird-sown around the countryside. Stace was not entirely convincedabout the value of these subspecies.
Sell & Murrell (2014) go a lot further into the variation, setting out a total of five subspecies(subspp. nordica, monogyna, leiomonogyna, brevispina and azarella), atotal of four varieties (including within subsp. nordica a taxon named var. splendensDruce (drupes 8-11 mm, dull purplish-red or red)) and seven forma in their critical Flora of B & I.
There are many decorative garden varieties in cultivation, Griffiths (1994) listing twelve cultivars andfour subspecies in the RHS Dictionary.
Fermanagh occurrence
C. monogyna is the 27th most frequently recorded vascular plant species in Fermanagh and has beenfound in 492 tetrads, 93.2% of those in the VC, making it our most widely distributed woody plant.Locally among our tree species it ranks second only to Alnus glutinosa (Alder) in terms of recordnumbers.
Due to their planted nature many modern hedgerow Hawthorn quicks are not of native Irish stock,but instead originate from English, Dutch or other European nurseries. This is both a historic fact anda continuing modern practice very greatly to be deplored on conservation grounds (Nelson & Walsh1993).
Tree longevity
C. monogyna plants in hedges are seldom very old, even the oldest ones generally being under ahundred years of age. In Ireland, they probably were planted after large farm holdings were split up andreallocated to previous tenants by the Land Acts at the end of the 19th century and the first fewdecades of the 20th century. Ancient, lone 'Fairy thorns' may be of much larger girth than usual, thesetrees being protected from depredation and development by the force of extremely strong localsuperstition, fear of bad luck and taboo right to this very day! Some of these trees may well reach 300years or so in age, but nobody dares try to sample their trunks for fear of what might happen. Very fewtrunk measurements appear to have been made and, for example, E.C. Nelson quotes only one of 80 cm atAbbey Leix, Co Laois (H14) listed in the Tree Register of the British Isles 1985https://treeregister.org/ (Nelson & Walsh 1993). Many old trees have hollow trunks anyway, sotheir age cannot be scientifically assessed.
Names and folklore
Hawthorn (known as 'sce', 'Whitethorn', 'May tree' or 'May bush'), was protected under the eighth centuryIrish 'Laws of Neighbourhood' as a, "commoner of the wood" (Kelly 1997, p. 380). The folkloreassociated with Hawthorn is vast and involves holy wells, cures and curses of all sorts, magicprotective and fertility powers, tales of luck both good and bad and many references in place namesthroughout B & I (Mac Coitir 2003).
British and Irish occurrence
C. monogyna subsp. nordica is the form that occurs widely andabundantly throughout B & I and also across the lowlands of N & C Europe. Two varieties of itare found throughout the B & I range of the species, var. nordica and var. splendens.Forma schizophylla Beck of subsp. nordica appears to have been grown in nurseries fromcuttings or uniform seed and identical bushes of it occur along long stretches of newly plantedhedgerows and it is the form commonly seen in recently planted English woodland funded by the WoodlandTrust and County Councils (Sell & Murrell 2014).
European and world occurrence
C. monogyna s.l. occurs throughout most of Europe from S Scandinavia to the Mediterranean islandsand coast of N Africa and extending eastwards to Afghanistan. It belongs to the European temperatephytogeographic element and has been introduced to eastern parts of N America (Hultén & Fries 1986,Map 1176).
Crataegus monogyna × C. laevigata (C. × mediaBechst.), Hybrid Hawthorn
Introduced, planted, very rare.
29 May 1988; Hackney, P.; roadside hedge near Belleek.
There are only two records of this fully fertile hybrid in the Fermanagh Flora Database and, as withC. laevigata (Midland Hawthorn) itself, it is very likely under-recorded. Both records were madeon 29 May 1988 by Paul Hackney in hedges near Farrancassidy crossroads on the Belleek-Garrison road and,again, closer to Belleek village. Vouchers exist in BEL. As both the parent species and thishybrid are so very variable, in practice it is advisable to identify as hybrids only plants with bothintermediate leaves and some flowers or fruits with one style or stone and others with more than one(Stace et al. 2015).
Under-recording of the C. laevigata component applies throughout the north of Ireland (including E& W Donegal, H34 & H35) and in England (Hackney & Hackney 1988; Williams 1989; FNEI3). Hybrid Hawthorn occurs occasionally to frequently in planted hedges in NE Ireland,originating from vegetative 'quicks' imported either from England in the second half of the 19thcentury, or more recently from Dutch and E European commercial horticultural sources (Hackney &Hackney 1988). A number of these non-native hawthorn stocks (mostly of decorative garden interest) havebeen commercially imported for gardens and for hedging in NI for quite some time (Hackney & Hackney1988). In conservation and ecological terms, planting native C. monogyna (Hawthorn) stock forhedging would be very much better practice. Very little horticultural material originates in NI and thisthreat to local biodiversity will persist as long as nurseries import hedging 'quicks'.
European studies of the genus concluded that Crataegus species hybridise wherever they overlapgeographically and hybrid swarms are common (J. Franco, in: Tutin et al. 1968). According toByatt (1975), the concept of two separate hawthorn species is largely irrelevant in SE England, sincemost populations exhibit varying degrees of introgressive hybridisation. Sell & Murrell (2014) gofurther and recognise C. media Bechst. as a fully fertile separate species in its own rightcontaining sufficient variation to warrant distinguishing and naming six sub-taxa as forma. Of thesesix, the white-flowered forma media is the most common and the white-flowered, cut-leaved formalaciniata is the next most frequent. Three of the remaining forma have pink or red flowers andthe fourth, forma aurea has white flowers and yellow haws (see the current author's C.laevigata species account).
In C & SE England, a number of isolated chalk scarps may provide the only refuge to the more dominantgenome, ie almost pure C. monogyna. Also in SE England, Williams (1986) referred toC. × media in the plural as, "Hybrid Hawthorns", since theyoccur as fully fertile individuals with a complete range of intermediate characteristics between theparents.
Given time and continuing introduction of foreign Hawthorn 'quicks' the same situation will eventuallyarise in NI. Habitat disturbance and soil type are important with regard to the balance between thethree taxonomic entities: woodland stability and less disruption favours the survival of C.laevigata genes, while heavier, clay soils, rather than lighter ones, also encourages theirbetter survival (Byatt 1975). The general ecological behaviour of C. laevigata indicates that itis much more shade tolerant than C. monogyna and it is characteristic of ancient, undisturbedwoodland, especially those standing on heavier soils (Williams 1986).
European occurrence
C. × media is said to be widespread in Europe, especially in thecentre and north, the common form being forma media. Forms with cut- or dissected-leaves appearin many hedges and planted woods, but it is not known how many of them belong to forma laciniata.The other four more decorative forms are commonly available through the horticultural trade and areprobably widely planted in gardens and amenity areas of streets and estates. However, it is not knownfor certain how many of the decorative forms really are C. × media(Sell & Murrell 2014).
Crataegus laevigata (Poir.) DC., Midland Hawthorn or Woodland Hawthorn
Introduced, neophyte, always planted, but very rare.
1934; Praeger, R.Ll.; Co Fermanagh.
Fermanagh occurrence
There are only two records of this deciduous multi-stemmed shrub or tree up to 12 m tall withshallowly-lobed leaves in the Fermanagh Flora Database (see Identification section below). In theirRevised Typescript Flora, Meikle et al. (1975) wrote, "… the species has notsince been seen in the county [by us], and the claim by Praeger must be questioned". However, RHNand the current author (RSF) feel that a botanist with the reputation of Praeger is extremely unlikelyto report this species, which is unusual in an Irish context, unless he had no doubt of it. RHN's 1997discovery of C. laevigata at the entrance gates of Castle Coole golf course (identificationfurther checked in 2001), proves that Midland Hawthorn is very occasionally or rarely planted inFermanagh, intentionally or otherwise.
Other Irish occurrences
In 1973, two other definite, vouchered specimens of C. laevigata were recorded in the RoI: anisolated tree in a hedgerow at Macroom, W Cork (H3) and the other in a shady, mature beech plantation atKilgobbin, Co Dublin (H21) (Synnott 1978). C. laevigata was also discovered in 1971 in a hedgenear Downpatrick, Co Down (H38) and isolated specimens then were found in Cos Antrim (H39) andLondonderry (H40) (Hackney 1986; Hackney & Hackney 1988).
The New Atlas hectad map indicates that C. laevigata has now been found in a total of nineIrish VCs, including Cos Sligo (H28), Cavan (H30) and Tyrone (H36).
British occurrence
Midland Hawthorn or Woodland Hawthorn as its English common names suggest is regarded as native ofrelatively undisturbed ancient woodland, the margins of woods and, more rarely, in old hedgerows andsloping banks, especially on heavy clay soils (Rackham 1980). The purest shrubs appear to be found inthe centre of ancient woods (Sell & Murrell 2014). However, it must be realised that both C.monogyna and C. laevigata can occur in ancient English woods and they vary from purespecies to a range of intermediate hybrid forms such that pure C. laevigata may be hard to find(Rackham 1980). The conditions for native stands of C. laevigata are best met in C & SEEngland (Huntingdonshire, Cambridge, NW Essex and SW Suffolk, respectively VCs 31, 29, 19 & 26)(Rackham 1980), although published distribution maps, including the New Atlas, indicate it isabsent from the area around the East Anglia Wash (D.J. McCosh, in: Preston et al. 2002). Inancient woods, C. laevigata is usually a member of the underwood (secondary canopy) and it formsmassive stools whereas C. monogyna is more often present as much younger, scrubby, single-stemmedtrees that may have only been coppiced once, if at all (Rackham 1980). This contrast in appearance andbehaviour very probably reflects on the one hand the much greater persistence of C. laevigata inshade and a long history of its coppice management, compared with the rather better dispersal andpioneer colonising ability of C. monogyna in such conditions.
The common, widespread and often abundant presence of C. monogyna across B & I is not entirelyartificial, but certainly human activities of various kinds (ie vegetation clearance, disturbance anddeliberate planting being the most obvious) have given the species innumerable opportunities to extendits distribution range and frequency beyond its previous natural habitat limitations of ash woodlandsand other wood margins on lighter soils (Rackham 1980).
Both beyond and within its supposed English native area, C. laevigata has been frequently andwidely planted and it also occurs bird-sown. Most of the more recent upsurge in recorded finds of itprobably reflect the results of both wider plantation and better recognition. C. laevigata can bevery difficult to distinguish from C. × media, its hybrid with C.monogyna, as the two taxa overlap considerably in many characters. Thus, some of the recordsmapped as C. laevigata in the New Atlas could very easily belong to C. × media Bechst. (Intermediate Hawthorn) (D.J. McCosh, in: Prestonet al. 2002).
Identification
C. laevigata can usually be readily distinguished from C. monogyna by the 2-3 styles in theflower as compared to one in the latter; the deepest sinus between its leaf-lobes reaches less than 2/3way to the midrib; the leaf-lobes are usually three in number and the lateral pair are obtuse in shape.The twigs of C. laevigata are also less stiff and less spiny than those of C. monogyna(Stace 2019). C. laevigata is better able to tolerate shade in woods and hedges than C.monogyna.
Most decorative garden cultivars with pink, red or double flowers belong either to C.laevigata or to the hybrid C. × media, rather than tostraight C. monogyna (Stace 2019).
The distinction of C. laevigata from its fully fertile hybrid with C. monogyna, C. × media Bechst., is much more difficult to achieve and some botanistsbelieve that through frequent back-crossing between the parent species and their hybrid, a high degreeof genetic introgression has occurred within the populations of hawthorn in C & SE England andperhaps beyond in planted areas, creating a continuous swarm of variation linking the three taxainvolved.
The recent critical Flora of B & I by Sell & Murrell (2014) goes further than most othertreatments of these three Crataegus taxa, recognising C. media Bechst. as a fully fertileseparate species intermediate between its parents. The same authors regard C. × media sufficiently variable that they distinguish and name six formswithin it. Most or all of the recognised forms are decorative and appear in garden and parklandcultivation. Some of these forms or cultivars are quite frequently planted, including in NI.
Again, as mentioned earlier, the identification difficulties mean recognition errors are probably veryfrequent, especially between C. laevigata and C. × media(whether regarded as hybrid or species) and published distribution maps are almost certainly unreliable.
European and world occurrence
The distribution of C. laevigata is confined to W & C Europe, stretching from England throughFrance and S Scandinavia to Estonia, Latvia, Lithuania, Poland, Romania, the Balkan peninsula and Italy.It is absent from the Iberian peninsula and all the Mediterranean islands. Most shrubs in the westernpart of these areas and in the lowlands are var. laevigata, but var. palmstruchii is theform found further east and in the mountains. The latter is also sometimes planted in hedgerows, whereits larger fruits and leaves make it more obvious (Hultén & Fries 1986, Map 1174; Sell & Murrell2014).
FABACEAE – Pea family
Native, quite frequent but local. Eurosiberian boreo-temperate, but widely naturalised including in NAmerica and previously in New Zealand.
1882; Stewart, S.A.; Carrick Td.
April to September.
Growth form and preferred habitats
This typically pale-yellow flowered, but extremely variable, polymorphic, creeping, decumbent or erect,usually perennial legume with a stout rootstock has lower leaves pinnate, often with a large terminaland 4-7 pairs of usually smaller lateral leaflets without tendrils. It is a characteristic indicatorspecies of calcareous, shallow, dry, rendzina soils of chalk and limestone grassland vegetation across B& I. As many as 24 subspecies are recognised in W Europe and five of them, plus several varieties,are recognised as occurring in B & I (Cullen 1986; Stace 2019). Only three of the five subspecies inB & I are considered native. Subsp. vulneraria is one of these natives and chiefly frequentsopen, dry, usually calcareous grassland, including on waste ground, and at the coast on sand dunes, inrock crevices and on cliff ledges and maritime heaths. The coastal form of this subspecies is sometimesreferred to as var. langei Jalas. A more mountain-based form of much more restricted distributionin B & I is subsp. lapponica (Hyl.) Jalas.
Fermanagh occurrence
A. vulneraria is a more important member of Fermanagh's plant community than might first appear.It has been recorded in 27 tetrads (5.1% of the total), spread largely across warm, well-drained,sloping limestone pastures, rock outcrops and south-facing screes to the W of Lough Erne.
At Monawilkin, on limestones of the Western Plateau, it is the essential, specific food-plant of themonophagous caterpillars of the only colony of Small Blue butterflies surviving anywhere in NI. A.vulneraria is also the principal larval food-plant of the much more common and very familiarSix-spot Burnet Moth. The only Fermanagh colonies of this insect are again found on the Monawilkin andKnockmore limestones (Thompson & Nelson 2006).
Flowering reproduction
Kidney Vetch flowers from June to September. The inflorescence is a dense cyme of around 18 flowers, upto 4 cm in diameter. Flower-heads are often paired and have twin, leafy, finger-like bracts at theirbase. The twin flower-heads are carried on a single, long stalk or peduncle. Flower colour is veryvariable, but with us they are usually pale yellow, or rarely reddish. Apart from the interestingbutterfly and the day-flying moth mentioned above, various bees visit the plant to feed on pollen andnectar and collect these for their brood. Only larger bees with a long proboscis are sufficiently heavyto effect successful pollination. Nectar theft is common however, short-tongued bees biting through thecalyx and corolla near the base to steal from the flower.
Unlike many other pea-flowers, the two keel petals do not separate to expose the style and stamens whenthe keel is depressed by the weight of a visitor. First the pollen and, subsequently, the stigma aresqueezed like tooth-paste through a tiny opening at the tip of the fused keel petals of the corolla,depositing and later collecting pollen grains transferred on the hairy abdomen of the visiting bees(Proctor & Yeo 1973).
The single- or two-seeded, dry, indehiscent legume fruit pod is surrounded by the persistent inflated,densely white-hairy calyx. Eventually the dry, brown flower-heads become detached and arewind-dispersed, representing a good example of a tumbleweed (Cullen 1986; Knight 1997). Seed germinationis delayed by the indehiscent fruit pod, but probably the species population of the following yearbenefits most from overwintering seedlings initiated in autumn, rather than from slower developingplantlets from spring germination (Knight 1997). Work is required to clarify this matter.
Population biology
Field populations of coastal and inland plants were studied in the Netherlands for several years by Sterk(1975). This showed that seeds germinated predominantly in the spring and seedlings grew on but did notflower until their second spring at the earliest. There was a period of high seedling mortality in bothinland and coastal populations, and plants took longer to reach flowering capability, and fewer plantsmanaged to flower, and each produced fewer flowers in the inland populations compared with coastalplants. The more open, less competitive vegetation in coastal dune grassland conditions allowed moreappreciable annual fluctuations in population density and in biomass production of generativeindividuals than in inland calcicole grassland populations on a loam soil. Coastal plants floweredearlier, more frequently and were more seed productive than the inland populations.
Generally, the seed production per plant in A. vulneraria is low, each flower producing only asingle seed (or very rarely two seeds). In favourable years, coastal plants produced a mean of 20 flowerheads, of which usually 13 were seed-forming, thus averaging 260 seeds per plant. Plants of the inlandpopulation in favourable years formed 4-5 heads, with ten productive flowers, producing around 50 seedsper individual. Predation and parasitism by a number of beetles, moths and gall midges reduced thenumber of viable seed produced per m² (Sterk et al. 1982). However, a small proportion of theinland plants survived longer than coastal plants did, rarely up to four years and very rarely five(Sterk 1975).
In open stands of vegetation along the coast, lack of moisture was a more important cause of mortality,whereas in the denser vegetation inland, biotic factors, especially competition and predation were moreimportant. In unfavourable years, not a single individual attained the generative stage in the coastalpopulations and local survival in such cases depended on dormant seed present in the soil (Sterk 1975).
British and Irish occurrence
In NI, A. vulneraria is predominantly a common coastal plant of rock crevices and sand and shinglehabitats, although a few inland stations on free-draining, base-rich, usually calcareous rocks and soilsdo exist in all six VCs (H33 and H36-H40).
The New Atlas hectad map shows that Kidney Vetch is much better represented at inland sites on thelimestones of the Irish Midlands than it is in inland NI. This is especially so on the bare limestonekarst areas of the Burren in Cos Clare (H9) and NE Galway (H17) and along the banks of the great inlandRiver Shannon.
In Britain, the species is very widespread in the lowlands, again principally occurring either at thecoast or on inland limestone and on other near-neutral, base-rich soils (Preston et al. 2002).
European and world occurrence
A. vulneraria s.l. is exceedingly polymorphic and is native throughout Europe, stretching east tothe Caucasus and south into N Africa and Ethiopia (Hultén & Fries 1986, Map 1252). J. Cullen (1968)in Flora Europaea 2 recognises 24 subspecies and there are numerous additional varietieswithin these. The subspecies each have rather distinct geographical distributions and some are endemicto restricted areas. Subsp. vulneraria is the typical form in B & I and it stretches across NEurope from Ireland to Finland and Latvia. The coastal form in B & I is referred to as var.langei Jalas and it occurs from B & I to the coasts of the Channel Isles, France, Belgium,the Netherlands, Denmark and Germany (Sell & Murrell 2009). A. vulneraria s.l. has beenintroduced to several areas in N America and also to New Zealand (Hultén & Fries 1986, Map 1252).
Names and uses
The names Anthyllis vulneraria ('vulnerary' means 'healing of wounds') and Kidney Vetch mightcertainly point to herbal medicinal usage, but there does seem to be a dearth of information to confirmthis in many English literature sources (eg Grieve 1931; Vickery 1995; Allen & Hatfield 2004). Thelatter says, "Despite a reputation throughout Europe as a vulnerary, the only allegedly folk usetraced of Anthyllis vulneraria has been in the Highlands [of Scotland], where, under two Gaelicnames, it is said to have been used in the past for cuts and bruises." Darwin (1996), The Scot'sHerbal, confirms this, briefly stating, "Kidney vetch was used to treat wounds and also wasadded to hay." Vickery (1995) mentions that on the Channel Islands the leaves were used to checkbleeding from wounds.
The most detailed herbal medicinal account found is Launert (1981) who describes the plant as an ancientremedy for eruptions of the skin, slow-healing wounds, minor burns, cuts and bruises. A decoction couldeither be added to bathwater or applied as a compress, or sometimes the bruised fresh herb was appliedto the affected area. A mild infusion could be taken against constipation or drunk as a spring tonic.The dried flower-heads might also be used as a substitute for real tea.
The genus name 'Anthyllis' is from the Greek 'anthos' meaning 'a flower' and 'ioulos' meaning 'down' or'downy', an obvious reference to the downy calyx (Johnson & Smith 1946).
Grigson (1955, 1987) lists no less than 20 English common names for the plant, many of them shared withor borrowed from Lotus corniculatus. The much smaller pods of A. vulneraria make theapplication of some of the names involving feet, fingers, thumbs or claws much less than appropriate inthis instance, although the reference might be to the paired, finger-like bracts beneath the twininflorescences.
Threats
'Improvement' or disturbance of limestone pastures.
Native, common. Euroasian southern-temperate, but widely naturalised.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This as a low, sprawling, sometimes carpet-forming, usually almost hairless, bright-yellow floweredperennial legume with solid spreading stems, round in cross-section at their base but square higher up.It is readily distinguished from the most commonly found related species, L. uliginosus Schkuhr(= L. pedunculatus Cav.) Greater Bird's-foot-trefoil, by its solid rather than hollow stem andits calyx teeth that are appressed to the corolla in bud, rather than spreading (Webb et al.1996).
L. corniculatus is common and locally abundant throughout B & I in unshaded situations onshort pastures and on dry to wet grassy places on a very wide variety of soils. These include thoseformed over both limestone and more acidic rocks, of sandy, clayey or peaty texture, and including veryinfertile situations on damp upland moors and wetter Sphagnum bogs. Common Bird's-foot-trefoilproduces its most vigorous growth on moist, heavy, fertile, lime-rich soils of pH around 6.5 (Smith1975; Turkington & Franko 1980).
Particularly in drier grassland conditions or in rock crevices, the plant develops a deep, robust taprootup to 100 cm long, with numerous laterals. This penetrative organ, which becomes thick and woody withage, allows the species to survive drought for several weeks. In wetter soils and those subjected towater-logging for 1-3 months or more, the 'rootstock' is short and thick, with the dense fibrous lateralroots being retained near the soil surface. The rootstock is a long-lived organ enabling perennation,since the aerial parts generally die down and become quite limited or entirely absent in winter.Die-back to a small rosette in the late autumn allows the new buds for next spring's growth the shelterand protection of the dead stems (Jones & Turkington 1986). The extensive root system explains thefrequency of the plant on dry rocky soils, eg screes, quarries and cliff ledges. When occurring incoastal or inland sandy soils, the deep roots help stabilise the substrate, enabling the frequentlyobserved large colonies to develop.
In meadows and pastures, the plant is favoured by regular cutting or grazing which reduces competitionfrom taller, more vigorous grasses and herbs, although L. corniculatus is capable of survivingand being drawn up to some extent with inherently taller species (Jones & Turkington 1986). CommonBird's-foot-trefoil has a weak stem and growth with a companion grass in pasture leys provides supportand prevents lodging. It does particularly well with slow-establishing grasses such as Phleumpratense (Timothy), which offer less intense competition at the seedling stage. Seedlings ofCommon Bird's-foot-trefoil are small, non-aggressive and slow-growing and are thus easily out competed.While it is light-demanding, L. corniculatus is otherwise decidedly stress-tolerant and ratheruncompetitive. Grime et al. (1988) categorised its established strategy as intermediate between astress-tolerator and C-S-R, a balance which really is non-committal on this issue.
However, when growing in infertile, species-rich grasslands or heaths, this legume, being self-sufficientin nitrogen and with its extensive root and shoot system, can manage to co-exist with up to 35 otherplant species (Atkinson 1973). In pastures in N America, once established, it can withstand heavygrazing and does not cause bloat in grazing animals, as do most other legumes (Smith 1975). The feedingvalue of Common Bird's-foot-trefoil in silage or hay has been reported as equal or surpassing that ofalfalfa and other good legume hays when fed to dairy cattle and sheep (Marten & Jordan 1979).
While it is always absent from shade in woodland or scrub, once established it can persist in more open,marginal or disturbed areas of these habitats, as well as in lower-growing, woody heath vegetation.
Flowering reproduction
Regeneration of this long-lived herb is primarily by seed, produced in large numbers aftercross-pollination by bees. Nectar can only be obtained by long-tongued insects of sufficient weight andstrength to open the pea flower and cling on to it. Unlike many other pea-flowers, the fused keel petalsdo not open, and a peculiar 'pumping arrangement' is used to effect cross-pollination. Before the floweris visited, pollen is discharged into the conical tip of the fused keel petals and held there by theelongated club-shaped filaments of the five outer stamens. When the bee lands on the flower, its weightpressing on the wing and keel petals acts like a piston on the contained stamen filaments, which pressforward and squirt the somewhat sticky pollen like a worm-like pasty mass through a very small hole orslit in the apex of the keel, to form a small coil of pollen that sticks to the hairy lower part of theinsect's abdomen. Later on, the stigma emerges from the tip of the fused keel petals and rubs on theabdomen of a subsequent bee visitor (Hutchinson 1972; Proctor & Yeo 1973). Pollen-collecting beesare said to be more effective in pollination than nectar-collecting bees (Bader & Anderson 1962).Self-pollination can occur, but experimental evidence showed it only occurred in 53% of flowers, ofwhich 46% set 0.01-0.49 seeds per flower and 7% set just 0.50-1.0 seeds per flower (Seaney 1964). Inother studies, wild collected British flowers were found to be totally self-sterile (Ramnani 1979).
Flowering begins in late May, reaches a peak in June and July and continues into August. In somesituations, a second flush of flowering takes place from mid-August to mid-October. The flowers, 15 mmlong, are borne in cymose clusters of 2-6 (usually five) on erect peduncles 3-10 cm tall. The corollavaries in colour from bright yellow to coppery or brick red and the standard petal is often streakedwith red lines. Frequent flower-shedding, and the failure of many flowers to develop seed pods, setlimits to seed-production capacity of the species (Bader & Anderson 1962).
Fruits develop from late June onwards. The cylindrical ripe pods, five or six together, are brown toblackish, 15-30 mm long and each is tipped with the persistent style, making them together look like theclaws of a bird's foot and hence the English common name. Each pod contains between 1-20 seeds (mean5.9) that are explosively released from the ripe legume when it suddenly splits along two sutures andtwists violently to eject the seeds. Measured as seeding distance from the parent plant, the meandispersal distance was 0.24 m and the maximum achieved was 1.75 m. Seed per plant can vary enormouslyfrom zero to over 18,000 depending upon site and season (Jones & Turkington 1986). Seeds germinatein the spring, but a portion of them may persist in the soil seed bank for at least five years (Thompsonet al. 1997). The seeds are sufficiently 'hard' (ie water impermeable) that they can pass throughthe digestive tract of sheep, cattle and birds unharmed (Grant 1967).
L. corniculatus is one of the few legumes that can produce adventitious buds and shoots from theroot when the crown is removed or the root sectioned (eg by cutting, grazing or heavy trampling) (Smith1975). Thus plants in some habitats may be capable of displaying a limited degree of vegetativereproduction through rooting of older, horizontally spreading stems (Jones & Turkington 1986).
Variation
L. corniculatus is tetraploid (2n=24) and is the most variable species in the entire genus ofaround a hundred species, displaying both great morphological and physiological variation, some of whichis genetic. The species is important as a legume in grass seed mixtures in agronomy in the making ofsilage and hay, and many named varieties have been developed by plant breeders, particularly in the USAand Canada since the 1950s (Turkington & Franko 1980). The cultivars tend to have an upright habit,unlike the more usual prostrate native forms of the species.
In Britain, the species gives rise to up to eight named varieties (Ellis et al. 1977; Jones &Turkington 1986; Sell & Murrell 2009), although Stace (2019) ignores them all except the distinctiveintroduced, weedy, roadside var. sativus Hyl. which is up to 50 cm tall, has large leaflets andsmall flowers.
Toxicity
The leaves and flowers of some L. corniculatus genotypes are cyanogenic at levels sufficient todiscourage species of molluscs and other selective herbivores (Ellis et al. 1977). However,numerous insects and their larvae have adapted to cope with the toxins. Jones & Turkington (1986)published a four page list of insects that feed on the plant.
Fermanagh occurrence
L. corniculatus has been recorded in 244 tetrads, 46.2% of those in the VC. As the Fermanaghdistribution tetrad map indicates, however, it is unevenly scattered in the VC, being most prevalent indamp grasslands around Lough Erne and on the limestones of the upland Western Plateau.
British and Irish occurrence
The New Atlas hectad map shows that this is one of the most omnipresent legumes on both islands,there being very few hectads where it has not been recorded.
European and world occurrence
The species is native and occurs almost throughout Europe to 71°N and to an altitude of 3050 m in theSwiss Alps, but it is absent from Spitsbergen. It has been introduced to Iceland. It is also indigenousin coastal N Africa and in parts of Asia. As it has been quite widely used as a forage crop for cattlein Europe since the 18th century and in N America since the 1950s, and also commonly occurs as a seedimpurity of White Clover (Trifolium repens) and low-grade grass seed, it has been widely spreadby man well beyond its native range. In Canada, it is classified as an adventive, ie introduced butimperfectly naturalised (Turkington & Franko 1980). It is introduced in E Africa (eg Ethiopia andKenya), very widely in N America and to a much lesser extent in S America, plus in S Australia and NewZealand (Turkington & Franko 1980; Hultén & Fries 1986, Map 1249).
Names and folklore
Although there is very little folklore associated with Common Bird's-foot-trefoil (Vickery 1995) and itdoes not appear to have much in the way of herbal uses either (Allen & Hatfield 2004), it does havea huge number of folk names attached to it. Grigson (1955, 1987) list over 70 names and even he finds itdifficult to explain this fact. In ancient herbal use, it was considered another vulnernarywound-healing herb, but the only more modern use traced by Allen & Hatfield (2004) was as an eyewashon South Uist in the Outer Hebrides. It does not feature at all in Grieve's (1931) comprehensive herbal.In Irish folklore, in the south of the country, children called it 'No blame' and regarded it as worthcollecting and bringing to school as a protection from the teacher's cane (Vickery 1995).
Threats
None.
Native, frequent. European temperate, but widely naturalised.
1884; Barrington, R.M.; Ely Lodge Forest.
May to December.
Taxonomic uncertainty
There appears to be some taxonomic debate regarding the correct name and status of GreaterBird's-foot-trefoil at present (Feb 2021). Stace (2019) uses the name above and gives L.uliginosus Schkuhr as its synonym. However, he mentions that there is some disagreement as towhether L. uliginosus and L. pedunculatus represent one or two species. If they areseparate species, then Stace takes the view that the B & I taxon is L. uliginosus. Sell &Murrell (2009) also regard the B & I plant as L. uliginosus and consider L.pedunculatus Cav. as a distinct, separate species that does not occur in B & I.
Growth form and preferred habitats
This very distinctive, 60 cm or up to 1 m tall, climbing or rather, scrambling on the support of otherplants, yellow or orange pea-flower grows in moist grassland and waterside habitats in late summer. Itis easily distinguished from the quite similar, but lower-growing, more compact, earlier flowering L.corniculatus (Common Bird's-foot-trefoil) by its hollow stems and calyx teeth recurved on theflower bud (Stace 2019). It is also more shallow-rooted than the latter and has a slender rootstock thatproduces numerous vegetatively spreading stolons that also help set it apart from L. corniculatus(Clapham et al. 1987).
Until a recently introduced name change, the previously applied Latin species epithet 'uliginosus' (=growing in marshy places) and its English common name Marsh Bird's-foot-trefoil reminded us that thisnitrogen-fixing legume occurs in a wide variety of damp to wet grassy habitats, sometimes indeedoccurring in stands of great abundance thanks to its vegetative spreading ability (Gilbert-Carter 1964).Locally in Fermanagh these wetland situations include rushy fields, margins of swampy fens orperiodically flooded lakeshores, marshy grassy waysides, on damp river banks and in damper hollows inwaste ground, a sand pit and on drier parts of bogs, including cut-over bogs.
In terms of soil preferences, L. pedunculatus occupies wet ground of similar nutrient status andrange of textures to L. corniculatus, but unlike the latter, as in the Burren in Co Clare (H9),it appears to be completely absent from the most lime-rich soils and, with rare exceptions (ie a coupleof cut-over bogs and along parts of the Finn Floods river), it is confined to moderately acid to neutralconditions between pH 4.5 and 6.5 (Webb & Scannell 1983; Sinker et al. 1985). It is largelyabsent from soils below pH 4.5 (Grime et al. 1988).
While common in periodically or seasonally flooded ground, L. pedunculatus is completely absentfrom permanently submerged, fully aquatic habitats and also from heavily disturbed sites. L.pedunculatus is the only common legume of wetlands in the B & I flora. The scarcity oflegumes on wet ground probably reflects the conflicting demands for a limited oxygen supply during theprocess of nodule nitrogen-fixation and also in its role in the detoxification of anaerobic conditionsaround the root in wet soils (Sprent 1984).
On the other hand, Large- or Greater-Bird's-foot-trefoil is much more tolerant of half-shade than L.corniculatus and, since it grows taller and has a larger leaf area, it is the strongercompetitor of these two legumes, growing with associated grasses and wet habitat herbs such asFestuca pratensis (Meadow Fescue), Holcus mollis (Creeping Soft-grass), Lychnisflos-cuculi (Ragged-Robin) and Succisa pratensis (Devil's-bit Scabious). The establishedstrategy of L. pedunculatus is described as intermediate between competitor and C-S-R by Grimeet al. (1988), on this measure one step up from the more stress-tolerant, less competitive L.corniculatus. It appears to colonise and grow best in relatively infertile, sub-optimalconditions where the growth of potential dominants is restricted.
Flowering reproduction
Flowering in L. pedunculatus is very similar to that of L. corniculatus. Anthesis occursbetween June and August, the deep yellow, hermaphrodite, pea flowers being borne 5-15 in number, inaxillary cymose heads on long slender peduncles. Like L. corniculatus, the 10-20 mm flowers areprotandrous, bee-pollinated and self-incompatible. The weight of the insect visitor is the importantfactor required to open the flower and squeeze or pump the previously released somewhat sticky pollenout of the tip of the fused keel petals onto the bee's hairy abdomen, rather like a coil of toothpaste.The stigma also protrudes through the keel tip whenever the flower enters the female phase, and aninsect revisit then achieves cross-pollination (Proctor & Yeo 1973).
The slender, dry, fruit pods, 15-35 mm in length, slightly longer than in L. corniculatus, maturein a star-like arrangement from August to October. They open in the same manner as in L.corniculatus, by sudden rupture and twist of two valves that explosively eject around 14 seedsfrom each pod (Grime et al. 1988). Seeds germinate mainly in spring, but a small proportion ofthem can survive soil burial for at least five years (Thompson et al. 1997). The 1 mm seed isrelatively large and although there is no obvious method of its long range dispersal, the species is aquite frequent colonist of vegetation gaps in sufficiently disturbed, damp grassy or muddy watersidesites.
Vegetative reproduction
L. pedunculatus frequently forms substantial clonal patches by growth of numerous spreadingstoloniferous shoots produced from the central rootstock of the original plant.
Fermanagh occurrence
Although it is just about a third as frequent and only about half as widespread in Fermanagh as CommonBird's-foot-trefoil, it is almost as widespread as the latter in the eminently suitable, lowland,periodically wet, 'water meadow' habitats around the shores of Upper Lough Erne. L. pedunculatusalso flourishes in the eastern half of the VC where, as the tetrad distribution maps illustrate, boththese species are much more thinly scattered. There are records of L. pedunculatus in theFermanagh Flora Database from a total of 150 tetrads, 28.4% of those in the VC.
British and Irish occurrence
The New Atlas map shows that this species is very widespread in Ireland, but much less frequent onthe Central Plain and in parts of the extreme west where ± constantly wet, strongly acidic peat bogspredominate. It is well distributed throughout Britain, except in N & NW Scotland and the Highlands,where again, presumably, the soils are just too acid, peaty and cold for it to manage.
Uses
L. pedunculatus contains as much protein and fibre as other common legumes used for fodder, and isor was therefore, a recommended component for cultivation in permanent grassland in periodically moisthabitats in Poland (Zimny 1965). It is not used for this purpose in B & I at present.
European and world occurrence
Widely distributed across W, C & S Europe, north to 60°N in Scandinavia and east to 25°E in Ukraine,plus in N Africa and the Canary Isles. It has been spread by man within and beyond this area and isoften a casual. It has certainly been introduced to N Fennoscandia and Iceland and to N America, Chile,Tasmania and New Zealand (Hultén & Fries 1986, Map 1250).
Threats
None.
Native, common. Eurasian boreo-temperate, but naturalised in N America and thus now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
April to November.
Growth form and preferred habitats
This climbing, trailing and twining, tangled, web- or curtain-forming perennial with its very beautifullydecorative, many-flowered, bright blue-mauve, one-sided, spike-like raceme of 10-30 pea flowers on along peduncle and pinnate leaves each with 5-15 pairs of ± parallel-sided leaflets and a branchedtendril is unmistakable, at least in Ireland. The base of the plant is a perennial rootstock or a shortrhizome (depending on which reference you consult) from which arise both a branched, annual, aerial stemup to 200 cm tall, plus spreading mycorrhizal roots with nitrogen-fixing nodules in the upper soillayers (Aarssen et al. 1986; Grime et al. 1988).
V. cracca clambers over tall supporting herbaceous plants in more open areas associated withwoods, scrub, cliffs, hedge-banks, walls and fences. It also grows in various rough grasslandsituations, including marshy or swampy river- and stream-banks, lakeshores, roadsides and waste places.While quite unspecialized in its substrate requirements, V. cracca appears to be most commonlyassociated with moist, sandy or gravelly soils. A good example of a mesophyte species, it grows best atpH 6.2 and really avoids only extremes of soil moisture, pH reaction and nutrient levels. It is absentfrom soils below pH 4.5 (Grime et al. 1988).
Tufted Vetch dislikes excessive disturbance (grazing, cutting and cultivation), deep shade, or exposureto strong winds or severe cold (Aarssen et al. 1986). It can tolerate light grazing and, later inthe summer, can survive being cut for hay (Duke 1981). In terms of its established strategy, thecolonising and competitive ability of V. cracca is rated as intermediate between competitor andC-S-R, which puts it on a par with Lotus pedunculatus (= L. uliginosus). In species richgrassland it becomes stunted and is eventually ousted (Grime et al. 1988).
Variation
There is considerable chromosome and phenotypic variation within what is described as the V.cracca complex, including polyploidy and aneuploidy (ie unusual non-multiples of the basicchromosome number). The common chromosome race in B & I is tetraploid (2n=28), plus there are otherraces with 2n=14 (diploid), 12, 27 and 30 chromosomes (Aarssen et al. 1986; Grime et al.1988). Four intraspecific taxa have been recognised and named as varieties in the critical Flora of B& I (Sell & Murrell 2009), as var. cracca, var. leptophylla Fr., var.sericea Peterm. and var. pulchera (Druce) P.D. Sell. They differ in degrees of hairiness,leaflet dimensions and flower colour. Stace (2019) makes no mention of any of these varieties.
Flowering reproduction
From June to August, the very numerous, 8-13 mm long, clear blue-mauve flowers, which areself-incompatible, attract various large bees as pollinators. The vetches (Vicia andLathyrus) have a secondary pollen presentation reminiscent of that of the brush mechanism of thegenus Campanula. The style is bent up sharply from the tip of the ovary and carries a dense brushof fine hairs just below the stigma. The anthers open and release their pollen while the flower is stillin bud and the pollen is shed on to the tip of the brush or into the tip of the corolla keel where thebrush sweeps it out as the keel is depressed. By the time the flower opens the anthers have retractedbut the stigma brush is fully charged with pollen as it comes up into contact with the lower abdomen ofa visiting bee.
The petals are relatively large and stiff and an insect has to be weighty, powerful and long-tongued inorder to penetrate the stamen tube and reach the concealed nectar. The flowers are, therefore, more orless restricted to bumble-bees, although other insects may steal the nectar by biting into the base ofthe flower and circumventing the designed mechanism (Proctor & Yeo 1973).
Later in the season (mainly August to September), the dry, mature, 10-25 mm, legume pod splits suddenlyat maturity, explosively hurling out 2-6 relatively large, hard, reddish-brown seeds a short distance.Birds and grazing animals may also eat the legume and eventually pass the intact seeds with theirfaeces, achieving long distance dispersal (Aarssen et al. 1986; Grime et al. 1988).
Vegetative reproduction
Shallow spreading roots assist the plant to reproduce by vegetative means: buds on the roots can giverise to new adventitious aerial shoots at intervals. Typically, V. cracca forms quite smallclonal patches in grasslands, indicating that this form of reproduction is not very significant orparticularly successful. The main method of increase is through seed production and dispersal.
Fermanagh occurrence
V. cracca is not quite as common and widespread in Fermanagh as the much less showy V.sepium (Bush Vetch). Yet with records in 310 tetrads, 58.7% of those in the VC, V. craccaremains a very familiar and widespread vetch in lowland Fermanagh.
British and Irish occurrence
The undemanding habitat requirements mean that V. cracca is very common and widespread throughoutB & I, scarce only in the Scottish Highlands and the wet, peaty NW of Britain (New Atlas).
European and world occurrence
Rather variable and partly spread by human activities including farming and trade in seed and fodder,V. cracca s.l. (including in more southern areas V. tenuifolia Roth (= V. cracca subsp. tenuifolia (Roth) Gaudin)) is widespread throughout most ofEurope plus Greenland, and Asia to Sakhalin and Japan. It is introduced in N America, S Africa, Tasmaniaand New Zealand and is now circumpolar boreo-temperate (Hultén & Fries 1986, Map 1200; Sell &Murrell 2009).
Threats
None.
Native, rare and declining. Eurosiberian, boreo-temperate.
1882; Corry, T.H.; Ballinamallard.
July and August.
Growth form and preferred habitats
In Britain, at least, this distinctive vetch, like its relatives V. sepium (Bush Vetch) and V.cracca (Tufted Vetch), is usually a trailing, scrambling or climbing, branched, rhizomatousperennial legume, 50-200 cm in height, leaves with 8-20 leaflets and a much-branched terminal leaftendril. It occupies habitats such as wood and scrub margins, rocky gorges and clearings, various formsof rough wayside grasslands and steep, dry, stony banks. It also grows in open, ungrazed conditions oncliffs, talus slopes and, at the coast, it grows on sea cliffs, lightly vegetated shingle and on screes.In some VCs in B & I, it is regarded as one of the indicator species of ancient woods (Rackham 1980,p. 54). V. sylvatica has always been very much more scarce, occasional and local in Ireland thanis the case in Britain, occupying a more restricted habitat range, principally in heathy scrub, onsometimes steep slopes, in coastal situations.
Variation
The species has two varieties, the type var. sylvatica is larger, with weaker, trailing orscrambling stems and leaves with 12-20 leaflets and tendrils 2-7 cm, 2- to 4-branched, the floweringraceme distinctly exceeding the leaves. The alternative form, var. condensata, is smaller withstems 20-50 cm, more rigid, procumbent to decumbent, forming compact patches or low hummocks. It occursin very scattered, local, coastal sites. Intermediate forms do also occur (Sell & Murrell 2009).
Flowering reproduction
Like the related vetches of the Section Cracca, reproduction is almost entirely by seed.The plant flowers from June to August, the inflorescence being a rather lax, one-sided (secund), racemeof up to 20 flowers, the corolla of each with a beautiful white standard and wing petals, tinged withlilac and striped with purple veins. Pollination is by bees and the resultant legume fruit is 25-30 mm,oblong-lanceolate and acuminate at both ends. When ripe it releases four or five black seeds (Claphamet al. 1987; Sell & Murrell 2009).
Experience of the species in Fermanagh suggests the seed is surprisingly mobile, the plant cropping up atleast temporarily and sporadically in several wayside areas within the upland Lough Navar Forest Park.Movement of the plant may well be assisted inadvertently by wheeled transport, or possibly the fruitsare eaten and the seeds internally transported by grazing animals. The truth is science does not knowhow it gets about, but clearly it does.
Fermanagh occurrence
V. sylvatica has been recorded in a total of ten tetrads (1.9%), although only two squares containpost-1975 records. As the tetrad distribution map demonstrates, Wood Vetch has definitely greatlydeclined in Fermanagh since the 1950s when Meikle and his co-workers recorded it at two completely newsites additional to the seven known from the turn of the 20th century. Their additional sites were:Mullylusty Td, in the Lurgan River Glen, 1947; and on the hill above Drummully Old Church, in the far SEof the VC, 1950. Five of the Victorian stations were discovered by Praeger, who remarked on the factthat Wood Vetch occurred on sandstone on the shore of Lough Fadd and on one of the Lough Navar scarps,but otherwise appeared to favour limestone, eg on the W & E ends of the Cliffs of Poulaphouca (alsoknown as the Cliffs of Magho) (Praeger 1892, 1901c & 1904). The red scarps in the Lough Navar ForestPark are in fact dolomatized sandstone, some of the minerals having been subjected to replacement andhence they are not or only scarcely acidic and are capable of supporting definite calcicoles such asAsplenium viride (Green Spleenwort).
Wood Vetch has only been seen by RHN and the current author (RSF) seven times in Fermanagh in the last 35years: once at Praeger's Bess Island site, where it was still growing in profusion in the middle of thewooded island, and on four occasions, one or two quite large patches were observed growing in full sunon loose gravel beside forest roads near Shean Lough in the Lough Navar Forest Park. Recently, it hasdisappeared again at this rather public, presumably vulnerable site; in 2009 and 2010 several patches(up to 10 m x 6 m in size) were found by RHN and HJN growing at the entrance to a quarry in the sameforest park.
Irish occurrence
An inspection of the New Atlas hectad map indicates that losses of Wood Vetch in Ireland appear tohave occurred in 26 pre-1970 sites, while the number of post-1970 hectads with records totals just 31,the majority of these being in NI.
British occurrence
Today in Britain, V. sylvatica is still thinly but widely scattered across the whole range oflatitude, although rather local and with no clear or obvious pattern to its distribution. The one thingdefinite about its occurrence is that the plant is declining and disappearing from previous stations innumerous areas throughout the whole island, as has been reported from many county floras in recentyears. Some of the decline in woodland margins may result from a widespread reduction in coppicing,reducing the light levels the plant requires for its survival (D.A. Pearman, in: Preston et al.2002). Another suggestion is that the marked decline of the species, measured by the Change in theBritish Flora survey of 1987-2004, may indicate its vulnerability to increased grazing pressure,and possibly to our rather rapidly changing climate pattern (Braithwaite et al. 2006).
The current author cannot offer any explanation for the decline of the species in Fermanagh, other thanthe fact that it has always been a rarity and, of course, the smaller and the more isolated anybiological population is, the more vulnerable it becomes to a multitude of deleterious factors. A searchof published literature shows very little is known of the biology and autecology of the species, whichin view of its significant decline in B & I, deserves urgent study.
European and world occurrence
V. sylvatica occurs thinly scattered across N, C & E Europe and W & C Asia as far east asSiberia, but is absent from the Iberian, Italian and Balkan peninsulas, most of France and all of theMediterranean basin (Hultén & Fries 1986; Map 1203). The latter authors indicate that its occurrencein, for instance, N Europe is not as continuous as they indicate in their published map.
Threats
Apparently none, but the species is definitely in decline and requires monitoring and possiblyrestoration if it is to survive long-term.
Native, very rare. European temperate, but so widely naturalised it is now circumpolar.
1892; Praeger, R.Ll.; Long Island, Lower Lough Erne.
Growth form and preferred habitats
A small, scrambling, slender annual with weak stems 30-60 cm long and the leaves with 6-10 pairs ofsmall, usually blunt leaflets that end in a branched tendril. The small flowers, 2-6 together on aslender peduncle, are pale blue. The hairy legume pods are very distinctive and contain just two largeglobose seeds (1.2-2.2 mm in diameter).
Once a troublesome weed of arable crops, typical habitats nowadays are dry banks, roadside verges andother forms of disturbed scrubby grassland, the margins of arable fields, as well as on coastal ground(Garrard & Streeter 1983; New Atlas). The plant appears to prefer light, well-drained, mildlyacid to calcareous soils and in the mid-19th century and earlier it was a significant weed of cornfieldand other cultivation, as it still is in many parts of the world. Entire crops were sometimes destroyedby its rampant growth, which earned it the feared name of 'Strangle Tare' or 'Tine-tare', the verb'tine' meaning 'to suffer loss or deprivation' (Grigson 1955, 1987). It appears the seeds were noteasily separated from the harvested grain and flour was liable to be made unpalatable by the level ofcontamination. At the time, V. hirsuta was also a frequent impurity of commercial Clover, Wheat,Oat and Rye seed (Salisbury 1964).
The established strategy of V. hirsuta was given as R/CR by Grime et al. (1988) meaning itwas intermediate between ruderal and competitive ruderal; it is certainly weedy in its behaviour (rapidturnover of growth, flowering and seeding). In terms of phenology, Hairy Tare is a winter annual (iehibernal), germinating in the autumn, overwintering as a small plantlet and flowering and fruiting inearly summer. Seed has been recovered from the dung of cattle, suggesting that if the plant is grazed inautumn pastures, seed may be transported internally (Salisbury 1964). As is very often the case,measures or estimates of buried seed survival in soil suggest a range of values from transient (lessthan one year) to long-term persistent (surviving at least five years) Thompson et al. (1997).
Irish occurrence
V. hirsuta is a widespread but very local species in Ireland, being most frequently found on fieldmargins and dry banks in the eastern half, but much rarer in the west (Parnell & Curtis 2012).
Fermanagh occurrence
Although possibly never very common anywhere in W Ireland, V. hirsuta has declined to great rarityin Fermanagh following the almost total demise of arable farming here during the last 60 or more yearsfollowing World War II. There are a total of just seven records of Hairy Tare in the Fermanagh FloraDatabase in six tetrads, four of which are pre-1950 in date. The details are as follows: Praeger's late19th century finds on Inishfree and Long islands, Lower Lough Erne (Praeger 1892), and two records byMeikle and co-workers on or near the Crom Castle estate on Upper Lough Erne – at Galloon Td in 1946 andin a potato field near Ports Lough in 1949 (Meikle et al. 1957 & 1975). V. hirsuta isstill a very rare plant in Fermanagh, having been seen at just three stations in recent years: twoadjacent sites on Lower Lough Erne islands in 1989 by Matthew Tickner (Muckinish West and Rosscor) andin the same year in the Correl Glen NR by members of an EHS Habitat Survey Team. The latter is a mostunexpected site for an annual species that is most often associated with dry, stony, disturbed groundand RHN and the current author (RSF) are a little wary of accepting it. However, the access paths inthis wet upland wooded glen are constructed of rough angular gravel and they may therefore have provideda suitable niche (however fleetingly) for this species.
British occurrence
In strong contrast to its much rarer occurrence in Ireland, the New Atlas map indicates that, atleast in lowland Britain, V. hirsuta is fairly frequent, widespread and locally abundant insuitable rough grassland or disturbed ground; the main exceptions to this lie in N, W & SW Scotland,NW England and parts of C Wales. At least in the Scottish areas, the predominant wet, acid peat, boggyground provides an obvious explanation for the absence of this species.
European and world occurrence
Crop weed species, like V. hirsuta, spread with agriculture around the world in both hemispheresand become almost circumpolar in the north and this can make it difficult to distinguish native fromintroduced occurrence. However, this nowadays very widespread species is considered native in W & SEurope, N Africa and parts of SW Asia. It is near-naturalised in many countries worldwide (Hultén &Fries 1986, Map 1206; Sell & Murrell 2009).
Threats
Unknown.
Native, common and widespread. Eurosiberian boreo-temperate, but widely naturalised.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Bush Vetch is an extremely common, familiar and widespread, vigorous, tap-rooted, perennial,nitrogen-fixing legume. The dull, pale purple-blue flowers, produced in clusters, are neither asnumerous nor as attractive as those of the other common, but less widespread vetch, V. cracca(Tufted Vetch). Aided by its branched tendrils, V. sepium climbs and clambers up to 100 cm ormore high on supporting species in practically every hedgerow one examines carefully enough. It is alsovery common climbing or trailing and decumbent in relatively unmanaged, lightly grazed, or infrequentlycut rough grassland, eg on woodland and scrub margins, moist to dry waysides, roadside verges and wasteground. It is less frequently found in tall-herb vegetation on lakeshores and river banks and in almostinaccessible, ungrazed areas on cliffs and steep screes, particularly in limestone areas of the country.
Bush Vetch appears to grow most luxuriantly and is able to compete well with other tall growing plants onmoist, neutral to basic soils of low fertility, especially where the ground is relatively undisturbed,but not completely so, thus restricting the vigour of this quite competitive species. The establishedstrategy of V. sepium is reckoned to be intermediate between competitor and C-S-R, placing it inthe same category as V. cracca (Grime et al. 1988). While it tolerates only mild levels ofacidity (restricted to soils above pH 4.5), but can cope with low nutrient levels, V. sepium israre or absent from permanent wetlands (Grime et al. 1988). It particularly avoids very acid bogsand wet, peaty, upland heaths and moors and arable land.
V. sepium forms apparently long-lived patches in suitable growing conditions, but it regularlydies down in the autumn, except in mild coastal areas where it often remains wintergreen.
Variation
The species has been split into two varieties, the widespread type var. sepium and the very muchrarer endemic coastal var. hartii Akeroyd which has much shorter, prostrate or decumbent,trailing or weakly ascending stems that usually forms mats or hummocks in sand dunes, but occasionallyclimbs on Ammophila arenaria (Marram Grass). This variety is confined to widely scatteredlocalities in B & I including the island of Coll in the Hebrides (VC 110), the N coast of Sutherland(VC 108) and Caithness (VC 109) and on the Mullet Peninsula, Co Mayo (H27) and at Kincashla Point, WDonegal (H35) in NW Ireland (Akeroyd 1996). Intermediates are also recorded between the two varieties(Sell & Murrell 2009).
Flowering reproduction
V. sepium flowers from May through to August or even later in milder areas. Indeed, at the coast,it can often still be found in flower in December. The inflorescence is a compact axillary raceme of 2-6flowers borne on very short or no peduncles. The 12-15 mm corolla is reddish purple, the standard petalmarked with dark purple veins. The wing petals are paler or bluish and the keel reddish. The wholeflower fades to a dull blue or greenish-blue colour. The flowers offer plentiful pollen andwell-concealed nectar and attract heavyweight bees (honey bees and bumblebees) that operate the stylarbrush pollen transfer mechanism of cross-pollination (Proctor & Yeo 1973, pp. 200-1).
The legume fruit pod, 20-35 mm long, black and hairless, ripens from July to September or later,splitting explosively to throw out 3-7 relatively large, hard-coated seeds (Grime et al. 1988;Sell & Murrell 2009). There does not appear to be any evidence for long-range seed dispersal or anyobvious mechanism to make it possible, Ridley (1930) remaining silent on the topic, although he doesallow that V. cracca, in somewhat similar habitats, may be carried in mud on boots or animalfeet.
It has been pointed out that as with most native legumes, there is little or no understanding of themechanism and significance of hard-coat seed dormancy under relatively mild B & I oceanicenvironmental conditions (Grime et al. 1988). The large seeds of V. sepium appear to beinefficiently dispersed and the species is seldom found colonising new, open artificial habitats.However, it remains such a common and widespread species all across B & I, and there does not appearto be any noticeable change in its distribution between the two BSBI Atlas surveys (D.A. Pearman, in:Preston et al. 2002), that the current author (RSF) cannot agree with Grime et al. (1988)that it is, "probably decreasing". Furthermore, there is definite evidence of spread andnaturalisation of V. sepium beyond our shores in Europe and elsewhere across the globe, stronglysuggesting widespread dispersal assisted by man through his trading and agricultural activities.
V. sepium has a limited capacity for vegetative spread, but seed production appears to be thedominant form of reproduction (Grime et al. 1988). The possibility of a persistent buried seedbank is uncertain: the survey of NW European soil seed banks listed a total of nine studies of thistopic, eight of which reckoned V. sepium seed was transient (ie survived less than one year),while the remaining study suggested it survived burial for at least five years (Thompson et al.1997).
Fermanagh occurrence
V. sepium is the 50th most frequently recorded vascular plant in Fermanagh and it has sites in 469tetrads, 88.8% of those in the VC.
British and Irish occurrence
Very common throughout B & I, except on exposed coasts and high ground. Other areas that show up evenat the hectad scale of the New Atlas map as being unsuitable for the species are wet groundaround the English Wash and in adjacent Lincolnshire (VC53 & 54) and Cambridgeshire (VC 29), whichtogether have the most heavily cultivated arable land in the country.
European and world occurrence
Very widespread throughout most of Europe but becoming more scarce or rare towards the south on theIberian and Balkan peninsulas, although present throughout Italy and even reaching Sicily and Sardinia.Absent from all the other Mediterranean islands and from N Africa and Macaronesia (Azores, Madeira, theSalvages, the Canaries and the Cape Verdes). To the north of Europe, it is present well within theArctic Circle in Scandinavia and also very rare and possibly introduced and naturalised in S Iceland andthe Faeroes and definitely introduced in S Greenland (Ostenfeld & Gröntved 1934; Böcher etal. 1968; Löve 1983). V. sepium is also considered indigenous in temperate Asia andKashmir and it has been introduced further east in Japan, S Australia and also in N America (Hultén& Fries 1986, Map 1208; Sell & Murrell 2009).
Threats
None.
Introduction, archaeophyte, doubtful numbers, perhaps occasional, but most probably it has declined toextreme rarity. European southern-temperate, but widely naturalised.
24 May 1992; Northridge, R.H.; N end of Devenish Island, Lower Lough Erne.
May to September.
Growth form and preferred habitats
This robust, annual vetch with its typically bi-coloured (claret and blue) or violet, purple or rarelywhite pea flowers was probably introduced to Britain by the Romans around 80-130 AD as a fodder legumecrop and, for a long period, it was popular with farmers in both B & I for this purpose and as aploughed in green manure to enrich soil fertility (Hollings & Stace 1978). Like other vetches, it isnot highly specialised in its substrate requirements, but it is generally associated with light, sandyor gravelly soils. V. sativa produces fibrous roots with nitrogen-fixing nodules and is capableof growing up to 150 cm tall on supporting vegetation. It persists to a certain extent on field margins,hedgebanks and amongst rough, occasionally-mown, tall grass on roadside verges and on various forms ofdisturbed waste ground.
Fermanagh occurrence
V. sativa s.l. is occasional and widely scattered in suitable disturbed ground in the Fermanaghlowlands and around 50% of the local records of it were generated during the exhaustive field-by-fieldscrutiny of the shores of Upper Lough Erne made by the EHS Habitat Survey Team recorders in the late1980s (see subspecies accounts below).
Variation
In the 1980s, when the Upper Lough survey was being carried out, this very variable plant was referred toas V. sativa subsp. sativa in all the available identificationFloras, including An Irish Flora 1977 and Clapham et al. (1962). Studies over the last 40years have found that the true nature of V. sativa is that of a taxonomically confusing andvariable complex or aggregate of at least six or seven subspecies. These taxa include wild types, weedyraces and cultivated derivatives (P.W. Ball, in: Flora Europaea 2, Tutin et al.1968, pp. 129-36; Hollings & Stace 1978; Aarssen et al. 1986). It is important to appreciatethe extent of the variation involved, which is such that at the variety and form levels, the number oftaxa described for the V. sativa species aggregate or s.l. comprise several hundred! A widevariation in chromosome morphology which underlies and helps to explain this situation has beenreported, with cytotypes existing of 2n=10, 12 and 14 chromosomes (Hollings & Stace 1974).
Further taxonomic work has now considerably clarified the variation within V. sativa found in B& I (Hollings & Stace 1978; Sell & Murrell 2009), so that with the recognition of subsp.segetalis (Thuill.) Gaudin, three, rather than two subspecies of V. sativa (ie subsp.sativa and subsp. nigra (L.) Eheh. (the latter = V. angustifolia L., Narrow-leavedVetch), are now listed in the New Flora of the BI (Stace 1991, 1997 & 2019; H.J. Killick, in:Rich & Jermy 1998, pp. 183-5).
The critical Flora of Great Britain and Ireland 3 takes taxonomic matters further and nowdescribes seven subspecies in these isles (Sell & Murrell 2009). The additional names to the abovementioned being subsp. uncinata (Rouy) P.D. Sell and subsp. bobartii (E. Forst.) P.D.Sell, both of which Stace (2019) together subsumes into (or includes within) subsp. nigra (L.)Ehrls.; subsp. cordata (Wulfen ex Hoppe) Arcang., which Stace (2019) regards as not confirmed forour flora; and subsp. macrocarpa (Moris) Arcang., which Stace (2019) regards as a rare casualonly.
Decline in planting vetches for fodder
The enormous decline of arable farming in Fermanagh and across Ireland in general, plus the generalousting of vetches by clovers and other legumes as preferred fodder and green manure crops, has seen thearea of subsp. sativa grown by farmers in B & I gradual drop from the early 1890s when it was216,000 acres [87,400 ha] in England and Wales, to near-rarity by the late 1950s (Killick 1975). Thislast author also found that even when it was being replenished by regular sowing, subsp. sativaseldom persisted for long and by 1974 it was becoming scarce in Britain.
Buried seed longevity
Measurements or estimates of the period of survival of V. sativa s.l. seed buried in soil listedin the NW European survey indicated a range from transient (less than one year) (eleven studies), toshort-term persistent (1-5 years) (seven studies), to more than five years (just two studies) (Thompsonet al. 1997).
V. sativa subsp. segetalis was also cultivated for fodder andmanure purposes, so it might also have suffered a similar decline, although like subsp. nigra, itmay perhaps maintain itself rather better than subsp. sativa does without additions from freshagricultural sowing.
Transfer of Fermanagh V. sativa records: The New Flora of the BI(1991, 1997, 2010 & 2019) regards subsp. segetalis as the commonest of the three subspeciesin B & I and in view of the inadequate identification treatment of this aggregate in An IrishFlora (1977, 1996), RHN and the current author (RSF) believe it is more sensible to transfer allthe Fermanagh records with dates between 1980-91 from subsp. sativa to subsp. segetalis(see below), rather than accept them as the former.
This leaves just nine definite records for subsp. sativa in the Fermanagh Flora Database in seventetrads, all but two of which lie between Enniskillen and Gublusk Bay, on the eastern shore of LowerLough Erne. Details of the other two records are: roadside between Enniskillen and Lisnarrick atGlenross, 21 May 1994, RHN; and Sand pit at Pubble Bridge, Tempo River, 20 August 1999, RSF & RHN.
Irish occurrence
The Reynolds's 2002 Cat Alien Pl Ir acknowledges this naming problem and it lists a number of1990s records of subsp. segetalis from 17 of the 40 Irish VCs.
Doubtfully native, very rare. Widely introduced and naturalised in both hemispheres.
1892; Praeger, R.Ll.; stony shore, Inishmacsaint Island, Lower Lough Erne.
May to September.
Identification and taxonomic confusion
V. sativa subsp. nigra is recognised as being a straggly, climbingor procumbent, slender-stemmed annual or winter annual with concolorous (ie single coloured) brightpurple, solitary or paired flowers only 10-20 mm long, with upper leaves having much narrower leafletsthan the lower leaves (ie it is markedly heterophyllous) and the legume is 30-50 mm long, smooth and notcontracted between the seeds (Hollings & Stace 1978; Aarssen et al. 1986; Sell & Murrrell2009). This vetch is so different from other variants of V. sativa that until quite recently itwas often regarded as a separate, though rather variable species, V. angustifolia L., thetranslated English common name of which still applies to this subspecies.
There has been much taxonomic confusion and change in nomenclature involving this particular taxon and ithas several recently used synonyms including: var. nigra L.; subsp. angustifolia (L.)Gaudin; subsp. uncinata (Rouy) P.D. Sell; subsp. bobarti (E. Forst.) P.D. Sell; V.forsteri Jord.; and V. angustifolia L. ssp. angustifolia (Sell & Murrell 2009;Stace 2019).
Fermanagh occurrence
V. sativa subsp. nigra grows on dry, sandy or gravelly places andthe Fermanagh database has a total of just six records for it in the VC. The first shown above was froma stony island shore and the others are: Enniskillen town, 1900, Praeger; sand pit, Pubble Bridge, TempoRiver, 11 September 1994, RHN; Drumcullion Lough, S of Tamlaght, 4 June 1996, HJN & RHN; top of awall, Old Crom Castle, 21 May 1999, RHN; and Rushin Point, Upper Lough Macnean, 18 May 2002, HJN &RHN.
Questionable native status
Recent taxonomic study has made subsp. nigra more readily identifiable and apart from the relatedbut more decidedly coastal V. lathyroides (Spring Vetch), which has never been found inFermanagh, subsp. nigra is the only form of the aggregate that is ever considered native. Thecurrent author (RSF) has doubts on this matter however, since the archaeological evidence for thepresence of Narrow-leaved Vetch quoted in Godwin (1975, p. 180), points back only as far as the Romanperiod in England. Furthermore, in B & I, subsp. nigra appears confined to lowland grassy andwayside waste places on dry, sandy soils and, if native, it is more likely so in coastal habitats suchas dunes, shingle, sea-cliffs and heaths. At its inland sites, subsp. nigra is very likely mostoften an introduction occurring in similar types of grassy places. Furthermore, it has been suggested(D.A. Pearman, in: Preston et al. 2002) that mis-identification for the widely planted, morerobust fodder and green manure form of the species, subsp. segetalis, is probably quite frequent.
European and world occurrence
This form of vetch is thought to have originated somewhere in Europe, W Asia and N Africa, to all ofwhich it is considered native by Hultén & Fries (1986, Map 1209). These authors regard Narrow-leavedVetch as the original form of the fodder plant that was very widely planted and spread by agriculturearound the globe. Their map shows the native area shaded and stretching from S Fennoscandia to the NWcoast of Africa, eastwards into W Asia and south into Turkey, Asia Minor and NE Egypt.
Beyond these areas, the distribution of subsp. nigra as an agricultural introduction is scatteredvery widely across Greenland, Iceland and from SW Asia to the Far East and to Australia, Tasmania, NewZealand and the South Sea Isles. It is also introduced in Africa in countries such as Ethiopia, EastAfrica and the Cape Province and also in both N & S America (Hultén & Fries 1986, Map 1209).
In phytogeographical terms, V. sativa is regarded as European southern-temperate (Preston &Hill 1997), but no phytogeographical element has been allocated to either V. angustifolia or toits synonym, this particular subspecies.
Introduced, occasional.
May 1978; Northridge, R.H.; Killyvilly, near Enniskillen.
May to September.
Identification
V. sativa subsp. segetalis is distinguished from other annualmembers of the V. sativa aggregate by its usually bi-coloured, 9-26 mm flowers, the standardpetal much paler than the wings and by its numerous narrow, smooth (unconstricted), usually glabrous,brown to black ripe fruit pods 28-70 mm long. The average plant is also an altogether much more robustform of the members of the V. sativa agg. and, unlike subsp. nigra, the leaves are not or onlyscarcely heterophyllous. V. sativa subsp. sativa, subsp.nigra and subsp. segetalis are not genetically isolated from one another and the complexpattern of variation they encompass is almost certainly the result of extensive inbreeding (Hollings& Stace 1978).
For quite a long period in the 20th century, V. sativa subsp.segetalis was an important, widely sown animal fodder and green manure crop. However, its use inagriculture has more or less ceased as it has been replaced by other legumes in grass seed mixtures(Killick 1975).
Fermanagh occurrence
This subspecies is occasional, widely scattered and sometimes apparently established on various forms ofdisturbed ground in the Fermanagh lowlands, including in lakeshore grasslands, a sand pit, a disusedquarry, roadside verges and urban and rural waste ground. There are records for this previously widelygrown fodder and green manure plant from a total of 22 tetrads, 4.2% of those in the VC. Around 75% ofthe records for the plant were generated during the exhaustive field-by-field study of the shores ofUpper Lough Erne made in 1986 by the EHS Habitat Survey Team and this obviously skews the distributiontowards this area of the VC as shown in the tetrad map.
Transfer of records
Due to inherent genetic and phenotypic variation associated with habitual inbreeding within the V.sativa species aggregate, hybridisation with closely related taxa and the breeding of largenumbers of cultivated strains, some of which have escaped into the wild and crossed with wild forms,there has developed an almost continuous spectrum of variation. This amount of variation, together withdiffering taxonomic treatments and nomenclature confusion dating right back to Carl Linnaeus, led tothis subspecies being erroneously identified as subsp. sativa at the time of the Upper Lough ErneEHS survey (Hollings & Stace 1978; Clement & Foster 1994).
Prior to Stace's New Flora of the BI (1991 and later editions), there was confusion betweensubsp. sativa, subsp. nigra and the more commonly plantedsubsp. segetalis. Consequently, RHN & the current author (RSF) have decided, on the balanceof probability, to transfer all the supposed subsp. sativa records dating between 1980-91 acrossto the more widespread, more persistent and very much more likely subsp. segetalis. This is a farfrom ideal position to be in, but we believe it draws the best picture of the likely Fermanagh situationof these three forms of V. sativa over the post-1975 period of the local recording effort and thecreation of the Fermanagh Flora Database.
Lathyrus linifolius (Reichard) Bässler (= L. montanus Bernh.),Bitter-vetch
Native, occasional to locally frequent. European temperate.
1881-2; Barrington, R.M.; Co Fermanagh.
April to January.
Growth form and preferred habitats
This variable perennial legume has a creeping, tuberous rhizome and produces erect, glabrous, branched,winged stems, 15-50 cm tall. The pinnate leaves have 4-8 leaflets, lack a tendril and end in just ashort point. Its shallow fibrous mycorrhizal roots also bear nitrogen fixing nodules.
L. linifolius is a scrambling, slow-growing, occasionally loose patch-forming plant ofcontinuously moist to damp, but well-drained, acid to neutral, but always fairly unproductive,nutrient-impoverished soils. Suitable substrates and growing conditions generally occur in rocky orstony, moderately to heavily grazed, heathy meadows and pasture grasslands. In both low-lying and uplandsituations these habitats can merge into slightly drier, blanket bogland slopes, or occur on the marginsof flushes where Calluna vulgaris (Heather) and other heathers tend to come to the fore (Grimeet al. 1988; D.A. Pearman, in: Preston et al. 2002).
Alternatively, L. linifolius also grows in light shade in low-growing vegetation on stony,moderately acid heaths – some of which are possibly in the process of degrading to unproductivegrassland under various pressures. Additionally, it appears on damp ledges of usually N-facing cliffs,or on the margins or more open areas in scrubby or woodland vegetation, often but not always in fairlyexposed situations. In this rather wide range of habitat situations, L. linifolius avoidspermanently wet ground, extremes of pH (ie it usually occurs in conditions above pH 4.0 and is rare atpH 7.0). In pastures, it avoids moisture stress, heavy grazing and other forms of excessive disturbance.In terms of its established growth strategy and competitive ability, L. linifolius is consideredintermediate between C-S-R and a stress-tolerator (Grime et al. 1988).
In lowland, semi-shaded situations in open deciduous woodland or scrub, Bitter-vetch makes use of its'semi-vernal' growth and reproductive strategy. Using energy resources stored in its overwinteringtuberous rhizome, the plant puts on a growth spurt in early spring producing its aerial stem and leaves.It then flowers as early as April onwards, although in contrast in more upland or full sun conditions ofother habitats, this may often be delayed until the end of May. (Grime et al 1988).
Probably on account of its predominantly early season growth, followed by very limited further shootextension throughout the summer and with the aerial parts then dying down in the autumn, L.linifolius tends to be absent from pastures that are regularly grazed in the spring, or whichare heavily grazed, or fertilised and 'improved' at any time of year (Grime et al. 1988).
Flowering reproduction
Flowering begins early in the season in April and continues into July. The inflorescence is an axillaryraceme with 2-6 flowers borne on peduncles longer than the leaves. The flower is 10-16 mm long, withpale, bluish-mauve petals, the keel deeper in colour and greenish towards its base. Lathyrus andVicia flowers share a secondary pollen presentation model similar to that of the genusCampanula in which pollen from the ten stamens is first shed while the flower is still in budinto the folded keel around the tip of the single style which is furnished with a brush of hairs nearthe up-turned stigma. The pollen ends up either shed onto the stylar brush, or into the tip of the keel,where the brush sweeps it out when the keel petals are depressed by the weight of a visiting honey- orbumble-bee. It takes a powerful insect with a long tongue (proboscis) to reach the nectar at the base ofthe ovary within the anther filament tube. By the time the flower is open and ready for pollination thestamens have shrunk and retracted, but the stigma brush is fully charged with pollen as it comes intocontact with the underside of the bee's abdomen when it hovers or sits astride the keel and reaches intothe flower for the well-concealed nectar (Proctor & Yeo 1973).
The flowers are self-incompatible, making cross-pollination essential for seed production. The legume podis 25-45 mm long, sub-cylindrical and contains 4-10 large seeds. Seeds are released explosively by therupture of two sutures or lines of weakness as the ripe pod wall dries and suddenly splits, hurling theseeds out. Seed is shed between July and October, but it does not travel far from the parent plant. Incommon with other legumes, L. linifolius seeds can germinate immediately or soon afterrelease in the autumn, but within days they rather quickly develop hard-coat dormancy. This thenrequires physical scarification or over-winter weathering before the seed coat weakens and allows waterand oxygen to be imbibed, so that subsequent spring germination may occur.
Vegetative reproduction
A balance of seed and vegetative reproduction occurs, but in many instances the latter ispossibly the most productive. Effective regeneration is achieved by lateral growth of the creepingtuberous rhizome in the upper layers of soil or under leaf litter on woodland margins or in canopy gaps(Grime et al. 1988).
Variation
There is sufficient genetic and phenotypic variation within L. linifolius for Sell & Murrell(2009) to describe three varieties: var. montanus (Bernh.) Bässler with leaflets 25-50 × 10-25mm, narrowly elliptical to elliptical, obtuse-mucronate at apex; var. linifolius with leaflets20-50(-100) × 0.5-3.0(-5.0) mm, long-linear, narrowly acute at apex; and var. varifolius(Martrin-Donos) P.D. Sell with leaflets 18-50 × 3-10 mm, narrowly elliptical, those of the lower leavesusually broader than those of the upper, mostly pointed at the apex.
The distribution and ecology of these three varieties in B & I is not properly understood as yet,although they are all regarded as widespread in continental Europe (Sell & Murrell 2009). Stace(2019) confines himself to remarking that, "our plant is var. montanus."
Fermanagh occurrence
On the more upland areas of Fermanagh limestone terrain, L. linifolius very often grows inassociation with Calluna vulgaris (Heather) in small pockets of acidic peaty conditions developedon top of heavily leached, generally shallow, 'ranker' soils, or over very shallow rendzina profiles.The current author (RSF) regards Bitter-vetch as a definite calcifuge species and it is occasional tolocally quite frequent and widespread in Fermanagh, having been recorded over 200 times in 92 tetrads,17.4% of those in the VC.
As the tetrad map displays, the Fermanagh presence of Bitter-vetch is scattered throughout most of theVC, but it is heavily weighted towards the Western Plateau, where the upland scarps and the limestonespredominate. Agriculture is also less intensive and human population is at its lowest in this region ofthe county. There is no evidence in the Fermanagh records of any decline in the presence of this legumein the VC, although the record details of early finds in the county are really too sparse and sketchy todisplay any trend.
British and Irish occurrence
Bitter-vetch is common and widespread throughout most of Ireland, although rarer or absent in much of theintensively managed agricultural Midlands where grassland improvement has been most extensive. It isalso absent in the unsuitable wet peatlands of the far west.
In Britain, although there has been a definite decline since the 1950s, L. linifolius is stillvery widespread, except in SE England and NW Scotland, presumably for similar reasons to the Irishexperience (D.A. Pearman, in: Preston et al. 2002). As the narrow range of suitable habitatsbecomes ever more encroached upon and modified, the decline of L. linifolius appears increasinglylikely (Grime et al. 1988, 2007).
European and world occurrence
L. linifolius is widespread in S, W & C Europe reaching north to the Baltic region and ERussia. It becomes scarcer towards the south in the Iberian peninsula and the Balkans, although it ispresent throughout Italy including the far south. It is absent from all the Mediterranean islands, buthas one isolated foothold in NE Africa (Hultén & Fries 1986, Map 1215).
Uses
The tuberous rhizomes were previously collected, dried and stored for famine food and for medicinal usein Scotland, and they were even used to flavour whiskey. Chewing dried rhizome was thought to sweetenthe breath after heavy drinking and prevent drunkenness (Garrard & Streeter 1983; Darwin 1996).
Threats
None.
Native, common and locally abundant. Eurosiberian boreo-temperate, but widely naturalised, including in N& S America and New Zealand.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
L. pratensis is a very variable, conspicuous, yellow-flowered perennial, its tendril-bearingleaves with just one pair of leaflets; it is a vigorous, patch-forming climber of open habitats. Theseinclude grassy roadside verges, hedgerow banks and other forms of semi-natural rough unimprovedgrassland, such as field margins, river banks, cliffs, screes and other similar open, sunny situations.As this clearly indicates, Meadow Vetchling has a very wide ecological amplitude, but like manyspreading, rhizomatous species it prefers moist, mesic fertile, moderately acidic, well-lit growingconditions. Despite this, its subterranean rhizome provides energy reserves that allow L.pratensis to tolerate rather drier, less fertile, half-shade, calcareous soils of a wide varietyof textures, from clay to sand or even mull humus conditions, although it may not persist for long insuch situations.
Despite its relatively high vegetative vigour, L. pratensis is not strongly competitive and canonly tolerate light grazing, very occasional mowing, or other mild forms of moderate-level disturbance.Thus it tends to be restricted to sites where the potential dominance of more vigorous species islimited by disturbance. It is seldom if ever found on substrates of pH below 4.5 and it completelyavoids waterlogged aquatic situations, wet peat and deep shade (Brunsberg 1977; Grime et al.1988). When not clambering over other taller-growing, coarse grassy or woody vegetation, L.pratensis is often found entangling and supported by sheep netting wire on fencing. Locally italso appears on lakeshores, cliffs, screes and in limestone pavement.
Flowering reproduction
Flowering takes place from May to August. The inflorescence is an axillary raceme of 5-12 flowers, eachwith a 10-20 mm, bright yellow corolla, the standard petal with greenish veins. Cross-pollination of thenectar-containing flower is by the stylar brush mechanism involving honey- and bumble-bee visitors (seeL. linifolius account for a description). In isolation experiments, less than 4% of flowersmanaged self-pollination, but clearly the barrier to autogamy is incomplete and self-fertilisation canoccur (Brunsberg 1977). Seed is the chief means of L. pratensis dispersal, yet Brunsberg foundthat in some populations its production was poor, only around 15% of flowers setting any seed at all.
Seed dispersal
The legume pod developed after cross-pollination is between 25-35 mm long, glabrous or finely pubescent,compressed and contains 3-6(-10) rather heavy seeds. Fruit pods ripen from August to October, splittingto release the seed, although as this is without an autochorous dispersal mechanism (ie there isno explosive release), they always fall near the parent plant. The seeds are not adhesive either, sothere really is no specialised seed dispersal mechanism of any sort (Brunsberg 1977). After the seed isshed, the aerial shoot dies down in the autumn. Shortly after their release, the seeds quickly develophard coat dormancy (see the current author's L. linifolius account).
In view of the lack of any obvious or efficient seed dispersal mechanism, the widespread British, Irishand world distribution is rather puzzling. However, man has very probably assisted dispersal in pastyears, through seed being accidently transported with hay, or eaten along with the plant by animals andtransported internally, or spread along with commercial crop seed as an impurity. Plants growing nearflowing water might also end up floating to new territory, and seed produced on scree or cliff sites inuplands, might be carried to lower levels by rainwater streamlets.
Seed germination and survival
Seed germination is controlled by the permeability of the seed coat and it is greatest in spring after anoverwintering period of dormancy which presumably weakens the seed coat (testa) and allows water andoxygen entry to the embryonic tissues. In cultivation experiments, using seed from all over Europe, onlyaround 10-15% germination was achieved. Accord to Brunsberg (1977), seeds can survive dry herbariumstorage and remain viable for up to 90 years. On the other hand, the seed bank survey of NW Europeshowed that the great majority of studies concluded that seed was merely transient in soil (ie itgerminated or survived less than one year), a small minority of just three studies suggested seed wasshort-term persistent (ie survived 1-5 years burial) and only a single study found seeds were long-termpersistent (ie remained viable for more than five years) (Thompson et al. 1997).
Vegetative reproduction
Under suitable habitat conditions, the creeping rhizome of L. pratensisallows it to spread horizontally and surface root. In Brunsberg's study, "rhizomes up to 7 m longwere developed during one vegetative period", presumably meaning one growing season, although tothe current author (RSF) this would appear to be quite incredible. However having said this, Brunsbergdid find the variation between individuals and populations was very wide in respect to this rhizomecapability. Vegetative reproduction is most significant in pastures or meadows where the timing ofgrazing or cutting prevents seed production and dispersal (Brunsberg 1977; Grime et al. 1988).
Variation
The basic chromosome number in L. pratensis is x=7 and diploid (2n=14) and tetraploid (2n=28)forms have often been recorded, plus a range of other chromosome counts including 21 (triploid) and 42(hexaploid). In a study of the L. pratensis complex in Europe, Brunsberg (1977) found the diploidform had the widest distribution on the continent, but it was not found in the most western parts, wheretetraploids replaced it. Although there was a zone of overlap, the tetraploids appear to be restrictedto W & C Europe. Triploids and hexaploids were found only exceptionally. Root tip studies discoveredaneusomatic plants with missing or additional chromosomes (generally one missing or extra in diploids,or two chromosomes in the tetraploids) (Brunsberg 1977). The two main cytotypes cannot be distinguishedin herbarium material and they have not been given any taxonomic recognition.
Sell & Murrell (2009), in Flora of Great Britain and Ireland 3, recognise threevarieties within L. pratensis differing in degree of hairiness and the size of leaflets andlegumes: plants of var. pratensis are glabrous or nearly so and have leaflets 12-35 × 4-10 mm andlegumes 25-38 mm long; var. velutinus DC. has plants hairy, leaflets 10-25 × 1-5 mm and legumes30-35 mm long; while var. speciosus (Druce) Druce has plants ± hairy, leaflets 20-40 × 5-7 mm andthe mature legume was not seen.
Fermanagh occurrence
In Fermanagh, Meadow Vetchling ranks the 57th most common vascular plant species in terms of recordnumbers and, with a presence in 431 tetrads, almost 81.6% of the VC total, it ranks 41st in termsof tetrad distribution.
British and Irish occurrence
The New Atlas map shows that Meadow Vetchling is widespread and common throughout the whole of B& I, with the exception of the wet acid boglands of N Scotland and W Ireland. All plants here aretetraploids (Brunsberg 1977).
European and world occurrence
L. pratensis is widespread and considered native in temperate areas of Europe, Asia and parts of NAfrica. It is common in almost all of Europe, becoming more scarce southwards and eastwards from theIberian Peninsula, to Greece, the Caucasus and SE into Iran, Armenia and Turkestan. In the Alps andother European mountain areas, it often grows above the timberline. In N Europe, it is introduced andnaturalised in Iceland and Greenland and is considered recently introduced in N Finland, the Faeroes andparts of Norway (Brunsberg 1977). Of the Mediterranean islands, it is present only on Sicily, Corsicaand Sardinia. In Asia, it is recorded from Siberia to the Arctic Circle, eastwards to Japan and south tothe Himalaya. In Africa, it is present on high ground in Morocco and Ethiopia. L. palustris isalso introduced in N America and New Zealand (Hultén & Fries 1986, Map 1216).
Threats
None.
Native, occasional. Circumpolar boreo-temperate.
1859; Moore, D.; Upper Lough Erne.
May to October.
Growth form and preferred habitats
Marsh Pea is a tap-rooted perennial climber that scrambles with the aid of its branched tendrils overtall-herb, low-lying, wetland vegetation. It has almost hairless, winged stems that can reach 60-120 cmtall and which bear alternate, compound, blue-green leaves with 4-6 pairs of 35-70 mm, narrow lanceolateleaflets. The whole plant has a glaucous hue to it.
In Britain, L. palustris is increasingly rare, but remains a characteristic species of tall, richfen and reed-bed conditions, preferring situations that are rather deficient in nitrogen, an element itcan supply with the aid of its root nodules. However, in continental Europe and certainly also inFermanagh, Marsh Pea is very much more typical of wet coarse, relatively ungrazed grasslands and haymeadows over sedge peat or clayey soils bathed in lime- or base-rich inflow waters (J.O. Mountfield, in:Stewart et al. 1994).
In the Revised Typescript Flora, Meikle et al. (1975) commented on this then Fermanaghrarity, "Still found in moist meadows (not in marshes or reed-swamps), about Upper Lough Erne, andgenerally to be found by lake shores where cattle have been excluded." This statement remains trueat the time of writing, although this perennial pea with its distinctive winged stem now alsooccurs rarely and sparingly under wet fen-carr Alnus-Salix scrub, along fringing vegetation ofditches, canals and river banks. Sometimes it grows rather luxuriantly, clambering on fences around hayor silage meadows on the wet, calcium-rich, muddy, sedge peat shores of Upper Lough Erne and more rarelyit appears along the banks and fences of feeding streams.
Sexual reproduction
L. palustris flowers from May to July, producing 2-6 pea flowers on an axillary raceme, thepeduncle longer than the stem leaves. The flower corolla is 12-20 mm long, a delicate mauve or palebluish-purple colour that fades to a more greenish tinge as the blossom ages. Pollination is by bees andbumble-bees and the pollen brush mechanism (Proctor & Yeo 1973, p. 200). The legume pod is 3-6 cmlong, hairless, compressed and contains 3-12 seeds (Sell & Murrell 2009).
Marsh Pea reproduces largely by seed which is freely set, but it is also a long-lived perennial (J.O.Mountfield, in: Stewart et al. 1994). We might therefore expect it to be spread by flotation inwater currents and be transported internally by birds and other animals consuming the seed.
Variation
Although this is a scarce and local species in B & I, three varieties are recognised by Sell &Murrell (2009), the most frequent being var. palustris, which is glabrous. The other twotaxa are: var. linearifolius Ser. which is also glabrous and has leaflets up to 3.5 mm wide; andvar. pilosus (Cham.) Ledeb., in which the plant is hairy in all its parts. Var.linearifolius is known from Cambridge (VC 29), Yorkshire (VCs 61-65) and Kintyre (VC 101) andvar. pilosus from Berrow in Somerset (VC 6) and Pembrey in Carmarthenshire (VC 44). The latter isa common plant in N America and N Asia (Sell & Murrell 2009).
Fermanagh occurrence
In the post-1975 period, L. palustris has been recorded in a total of 23 Fermanagh tetrads (4.4%),so it is no longer described as rare, but instead pleasure is taken in upgrading its frequency to'occasional'. In quantitative terms, Marsh Pea can regularly be found in considerable abundance in someFermanagh fields and, in an exceptionally good year, it has been plentiful over a hectare or so to the Wof Lough Digh. It has also formed very large patches at Corraslough Point on Upper Lough Erne.
Irish occurrence
Further east in NI, L. palustris used to occur in four of the five VCs surrounding Lough Neagh,but it has declined and survives there now only on the Armagh and S Antrim shores (H37 & H39)(Harron 1986). The NI Flora Website (2005) hectad map displays post-1986 records on the Armaghshore. Elsewhere in Ireland, Marsh Pea occurs in several VCs along the basin of the River Shannon, plusat two outlying stations in Co Wicklow (H20). The New Atlas map suggests that it has not beenseen at one of these recently. After their Irish Red Book survey of vascular plants, Curtis &McGough (1988) felt that recent widespread drainage operations of wetlands throughout the island mightsomehow favour Marsh Pea and allow it to increase. Happily this prediction has proven accurate andseveral completely new stations have been discovered in Sligo (H28), where three hectads are now plottedin the New Atlas (Preston et al. 2002).
Lathyrus palustris has Schedule 8 Conservation status in NI.
British occurrence
This lowland species lost many of its sites in E England by the end of the 19th century and it hascontinued to decline there due to excessive drainage, 'grassland improvement' and long-runningdeficiencies in conservation management. However, there have been a few compensatory new finds since1970 in coastal Wales and in Kintyre in W Scotland. One new station (of var. pilosus) in a duneslack in Wales may result from trans-ocean drift or jump-dispersal of American seed (Vaughan 1978).
European and world occurrence
L. palustris is native and widespread in temperate Europe but rare in the Mediterranean basin. Itis also widespread in temperate and arctic Russia and Siberia and then eastwards to Japan and N Americawhere var. palustris gives way to var. pilosus (Hultén & Fries 1986, Map 1217). Inphytogeographical terms, it belongs to the circumpolar boreo-temperate element (Preston & Hill1997).
Threats
Probably none, though individual stands of the plant are occasionally decimated during silage gatheringoperations.
Native, probable mis-identifications. Southern sub-Atlantic.
4 September 1990; Waterman, T., Farren, J. & Montgomery, J.; Curragh More (a bog and heath).
May and September.
The two records of this pink pea-flowered, rhizomatous sub-shrub made by members of the EHS HabitatSurvey Team must be clerical errors on the field recording cards involved. The first record is detailedabove, the second is: Correl Glen (woodland), 7 May 1992, J. Farren & T. Waterman. The habitats(especially the upland woodland of the Correl Glen) are completely and utterly unsuitable for the plant,even as a casual. O. repens is associated with rough grasslands, scrub, quarries and similardisturbed situations on well-drained, dry, base- or lime-rich, generally light soils (or veryoccasionally on calcareous boulder clays). The typical habitats are often stony or gravelly, as oninland roadsides, coastal sand dunes and shingle beds. A glance at the New Atlas hectad mapindicates that it would not be impossible for O. repens to occur in inland Co Fermanagh, but notat either of the sites so far recorded.
O. repens is familiar enough to most local NI field botanists from sites on the east and northcoast where it is common, and it is so distinctive the current author (RSF) cannot imagine anidentification error being made. The only sensible explanation is that a slip of the pencil was made onthe recorder's field card, which was subsequently overlooked and became accepted.
Introduction, neophyte, casual. Probably Eurasian temperate, but native distribution obscured bywidespread naturalisation in both hemispheres.
23 June 2003; Northridge, R.H.; on gravel dumped on lakeshore, S corner, Holme Bay, Lower Lough Erne.
In Ireland, Ribbed Melilot is an infrequent or rare casual, most likely introduced as acontaminant of grain or pasture seed mixtures. In Britain, this biennial, which probably is a native from C & S Europe and SAsia, is occasionally abundant on open, disturbed habitats such as sand dunes, roadsides, railwayembankments, waste ground and rubbish tips. The Cen Cat Fl Ir 2 lists ten Irish VCs in whichrecords have occurred and Reynolds in Cat Alien Pl Ir adds post-1990 records mainly from aroundthe docks of Dublin and Belfast.
The solitary Fermanagh record found recently by RHN was in a most isolated lakeshore station and it isonly possible to imagine the seed source in terms of perhaps reseeded pastures nearby, or contaminatedgrain used for animal fodder.
The New Atlas hectad map shows that all the previous NI records were around Belfast or on thecoast, so the Fermanagh occurrence is a most interesting find. A voucher specimen that was sent toSylvia Reynolds for confirmation is deposited in DBN.
Native, occasional. Eurosiberian temperate, but very widely naturalised in both hemispheres and nowcircumpolar.
1884; Barrington, R.M.; Co Fermanagh.
May to November.
Growth form and preferred habitats
This variable little creeping annual, biennial or short-lived perennial legume produces a slender taprootthat branches and may penetrate 60 cm deep. Angular stems can be prostrate, decumbent, ascending orerect and vary from 15-80 cm in length and from hairless to densely hairy. Leaves are stipulate,petiolate and trifoliate and the terminal leaflet is stalked. Leaflet shape is variable, even on thesame plant. Two cytological forms with differing chromosome numbers have been reported: diploid (2n=16)and tetraploid (2n=32) (Turkington & Cavers 1979).
The remarkably wide distribution of M. lupulina across the globe suggests it has become adapted toa huge range of growing conditions. It typically colonises bare areas in fairly open, dry, lowland,permanent calcareous grassland and it competes and survives best in distinctly infertile conditions. Insuitably open conditions, such as wasteland or recently cleared or dug ground or forest margins, itbehaves as a ruderal colonist growing extremely rapidly and sometimes even become dominant for a shortperiod. In C England, the established strategy of the species is classified as being intermediatebetween Ruderal and C-S-R, which recognises it can behave rather differently across the wide range ofgrowing conditions it occupies (Grime et al. 1988, 2007).
M. lupulina is frequently found in moderately disturbed sandy or gravelly soils, where grazing,cutting or trampling curtails the growth of taller, more aggressive species. Black Medick is a typicallegume of lowland mown grass on roadside verges, lawns and grass paths, particularly over limestone. Thespecies withstands drought rather well, either surviving vegetatively or as seed, and it avoids stronglyacidic or very wet soil conditions (Turkington & Cavers 1979; Grime et al. 1988, 2007). Themost acid soil so far reported for this species was pH 4.8 in New Zealand. In moist, somewhat morefertile conditions, individual plants probably have a lifespan of at least four years (Grime etal. 1988, 2007).
M. lupulina can be distinguished from the similar small, yellow pea, Trifolium dubium(Lesser Trefoil), by its bright, as opposed to pale, yellow flowers and by its three apiculate ormucronate leaflets, the terminal one of which is stalked.
Variation
Black Medick is sufficiently variable for five varieties to have been recognised in B & I by Sell& Murrell (2009), all but one of them annual. Of these, var. eriocarpa (Rouy) P.D. Selloccurs in sandy places near the sea and on sandy heaths inland and is considered the most probablynative of all five varieties. It also occurs in similar habitats in continental Europe. Var. lupulinaprobably occurs throughout the range of the species and is a plant of rough grassland on roadsides andwaste-ground. It may not be native in B & I. Var. major G. Mey. is a cultivated formpreviously used for making hay. Today it is mostly found in wild flower seed for planting on waysides.Var. willdenowiana (Boenn.) W.D.J. Koch is also found in cultivated wild flower seed mixtures,although in some habitats it might be native. Var. cupaniana is a perennial variety thatoriginated in S Europe and is a rare introduction in Britain (Sell & Murrell 2009).
Flowering reproduction
M. lupulina regenerates only by seed. The inflorescence is a small, globose or shortly cylindricalflower-head up to 1.5 cm long, containing 20-50 tiny, yellow pea flowers, 2-4 mm long. Well establishedplants flower continuously right through the summer from April to August. The flowers are usuallyself-pollinated, but they may also attract bees and other insects that can cross-pollinate them. Theblack, slightly curved, 3 mm fruit pods each contain a single seed, yet a single robust plant in goodgrowing conditions may produce 2,000 or more seed in a growing season. The pod is indehiscent, the seedand pod being dispersed together as a unit (Turkington & Cavers 1979). In mown lawns and pastures,the plant becomes completely prostrate, most flowers are then borne very close to the soil surface andtend to avoid damage from mowers and larger grazing animals.
Seed dispersal
Dispersal involves birds and other animals ingesting the hard-coated seed along with the plant andinternally transporting it. Although they are not adhesive, the small seeds can attach externally toanimal coats, human clothing and to mud on machinery. Seeds can also float for up to five days and thusbe transported in flowing water (Ridley 1930). As with other grassland legumes, man has also assisteddispersal by transporting seed as impurities in seed mixtures and in fodder crops, so that whatoriginally was a Eurosiberian temperate plant has now become circumpolar.
Germination and phenology
In suitable, bare patch growing conditions in lawns and pastures, seeds can germinate immediately afterrelease from their pod. However, after around ten days, those that mature above the soil surface quicklydevelop the hard seed dormancy that is so very characteristic of the family. The tough, impermeable seedcoat then maintains dormancy through several unfavourable seasons, or for many years of soil burial. Thevast majority of individuals overwinter as dormant seeds and germinate in spring, often after adisturbance event like cultivation, animal digging or frost heaving of soil. Under favourable growingconditions, M. lupulina plants will flower within six weeks of seedling emergence and up to three± distinct generations can arise in a single growing season. In most habitats, the majority of plantsdie during their first winter, but a small minority may survive up to three years, chiefly in lawns(Turkington & Cavers 1979).
In a comparative study, the presence or absence of moss and the degree of moss in a patchy abandonedgrassland habitat in Canada, considerably assisted M. lupulina seedlings to establish, reachflowering capability and survive for more than one season. Moss ground cover reduced evaporation,thereby presumably reducing mortality resulting from drought. It also enabled plants to reproducerepeatedly, whereas without moss, plants simply did not survive long enough to reproduce more than once(Pavone & Reader 1985).
Fermanagh occurrence
M. lupulina is only occasional in Fermanagh, having been found in 29 tetrads (5.5%). It is thinlyscattered through the lowlands around Lough Erne, but is slightly more prevalent in the SE of the VC. Ittypically occurs in grassland habitats, plus in disused quarries and along disused railway waste ground– where presumably thanks to a persistent seed bank and regular disturbance from trampling cattle, itstill manages to survive. It also occurs in drier areas of grassland near lakeshores and in urban areasin and around Enniskillen. One odd habitat does occur, however, as it features in a strange admixture ofspecies on a cut-over bog just east of Clonnagore, near the old disused Ulster Canal. The range ofspecies on this peculiar, rather dryer than normal bog site included: Potentilla anserina(Silverweed), Stachys sylvatica (Hedge Woundwort), S. palustris (Marsh Woundwort),Petasites hybridus (Butterbur), Stellaria graminea (Lesser Stitchwort), Ligustrumvulgare (Wild Privet), Barbarea vulgaris (Winter-cress) and Equisetum arvense(Field Horsetail).
British and Irish occurrence
M. lupulina is rather uncommon in both the NW and the far W of Ireland in comparison with the restof the island. In Britain, it is widespread and common in most of lowland England and Wales, but itbecomes more scarce and coastal in Cumbria and Scotland, declining both northwards and westwards(Walters & Perring 1962; Preston et al. 2002).
In both these geographical areas the prevalence of wet, very acid soils is probably the only explanationrequired for the low presence of this species.
European and world occurrence
This rather variable species is widespread and native in most of Europe, Asia and N Africa. It has beenvery widely introduced, cultivated and naturalised throughout temperate and subtropical regions of theworld including N Europe, Iceland, Greenland, parts of Africa, N & S America, Australia, New Zealandand many other places (Hultén & Fries 1986, Map 1230).
Threats
None.
Native, very common, widespread and locally abundant. Eurosiberian boreo-temperate, but very widelynaturalised and now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year, peaking in May.
Growth form and preferred habitats
Very common, extremely variable, locally abundant and geographically widespread, White clover is one ofthe compound, three-leaflet species which in Ireland is regarded as St Patrick's Holy Trinity teachingaid and Irish emblem, the Shamrock. It is a stoloniferous perennial of indeterminate height, rooting atthe nodes of procumbent, much-branched creeping stems, up to 50 cm long. Most of the wiry, fibrous rootsare shallow, although it has a taproot that can reach depths of 60 cm. Plants are ± hairless (glabrousor glabrescent), with trifoliate leaves on erect petioles usually 7 cm or longer, leaflets with awhitish or reddish angled band towards their base. The inflorescence is a globular head of 20-40 whiteor pinkish, sweetly scented pea flowers on a peduncle that usually is longer than the leaf petioles(Burdon 1983).
The most important pasture nitrogen-fixing legume in many parts of the temperate zone, native forms andcultivar escapees of T. repens are abundant in lowland, moist, open, roadside grass verges, roughgrassy waste ground and other ruderal and wayside habitats. It is rarely a pioneer colonist as it hasrelatively high nutrient requirements. At the coast, it colonises damp pasture areas and dune slacks infixed and late stage sand dunes, despite having a low tolerance of sodium chloride (sea salt) in thesoil: it is seldom abundant on dunes and absent from salt marshes for this reason (Burdon 1983).Preferring flat or gently sloping, moist, fertile ground, T. repens avoids only the wettest andmost acid soils, but does occur to a lesser extent on grassy heaths and upland moors. Like otherlegumes, it also abhors shade and is almost totally absent from woods, scrub and tall vegetation of anykind (Grime et al. 1988, 2009). T. repens is also intolerant of prolonged drought andsevere frost, the latter potentially most damaging to stolons of clones with long internodes that rootless frequently (Ronningen 1949, quoted in Turkington & Burdon 1983).
The creeping shoots of T. repens enable it to colonise vegetation gaps in meadows and pastures,especially those involving Lolium perenne (Perennial Rye-grass), with which clover is verycommonly sown. As an efficient nitrogen fixer, T. repens holds a key position in the economy ofmany agricultural grasslands and may provide a driving force for the cyclic changes occurring between itand associated grass species in many grassland communities (Turkington & Harper 1979 a, b).
Grazing, mowing and trampling pressure
The effect of grazing on T. repens abundance in pastures is determined by its frequency, timingand intensity. In general, frequent and intense grazing encourages the growth of T. repens andtends to lead to the development of simple pastures entirely dominated by T. repens and Loliumperenne. Lighter, less intensive grazing pressure allows the entry or survival of taller grassessuch as Dactylis glomerata (co*ck's-foot) and Holcus lanatus (Yorkshire-fog), and this isassociated with a decline in T. repens abundance.
Differing seasonal growth rhythms of clover and grass species are also significant in terms of the effectof the timing and duration of grazing pressure and the effect observed on species composition of thesward. This is especially the case when selective grazing by sheep is involved. Close, heavy grazing ofa T. repens/Lolium perenne sward in March to May led to a substantial increase in cloveras the Rye-grass was grazed at a critical growth period. A slightly later and lighter grazing regimebeginning in mid-April resulted in a clover decline as L. perenne was able to maintain its vigourand remain dominant (Jones 1933).
A similar pattern of botanical changes in clover-grass mixtures occurs with variations in timing andfrequency of mowing. T. repens/L. perenne swards cut three times per year quickly becomegrass dominated, while swards cut six times per year become clover dominated (Kishi 1973, 1974).
White Clover is one of the most trampling-resistant common pasture legumes known. Again, as with grazingpressure, heavy trampling leads to a simplification of the species mixture present and the predominanceof the familiar T. repens and L. perenne partnership (Brown & Evans 1973).
Flowering reproduction
T. repens reproduces both sexually and asexually.
Reproduction by seed allows the rapid colonisation of newly available habitats at a distance from theexisting breeding population, while vegetative reproduction is important in maintaining individualgenetic individuals (genets) in the dynamically changing environment of long-term pastures. Disturbanceof the existing field vegetation appears necessary to create a gap allowing seed to germinate or avegetative genet to colonise the ground. Seedlings of T. repens are very infrequently observed inundisturbed pastures and even more rarely are seen to develop into established plants (Burdon 1983).However, this is almost certainly true of very many other pasture and meadow species.
Flowering takes place from June to September peaking in July, the small, honey-like scented pea flowerswith corollas 7-10 mm long, offer pollen and nectar to attract pollinators. They are visited by flies,solitary-, honey- and bumble-bees, particularly the last two mentioned (Proctor & Yeo 1973; Burdon1983). Nectar is secreted between the base of the stamen tube and the ovary and is accessible to insectswith quite a short proboscis, although they have to use their weight to open the flower by separatingthe wing and keel petals. Other insects frequently bite and puncture the base of the flower and stealthe nectar without effecting pollination.
Plants may flower in their first year of growth and continue to flower yearly thereafter. In establishednative populations, however, the entire sexual reproductive effort of the population may be concentratedin a small number of the genets present on a site. Burdon (1980) found that of 50 clones collected fromsuch a population, 20% produced 81% of the flowers, 40% produced no flowers at all and a further 10%produced only a few flowers.
The flowers are basically self-incompatible, although very rarely a minute amount of self-pollinated seedmay be produced and very rarely a few highly self-fertile plants have been found. After fertilisation,the flowers wither and droop, becoming reflexed. Seed matures about 28 days after pollination. Theresultant oblong legume pod develops, 4-5 mm long, constricted between the 2-6 seeds (usually just twoseeds) and it protrudes from the calyx tube while still surrounded by the dry, withered corolla (Burdon1983). The pod is indehiscent, but it eventually releases the seeds, although no special dispersalmechanism exists for them. By the time the seeds are released, hard seed dormancy has developed due tothe outer seed coat becoming impermeable. Most seed is probably incidentally spread by the movement ofgrazing and other passing animals, some being ingested and carried undamaged in the gut of cattle, deerand horses (Ridley 1930). Seed may also be transported with dried animal fodder such as hay. Hard seedscan survive soil burial for more than five years and possibly up to 80 years, although the density ofviable seed in soil is often low. Germination occurs in spring and early summer (Burdon 1983).
Hybrids with other Trifolium species do not occur naturally, although with great difficulty theyhave been artificially created (Burdon 1983).
Vegetative reproduction and genet longevity
Vegetative reproduction by the lateral extension of stolons is very important in T. repens. Thefrequency of new stolon branch initiation by lateral nodes varies with the genotype and through thegrowing season. It is always at a maximum in May and minimum in October and is strongly governed bytemperature. Under competitive field conditions, an annual extension of around 18 cm seems usual forstolon growth, but this is very variable with respect to local environment and the particular genotype(Burdon 1983).
The profuse vegetative growth and frequent intermingling of clones of varying genome makes any estimationof individual genet longevity very difficult. Potentially the lifespan of any genet is indefinite,particularly after the eventual break up of the original individual into a number of geneticallyidentical ramets. For instance, in old dune slacks Harberd (1963) estimated that a number of WhiteClover clones were 20 years old, while a particularly fragmented clone had a minimum age of 60 years anda possible maximum age in excess of a century. This contrasts with a typical plant of the lesspersistent Ladino crop variety that behaves as a winter annual: germinating in the autumn, floweringvigorously in the spring of the following year before dying (Burdon 1983).
Fermanagh occurrence
Together with T. pratense (Red Clover) this is one of the two most common and the most widespreadlegumes in Fermanagh and indeed throughout B & I. Locally, T. repens is slightly the morefrequent of these two clovers, although T. pratense is slightly more widespread – ie the latteris present in 89% of the Fermanagh tetrads, while T. repens is recorded in a mere 86.4% of them(462 tetrads)!
Unlike tap-rooted T. pratense, T. repens also grows quite frequently on upland moorland andthus it is likely that surviving, potentially untainted native populations of White Clover are only orchiefly found in remote sites. In Fermanagh, these include higher altitudes on Cuilcagh and Belmore andon obviously completely unsown ground, eg many wet or unapproachable lake shores.
Variation
A highly variable species with considerable differences within and between populations over a wide rangeof morphological characters, over the years T. repens has presented many difficulties in taxonomyand nomenclature. Genetically, T. repens is so extremely variable that it has been described as,"the Drosophila of plant ecology" by Turkington & Burdon (1983). T. repensis an allopolyploid species with chromosome number 2n=32.
Tutin et al. (1968) in Flora Europaea 2 and Burdon (1983) recognised six subspecies,of which only subsp. repens is considered native in Britain, the other five subspecies beingchiefly found in the mountains of S Europe. When it comes to varieties, some older ones claimed asnative in Britain (Erith 1924), such as var. rubescens Ser. ex DC. (with pink flowers) and var.sylvestre Alef. (white flowers), do not appear in any standard B & I Floras and fail toequate with other taxa listed by Sell & Murrell (2009). In their critical Flora of B & I, thelatter recognise four T. repens varieties: var. carneum Gray, corolla slightly or deeplytinged pink, a native found on heaths and by the sea; var. townsendii Beeby, corolla purple, arare form from the Isles of Scilly; var. repens, inflorescence up to 25 mm, corolla white, awidespread mix of native wild and escaped cultivated forms; and var. grandiflorum Peterm.,inflorescence 30-35 mm, corolla white, formerly grown for hay, now planted with wild flower seed.
Cultivation and agricultural benefits
Its vigorous growth, efficient nitrogen-fixing ability over a very wide range of ecological conditions,reliable long-season availability and nutritious quality as animal fodder, all taken together, have madeT. repens by far the most important cultivated pasture legume in these islands. The beneficialcharacteristics of T. repens include the very high quality of herbage protein and mineral contentand its high levels of acceptability and digestibility by grazing animals. These characteristic benefitsare retained throughout the growing season significantly better than all grasses, and also better thanother tap-rooted legumes including T. pratense (Red Clover) and Lucerne (Medicago sativa subsp. sativa) (Williams 1970). As forage for animals, T.repens compares favourably with grass, being higher in protein and certain key minerals and lowin structural fibre. Unlike grass it maintains a high level of digestibility as it matures (Thompson1984). Since it is easily and quickly digested and is very palatable, stock animals preferentially seekit out and eat a lot of it. However, if the proportion of White Clover present in clover/grass leyherbage exceeds 50%, there is a danger of cattle developing bloat, as the rapid fermentation that takesplace in normal ruminant digestion may release dangerous amounts of gas, which the animal cannot get ridof quickly enough (Davies 1992).
In the past, White Clover growth patterns and the nature of the interaction with grass have tended tocause significant seasonal variation of clover content in swards – from as little as 5% in the spring upto 60% in summer – but clover breeding is producing varieties that are more compatible with modernrye-grasses and have more even seasonal growth curves. Accumulated experience and scientific evidenceindicate that the optimum balance is achieved with a White Clover content of 30–35% of the total annualdry matter yield of the sward.
The most notable agricultural advantage of White Clover is its contribution to the nitrogen balance ofsoils. Agronomists reckon T. repens can contribute roughly the same annual amount of nitrogenelement (150-200 kg N/ha) to pasture soils as the national average application of fertiliser N tosheep-grazed pastures in the UK. Clearly this is dependent on the clover content of the sward however,which can vary greatly (Davies 1992).
Since about the 16th century onwards White Clover has been and remains extremely commonly sown in pasturemixtures with grasses. Over 400 metric tonnes of White Clover seed are sown annually in the UK (DEFRASeed traders annual return – year ended 30 June 2002). Unfortunately up-to-date comparison figures arenot available, since some years ago DEFRA stopped collecting annual measurements of agricultural seeduse. Around 75 different cultivars are listed for Europe (Burdon 1983).
Crossbreeding and its effects
T. repens is mainly outbreeding, so hybridization and introgression have taken place between theold, presumably native forms and the multitude of introduced cultivated varieties, crosses very probablycommonly occurring in areas of overlap. As a result we may never know the distribution of the'unpolluted' native species. Cultivated strains tend to be larger in all their parts and individualWhite Clover plants persist for only one or two years. These clover varieties and differing forms canseed themselves profusely however, even when growing in heavily grazed or disturbed ground and theresultant seeds become added to the long-persistent fraction of the soil seed bank present in improvedpastures and meadows (Duke 1981; Burdon 1983).
The seed of T. repens is small and light and may be transported both by wind (perhapsinsignificant) and internally by grazing animals which is probably highly significant (Ridley 1930).Thus any surviving native strains of clover would need to be extremely remote from farmland to avoidinterbreeding or competing with the widely used cultivated forms.
There is an enormous scientific literature dealing with many aspects of T. repens, especially onthe agricultural properties of the species since it offers unrivalled flexibility in its utilisation asa quality animal fodder (Turkington & Burdon 1983). The latest breeding development in White Cloverhas been the introduction of a hybrid variety between T. repens and Caucasian clover(Trifolium ambiguum), which is a drought tolerant, rhizomatous species. This new variety has bothstoloniferous and rhizomatous root structures that aid grazing tolerance, drought tolerance and winterhardiness (Marshall et al. 2004).
British and Irish occurrence
T. repens is almost ubiquitous throughout both B & I, but as noted above, distinguishingnative populations as opposed to cultivated forms and crosses between the two in the field is verydifficult or almost impossible (New Atlas).
European and world occurrence
The postulated centre of origin of T. repens s.l. is in the Mediterranean mountains of Europe fromwhere it has spread across the whole continent up to about 71°N and it now occupies around 97% ofEuropean territory, although in warmer southern regions, including the Alps and the Mediterranean basinit is confined to upland pastures and meadows. It is also considered native in NW Africa and N & WAsia and in phytogeographic terms is considered Eurosiberian boreo-temperate (Davies 1992). However, ithas been spread almost worldwide by its use from the 17th century onwards in agricultural grass seedmixtures. T. repens s.l. has become partly naturalised in most parts of the world, especially inN America and thus can now be recognised as circumpolar boreo-temperate (Hultén & Fries 1986, Map1235; Preston & Hill 1997).
Threats
Hybridization between native and cultivated populations will be increasingly likely, but selectionpressure in non-agricultural sites will continue to favour persistence, with the ability to coloniserelatively fertile, rather drier soils than the agricultural norm.
Introduction, neophyte, a very rare casual.
1901; Praeger, R.Ll.; Co Fermanagh.
Growth form and preferred habitats
Alsike Clover is a short-lived alien perennial with globular heads of white flowers that turn pinkas they age from the bottom of the inflorescence upwards and then go brown as they fade. In otherrespects, T. hybridum, which is a full species – and is not a hybrid between White Clover (T.repens) and Red Clover (T. pratense) as Carl Linnaeus erroneously thought – resemblesT. repens, but it has ± erect stems up to 60 cm that do not root at the nodes. In case you arewondering, the English common name 'Alsike Clover' refers to a small Swedish village where Linnaeuscollected the plant he named.
For several centuries, certainly from the 18th century onwards, Alsike Clover was regularly included inclover-grass seed mixtures planted for forage. However, when it was realised that it was far lessnutritionally valuable than T. pratense (Red Clover), its use rapidly declined from the 1930sonwards. T. hybridum is scarcely used nowadays in agriculture in B & I, probably onlyoccurring in pastures and meadows as a seed impurity.
Ecologically, Alsike Clover resembles T. pratense (Red Clover) in many respects, but is somewhatmore adaptable and better able to tolerate wet, acid, infertile conditions. It is also more resistant todiseases like Clover Rot and Stem Eelworm than is T. pratense; in grass-clover mixtures it isconsidered non-aggressive, although it produces lower yields than Red Clover (A. Smith, in: Spedding& Diekmahns 1972, p. 419). The established strategy of T. hybridum as measured by the indexdeveloped by (Grime 1979) is described as C-S-R, meaning that it demonstrates a balance of thecharacteristic behaviours of Competitor, Stress-tolerator and Ruderal forms of plant natural selectionaffecting growth, reproduction and survival (Grime et al. 1988, 2007).
In comparison with most other legumes, T. hybridum does poorly on dry, sandy or gravelly soils,but does very much better than most on heavy silt or clay where there is plenty of moisture. T.hybridum can also survive flooding that would kill most crops and has been known to survive fora year and make extensive growth in water-covered soil (Duke 1981).
Perhaps on account of these tolerant properties, planting of Alsike Clover in Britain and Ireland hasrevived considerably in the last few decades and it has become a frequent constituent of amenitygrass-legume mixtures and wildflower seed mixtures that are widely used for reseeding open habitats,including on roadside banks or for reclaiming less-than-ideal land (Sinker et al. 1985; Reynolds2002; Crawley 2005). The form used is the hollow-stemmed subsp. hybridum and much of the seedsown in the 20th century and at present is imported from Canada where the plant is still used as a crop(Chater 2010).
Variation
There are two subspecies recorded in B & I, of which subsp. hybridum is the much commonercultivated from: it has hollow stems, grows erect, is sparingly branched and has the largerinflorescence, over 20 mm in diameter. The other rarer taxon is subsp. elegans (Savi) Asch. &Graebn., which in contrast has solid, decumbent, much branched stems and bears an inflorescence lessthan 20 mm in diameter (Sell & Murrell 2009).
Flowering reproduction
Since the species has no means of vegetative spread or reproduction, it relies entirely on seedproduction for its increase, dispersal and survival. T. hybridum flowers from June to Septemberand the flowers, being self-incompatible, require cross-pollination by bees. The legume pods, 3-4 mmlong, are ± included within the calyx and they contain 2-4 seeds of varying colour. The pods areindehiscent, eventually falling off the inflorescence head to land at the base of the plant.
Although the individual plant is usually considered casual, failing to generate established populations,any seed that it manages to produce is capable of long-term survival buried in the soil (Thompson etal. 1997). The life-span of the individual Alsike Clover plant, which possesses a shortroot-crown, is variously quoted as being between three and six years (A. Smith, in: Spedding &Diekmahns 1972, p. 419). In agricultural sowings, however, T. hybridum is generally treatedeither as a biennial legume (Duke 1981, p. 243), or as an annual. Persistence in any event requiresrepeated sowing (D.A. Pearman, in: Preston et al. 2002).
Fermanagh occurrence
There are only three records of T. hybridum in the Fermanagh Flora Database. Apart from Praeger'scompletely site-unspecific record above, the other two were made by Ian and David McNeill in the NE ofthe county. The record details are: Drumbrick Td, 2 km N of Ederny, 1985, I. McNeill; Glen Lodge, 3 km Nof Ederny, 1986, D. McNeill.
Irish occurrence
T. hybridum is still fairly widespread in Ireland, especially in the north on open road- andtrack-sides and on waste ground and it seems to be reduced to the status of an uncommon casual of suchhabitats, typically occurring as single plants or small patches (Cat Alien Pl Ir). However, itcan also feature (again as a casual), in more damp or wet situations along other linear habitats,including beside rides and paths in conifer plantations and along river banks (Green 2008). In the late19th and early 20th centuries, however, thanks to repeated agricultural sowings it was a frequent alienescapee in waste places in much of Ireland (Irish Topographical Botany). Occasionally it becameestablished (particularly in the NE) and naturalised itself in open, dry, disturbed, wayside soils(FNEI 2; FNEI 3).
In the 1980s T. hybridum was much more often reported in the S & E of Ireland than elsewhere,while in the north at this period it became reduced to a rare casual, possibly of bird-seed and grainimpurity origin, confined to open, sunny, ruderal habitats (FNEI 3).
The botanical survey of urban Belfast recorded it from 18 1-km squares in the city. It was most common inthe central area of the city, nearest the docks, where grain and other imported seed was beingtransported in open-topped lorries (Beesley & Wilde 1997).
British occurrence
Frequent and widespread throughout lowland England and Wales, becoming more thinly scattered,scarce to rare further north, especially in N & W Scotland (New Atlas).
European and world occurrence
T. hybridum is regarded, possibly erroneously, as a native of N Europe, apparently simply on thegrounds that Linnaeus (1745) mentions it in his Flora Suecica, as growing abundantly in theparish of Alsike, about ten English miles S of Upsala (Britten & Holland 1886). Whatever the site oforigin, which is disputed, the species now occurs throughout Europe, reaching C Asia and Asia Minor. Ithas been introduced worldwide with agriculture, either deliberately or as a crop seed impurity (Duke1981; Gillett 1985) so that now it is circumpolar (Hultén & Fries 1986, Map 1236).
Threats
None.
Native, very rare and probably only casual. Eurosiberian southern-temperate, but widely naturalised,including in N America and New Zealand.
1901; Praeger, R.Ll.; Co Fermanagh.
Growth form and preferred habitats
Hop Trefoil is a very variable, small, low-growing, winter-annual clover with fibrous roots. It variesparticularly in habit, leaf and inflorescence characters. The pale green stems can vary enormously inlength from 3-30(-50) cm and their posture ranges from prostrate or sprawling to erect. The stems arethinly clothed with white appressed hairs and are usually much branched, especially towards the base,but they fail to root at the nodes so the species has no means of vegetative spread and reproduction.
Essentially, T. campestre is a ruderal, stress-tolerant species of relatively infertile,unproductive grassland on dry, sandy or gravelly, often lime-rich, near-neutral soils. It is chieflyfound in grasslands and dry waste places. In NI, the majority of its stations are coastal, or occuraround Lough Neagh, which is the largest expanse of inland freshwater in the whole of B & I.
Variation
Sufficient variation exists for Sell & Murrell (2009) to recognise and list three varieties basedchiefly on stem length and inflorescence diameter: var. minus (W.D.J. Koch) Gremli has stems upto 12 cm, spreading or prostrate with short branches and inflorescences up to 10 mm in diameter; var.campestre has stems up to 30 cm, ascending or erect, with longer, slender branches andinflorescences again up to 10 mm in diameter; var. majus (W.D.J. Koch) P.D. Sell has stems up to50 cm, usually erect and usually much branched with larger inflorescences up to 17 mm in diameter. Noneof these have been seen or distinguished in Co Fermanagh.
Flowering reproduction
Hop Trefoil is characterised by very rapid spring growth, yet it does not begin flowering until late Mayor June, from when it continues for four months into September. In common with most small cloverspecies, the tiny, 4-7 mm, pale yellow flowers borne in inflorescence clusters of 20-40 flowers, withpedicels about as long as the calyx tube can to some extent self-pollinate. However, they also attractflies, bees, butterflies and moths, enabling some degree of out-breeding (Fitter 1987). When the flowersare fertilised the petals wither, turn brown and scarious (dry, membranous) and persist, covering thedeveloping legume pods. The pod is c 1.3 mm long, oblong-ellipsoid in shape and compressed. Each podcontains just one yellowish-brown, oblong seed (Meikle 1977; Webb et al. 1988; Sell & Murrell2009).
The ecological established strategy of T. campestre is described as R/SR by Grime et al.(1988, 2007), meaning it has properties intermediate between a straightforward ruderal and astress-tolerant ruderal, which means it has a rapid growth rate and population turnover in bare ground,often under physical and chemical stress, without it having much in the way of competitive or colonisingability with respect to accompanying plant species. Hop Trefoil is somewhat more ecologically demandingthan T. dubium (Lesser Trefoil) and, being the larger plant of the two, as a component of afodder crop it provides a greater yield and produces a more nutritious animal feed than Lesser Trefoil.
Seed dispersal
The solitary seed is slightly smaller and considerably lighter than that of T. dubium (Gillett1985). Dispersal is sometimes assisted by the activities of Harvester Ants, but observed clusters ofseedlings suggest that seed dispersal is inefficient since it usually lacks transport away from theparent plant. One exception to this is internal transport of T. dubium seed in the gut of cattle,along with a long list of other species reported from Sweden by Heinitz, quoted in Ridley (1930, p.361). The main method of species dispersal around the globe is, of course, as part of agriculturalfodder crop seed mixtures, which has enabled T. campestre to travel widely across bothhemispheres (see below).
Fermanagh occurrence
The first report of T. campestre (as T. procumbens L.) occurring in Fermanagh appearsnon-specifically in Irish Topographical Botany (ITB), where Praeger (1901) described itsoccurrence as, "Divisions all, except 29 Leitrim, where it, no doubt, will be found.Frequent." The reference to 'divisions' here refers to Praeger's 40 vice-county subdivision of theisland still used today, and not to the twelve botanical divisions previously used in both editions ofCybele Hibernica (More & Moore 1866; Colgan & Scully 1898).
For Fermanagh, the basis of Praeger's ITB statement apparently derives from a Royal IrishAcademy paper by Dr G. Sigerson (1871), listing, 'Additions to the Flora of the TenthBotanical District, Ireland'. This tenth division comprised five counties including Fermanagh,but as stated in his introductory remarks, Sigerson was mainly interested in recording Tyrone (H36).This is the context in which he first records Hop Trefoil on, "gravelly and dry banks". Thefollowing year, District 10 is listed for Hop Trefoil in the first supplement of Cybele Hibernica(More 1872), adding, "Frequent in Tyrone; Dr Sigerson". Praeger (1901) must have believed thatSigerson found T. procumbens (= T. campestre) in all five counties of 'District 10', butas indicated here, there is no real basis for this notion. Dr Sigerson may have been correct about HopTrefoil in his day, but nowadays T. campestre is rare in Tyrone and is confined to the east ofthe VC, towards Lough Neagh. In his 2010 The Flora of County Tyrone, Ian McNeill lists just threerecords from: "Roan (near Coalisland), IMcN & RI 1998; Aughlish (W of Castlecauldfield), IMcN2003; and Killens (ENE of Aughnacloy), IMcN 2008."
Despite the very confident, rather sweeping claim by Praeger in ITB, only three other Fermanaghrecords of this species exist. Of these, the first two were made by Meikle and co-workers and date from1951: "Roadside near Derrysteaton Lough" and "Roadside near Springfield, NE of Boho"(Meikle et al. 1975). The third record is the Kilmore South-Derrychaan Td shore of Upper LoughErne, 6 August 1986, recorded by S.J. Leach and A.S. Mullin, then surveying for the EHS Habitat SurveyTeam.
R.H. Northridge and the current author (RSF) have never seen T. campestre in Fermanagh. Thepossibility of confusion with the much more common T. dubium (Lesser Trefoil) cannot be totallyruled out, but it seems likely that the Fermanagh recorders concerned were not mistaken and that T.campestre really is a very rare, casual species in the county.
Irish occurrence
It is sometimes difficult to make any valid comment about a species with very few Fermanagh records, butin this case RHN and the current author (RSF) can say that T. campestre is always a rare speciesat inland sites this far north in Ireland. The Cen Cat Fl Ir 2 (Scannell & Synnott 1987)follows the information given in Cybele Hibernica 2 (Colgan & Scully 1898) in suggesting thatT. campestre is common and widespread, having been found in every Irish district and VC. Thistreatment gives a completely wrong impression of its current distribution, since in reality there arevery few Fermanagh records and, indeed, apart from around Lough Neagh, few inland records in existencein NI, the species being decidedly coastal in N & W parts of Ireland. It is only in the Midlandplain and in lowland areas of SE Ireland that T. campestre has anything like a frequentoccurrence and, even then, it is scattered and much less continuous than appears the case in England andWales (New Atlas).
British occurrence
T. campestre is widespread and common in lowland England and Wales, absent from mountain areas andbecoming more confined to coastal and disturbed ground further north and west in England and especiallyso in Scotland. The species has been in decline for many years for reasons that are not clear. The indexof change between the two BSBI Atlas surveys 40 years apart is calculated as -0.45, indicating a declinethat is particularly noticeable in the N & W (D.A. Pearman, in: Preston et al. 2002). Theoverall B & I distribution suggests the species is limited by low winter temperatures and a similarpattern is apparent in continental Europe, although at least as an agricultural introduction, the plantextends much further N & E than one might expect (Hultén & Fries 1986, Map 1240).
European and world occurrence
T. campestre is considered native and is widespread throughout most of Europe withthe exception of the N & E. As a native, it reaches S Norway and S Sweden, but it has beenintroduced further north in Scandinavia and eastwards into NW Russia. T. campestre is alsopresent and considered native in Iran, W Asia, N Africa and Macaronesia. It has been widely introducedwith agricultural seed around the world, including in N America, Chile, parts of Africa includingEthiopia and Cape Province, Madagascar and New Zealand (Hultén & Fries 1986, Map 1240). It will havenaturalised in some of these regions (Meikle 1977).
Threats
None.
Native, common, widely scattered. European temperate, but widely naturalised in both hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
April to November.
Growth form, recognition and preferred habitats
A frequent to common, small, creeping, ruderal, winter-annual with fibrous roots and stems varying from3-30 cm in length, T. dubium is another of the diminutive, trefoil-leaved plants commonly used on17th of March (St Patrick's Day) as the 'Shamrock'. This little nitrogen-fixing legume is typicallyfound in short, disturbed grassland such as lawns, roadside verges and banks, waste ground and other dryor well-drained, open, sunny, mainly lowland sites including in quarries and on the tops of walls.
In freshly sown lawns, T. dubium can be a troublesome weed in the early days as it forms aninterlacing mat of thin wiry stems prior to the grasses tillering up and forming a proper turf that willsoon competitively exclude it. Having said this, T. dubium is renowned for its ability totolerate mowing and trampling. Like its close relative, T. campestre (Hop Trefoil), theecological established strategy of T. dubium is described as R/SR (Grime et al. 1988,2007), meaning it is intermediate between a straight ruderal and a stress-tolerant ruderal, ratherlacking in competitive ability, probably on account of the usually rather dry, low nutrient soilconditions it appears to mainly tolerate and frequent.
Despite its preference, or rather its tolerance for drier, sometimes droughted, infertile soils, LesserTrefoil does also occur quite commonly in Fermanagh on damper, winter-wet, lakeshore grasslands, whichmay sometimes be moderately fertile. In any such pasture conditions, T. dubium is also remarkablytolerant of grazing and trampling pressures (Sinker et al. 1985).
In flower, T. dubium could be confused with Medicago lupulina (Black Medick), butvegetatively the latter has leaves that are usually densely hairy and apiculate, ie with a small, abruptpoint in the notch at the leaf tip, whereas T. dubium has sparsely hairy to glabrous leaves thatlack any apiculus (Rich & Jermy 1998).
T. dubium might possibly also be confused with T. campestre, but it is smaller in all itsparts than the latter and has fewer flowers per head (only 10-15 in T. dubium compared with 20-40in T. campestre). Also T. campestre is mainly coastal in NI and it is very rare inFermanagh, while by comparison T. dubium is extremely common and widespread.
Clover evolution and variation
In a review of the question of pasture clover species origins, Abberton (2007) commented, "Thereappears to be no significant role for interspecific hybridization in the evolution of the genus."Interspecific hybridization in the genus Trifolium by conventional crossing techniques has beenlargely unsuccessful and there currently are no known clover hybrids anywhere in the flora of B & I(Stace et al. 2015). Post-zygotic barriers appear to be a primary cause of the reproductiveisolation and these are associated with endosperm disintegration and consequent abnormal differentiationand starvation of the hybrid embryo. Evans (1976) postulated that the rarity of wide hybridization inthe ten main Trifolium forage legume species may well be associated with their predominantadaptation to insect pollination, although the annual clovers at least are often capable of additionalself-pollination. Ellison et al. (2006), in a comprehensive DNA based phylogenetic analysis,found only five or six instances of apparent hybrid speciation in fodder clover species.
Various chromosome counts have been obtained for T. dubium ranging from 2n=14, 16, to 28 orhigher. Other work suggests Lesser Trefoil is an allotetraploid with a chromosome number 2n=32 (Sell& Murrell 2009). A similar chromosome count of 2n=30 is given in a study testing the idea that T.dubium arose from the crossing of two diploid clover species of similar world distribution,T. campestre (2n=14) and T. micranthum (Slender Trefoil) (2n=16), followed by chromosomedoubling by means of unreduced gametes (Ansari et al. 2008). T. dubium is said by theseworkers to be morphologically intermediate between these two parent species.
The current author (RSF) could only access a summary of the Ansari et al. (2008) paper since it isprotected by a paywall, but it appears to involve a number of the same research workers and followssimilar genetic phylogeny analyses using evidence from DNA sequence analyses, molecular cytogenetics,interspecific hybridization and post-hybridization experiments as a subsequent freely available study onthe origin of T. repens which indicated a hybrid origin for White Clover that involved multiplecrossings of T. pallescens Schreb. (Pale Clover) and T. occidentale D.E. Coombe (WesternClover). These crosses were probably associated with the major species migrations southwards intorefugia during past glacial climatic phases (Williams et al. 2012). It appears, therefore, fromthe two examples mentioned, that hybridization between clover species was possible and did occur somethousands of years ago, although this is no longer the case since barriers to embryo survival havesubsequently arisen.
In B & I, T. dubium is sufficiently variable for two varieties to be recognised by Sell &Murrell (2009): var. microphyllum (Ser.) P.D. Sell is a dwarf form with stems up to 10 cm;spreading or prostrate leaves small, mostly under 5 mm; and inflorescence also small, up to 7 mm indiameter. The alternative form, var. dubium has stems up to 30(-45) cm, ascending or erect,leaves up to 12 mm, and inflorescence up to 10 mm in diameter.
Flowering reproduction
T. dubium seed germinates in the spring and the species flowers, often profusely, from May toOctober. The individual plant produces large numbers of axillary, subglobose, inflorescence heads, 5-10mm in diameter, each usually containing 3-15 yellow pea flowers, the corolla 3-4 mm in length. Afterfertilization, either by insect-pollination (Fitter 1987) or, as is more usual in small clover species,self-pollination (Proctor & Yeo 1973), the corolla turns brown and the standard petal folds itselfdown to cover and partially conceal the developing legume fruit pod. The legume is 2.5-3 mm in length,ovoid in shape and it is usually single seeded (Clapham et al. 1987). Like related legumes, seedsdevelop hard coat dormancy and can survive soil burial for many years (Thompson et al. 1997).
Fermanagh occurrence
Lesser Trefoil has been recorded in 181 Fermanagh tetrads, 34.3% of those in the VC. As the tetrad mapshows, it is widely scattered everywhere around Fermanagh except on high ground. Although most typicalof open, disturbed sites on damp to dry, relatively infertile soils, T. dubium does also occur onlimestone cliffs, as at Knockmore Hill and Trien Mountain and by tracks in lime-flushed areas of uplandblanket bogland, eg at Black Bridge in the W of the VC.
British and Irish occurrence
T. dubium is common and widespread throughout B & I in suitable lowland grassymeadows, lawns and waysides, open waste ground and rocky sites. It is absent mainly from acid, peatymountain sites in Scotland and is introduced and naturalised in Shetland (VC 118). The New Atlashectad map shows there is no significant change in T. dubium presence in the 40 years between thetwo BSBI Atlases (D.A. Pearman, in: Preston et al. 2002).
European and world occurrence
Lesser Trefoil is widespread in Europe stretching from the Atlantic Isles eastwards to N Iran andnorthwards to S Scandinavia. It has been widely introduced with other agricultural clover and grass seedmixtures both in Europe and beyond to places as far apart as Iceland, the African Cape Province, SAustralia, Tasmania, New Zealand and N America. Thanks to its long-lived seed, it is naturalised in someof its European and more outlying areas (Hultén & Fries 1986, Map 1241).
Threats
None.
Native and introduced, common and widespread, locally abundant.
Eurosiberian temperate, but widely naturalised and now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year, peaking in April.
Growth form and preferred habitats
Red Clover is a wintergreen, deeply penetrating, tap-rooted, short-lived perennial legume of leafy,tufted habit, 5-100 (usually 30-70) cm tall, common and widespread on grasslands on and off thelimestone, including pastures and in silage and hay meadows. It is also common but sometimes sparse inrough, open areas on roadsides and waste ground, on damp or marshy lakeshores, rock outcrops andlightly-shaded, artificial, disturbed sites and waste ground below about 360 m. Despite the wide rangeof habitats occupied, T. pratense is most frequently found on ± unshaded, productive, lowlanddisturbed ground and it reaches its highest altitude in B & I at around 850 m in the ScottishHighlands (D.A. Pearman, in: Preston et al. 2002). T. pratense can grow on all types ofsoil except strongly acidic peatland (ie below about pH 4.5), waterlogged ground, or in permanentwetlands.
This was an extremely important agricultural grassland legume for pasture, silage, hay and green manuresoil improvement and it is second only in importance to T. repens (White Clover). Clovers arecomponents of natural grasslands and they are almost always cultivated in association with companiongrasses or with crop species such as Brassica rapa (Wild turnip) in simple or complex seedmixtures supplied by the agricultural trade. The proportion of clover is usually between 30-50% in theseseed mixtures. The principal ley grassland species used with Red Clover are Timothy (Phleumpratense), Meadow Fescue (Festuca pratensis) and Perennial Rye-grass (Loliumperenne).
The main value of clovers to the farmer lies in their ability to fix atmospheric nitrogen through theRhizobium bacteria which inhabit their root nodules (Evans 1976). This fixed nitrogen is the maindriver for the growth of the accompanying grass or crop species. The nodules of T. pratense var.sativum are such efficient nitrogen-fixers, the species becomes heavily suppressed if additionalnitrogen fertiliser of any sort (organic or inorganic) is unnecessarily applied (Grime et al.1988, 2007). In addition to their nitrogen capturing ability, clover plants are also rich in protein,minerals and trace elements contributing to their fodder value (Evans 1976). Around the world, about 25Trifolium species are of significance as food for grazing animals and, of these, about ten areagriculturally important.
Variation
T. pratense shows considerable genetic variation and as a species has wide ecological tolerances,although it is not strongly competitive, especially in fertile, wet, or shaded conditions. In commonwith most VCs in B & I, the Fermanagh database records do not distinguish the two widely recognisedvarieties of Red Clover, the native var. pratense (diploid with 2n=14 chromosomes) and thecultivated var. sativum Schreb. (of which there are at least ten strains currently inagricultural use in these islands). It is likely, however, that the larger, more vigorous, erect,short-lived (surviving 2-3 years only), cultivated variety (which is hollow-stemmed and has less heavilytoothed leaflet margins) is nowadays more common in all types of lowland grassland compared with thelonger-lived native var. pratense (Stace 1997). The numerous cultivars of var. sativuminclude both diploid and tetraploid chromosome forms (2n=14 & 28) and, in terms of flowering, theyvary from early to intermediate and late-season types. Polyploidy is not common inTrifolium and most species are diploid with 2n=14 or 16. T. repens is, however, atetraploid with 2n=4x=32, but it shows regular bivalent formation at meiosis and thus is anallotetraploid, formed by spontaneous chromosome doubling after hybridisation.
Both of these T. pratense varieties are less tolerant of heavy grazing and trampling pressure thanT. repens and T. pratense var. sativum is probably the more sensitive variety dueto its erect growth habit and hollow stems (Grime et al. 1988, 2007). Before the expansion ofsilage making, Red Clover was used in ley mixtures with several grass species, mainly for cutting as ahay legume. While it is quick to establish and releases nitrogen to the grasses in its first year andsubsequent years, it is a tall plant and only survives for two or three years. On the other hand, whensown with grasses, White Clover is slower to establish and does not release much nitrogen until itsthird or fourth year of growth. However, on account of its low, creeping nature, it persists and canbecomes a long-lived meadow or pasture legume in permanent grassland (Dr John Orr, pers. comm., 2 May2021).
Alsike Clover (T. hybridum) is another perennial that can replace T. pratense on more acidsoils in N Europe and in Canada. T. dubium (Lesser Trefoil or Yellow Suckling Clover) is anannual that is sometimes included in clover-grass seed mixtures in B & I for short leys, where itmay persist by self seeding. The three perennial clovers mentioned are all mainly or almost entirelyself-incompatible and they require insect-pollination to set seed (Evans 1976).
The decline of Red Clover on most (but not all) farms: Before the widespread use of nitrogenfertilisers, ie post-World War II, Red and White Clover were the main source of nitrogen for grassswards (the soil releases a small amount into the atmosphere each year) and both clover species wereincluded in commercial grass seed mixtures to give 'mixed swards'. Red Clover grew well in the first twoyears and its upright growth habit made the sward suitable for haymaking.After the second year,the Red Clover died out and the White Clover took over the provision of nitrogen as the driver of grassgrowth.T. repens was less suitable for haymaking because the tall grasses tended to shadeit out, so the sward was grazed and was converted into long-term pasture.These clover/grass seedmixtures were known as 'hay and grazing mixtures' and they were the basis of grass farming for manyyears (Dr John Orr, pers. comm., 2 May 2021).
Post-World War II, the introduction of 'artificial' nitrogen fertilisers (NPK), meant that intensivefarmers (mainly dairy farmers) could get a 'quick fix' for growth in the first year of the sward andsilage making meant they did not need hay, so the use of Red Clover declined rapidly.White Cloverwas retained because its spreading growth habit meant it filled in spaces in the sward and gave someyield, but its nitrogen fixation was very little since it was suppressed by the NPKfertiliser.Less intensive farmers (mainly cattle and sheep) used less nitrogen fertiliser andstill used White Clover for some nitrogen fixation.
In the first eight decades of the 20th century, vast quantities of commercial strains of Red Clover seedwere sown by farmers each year in B & I. In Britain and NI, the Ministry of Agriculture, Fisheriesand Food (MAFF) estimated, in figures released in 1984, a seed weight use of around 200 metrictonnes/year. As a result of its heavy sowing rates, var. sativum was a frequent escapee fromcultivated fields and was most commonly found on fertile disturbed ground near the agricultural fieldswhere it was being repeatedly sown (Grime et al. 1988, 2007). However, from the late 1940sonwards, there was a gradual move on farms towards developing intensive managed and fertilised pasturesand meadows using applied agrochemical NPK fertilizers. This was accompanied by a move away from hayproduction towards the use of fermented grass silage as animal fodder. This happened in tandem with anincreasing application of farm-produced cow and pig slurry, sprayed on grassland as a cheaperalternative to expensive chemical fertiliser mixtures produced by the petrochemical industry. Cloverspecies, and especially short-lived Red Clover, could not tolerate additional N fertiliser being addedto the soil as it heavily depressed their growth. Furthermore, T. pratense fermented poorly insilage, the leaves breaking down excessively, so it quickly became redundant (Morrow 2010; pers. comm.J. Crea, May 2021).
The increase in organic farming in recent decades renewed the interest in Red Clover.Organicfarmers cannot use nitrogen fertiliser (its manufacture uses a massive amount of energy) and theytherefore rely on clovers for their grass production. They found that Red Clover, mixed with suitablevarieties of grass and properly managed, could produce large quantities of silage (as much as farmersusing large amounts of sprayed chemical or organic fertiliser). The organic clover/grass ley was also ofhigher nutritive value than ordinary silage, because of the additional protein provided by the T.pratenseand it was made without any nitrogen fertiliser.Some conventional farmerstried to grow T. pratense, but they often did not know how to properly manage it, so there hasbeen very limited uptake outside organic farming (pers. comm. J. Orr, May 2021).
The cultivated clover strains most often grown by farmers are early-flowering ones and they tolerate orsurvive at least two hay or silage cuttings per growing season (Proctor & Yeo 1973).
The downside with Red Clover swards is that they generally only last around three years and expensiveploughing and cultivations are required to establish new ones.As a result, organic farmers have toplough more often than traditional grass farmers. However, there has been sufficient demand for T.pratense seed in recent years that one seed supply company has developed a variety called'AberClaret' that can persist for up to five years. This appears to be a good step forward and is anindication of expected growth in this seed market (pers. comm. J. Orr, May 2021; https://germinal.co.uk/top-tips-for-growing-red-clover/; https://www.dlf.co.uk/forage-grass-seed/species/dlf-uk/clovers, both websitesaccessed by RSF in May 2021).
Clover-Herbal leys
Another innovation which should increase demand for Red Clover seed is the development of herballeys.These are complex mixtures of deep rooting pasture grasses, clovers and herbs, principallyChicory (Cichorium intybus) and Ribwort Plantain (Plantago lanceolata) and they havepromising features including resilience to drought, improved animal health and greater biodiversityabove and below ground.
The mixture of plants contained in the herbal ley brings multiple benefits. Red Clover, for instance, hasa deep root structure and can fix nitrogen into the soil. A potentially large herb like Chicory providesan even deeper root, mining the soil for minerals, improving the life and health of the soil. Mixescontaining Sainfoin (Onobrychis viciifolia), Bird's-foot-trefoil (Lotus corniculatus) andChicory also have natural anthelmintic properties and the growing times of the different plants alsohave the great advantage of providing year-round forage. It is standard agricultural practice to lightlygraze a herbal ley in the year of sowing to control annual weeds and to thicken the sward. It is alsoimportant not to over-graze too early after sowing and to leave plenty of leaf on the sward to provideground cover and prevent weed colonisation of vegetation gaps.
Over the years, there has been a commitment to adapting the rotation to fully incorporate herbal leys,which are left in place for 2-4 years to really build root mass and organic matter in the soil. Thegrasses in the herbal ley typically include Dactylis glomerata (co*ck's-foot), Festucaarundinacea (Tall Fescue) and Festuca pratensis (Meadow Fescue), that have very good rootsystems which create pore spaces and a large amount of root mass in the soil that breaks down over along time. Plants in the mix that have anthelmintic properties (ie natural wormers) benefit livestockhealth. The legumes in the mix – Clover, Bird's-foot-trefoil, Lucerne (Medicago sativa) andSainfoin – are used to fix atmospheric nitrogen, reducing the reliance on inputs. Forage herbs with deeproots like Chicory, Burnet (Sanguisorba minor), Yarrow (Achillea millefolium) and RibwortPlantain mine trace elements from further down the soil profile. This system of soil improvement hasmade land easier to work, requiring less cultivation. It also increases the soil's resilience toextremes of weather, causing it to act as a sponge after heavy rainfall, yet hold moisture in drysummers. The herbal leys are usually undersown into a cereal such as winter rye or spring barley toprovide a cash crop whilst the ley develops, which reduces the amount of time land is left out ofproduction. The herbal ley is left down for between two and four years – longer when required to controlweeds, improve soil structure and drainage, before returning to three years of cereal crops.
It is important to choose the right combination of species for the right situation. The diversity ofingredients in a herbal ley means that mixtures can be tailored to suit individual soil types andcirc*mstances and can be matched to a farmer's particular needs or preferences. For instance, SimpleHerbal is an entry level herbal ley which can be used as a stepping stone to more complex mixtures,while the Heavy Land Herbal is tailored to wetter and heavier soils.
Although Red Clover can be grown as a monoculture, most farmers incorporate it into a mixture withLolium perenne (Perennial Ryegrass). This combination ensures both the clover and grass reach theoptimal stage to silage at the same time. As Red Clover can in some circ*mstances be slower to establishand grow, it is not recommended to mix it with fast growing L. multiflorum (Italian Ryegrass(IRG)), as this can result in the T. pratense being overwhelmed. IRG will also be at or past itsbest when the clover is ready to ensile, reducing the grass quality. Leys which include Red Clovershould always be part of a rotation to control stem eelworms and Sclerotinia (Clover Rot). Theseparasites and disease are always present in the soil, but as Red Clover is a host plant, regular sowingcan result in a substantial build-up. To prevent this, a six- to seven-year rest period from Red Cloveris required, the break allowing the populations of these pests to reduce before reintroducing T.pratense.
Harvesting leys that include Red Clover must be done carefully. Much of the protein content of Red Cloveris in the leaf, so it is important to prevent leaf damage during harvest in order to maintain thenutritional value of the silage. Mowing needs to be done as gently as possible since the clover leavescan be brittle and may shatter. Turning the crop when there is moisture on it, such as during earlymorning dew, also helps prevent leaf breakage. The cutting height is also very important, going no lowerthan 7-10 cm avoids taking out the crown of the plant. If the crown becomes damaged, the cloverpersistency in the field is severely compromised (https://www.cotswoldseeds.com/ accessed by RSF in May 2021).
Species variation recorded in wild habitats in B & I. Five varieties of T. pratense arelisted by Sell & Murrell (2009) as occurring beyond cultivation in B & I, the other three notyet mentioned here being var. parviflorum Bab., stems 15-35 cm, with few appressed hairs; var.villosum Wahlb., stems up to 20 cm, with dense, long, appressed hairs on the upper stem; and var.americanum Harz., stems up to 80 cm, with numerous stiff, spreading hairs. These are quite rareforms, but they occasionally appear in so called 'wild flower seed mixtures' commonly sown by councilson roadsides and increasingly by gardeners.
The degree of persistence of any of these escaped cultivars is not known, nor the extent of hybridizationand introgression between the two most common varieties. As with T. repens, these are examples ofpossible research studies the results of which would certainly be interesting to know.
Fermanagh occurrence
In Fermanagh, T. pratense is slightly less frequent than T. repens in terms of recordnumbers. However, in contrast, it is slightly the more widespread of these two commonly cultivatedclover species in terms of tetrad distribution, being found during the Fermanagh survey in a total of470 squares, representing 89% of those in the VC. In fact Red Clover ranks as the 31st most frequentlyrecorded vascular plant in the local Flora Database.
Phenology and flowering reproduction
Being a tufted perennial, Red Clover does not reproduce vegetatively, but instead relies completely onseed for long-term survival and dispersal to fresh sites. T. pratense is wintergreen, but it islate spring before the plant puts on any growth and flowering does not begin until May. However,anthesis may continue into September, depending upon any grazing or cutting it undergoes. The flowersare borne in large, round, or subglobose, dense sessile inflorescences 15-40 mm in diameter. Eachinflorescence consists of up to 100 reddish-purple to pink (rarely cream or white) almost stalklessflowers, subtended by two sub-opposite, reduced, bract-like leaves with very broad stipules (Sell &Murrell 2009). The flowers have a lovely honey-like fragrance and the foliage give off a pleasant cloverscent.
The flowers are highly self-incompatible and require insect pollinator visits to effect fertilisation.Insect visitors to Red Clover blossoms need a long proboscis to reach the nectar concealed at the baseof the tubular flowers. The proboscis must be longer than that of the European Honey-bee (Apismellifera), which cannot reach the nectar, but can nevertheless transfer pollen which itcollects to feed its larvae and, in doing so, effect flower pollination.
The natural pollinators of Red Clover are bumble-bees of various species, whose importance seems to varywith the corolla tube length of the clover cultivar being grown: short, medium or long. In an Englishstudy, the worker bees of the short-proboscis Bombus terrestris group were found to be persistentcorolla-biting, nectar robbers, though less troublesome in this respect on the shorter-tubed Red Cloverflowers than on the longer-tubed strains (Hawkins (in Mittler 1962) in Proctor & Yeo 1973, p. 346).While bumble-bees are essential for the pollination of the longer-tubed cultivars of Red Clover andparticularly for the valuable tetraploid cultivars which have larger flowers than the diploids, honey-bees may be used in some countries as pollinators of the remaining clover cultivars, and hives are hiredfor this purpose. They are more effective on the flowers of the second flush of the season, since theblossom of later inflorescences tend to have shorter corolla tubes and perhaps the nectar is alsosweeter and more abundant in warm, dry weather conditions (Proctor & Yeo 1973).
After fertilisation, the one- or two-seeded fruit pod develops (usually a solitary seed) and iscompletely concealed by the persistent calyx and withered brown corolla (Meikle 1977). The podeventually splits (dehisces) to release the seed, or else the fruit is consumed by a grazing animal,such as a horse, cow, elk or bird (eg members of the Crow family) (Ridley 1930) and the seed is guttransported and released in dung. Seed is long persistent in the soil, some surviving for five or moreyears (Thompson et al. 1997).
its most usual modern day a
T. pratense is frequent to common and widespread throughout both B & I,except in the Scottish highlands and on the most acidic boglands of western Ireland and N & WScotland. There has been no significant change in the hectad T. pratense distribution (includingboth the wild form and cultivated varieties) between the two BSBI atlases of 1962 and 2002 (D.A.Pearman, in: Preston et al. 2002). T. pratense var. pratense is the commonest formon non-agricultural ground in B & I, while var. sativum has spread out beyond the fieldboundaries where it was previously sown and became widely naturalised in both islands. In agriculture,despite something of a revival on organic farms, T. pratense is not quite as much used now aspreviously was the case. Beyond the farm gate it is probably in decline to some extent, althoughnowadays it is introduced in so-called wild flower seed mixtures that are frequently and increasinglysown by local councils on roadside banks, verges and in also in gardens by private individuals keen toconserve insect visitors (Sell & Murrell 2009).
s for both colour and flavour
As a polymorphic species with many varieties, T. pratense was originally restricted to Europe andadjacent parts of Africa and Asia until it became a forage crop widely introduced worldwide. It is nowto a varying extent naturalised in S Africa, E Asia, N & S America, S Australia and New Zealand.Taken in the broad s.l. sense, it is now circumpolar temperate (Hultén & Fries 1986, Map 1245).
Other uses
Apart from its most usual modern day application in agriculture as nutritious animal fodder and soilnitrogen supplier and green manure conditioner, T. pratense has an ancient reputation in herbalmedicine in B & I as a fluid extract of it was used as an antispasmodic and for the treatment ofbronchial and whooping coughs. In addition, "Fomentations and poultices of the herb have been usedas local applications to cancerous growths." (Grieve 1931, p. 208).
Back in the mid-17th century the juice of the plant was applied to adder bites and to clear the eyes ofany film obscuring them, or to soothe them when hot and bloodshot. Infusions of the plant were alsoapplied to rashes and used to treat coughs and colds in Norfolk and Cumbria. The leaves were chewed inthe Isle of Man to relieve toothache and in Offaly, Ireland they were applied to soothe bee stings(Allen & Hatfield 2004). Red Clover contains isoflavones and a herbal product sold in tablet form istaken by women during and after the menopause.
The plant is also edible and young leaves and flowers have been used in salads for both colour andflavour, or cooked and made into soup or used as spinach substitutes. The flowers in the past haveprovided not only popular sweet snacks for children, but were used to make both tea and wine (Darwin1996).
Threats
None.
Native or possibly introduced, rare and very local. Eurosiberian boreo-temperate, but widely naturalised.
1884; Barrington, R.M.; Lower Lough Erne shore near Circle Hill.
June to September.
Growth form and preferred habitats
In B & I, this is generally regarded as a species of unmanaged grassland on heavy clay, neutral soilsof intermediate or richer fertility. It is a robust, long-lived, rhizomatous perennial, tolerant ofmoderate shade and therefore capable of emergence through the tall canopies of derelict, neglected orlightly-grazed rough marginal grasslands and grassy banks at the base of hedges, along woodland marginsand developing scrub, including on heaths. Typical habitat for T. medium is among medium-sizedperennial herbs at the transition between unmanaged, rough grass and invading scrub (Sinker etal. 1985).
T. medium can also tolerate both non-calcareous and limestone soils of intermediate or nearneutral pH (ie pH=7.0), provided they are neither regularly droughted nor waterlogged (Grime etal. 1988, 2007). It is typically absent from chalk (Crawley 2005). T. medium can alsooccupy more ruderal habitats such as along railway lines and occasionally in old quarries. In Ireland,it is sometimes found on heathland scrub and it can also occur in B & I on upland stream banks andin tall-herb communities on rock ledges (D.A. Pearman, in: Preston et al. 2002). Although afterflowering it usually dies down in winter, in B & I, T. medium is described as being tolerantof frost and is said to be winter hardy (Duke 1981).
While it demands light and warmth for good growth and does not survive in gloomy woodland conditions, infully open situations and on bare ground, T. medium finds it difficult to survive competitionfrom better adapted colonising and meadow species. In Finland, it thrives in more open areas withinhillside forests and does particularly well both in storm damaged woodland and in other areas open dueto either logging or fires. Under Finnish conditions, T. medium appears to have exploitedsituations involving human activity, such as forest clearance, and the moderate rate of change involvedin hillside forest conversion into grazing land. It has spread there from its original habitats tobecome a companion of people, including in neglected grass and ruderal situations (NatureGate website athttps://luontoportti.com/suomi/en/kukkakasvit/zigzag-clover).
The established strategy of Zigzag Clover is described as intermediate between stress-tolerant competitorand the more general ecological role balance of C-S-R by Grime et al. (1988, 2007). It isimportant to bear in mind that Grime (1979, 2001) placed low nutrient availability at the core of histhinking on plant adaptation to survival in stressful environments (Craine 2009).
Identification
T. medium is rather similar in appearance to T. pratense (Red Clover), but it has narrower,oblong leaflets without white markings and stipules that taper evenly to a point, but do not possess thehair- or bristle-like leaf-tip characteristic of Red Clover. The English common name 'Zigzag Clover'refers to the appearance of the stems. The inflorescences are also stalked (not sessile) and the flowersare of a somewhat brighter red colour than those of T. pratense (Parnell & Curtis 2012).Although it is sometimes possible to distinguish T. medium from T. pratense by theleaflets of the former being longer and narrower, the only really reliable distinguishingcharacter between the two is the appearance of the tip of the basal stipules of their stem leaves. InT. medium, the free part of these stipules is linear-lanceolate (ie like the blade of amedium-length sword), tapering evenly and coloured green almost to the tip. The free portion of T.pratense stem leaf stipules are triangular-ovate, narrowing abruptly to a brown, bristle-liketip (Webb et al. 1996; Stace 1997; Crawley 2005).
Although Zigzag clover looks very like what some refer to as the 'King' of useful plants, Red Clover,T. medium has never itself been a significant fodder crop.
Flowering reproduction
T. medium flowers from June to September. The numerous, globose, bright, purple-redinflorescences, 25-35 mm in diameter are borne on short stalks subtended by a pair of leaves. As withmost clover species, T. medium must be cross-pollinated for any seeds to develop. The flowers arelong-tubed, the corolla 12-20 mm long and the stamen filaments are fused from the base for most of theirlength, so that only quite large bumble-bees, honey- bees and certain butterflies and moths with aproboscis long enough can reach the nectar secreted at the base of the tube. Sometimes a small hole canbe found at or near the bottom of the calyx-tube that indicates a visit from an insect with a shortproboscis that has bitten a hole, broken into and robbed the nectar store. Obviously this robberactivity does nothing to pollinate and fertilise the flower.
Fruit and seed dispersal
After fertilisation the legume pod develops, surrounded by the persistent fading corolla and the nowspiny calyx. As in T. repens and some other clovers, the dead corolla plays a part in the seeddispersal by acting as a wing. In both T. medium and T. repens, the corolla does notenlarge, nor is it modified in any way as a flying organ, but it persists in a withered state, dry andscarious and enclosing the pod which contains one (or more) small, globose seeds. The withered flowerbecomes detached from the head of flowers and is blown away across the ground by the wind (Ridley 1930,p. 117). Eventually the 2.0 mm long membranous pod splits to release the solitary seed.
Apart from this and the rather remote possibility of the fruiting Zigzag Clover plant being grazed andthe seed carried in an animal gut, there is no other form of seed dispersal mechanism. While it might beimagined that the seed probably has the usual legume hard coat dormancy and associated prolonged buriedsurvival in the soil, the NW European survey of soil seed banks found that all five references consultedhad concluded T. medium seed is transient, surviving for less than one year (Thompson etal. 1997).
Vegetative reproduction and population decline
The underground rhizomes of Zigzag Clover branch and spread to form clonal patches of the plant, butsince the flowers are largely self-incompatible, many clones produce little or no seed. Fragmentation ofthe rhizome creates separate daughter colonies of identical genome, but a possible over-reliance onvegetative reproduction for population survival helps to explain why this clover is associated witholder, neglected, undisturbed grasslands, such as occur on Lough Erne lake islands and on moreinaccessible rocky shore headlands around the larger lakes in Fermanagh. It also suggests a reason forthe current decline in this legume's occurrence throughout the whole of B & I, since T.medium was always fairly thinly scattered and was never a widespread, common species with acrop-based reservoir of repeatedly sown seed like other clover species. This is particularly thesituation in Ireland, where the decline of the species from a low peak population is all the moreclearly marked.
Fermanagh occurrence
Half of the 23 records of T. medium in the Fermanagh Flora Database are of pre-1950 date and RHNand the current author (RSF) consider it a very local, decidedly rare and casual, more-or-less ruderalspecies in the VC. The status of the plant is unknown, but as it previously would very likely have beenincluded in agricultural clover seed widely and frequently sown for fodder, even as a contaminant, itmight well be an undetected introduction. The Fermanagh records are scattered across a total of 20tetrads, 3.8% of those in the county.
Irish occurrence
The New Atlas map shows T. medium widespread in Co Antrim (H39) and scarce in the otherfive VCs of N Ireland. Time will tell whether this indicates under- or over-recording or both! In theRepublic of Ireland, nowadays, T. medium is a rare or very rare plant confined to damp roadsideverges and banks (Doogue et al. 1998; Green 2008), or recently disturbed grassland in damplimestone pasture on the turlough-like bottom of shallow valleys (Reynolds 2013).
Previously, in the Burren, Co Clare and Connemara, Webb & Scannell (1983) regarded it as a ratherrare species of roadsides, waste places and rocky ground and, "in all cases apparently a relic ofcultivation." These authors also considered it, "obviously transient in many of itsstations" [they listed a total of just nine sites], "in a few, however, it seems wellestablished."
British occurrence
Zigzag Clover is much more generally distributed in lowland areas in Britain than in Ireland, although itremains a frequent to uncommon plant of heavy basic or clay soils. There are definite distribution gapswhere it becomes local or rare, for instance in C Wales and around the English Wash. In Scotland, itbecomes rare or absent in the N & NW, most likely due to the prevalence of unsuitable soils. T.medium reaches its maximum altitude in Britain at 610 m on Helvellyn in the English LakeDistrict. The BSBI Monitoring Scheme Survey of 1987-88 concluded that none of the widely reported lossesof T. medium populations across B & I that varied from -4% to -35 % were statisticallysignificant (Rich & Woodruff 1990, map and figures in 2, p. 87). The New Atlas hectad mapsuggests possible under-recording by modern botanists, or perhaps equally probably, mistakenover-recording of it in past years. Analysis of the New Atlas survey data indicated a markeddecline in S & E England since 1950 (D.A. Pearman, in: Preston et al. 2002). Evidence gleanedby the current author (RSF) from the biology and ecology of T. medium indicates there is littledoubt that populations are declining in abundance and in some areas also losing ground across both B& I, most probably due to the near disappearance of lowland permanent grassland habitats in the last60 years, together with progress in producing cleaner clover seed for agricultural grassland ley sowings(Grime et al. 1988, 2007; D.A. Pearman, in: Preston et al. 2002). However, the 'LocalChange' monitoring sample survey of Britain, carried out from 1987-2004, did not detect any furtherdecline (Braithwaite et al. 2006).
European and world occurrence
T. medium occurs throughout most of Europe except the extreme N & S. There is more variationwithin the species on continental Europe than exists in B & I, Flora Europaea 2 listing foursubspecies, with intermediates occurring. Two of the subspecies are very local: subsp. sarosiense(Hazsl.) Simonkai is restricted to the foothills of the Carpathians; and subsp. balcanicumVelen., to the Balkan peninsula. The other two are subsp. medium, which is found throughout therange of the species, possibly excepting Greece; and banaticum (Heuffel) Hendrych, which isrecorded from Czechoslovakia, Hungary and Romania (Tutin et al. 1968).
This polymorphic species was originally confined to Europe and adjacent parts of Asia but has been spreadby man and his activities, although not to anything like the same extent as T. pratense, T.repens and a few other grassland ley clover species. T. medium has been introduced,however, to N America, north central China, Japan, Tasmania and both islands of New Zealand (Hultén& Fries 1986, Map 1246). In New Zealand it is regarded as, "a rare casual [introduction] ofwaste places and cultivated land, formerly more widespread" (Webb et al. 1988).
Threats
Older undisturbed grasslands are vital to its mere survival, and the NI Government EnvironmentallySensitive Areas (ESA) policy should have assisted its maintenance.
Introduced, neophyte, a garden escape, very rare, but very probably under-recorded.
5 September 1998; Northridge, R.H.; roadside bank near Enniskillen Town.
Growth form and preferred habitats
This small to medium sized, deciduous, ornamental tree, 7-9 m high with a trunk rarely exceeding 20 cm indiameter has low, spreading arching branches when old. A native of the mountains of C & S Europe, ithas for centuries been very popular in gardens everywhere and is known to quite frequently self-seed andnaturalise in lowland parts of B & I. Laburnum usually occurs as a persistent, isolated gardenescape, typically in acid soil on waste ground or in other marginal, occasionally or regularly disturbedhabitats, eg along roadside verges or on railway embankments. It is, however, ecologically veryundemanding and can occupy and survive in any type of soil (Hadfield 1957).
Flowering reproduction
Flowering occurs in May and June, the long, lax, dangling, bright yellow racemes of numerous pea flowerswith a large standard petal, making the tree very conspicuous. Pollination is effected mainly bybumble-bees and the fruit is a narrow, pendulous, legume pod, 30-80 × 7-8 mm that hangs on the tree forsome time (sometimes all winter), before splitting and twisting to explosively eject at least some ofthe dark brown or black seed (Cooper & Johnson 1998). Seeds are abundantly produced and germinatefreely, and for this reason the plant is often used as a stock upon which to graft or bud otherleguminous trees and shrubs. Laburnum trees have the reputation of being not long-lived (Hadfield 1957).
Toxicity
All parts of the tree, but especially the bark and the seeds contain a quinolizidine alkaloid toxincalled cytosine (named after the old name of the plant – Cytisus laburnum) which resemblesnicotine (Hadfield 1957; Cooper & Johnson 1998). The quantities of cytosine found in Laburnum andsaid to be dangerous or even fatal vary greatly (Cooper & Johnson 1998). The seeds are regarded asparticularly poisonous and have killed children and stock, though it is said the leaves can be grazedwithout harm. Laburnums should not be planted near small garden ponds as the seeds and leaves may poisonfish (Hadfield 1957). In general, Laburnum poisoning is rare, but a few fatal cases have been reported,including large animals such as horses from eating leaves and seed and dogs from chewing sticks brokenfrom the tree.
With regard to human poisoning, it is very rare that anyone consumes a fatal dose, but treatment isconsidered necessary if more than five seeds are ingested by a child or 15 by an adult. Theadministration of activated charcoal, together with fluids, especially milk in the first six hours afteringestion, is the recommended treatment (Cooper & Johnson 1998).
Fermanagh occurrence
The solitary Fermanagh record listed above is of an individual tree which originated either from a nearbygarden, or very much less likely, it might have been deliberately planted.
Species and hybrid variation
Sometime after its introduction to gardens in Britain towards the end of the 16th century, L.anagyroides formed a partially fertile hybrid (L. × watereri (Wettst.) Dippel),with the closely related L. alpinum (Mill.) J. Presl, the so-called 'Scottish Laburnum' althoughthe latter has no real, valid origin connection with Scotland. The hybrid is a more vigorous andfloriferous medium-sized tree than either of its parents and is the preferred garden Laburnum specimentoday, although all three forms (the two species and their hybrid) are cultivated. The hybrid is oftensold in the horticultural trade under the name L. × vossii hort. Asone would expect, it is intermediate between its parents in degree of pubescence and in leaflet andfruit shape. The hybrid combines the more floriferous longer racemes (up to 50 cm long) of L.alpinum, with the larger (15-21 mm) flowers of L. anagyroides. The fruits of the hybridare much fewer than the species, malformed, usually containing no, or just one or two seeds per pod and,remembering Laburnum toxicity, it the safer garden option to choose (Hadfield 1957; Stace et al.2015).
The under-surfaces of the leaves and the young shoots of L. anagyroides are densely covered inappressed, silvery, silky hairs, while L. alpinum is almost glabrous and the hybrid isintermediate, ie it is only sparsely hairy. The upper suture of the pod is sharply ridged in both L.alpinum and the hybrid, but it is blunt and greatly thickened (ie much broader) in L.anagryoides (Hackney, C.R. 1989, Fig. 1.).
British and Irish occurrence
L. anagyroides is a quite frequent and widely scattered introduction across Britain from the farsouth and the Scilly Isles to Shetland. In Ireland, it is much less frequent, being rare and widelyscattered in the RoI, although quite a lot more frequent in NI. Many of the records for L.anagyroides shown in the New Atlas hectad map could well be mis-identifications resultingfrom the hybrid being regularly overlooked (D.A. Pearman, in: Preston et al. 2002). The hugeincrease in records of L. anagyroides in Britain and in NI since the 1962 BSBI Atlas,probably merely reflects increased interest in recording alien trees in these areas.
While L. × watereri is capable of setting a very limited amount ofseed, it is very much less likely to spread itself than L. anagyroides or L. alpinum.Seedlings of the latter are apparently very rare, even when many of the parent trees are planted as ahedge, as in the Slieve Gullion area of S Londonderry (Hackney, C.R. 1989). L. anagyroides hasalso been recorded more rarely in use as hedging on farm lanes in NE Ireland (Hackney et al.1992). On the other hand, the presence of L. × watereri sometimesindicates deliberate hedge planting, as has been shown to have taken place in Cos Tyrone and Londonderry(H37, H40) (P. Hackney, Northern Ireland Flora Website 2005; McNeill 2010). The hybrid has also beenused for hedging in three VCs in S Wales (VCs 44-6) (Chater 1996; Stace et al. 2015).
Uses
Laburnums are of no economic value although the wood is occasionally used as it is hard and durable andis similar to that of Robinia pseudoacacia (False-acacia). It takes a very fine polish and issometimes used for inlays in furniture, for turning small objects and for veneers. The dark heartwoodcan be substituted for ebony. Older writers record that on the Continent it was regarded as supplyingthe longest lasting bows as weapons (Hadfield 1957).
Threats
None.
Native, frequent. European temperate, but widely naturalised including in N America and New Zealand.
1892; Praeger, R.Ll.; Inish Doney island, Lower Lough Erne.
Throughout the year.
Growth form and preferred habitats
The bright lemon-yellow flowers, deciduous, both simple and trifoliate leaves, borne alternately onerect, evergreen, pliable, 5-angled, unarmed photosynthetic stems, 2.5-3.0 m tall, that become woody asthe plant matures, together make this variable shrub easily and instantly recognisable. It is present inFermanagh in a wide variety of open, sunny or lightly-shaded, more-or-less disturbed ground sites and inunmanaged grassland habitats, on suitably acidic, low-nutrient, infertile, lime-deficient soils. C.scoparius thrives on a wide range of acid to neutral soils and multiplies on disturbed sites,preferring nitrogen-medium substrates and strongly drained, water-shedding sites (Peterson & Prasad1998). It is practically impossible to exaggerate the importance of disturbance, natural or generated byman, to the spread and colonising ability of a plant like Broom.
Typically, C. scoparius is strongly calcifuge and avoids lime which puts a limit on itsdistribution (Clapham et al. 1987), but in Fermanagh it is recorded at Knockmore in the heart ofthe local limestone district and also in the Claddagh River Glen (otherwise known as the Marble Arch),where regionally famous limestone caves are situated. It is presumed the plant must have located pocketsof acidic, low-lime conditions in these areas, but they are the exception in terms of the speciesdistribution.
The shrub develops a deep taproot that allows it to tolerate considerable levels of drought. ItsRhizobium root nodules fix nitrogen efficiently and while its leaves are deciduous, in mildclimates the evergreen stems continue to photosynthesise in winter, facilitating year-round nitrogenfixation and prolonging the growing season. Nitrogenase activity ceases, however, if there is heavyfrost and severe levels of frost can kill the plant outright (Peterson & Prasad 1998).
Flowering reproduction
Flowering takes place in May and June; plants flower from their second or third year of growth andcontinue doing so for the 10-15 year lifespan of the typical individual. Pollination is carried out byboth honey- bees and bumble-bees in a mechanism first described by Müller (1883). Essentially,pollination is an explosive mechanism which depends for its effectiveness on the bursting of the anthersand the shedding of their pollen over the immature style and stigma while the flower is still in bud.The style then elongates so that when the flower is completely open, it is held under tension inside thefilament tube between the fused keel petals. The pollen covered style is suddenly released by the weightof the visiting bee, bursting the sutures of the keel, throwing out a cloud of pollen above the insectand dusting its hairy back with it.
The stigma then rapidly describes a spiral path and in doing so first strikes the insect's back, where itmay pick up pollen from the previous flower visited. It then continues in its path to strike theunderside of the insect's abdomen and possibly pick up pollen deposited there by the short stamens ofits own flower (Gill & Walker 1971; Proctor & Yeo 1973, pp. 200-1).
The legume pod is 25-50 × 10-12 mm, oblong and strongly compressed and becomes dry and dark brownish greywhen ripe. It is composed of two valves that have long brown or white hairs on the sutures. When fullyripe, which can be as early as mid-July, the pod audibly and explosively splits apart, the two valvestwisting and ejecting the numerous (up to nine) blackish brown seeds into the air and onto the groundaround the parent plant.
Secondary seed dispersal
The seed coat has an edible protein elaiosome outgrowth attached to the hilum scar, similar to that foundin the genus Viola (see the Viola species accounts on this website). The elaiosomeattracts ants that carry the seeds towards their nests and assist in their dispersal. Experimentalelaiosome removal had no significant effect on seed germination of C. scoparius in a Californianstudy (Bossard 1993). Other studies along riverbanks in New Zealand strongly suggest seed is alsodispersed by flowing water (Williams 1981).
Seed germination and survival in soil
Seed production is prolific and the hard seed coat (or testa) allows buried survival for 30 years orlonger. A study in California concluded that 66% of seed from one year's crop germinated during thefirst year. The same study showed that at least 7% of seeds remained un-germinated after three years inthe soil, allowing the development of a large seed bank. Variability in the duration between seeddeposition and germination provides C. scoparius with considerable flexibility for coping withthe intra- and inter-yearly fluctuations in precipitation and temperature typical of California'sMediterranean climate. Such climate factors also affected the degree and duration of hard-seeded-nessand thus helped determine levels of dormancy. However, no genetic basis for environmental differences inseed germination characteristics between widely separated countries should be assumed to exist withoutfurther research (Bossard 1993).
Lack of vegetative reproduction
There are no reports of vegetative reproduction occurring in C. scoparius (Peterson & Prasad1998).
Variation
This species has been through several generic name changes in the not too distant past, synonymsincluding Spartium scoparium L., Genista scoparia (L.) Lam. and Sarothamnusscoparius (L.) W.D.J. Koch. Today, two subspecies are recognised: the erect, non-littoralshrubby form is subsp. scoparius, the most commonly quoted chromosome count for it is 2n=46, butother sources suggest 2n=48 (Morton 1955; Peterson & Prasad 1998). The prostrate or procumbent dwarfalternative coastal form of the plant is subsp. maritimus (Rouy) Heywood grows only up to 10 cmabove ground and it has been variously listed as 2n=24, 46 & 48 (Gill & Walker 1971).
Morton (1955) found the prostrate plant had 2n=24 and the metaphase chromosomes were of twelve types,indicating a base number of 12 in the genus. He suggested that since the species had 48 chromosomes inthe same twelve chromosome types in quadruplicate as the prostrate form, it might have arisen as anautotetraploid from the subspecies that he called Sarothamnus scoparius subsp. prostratus, now called Cytisus scoparius subsp. maritimus (Morton 1955).
Gill & Walker (1971) found most chromosome counts they made from material across England and Walesgave 2n=46 and they could not reconcile this with Morton's count of 2n=24. Not only was his count atvariance with all other counts published, but the chromosomes he illustrated were much larger than thosefound by Gill & Walker, raising doubts about the identification of Morton's material.
There are also over 80 named garden cultivars and hybrids of C. scoparius, the crosses ofteninvolving C. multiflorus (L'Hér. ex Ait.) Sweet, White Broom, or C. × dallimorei Rolfe. (C. multiflorus ×C. scoparius) (Griffiths 1994).
Fermanagh occurrence
In Fermanagh, C. scoparius has been recorded in 95 tetrads, 18.0% of those in the VC. Eight ofthese tetrads contain only pre-1975 records, which suggest some decline in available habitats. As thetetrad map indicates, Broom is scattered quite widely across Fermanagh, but the great majority of itssites are found on the better drained, non-calcareous soils in the east of the VC. Habitats include dryor well-drained heathy banks or slopes on acid, often sandy or stony soils. It also invades open woodsor their margins, roadsides, river banks, quarries, old railway lines or embankments and other forms ofdisturbed ground. Broom is sometimes very prevalent on estate land, where it is probably derived fromplanted material.
In the wild, Broom is generally found as solitary bushes, seldom in quantity, but C. scoparius iswell known to be an opportunistic pioneer colonist of open soils and is especially successful when thereis little or no grazing pressure. Thus Broom can occasionally form large, dense, temporarily dominant,pure stands, as it does locally at the neglected sand pit near Pubble Bridge on the Tempo River, an opensite which this leguminous shrub has rapidly colonised and where it currently comprises a very large,dominant stand. It remains to be seen if it will be capable of regeneration and of keeping at bayaggressive secondary colonists such as Birch and other larger trees and shrubs that could shade it out.
Irish occurrence
The New Atlas hectad map shows C. scoparius is common in lowland areas throughout NI andagain further south in coastal counties from Dublin to Kerry, but is very much more scattered andoccasional in C & W regions of Ireland. It is presumed that soils in the areas where it is scarceare principally limestone, or wet and peaty, neither of which would be suitable to support Broom. TheCentral Plain of Ireland also suffers the coldest winters in the country and a single very cold nightcan destroy Broom populations.
British occurrence
Broom is widespread and common throughout lowland Britain, but scarcer further north in Scotland and incalcareous areas of the country for the reasons already mentioned (Preston et al. 2002). In thedistant past, C. scoparius must have been much more abundant than now since it features socommonly in English place names. Broom place-names can be counted by the score. A familiar one isBromley in Essex, the name meaning 'broom clearing'. Another example is Brompton in Middlesex, the'broom tun' or 'farmstead'.
Many cultivars of this ornamental shrub are very commonly planted in gardens, although it is notlong-lived, tending to flower itself 'to death' within 10-20 years at most. Forms of it are oftenplanted along new roadways to help stabilise steep embankments, or it is used as a nurse to young treesin plantations, as is, or was, the case in India where it quickly became a significant problem weed.
European and world occurrence
C. scoparius is native and widespread throughout lowland temperate Europe northwards to SSweden and eastwards to C Ukraine – although not mapped in most of the Balkans (Hultén & Fries 1986,Map 1180; Sell & Murrell 2009). Originally it was restricted to Europe reaching south to Sicily andincluding the Azores and the Macronesian Islands, but it has been introduced to N America, New Zealandand India. Abroad, the vigorous colonising ability of the species is such that it has become awidespread and troublesome woody weed in waste land, river beds, native grasslands and previouslyforested hillsides across New Zealand and parts of Canada and California. It does not survive in moreinland continental conditions in N America, being ± confined to lowland Oceanic or Mediterraneanclimatic regions by its sensitivity to frost and winter drought. The shrub thrives and grows rapidly inthe conditions found in New Zealand, however, and is more vigorous there than in Europe, partly becauseof the absence of its major invertebrate predators (Williams 1981).
Toxicity
Broom contains several toxic alkaloids, but it is not dangerously poisonous. It contains small amounts ofthe toxic quinolizidine alkaloids sparteine and isosparteine. Other toxins present include cytosine,genistein, lupinidine and sarothamnine, although some of these names may refer to the same compound(Cooper & Johnson 1998). The alkaloids can depress the heart and nervous system, sometimes withparalysis of motor nerve endings. It is unlikely Broom would cause threatening poisoning as insufficientamounts of the toxins are present. The leaves being small and the stems wiry, sheep are not attracted toit, although goats will browse even this level of herbage. A horse would need to consume a very largeamount of Broom (over 11 kg) to be at risk (Cooper & Johnson 1998; Peterson & Prasad 1998).
Names
The majority of the English common names in numerous, varied dialects refer to 'Broom' as in the brushused to sweep floors and other areas, eg 'Basom' in Cornwall and the W of England, 'Beeson' in Devon and'Brushes' in Dorset. Grigson (1955, 1987) describes it as, "one of the great landscape plants"and, as it flowers early in the year in May and June, it is considered one of the plants indicative oflove and romance in European poetry. He also refers to it as one of the best of all sweeping plantsalong with birch and heather. The long, slender, tough yet flexible branches, carried in tight fasciclesmake it ± ideal for sweeping. The English regal house the Plantagenets take their name from the broom,the Planta genista, a fact that Grigson thinks "everyone knows!" The Plantagenet familyoriginated in Brittany and in Anjou on the Loire, where C. scoparius is a common shrub to thisday.
Uses
Apart from its use to sweep floors, the tips of Broom branches have diuretic and cathartic propertiesthat are widely and frequently used in herbal medicine. The sparteine they contain is a powerfuldiuretic which triples renal elimination (ie urine flow) and has been used for dropsy and kidneyailments of all kinds. In modern herbalism, Broom is used to slow and regulate the heart rate. While thelist of other recorded uses from around B & I is long, varied, very local and minor, the two otherprincipal uses in folk medicine are for the treatment of rheumatic complaints and as a purgative forliver troubles, jaundice and piles (Allen & Hatfield 2004).
In times of fodder shortage, Broom branches were resorted to as winter feed for sheep in the same waythat Gorse was used, and it was allowed to grow around the margins of fields in preparation for thiseventuality (Grigson 1955, 1987). The green branches were also supposed to prevent rot and dropsy in thesheep that ate them (Grieve 1931).
Young flower buds are edible and once were a favourite delicacy as they appeared on three separate tablesat the Coronation banquet of James II. Grieve (1931) added that the flower buds served the doublepurpose of an appetizer and a corrective. Buds were collected and laid in pickle or salt until required,"which afterwards being washed or boiled are used in sallads as capers be and be eaten with no lessdelight" (Gerard 1633).
Threats
None.
Native, common, widespread and locally abundant. Oceanic temperate, but very widely naturalised in bothhemispheres.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
A much branched, large evergreen, profusely yellow-flowered, conspicuously armed shrub, usually around 2m tall, but capable of reaching 4 m in height if left to grow once established. All leaves beyond theseedling stage are reduced to prickly, green, acicular, spine-like phyllodes that easily detach and forma persistent spiny litter beneath the shrub. The majority of branches are reduced to form very sharp,rigid, green, deeply grooved spines, 15-30 mm in length, that thickly cover the woody main stems. Largerprimary spines branch to form secondary and tertiary spinesand they may also bear flowers a considerable distance from their bases. Each bush produces ataproot that usually manages to penetrate at least 30 cm in deeper soils, but most roots are superficialin the top 10 cm of soil (Grubb et al. 1969). The roots have both VA (vesicular-arbuscular)mycorrhiza and nitrogen-fixing nodules that are unusual in being perennial (see below).
The established strategy of U. europaeus is as a stress-tolerant competitor (Grime et al.1988, 2007). It is particularly abundant and rapidly develops dense thickets in neglected ground whichhas previously been disturbed, such as on the banks of ditches, streams, rivers and lakes, in hedgerowsbeside tracks or roads, on woodland margins, in woodland clearings and around human settlements ingeneral. It is frequent on rocky maritime cliff-tops and often forms dominant dense thickets on greysand dunes (Grime et al. 1988, 2007).
U. europaeus thrives in dry or well-drained, highly disturbed areas and it can invade and growwell in shallow, stony conditions, or in peaty nutrient-poor heathland or bog margin soils, although foroptimum development it really requires moderately fertile soils around 20-50 cm deep (Clements etal. 2001). Gorse is more tolerant of soil acidity than most other legumes (Hill 1949) and itproduces optimal growth at a soil pH between 4.5 and 5.0 (Meeklah 1979). Gorse does not grow all thatwell in calcareous soils and, therefore, is much less frequently met in limestone districts, although itis certainly not excluded from them. It can even grow surprisingly well on thin, shallow rendzina soilsover chalk and limestone (Proctor 1965). U. europaeus is absent from wetlands, managed meadowsand pastures, or cultivated ground.
The distribution and abundance of Gorse is strongly correlated with human disturbance, including allforms of agriculture, forestry and transport, together with the plethora of uses to which the plant hasbeen pressed by man in past times (Lucas 1960; Mabey 1996).
On heathland, there is evidence that U. europaeus is much more exacting in its requirements forcertain plant nutrients and mineralizeable salts and it demands a lower carbon:nitrogen ratio than otherheathland species, eg Calluna vulgaris (Ling) and Erica species (Tubbs & Jones 1964).Disturbance of the ground in heathland may encourage Gorse growth through the inversion of the soilhorizons, releasing plant nutrients from the layered podsol conditions which commonly form in acidic,heavily leached soils. In other soils occupied by Gorse, additional nutrients are made available whensoil is heavily trampled and manured by grazing stock.
Gorse and Bracken (Pteridium aquilinum) have long been used as indicators of above-average qualityin heathland soils. For example in the New Forest, Hampshire, in the past soils were chosen for croppingon the basis of the old saying, "Under Bracken lies gold, under Gorse lies silver, and underHeather lies lead." (Tubbs & Jones 1964). Another Irish version from Co Kerry has it as,"Gold under furze, silver under rushes and famine under heath." (Lucas 1960, p. 186).
The very presence of gorse acidifies soil through the addition of its litter and the removal of calcium.The high density of living evergreen shoots in Gorse thickets also reduces through fall of incomingprecipitation. In one study in New Zealand, precipitation penetration was reduced by 35-50%, withconsequent reductions in nutrient run-off and soil moisture levels (Egunjobi 1971).
U. europaeus is generally associated with relatively sparse grassland or scrub communities whichallow it to compete for light (Grubb et al. 1969). It is sometimes associated with similar andrelated shrubs, such as Cytisus scoparius (Broom), but only a few woodland species can grow underthe dense canopy of gorse, eg Hedera helix (Ivy) and Rubus fruticosus (Bramble).
Perennation of stems and root nodules
The evergreen stem and spines of gorse photosynthesise all year round and thus actively perennate theplant in conjunction with continuous nitrogen fixation, although as one would expect, growth and noduleactivity of the plant is much reduced in winter (Radcliffe 1986; Clements et al. 2001). Unlikemany other legumes, Gorse and Cytisus scoparius in N Ireland have been shown to possess trulyperennial root nodules, which are branched and can weigh up to one gram. Nodule perennation may onlyoccur during a succession of mild winters and possibly only in soil conditions where decay of the olderparts of the nodule becomes arrested (Pate 1961). In one study, Egunjobi (1971) estimated that in astand of seven year old Gorse, 65% of the nitrogen taken up and fixed by the species was returnedannually to the soil.
Flowering reproduction
Under favourable growing conditions, young plants of this spiny polycarpic perennial shrub grow extremelyrapidly and they are capable of flowering just two or three years after their germination (Rees &Hill 2001).
U. europaeus flowers in super-abundance, the blossom crowded in spire-like clusters at the tips ofthe branches. The large, showy, pea flowers vary in degree of yellowness from a soft buttercup to astrong orange, all of them heavily scented of coconut oil. The Gorse floral display continues all yearround in B & I, but is at its most impressive from March to early June. There is a legend, loved andoften repeated by the Victorians (but very possibly apocryphal), that when the famous Swedish botanistCarl Linnaeus visited England in August 1736 and saw gorse flowering for the first time on someheathland, he fell down on his knees and thanked God for the beautiful display (Blunt 1971, p. 90).
Honey-bees and bumble-bees visit some of the flowers, but there are far too many compared with the numberof insect pollinators (Knight 1996). The flowers lack nectar, but bees, if they are sufficiently heavyand vigorous, can trigger the explosive release of copious pollen which then coats the body hairs on thevisitor, who then transfers it to the special 'pollen basket' hairs on its hind legs. Once it has beentriggered in this way, the flower hangs limply and thus it is most unlikely to be revisited (Proctor& Yeo 1973).
Although gorse flowers are self-compatible, out-crossing is said to result in higher fertility, whichwould be the expected genetic outcome (Clements et al. 2001). The few quantitative studies thathave been made suggest that a large percentage of the flowers fail to set fruit (possibly up to 80%reproductive failure) (Clements et al. 2001). In a coastal district in Wales, however, Knight(1996) estimated that billions of Gorse flowers were unvisited (ie they were unsprung) and thiscertainly is the case also in Ireland. Knight found that the level of fertility appeared unaffected bybeing unvisited by bees, since the unsprung flowers all self-pollinated.
The number of seed set was equally low in both instances (insect pollinated and selfed), withapproximately twelve ovules per ovary setting a mean of just 2.2 seeds (Knight 1996). Low fertility inGorse has been shown to be related to the production of poor pollen (ie meiotic abnormalities occurringduring microsporogenesis) (Misset 1992). The annual seed production has been estimated in New Zealand as500-600 per m2 (Ivens 1978).
Seed dispersal
Most Gorse seeds fall beneath the parent plant, but some may be ejected explosively when the pods audibly'pop' and seed can travel in this manner up to 5 m from the parent plant. Longer distance dispersalinvolves water, ants (and perhaps birds), and also vehicles. The tough seed coat is water repellent andcan resist abrasion in stream gravel. Each seed bears a white nutritive outgrowth from the seedcoat (anelaisome), which attracts ants and perhaps also birds like Quail, where they occur (Chater 1931). Sincethe plant distribution closely follows agriculture workings, it is presumed that seeds frequently hitchlifts on machinery (and boots) in mud.
Seed and plant longevity
The seed has considerable longevity, persisting in large quantity (up to 20,000 per m2. in thetop 6 cm of soil), for as long as 30 years (Clements et al. 2001). It is the ability to produce alarge seed bank which helps U. europaeus to persist in many areas since the actual lifespan ofindividual plants is relatively short, maximum active survival being around 15-20 years. Older plantslose vigour, become top heavy and eventually keel over under their own weight and uproot themselves.
Vegetative reproduction
Vegetative spread of Gorse occurs by means of creeping roots, the species being capable of formingadventitious roots following major disturbances such as cutting of branches or of whole stems(Zabkiewicz & Gaskin 1978). Unfortunately, from the point of view of control or eradication of theshrub, both roots and shoots are also capable of re-sprouting after fire (Clements et al. 2001).
Fermanagh occurrence
Gorse is extremely common and widespread in Fermanagh, occurring in 417 tetrads, 79% of those in the VC.It is less frequent on limestone, but absent only from the highest ground, aquatic habitats and the bestmanaged and most fertile farmland.
British and Irish occurrence and status
U. europaeus is very common and widespread throughout B & I, except on ground aboveapproximately 640 m (D.A. Pearman, in: Preston et al. 2002).
Status in Britain and Ireland
Gorse is generally considered native in B & I, but all of the fossil records of the currentinterglacial period (known as the Flandrian in England and the Littletonian in Ireland) are from ZoneVIIb or later and they are from settlement areas of Neolithic or younger date (Godwin 1975, pp. 177-8).It might be better, therefore, to regard it as a possible archaeophyte (ie an early, pre-1500 ADintroduction). The close association of U. europaeus with disturbed sites, its known plantationfor game cover or as hedging along roadsides, together with its many uses including as fodder and fuel,certainly indicates it is not naturally occurring everywhere in these islands. For instance, it is shownin the New Atlas as an introduction on the Isle of Man, Islay and Jura, the Outer Hebrides and inOrkney and Shetland.
European and world occurrence
Ulex europaeus was originally restricted to W Europe but it has spread to C & S Europe and NAfrica. It remains most prevalent along the Atlantic coastline and near the Mediterranean basin, whereit extends eastwards to mainland Greece. It has gradually spread northwards and eastwards from itsnative range in Europe and become naturalised and now occurs in southern parts of several Scandinaviancountries (Tutin et al. 1968; Hultén & Fries 1986, Map 1185).
It is clear that the distribution of U. europaeus, like that of its relative, Cytisusscoparius (Broom), is very much governed by temperature. Gorse requires more or less oceanicconditions and it cannot survive in arid climates or in continental regions where there are seasonalextremes of heat and cold. Temperature also restricts the species to lower altitudes and while matureplants can tolerate quite severe frosts, in general they show a definite preference for habitatssheltered from cold winds. Day-length may also affect the latitudinal distribution of Gorse, asshort-day conditions inhibit maturation of the plant and prevent both thorn formation and flowering(Zabkiewicz 1976).
Gorse has also been purposely introduced by man (eg for hedging, fodder or as an ornamental) to more than15 countries or island groups throughout the world where maritime climates occur similar to that in itsnative distribution area,
including Australia, New Zealand, Chile, the Hawaiian Islands, Costa Rica and to both E & W coasts ofN America (Hultén & Fries 1986, Map 1185; Clements et al. 2001). In the latter, for example,a few plants were brought into Marin County, California before 1912 as, "a bit of ol' Ireland"(Pryor & Dana 1952). In many of these areas of introduction, the plant remained localised andnon-invasive, but after about 100-150 years residence in New Zealand, Tasmania and California, Gorsebegan to spread and it has now become a major weed problem, ousting native species from natural orsemi-natural vegetation. Little (1960) commented that, "The invasion of New Zealand by this speciesis more spectacular than (that by) any other plant."
Fodder and other uses
Despite its spiny and intractable nature, U. europaeus is grazed by a variety of large herbivoresand can be a valuable and highly nutritious source of fodder (Edwards & Ekins 1997). Much more couldbe made of this fact and a revival of this feeding practice should be encouraged since the necessarybruising treatment of cut gorse is very easily achieved. In past years, young plants were often severelygrazed by rabbits, but the myxomatosis epidemics of the 1960s and later produced a prolonged rabbitdecline, with the result that Gorse cover has often increased, especially in rough, ungrazed and moreinaccessible ground (Grime et al. 1988, 2007).
Gorse hedges about Irish farmsteads were and still are much prized, not only for the shelter they affordto stock – they are dense and do not drip rainwater – but because they form an admirable clothes line onwashing day and provide a ready source of kindling wood (Estyn Evans 1967, p. 42). Until the 1950s,young Gorse branches were fed to horses and other stock animals. Being so heavily spined, the brancheswere prepared by pounding them on knocking stones – flat slabs or stone basins where the shoots were'melled' with a wooden mallet referred to as the 'whin-bruiser'. Alternatively, they were crushed by alarge round, wheel-like grindstone on a long pole, powered by a donkey walking round a stone-lined'course' as described by Ritchie (1930). These whin stones can sometimes still be found lying around oldfarmyards in Ireland and Scotland. There were even a few water-driven 'whin-mills' in Ulster, althoughas Estyn Evans (1957, p. 110) commented, "their use is almost forgotten, as is the custom ofgrowing a field of whins to provide fodder. This was done in Co Cavan and it is a recognised method ofland utilization in the poorer parts of Brittany."
A survey of the many and ingenious uses to which U. europaeus has been put in Ireland filled a 204page book (Lucas 1960). The uses described by Lucas included: i. Fuel for various purposes. Since Gorseshoots contain a high concentration of oil, they make ideal kindling, but equally they pose a very realfire hazard in some sites. Gorse provides a quick hot blaze suitable for heating ovens and its use asfuel was common down to the 19th century and probably later and it was frequently traded for thispurpose (Rackham 1986, p. 295). It was such an important source of fuel in some parts of B & I, thatcutting of it was carefully shared and regulated in poorer communities (Mabey 1996, p. 230-2); ii.Construction – ie in roofing (as a framework for thatch), as bonding for mud walls and as abrushwood foundation for paths or roads across marshy or boggy ground; iii. Fencing, fodder and beddingfor farm animals; iv. As brushes for brush-harrowing, a means of aerating grassland, or for cleaningchimneys or wells; v. As sticks for the Irish ball game of hurley, or for walking sticks; vi. For dyeingfabrics – giving yellows and brown. It was also used for colouring eggs at Easter (Lucas 1960; Mabey1996).
Weed control
Since gorse is such a powerfully invasive woody weed of disturbed ground, capable of suppressingplantation forests and excluding grazing animals from pasture, there has been considerable research(especially in New Zealand) into improved methods of control using herbicides, grazing management(particularly successful when it involves goats or chickens) and fire, and various attempts at finding ameans of biological control (Krause et al. 1988; Clements et al. 2001; Rees & Hill2001). A very substantial body of research has been published and continues to grow on the whole subjectof Gorse control. As an indication of the scale of this study, an annotated bibliography on the biology,ecology and control of gorse published 40 years ago ran to 266 citations (Gaynor & MacCarter 1981).In the search for a suitable agent for biological control, a survey was conducted of insects feeding onthe shrub in its native European range. This found that many insect populations fed on the plant but didso at very low densities, so that gorse is scarcely affected by their presence. A rust fungus similarlyattacks the plant quite frequently, but it does insignificant damage overall.
The current position regarding control of gorse recognises that a combination of control treatments isoften required, eg laborious manual cutting, prescribed burning, or spot or broadcast herbicidetreatment, plus reseeding with desired species, and/or carefully managed grazing, preferably usinglightweight animals such as sheep and goats, since hoof damage of the soil surface by cattle or horsesfrequently provides gorse with ideal sites for fresh seed germination and re-establishment. A veryuseful and detailed review of the ecology of gorse and its control by Marc Hoshovsky is available on theInternet as an Element Stewardship Abstract of the Nature Conservancy, Virginia, USA athttp://tncweeds.ucdavis.edu/esadocs/ulexeuro.html (viewed May 2021).
Names
The genus name 'Ulex' is Latin and originated from the pen of the ancient Roman writer Pliny. However,all we know is that it was a name he applied to a shrub, possibly spiny, possibly ericaceous and, byanother source, said to resemble Rosemary (Gilbert-Carter 1964; Hyam & Pankhurst 1995). The Latinspecific epithet 'europaeus' is purely geographical and this time the current author (RSF) will notinsult readers by offering to translate it!
English common names are rather plentiful, Britten & Holland (1886) listing 25: however, many of themare related and fall into categories. The three most common are 'Gorse' (from the Anglo-Saxon or OldEnglish 'gorst'), Furze (Old English 'fyrs') and 'Whin'. The latter is conjectured to be from Old Norse'hvin', partly because it appears in place names in the N & E of England, eg 'Whinburgh', inNorfolk, and it is the English common name most frequently applied in the Scandinavian influenced partsof B & I (Grigson 1974). A dialect variant of it is 'Whuns' (Britten & Holland 1886).
'Gorse' is possibly derived from 'Gorst', or in Welsh, 'Gores' or 'Gorest', all words meaning orreferring to 'a waste', ie perhaps because the plant frequently grows on disturbed or waste ground.Other spellings are 'Gorze' 'Gost' and 'Goss' (Britten & Holland 1886). 'Gorse' might alternativelybe derived from the Medieval Latin 'gorassi' or 'gorra', meaning, 'brushwood', since fire making wasoften the major use of the species (Prior 1879).
'Furze', sometimes spelt 'Furres', or more rarely 'Furre', or 'Furrys' (or in dialect variants, 'Furzen','Furzen Bushes', 'Fuzz', 'Fuz', 'Fuzzen', 'Furra', 'Frez', 'Firsun', or even 'Vuzz', 'Vuz' or 'Vuss') isthe most commonly used name for Ulex europaeus in Ireland and in SW England (Britten &Holland 1886; Grigson 1987). It appears to have come from the Anglo-Saxon or Old English 'fyrs', a worddescribed by Prior (1879) as, "of obscure derivation, as are those of so many of our commonestplants". Both Prior and Grigson (1974) suggest the name is in some way related to 'fir' (or in OldEnglish 'fyrh' or 'furh'), meaning 'a fir-tree', the implication again possibly being that Furze, likethe coniferous fir tree, was a very common firewood or fuel and thus the name carries the same firemaking connotation as Medieval Latin 'gorassi' and 'gorra' does for 'Gorse'.
As if these names were insufficient, U. europaeus has also been referred to as 'Thorn Broom','Prickly Broom', 'Fingers and Thumbs' (Wiltshire), 'Thumbs and Fingers' (Somerset) and 'Pins andNeedles', all presumably on account of its spiny-ness.
Several other names given to U. europaeus suggest an association with different spine-armedplants, such as 'Hawth' or 'Hoth' (perhaps a mental link with Crataegus monogyna (Hawthorn)?) and'Qwyce' or 'Quyce', perhaps suggesting the neophyte Quince (Chaenomeles speciosa (Sweet) Nakaiand C. japonica (Thunb.) Lindl. Ex Spach), although these were only introduced in 1796 and 1869respectively. Further names given by Britten & Holland (1886), which cannot be rationalised in thisway, are 'Ruffet', 'Turr' and 'Ling', the latter a name widely applied to numerous heathland plants, butalmost always to members of the Heather family.
Names like 'Great Furze' and 'French Furze' appear to be used to distinguish the different species ofUlex, ie separating U. europaeus from the much smaller U. gallii and U.minor, or perhaps more likely, just the differing ultimate sizes of these plants.
Folklore
As one might expect of such a prominent landscape plant with so many folk uses, Gorse has numerousattached folklore beliefs and it was sometimes pressed into medicinal services also, eg for jaundice (onaccount of its yellow 'signature'), against snake bite and as an insecticide (Vickery 1995). Grieve(1931) mentions that an alkaloid was obtained from the seed which had a powerful purgative action.However, it turned out to be chemically identical to one obtained from Cytisus scoparius (Broom).
Threats
No threats to it, but it is so invasive it can itself become a weedy pest, especially when it isintroduced abroad, eg to New Zealand.
Native, very rare but possibly under-recorded. Oceanic temperate.
4 July 1997; McNeill, I.; Tempo River at Tonyglaskan Bridge.
The solitary Fermanagh record along the Tempo River indicates the possibility that this much smaller,more compact, rather dwarf form of gorse may have been overlooked elsewhere in the VC. Like its muchtaller relative U. europaeus (Gorse), this species typically occupies infertile, acidic soils onneglected or abandoned, stony pastures, scrubby or heathy banks and slopes, cliffs and waste ground.
Western Gorse differs from Gorse not only in its size, but also in its main flowering period, which is inlate summer and autumn. Ulex europaeus has its main flush of flowers in the early spring fromMarch onwards and although it continues to bear a smattering of blossom in a very sporadic mannerthroughout the rest of the year (proving conclusively that kissing is never out of season!), it does nothave any secondary major flush of flowering at any later period of the year.
Although the English common name of this leguminous shrublet is 'Western Gorse' and the New Atlasmap shows a very pronounced overall western predominance to the records in Britain, the main area forU. gallii in NI very definitely lies in the Mourne mountains and the nearby Slieve Croob areas inSE Co Down (H38), where the species is really very common (Preston et al. 2002). Elsewhere in NI,U. gallii is frequently met chiefly on exposed coastal heaths and it appears inland in countiesTyrone (H36), Londonderry (H40) and Fermanagh (H33), but in Tyrone to a much greater extent than to datein Fermanagh.
Threats
None.
Introduction, neophyte, deliberately planted, very rare. European boreo-temperate, but widely naturalisedbeyond its native occurrence.
1948; MCM & D; Crom Castle Estate.
Fermanagh occurrence
This extremely thorny, much-branched, thicket-forming, root-nodule bearing, dioecious shrub, 1-11 m inheight was deliberately planted on the shore of Upper Lough Erne in the Crom Castle estate, which is thesolitary station for the plant in Fermanagh to date. It was recorded as still present by the staff ofthe National Trust sometime between 1980 and 1990.
Vegetative reproduction
H. rhamnoides, which is deciduous, has very vigorous growth and strong powers of vegetative spreadby means of both a suckering rhizome and by layering, especially when growing in suitably open soils(preferably sand or gravel) and in the well-lit, wind-sheltered habitats the shrub requires (H.Ainsworth, in: Stewart et al. 1994).
Flowering reproduction
Sea-buckthorn bushes flower in the winter and early spring before the leaves appear, a feature thatassists its wind-pollination. Both male and female shrubs bear their flowers on the previous year'sgrowth (Tutin et al. 1968). The male flowers are borne in small spikes or catkins of 4-8 reducedflowers, each consisting of four stamens and two sepals borne on a short receptacle (Pearson &Rogers 1962). The female bushes produce their flowers just as the first leaves open and are borne inshort axillary racemes. The solitary, sessile ovary at the base of the perigynous receptacle contains asingle ovule. The fruit is a solitary seeded achene, enclosed in the orange swollen fleshy receptacleand they appear as dense clusters of 6-10 mm diameter orange berries in the autumn (Pearson & Rogers1962). They are attractive to birds, especially those of the Thrush family and the Magpie, wheneverother food is limited (Lang 1987; Snow & Snow 1988).
The size of the introduced colony of Sea-buckthorn at the Crom estate has not been ascertained by us, butthe plant has the ability to spread, naturalise and become an invasive nuisance in other plantcommunities, as it has done in several coastal dune systems around NI (FNEI 3). On account ofthis its distribution should be kept under review.
An important, briefly dominant, shrubby colonist of open areas in the Late-glacial period prior to itsreplacement by taller, shading trees (Godwin 1975), H. rhamnoides is considered native in B &I only on some stretches of the E coast of England from Bamburgh in Northumberland (VC 67) to Hastingsin E Sussex (VC 14). As the New Atlas map indicates, it is introduced in very many coastal and amuch smaller number of inland sites in Britain, and to a lesser extent in Ireland, although here it isalmost exclusively coastal. The most inland Irish station mapped is at Crom in Fermanagh.
Typical habitats in B & I include stabilised sand-dunes, coastal banks and sea cliffs, although incoastal situations it always grows best when sheltered from the wind. The soil reaction where it growsis always neutral or basic and the root nodules probably fix nitrogen. The fact that nettles(Urtica spp.) are very frequently associated with Sea-buckthorn stands, they being stronglynitrophilous, does suggest that N-fixation is taking place (Pearson & Rogers 1962).
European and world occurrence
Hultén & Fries (1986) describe this is a Eurasiatic species that includes as many asnine subspecies. Apart from coastal habitats, as in B & I, H. rhamnoides occurs on thecontinent on river gravel, alluvium and lateral moraines in mountain regions in Europe and Asia. It isconsidered native throughout a considerable portion of Europe from 68°N in Norway to N Spain, C Italyand Bulgaria, and from NW France to Finland and Moldavia. It is, however, local in its occurrence andabsent from wide areas of the continent. At the same time, it is frequently planted for ornament ingardens and parks, or to stabilise sand or gravel and it has become naturalised in many places (Tutinet al. 1968).
The overall distribution as mapped by Hultén & Fries (1986, Map 1310) shows H.rhamnoides s.l. stretching from the Pyrenees eastwards through C Europe, the Caucasus and the CAsiatic mountains to the highlands of N China.
Threats
The colonizing ability of this introduced spiny shrub and the difficulty and expense of eradicating orcontrolling it mean it should be regularly monitored, which is probably not happening at present.
Native, occasional. Circumpolar temperate.
1939; Praeger, R.Ll.; lakelet west of Lough Galliagh.
May to September.
Growth form and preferred habitats
Whorled Water-milfoil is a submerged aquatic perennial with long, flaccid leaves that are markedly longerthan the stem internodes that bear them. Mid-stem leaves are 15-45 mm long, 1.4-4(-5.5) times as long asthe adjacent internodes and are borne in whorls of four or five. M. verticillatum grows best instill to slow, sheltered waters, over fine inorganic or peaty soils at shallow depths down to 100 cm(Haslam et al. 1975; C.D. Preston, in: Rich & Jermy 1998). It rapidly becomes scarce indeeper waters down to about 3 m, depending upon levels of clarity and light penetration. M.verticillatum can tolerate some degree of water turbidity in shallow situations, but it grows ina much more restricted pH range than M. spicatum (Spiked Water-milfoil), being more or lessconfined to moderately acid to near-neutral, moderately productive, meso-eutrophic conditions (Preston& Croft 1997). When water levels drop during drier periods in summer, provided this is a gradualprocess, Whorled Water-milfoil very readily develops an emergent, dwarf, terrestrial growth form thatgradually becomes stranded on the exposed mud, and it persists in this form until re-submerged (Aikenet al. 1979).
Fossil evidence and status
In common with the other two Water-milfoil species in Fermanagh (M. spicatum and M.alterniflorum (Alternate Water-milfoil)), fossil evidence suggests that M. verticillatumis a very ancient native species, probably having survived throughout the last full-glacial period insitu in B & I (Godwin 1975).
Vegetative reproduction
As is general in all aquatic species, reproduction of M. verticillatum is heavily weighted towardsasexual means, involving in this case the detachment of specialised autumn-produced, robust,club-shaped, dark green, leafy vegetative buds (turions), borne either in leaf axils or terminal onstems, plus stem fragmentation. Detached turions demonstrate dormancy which can be broken by a coldtreatment (4°C) (Weber & Noodén 1976). Both of these vegetative methods enable potential rapidincrease and an easy means of dispersal in already colonised water systems.
In Ireland, boating and fishing on the more northerly Royal Canal can be affected by dense stands of thisaquatic and weed control involves mechanical removal in the autumn, after the overwintering turion budshave been formed on the lower parts of the stems (Caffrey & Monaghan 1995).
Flowering reproduction
Flowering and seed production certainly occurs, but as with other aquatics, seedlings are not readilyobserved. M. verticillatum, like the other two native members of the genus already mentioned, hasa terminal inflorescence, 7-25 cm long, of small, whorled, unisexual and bisexual flowers held above thewater surface (ie emergent). The unisexual flowers have reduced petals and sepals that sometimes areshed and they are thus well adapted for wind pollination. The bisexual (perfect) flowers have all theirparts present and usually occur between the male and female flowers (Sell & Murrell 2009). Powderypollen is released on the breeze from anthers on long filaments (Cook 1988). Flowering takes place inJuly and August. The fruit is a sub-globose schizocarp 2 mm across, that splits into four 1-seededmericarps (Sell & Murrell 2009). Very little is known about the breeding systems of aquatic species,so it really is not possible to generalize on the question of in- or out-breeding in relation to themode of pollination of these plants (Cook 1988).
Fermanagh occurrence
With the exception of one site in Co Down (H38) and a few scattered up along the Co Antrim (H39) coast,all recent records of this aquatic for NI are from the Lough Erne and Lough Melvin basins in Fermanagh.This species declined from rarity to local extinction at sites around Lough Neagh, possibly sometimeafter the late 1940s (Harron 1986) or perhaps even earlier in the 1920s (Hackney et al. 1992).
As the tetrad distribution map demonstrates, M. verticillatum remains widespread, but is stillonly occasional in Fermanagh, confined to the larger lowland lakes, and scattered in the S and W of thecounty. Locally it has been recorded in a total of 27 tetrads (5.1%), 23 of which have post-1975records. Almost all of its sites are in drains, or on fen-fringed, or muddy lakeshores, washed bylime-rich waters around the basins of the larger lakes in the VC.
M. verticillatum is the least frequent of the three species of Myriophyllum found inFermanagh. As an indication of scarcity, M. spicatum (Spiked Water-milfoil) is over nine timesmore likely to be found than M. verticillatum, while M. alterniflorum is also over threetimes more frequent than it in the VC.
Irish occurrence
In Ireland as a whole, Whorled Water-milfoil is most frequent in lakes, ponds, ditches and canals on theCentral Plain in Leinster and Connaught. Here, Carboniferous limestone underlies inorganic glacial tillsand more recent accumulations of raised and blanket bog peat, but drainage waters in the region arestill essentially lime-rich. During the last 50 years, with the advent of mechanised peat extraction andpeat-burning power stations, many areas of very extensive, deep peat deposits have been almost entirelyremoved in this region of Ireland, further exposing the underlying limestone geology.
Both editions of the Flora of the County Dublin (Colgan 1904; Doogue et al. 1998) and theintervening Supplement (Brunker et al. 1961), indicate the highly significant role theRoyal and Grand Canals linking Dublin with the great River Shannon and its lakes and tributaries furtherwest have played in the spread and persistence of this always rare aquatic species.
Changes in British and Irish occurrence
There are huge differences in the extent of the recording of this species between the two BSBIAtlas surveys produced in 1962 and 2000. The large number of older, pre-1970 symbols on theNew Atlas map indicate that the species decline observed in the earlier BSBI Atlas surveyin England has continued and the plant is now regarded as scarce there (Walters & Perring 1962;Stewart et al. 1994). Factors such as widespread eutrophication from agricultural run-off andatmospheric nitrogen pollution, or the deepening of water channels during drain clearance, very readilydestroy suitable habitats for this species.
Meanwhile in Ireland, an increased recording effort, assisted by better knowledge of aquatic species ingeneral, means that C Ireland now maintains the best M. verticillatum populations anywhere inthese islands (C.D. Preston, in: Preston et al. 2002). Although M. verticillatum doesextremely well in the Irish canal system and can grow to nuisance proportions, under-recording is stilla possibility here and it may well be that some of the local populations are small and not easilynoticed. They certainly are very scattered, reaching even a few isolated hectads on the N Antrim coast(H39).
European and world occurrence
M. verticillatum is widely distributed in temperate parts of Eurasia and N America and iscircumpolar. Records from Argentina and Chile, indicated by Hultén & Fries (1986, Map 1373), areerrors (Preston & Croft 1997).
Threats
Factors such as widespread eutrophication from agricultural run-off and atmospheric nitrogen pollution,or the deepening of water channels during drain clearance, may very easily destroy suitable habitats.
Native, locally frequent. Eurasian temperate, but naturalised in N America and now circumpolar.
1900; Praeger, R.Ll.; around Enniskillen.
May to December.
Growth form and preferred habitats
This submerged aquatic rhizomatous perennial has whorled, feather-like leaves and branched shoots thatvary from 0.5-7.0 m long. It often forms extensive mats at the water surface. M. spicatum usuallygrows on sand or gravel bottoms with an admixture of organic silt in lowland lakes and flowing waters,which can be up to 4 m or more in depth, but are often shallower, and which can be mildly acidic,near-neutral, or more usually calcareous and moderately to richly productive (ie mesotrophic toeutrophic) (Preston & Croft 1997). M. spicatum is absent from acidic waters and almost sofrom pure sandy bottoms (Reed 1977). In highly calcareous waters, plants precipitate lime and becomewhite and marl encrusted. M. spicatum can grow and survive in water down to 17 m deep,although this is very exceptional, the normal maximum depth being closer to 5 m. The success of theplant depends much more on light, temperature and nutrient levels than the actual water pressure atdepths below about 5 m (Dale 1981). Most plants are found attached in water 65-150 cm in depth (Reed1977).
M. spicatum can also tolerate brackish water in estuarine conditions and thrives in water with asalinity up to 10 parts per thousand, but grows more slowly at a salinity of 15 parts per thousand(Beaven 1960). This level of salt tolerance explains how the species might be transported around theworld in ship's ballast, as appears might have been the case at Chesapeake Bay in its originalintroduction to N America. Propagules might have been either seed or vegetative stem fragments (Aikenet al. 1979).
The light compensation point in M. spicatum is approximately 1-2% of surface light (Grace &Wetzel 1978), but it has nearly optimal configuration of photosynthetic tissue because it concentratesits leaf biomass near the water surface (Adams et al. 1974). M. spicatum displays a verywide tolerance of water reaction, indeed ranging from pH 5.4-11.1, and in wide lakes like Lower LoughErne and in river estuaries, it can withstand considerable wave action, although it really grows best inmore sheltered coves.
M. spicatum grows vigorously and can often shade out other aquatics, including Potamogetonspecies like P. crispus (Curled Pondweed) (Nichols & Shaw 1986). However, submerged aquaticsare relatively unproductive when compared to the productivity of terrestrial and emergent plants. Onereason for the low productivity is believed to be the limited carbon availability in the aquaticenvironment, which is explained by the slower rate of diffusion of carbon dioxide in water compared toin air. The anatomy of M. spicatum, however, provides a large internal lacunal system that actsas a gas reservoir that is capable of retaining respired CO2, allowing diffusive gas exchangeto take place between roots and shoots (Grace & Wetzel 1978).
While M. spicatum can also grow on a wide variety of sediment types, like other aquatics it growsbest on fine sediments where organic matter ranges from 10-25% (Pearsall 1920). Coarse substrates suchas larger gravels probably do not offer good anchorage and they may be nutrient poor. However, finebottom sediments can become too soft and flocculent to support plant growth, plants then being unable toroot and anchor sufficiently in soft, moving bottom substrates (Nichols & Shaw 1986).
Emergent plants
As in M. verticillatum, emergent aerial shoots develop in M. spicatum if the water bodytemporarily dries out or water level drops slowly and sufficiently to expose a shoreline. The leaves ofthe land plant are smaller, stiffer and they are less divided. If re-submerged, new growth withtypically divided aquatic leaves redevelops in around 7-10 days (Aiken et al. 1979).
Flowering reproduction
Except in deep or turbulent water, Spiked Water-milfoil populations flower and set seed regularly. Theinflorescence is a terminal spike, 5-20 cm long, of reduced flowers in whorls of four, often pink incolour. The stem, 5-20 nodes below the spike, is double the rest of the stem in width, very rigid anddistinctly curved so that this inflated portion lies parallel to the water surface. The stigmas ripenwell before the anthers do, favouring cross-pollination. Cook (1987) notes that M. spicatum isself-compatible, but says that there is a good chance of cross-fertilization. The anthers are linear,1.8-2.2 mm long producing large amounts of dry pollen. The spike is erect when flowering takes place inJune and July and is held above the water surface. After wind pollination takes place, it lies parallelto the water surface during fruit set in August and September. The fruit is a 4-lobed, 3 mmsub-orbicular schizocarp. The ripe fruit eventually breaks up into 1-seeded mericarps and the 'seeds'float for a short time, perhaps for 24 hours or so, and may disperse in any available water flow. Theactual true seed has a stony endocarp and shows prolonged dormancy which can be broken by chilling or byabrasion (Aiken et al. 1979).
In common with other aquatic species, despite vast numbers of seed being produced (eg four million perhectare in one Canadian study), seed germination is extremely erratic and field records of seedlings areeither non-existent (Aiken et al. 1979; Preston & Croft 1997) or rare (Hartleb et al.1993).
Unlike reports from elsewhere in Britain, none of the Fermanagh records of M. spicatum are fromnewly created water bodies, such as reservoirs, flooded quarries, gravel or sand-pits, and thus there isno evidence of jump-dispersal which requires birds or other vectors to transport the seed eitherinternally, or more likely, externally, in mud on their feet or plumage (Ridley 1930, p. 546). Nor arethere any examples of dumping of unwanted, excess, aquarium or pond plant material which, apart fromshipping ballast, is another method by which this species might have been transported beyond its nativeEurasian distribution (Reed 1977; Preston & Croft 1997).
Overwintering perennation, vegetative reproduction and dispersal
The plant is a rhizomatous perennial that dies back to the roots in winter. Unlike M.verticillatum, it does not develop specialised turion overwintering buds, but it does regeneratefrom axillary buds borne on the previous year's stem base that are easily detached (Preston & Croft1997). It can also spread locally by the lateral growth of the rhizome. Thus its effective reproduction,as with all aquatics, is largely asexual and highly dependent on detached vegetative buds, stemfragmentation and rhizome spread, all of which means of increase are highly developed in M.spicatum, allowing its rapid dispersal and expansion in the water bodies it colonises.Auto-fragmentation of M. spicatum plants typically occurs after its flowering period in theautumn, each upper stem segment being capable of forming roots and starting a new colony the followingsummer (Reed 1977; Nichols & Shaw 1986).
Strong waves and water currents, plus human activities such as motor boating and mechanical weedharvesting, all help to produce and distribute stem fragments (Aiken et al. 1979; Preston &Croft 1997). Colonisation success of M. spicatum propagules has been shown to be best in latesummer in shallow (0.5 m) water, on rich organic sediments; mortality was highest in deep water withcalcareous, nutrient poor sediments in early autumn (Kimbel 1982).
Relationship with algae
The relationship between aquatic macrophytes and algae appears to depend on whether the algae are freeliving or epiphytic on the macrophyte and the nutrient status of the water body they both inhabit.Macrophytes can alter local water chemistry around them and their presence provides a substrate forperiphyton growth (ie on and around them). By the same token, planktonic and filamentous algae may shademacrophytes, but they also provide easily available food for grazing invertebrates that make use of theshelter provided by macrophytes (Nichols & Shaw 1986).
Dense growths of large aquatic plants may have an inhibitory effect upon phytoplankton and rotiferplankton in small water bodies and, in larger lakes, aquatic weeds may display antagonistic activitytowards phytoplankton. Fitzgerald (1969) observed that areas of Lake Winga, Wisconsin with an unusuallyextensive weed bed of mainly Myriophyllum sp., were quite plankton free. The current author (RSF)cannot discover whether more than one Milfoil species was involved (M. spicatum certainly was thedominant species of the genus present), as after all these years, Fitzgerald's 1969 paper is stillhiding behind the publisher's paywall. Milfoil species form an excellent host for periphyton becausetheir finely dissected leaves provide a large surface area for colonisation; the epiphytes also benefitfrom the organic nutrients that are excreted by macrophytes (Wetzel 1975). It has been suggested thatepiphytes may provide an easily grazeable food supply, thus protecting the macrophytes they grow uponfrom invertebrate predation (Hutchinson 1975). However, Milfoil plants from declining populations areoften coated with algae or a brown slime of periphyton (Nichols & Shaw 1986). Only one species(sp.)? or more than one (spp.)?
Relationship with waterfowl
The primary importance of macrophytes like M. spicatum to waterfowl is as food. However,information on the topic of food uses is sparse. For instance, Tamisier (1971) found thatMyriophyllum and Potamogeton seeds formed less than 2% of the diet of Teal ducks (Anascrecca). Invertebrates associated with macrophytes are important duck foods, especially whenyoung ducklings need a protein-rich food source (Krull 1970). Even hydrophytes that are themselvesconsidered poor waterfowl food plants are believed to be of indirect importance to birds because theyharbour large quantities of macroinvertebrates (Nichols & Shaw 1986).
Fermanagh occurrence
M. spicatum is by a considerable margin the most common of the three native Water-milfoil speciesin Fermanagh, being recorded in 91 tetrads, 17.2% of those in the VC. It persists not only in both partsof Lough Erne, the major water bodies of Fermanagh, but also in managed ditches and riverbeds drainingthe land around these lakes or lake systems, since the shoreline of Upper Lough Erne, in particular, isextremely heavily dissected. As the Fermanagh tetrad map shows, M. spicatum is very much confinedto the Lough Erne area and along the marl lakes on the River Finn. The feeding waters in these areasderive from catchments of limestone geology.
The observed linkage between the distribution of M. spicatum and calcium levels might be partiallyexplained by the correlation with water colour which may limit photosynthesis in submerged species.Gibson (1988) found all calcium-rich waters in Fermanagh had clear water and all highly coloured waterswere low in calcium. The patterns of human settlement and of farming mean that both parts of Lough Erneare much more subject to eutrophication than the huge number of smaller lakes and ponds widely scatteredacross the VC (Gibson et al. 1980, 2003; Gibson 1988).
In the summer of 1948, Meikle and colleagues recorded M. spicatum in Lough Navar on the westernplateau, a site where it has not been seen since, although M. alterniflorum, with which it cancoexist, is still present there. The creation of the coniferous Forest Park around Lough Navar hasundoubtedly led to acidification of the waters and possibly substantial phosphorus enrichment (Gibson1976), but M. spicatum tolerates a wide range of both these factors (Aiken et al. 1979),so the current author (RSF) cannot easily explain the apparent demise of the species here.
Irish occurrence
M. spicatum is well, but very unevenly distributed throughout Ireland. The majority of more recentIrish records are from the north of the island – the outcome of several large-scale lake surveys carriedout in NI by the DOENI from 1985 onwards.
British occurrence
Spiked Water-milfoil is widespread throughout all of Britain, from the Channel Isles, thinningnorthwards, but still managing to reach the southern tip of Shetland (VC 112). In the SE of England thespecies is much more frequent than elsewhere, and especially so to the east of a line drawn betweenSwansea and Hull (New Atlas). Like the majority of aquatic plants, M. spicatum is nowbetter recorded than in the first BSBI Atlas (Perring & Walters 1962), and it is the mostcommon Myriophyllum species colonising the eutrophic lowlands of the country (C.D. Preston, in:Preston et al. 2002). It has probably increased in abundance across Britain in the last 150 yearsthrough a combination of eutrophication of existing grazing marshes and the expanded availability ofother suitable man-made habitats including sand- and gravel-pits and disused quarry pools (Preston &Croft 1997).
European and world occurrence
M. spicatum is widely distributed (almost ubiquitous) in Eurasia and extends north-westwards toIceland and Greenland. It includes subsp. spicatum in Eurasia and subsp. exalbescens(Fern.) Hult. in N America. Variation between these two subspecies certainly overlaps and there islittle cause to regard them as separate species (Hultén & Fries 1986). The species sens. lat.is widespread in boreal and temperate regions of the N Hemisphere and belongs to the Circumpolar plants(Hultén 1974, Map 171). From its centre of origin in Eurasia it extends southwards into N, C & SAfrica (Swaziland, Transvaal, Natal, the Cape) and eastwards to S Asia, including the Himalaya, Sumatra,the Philippines and Japan. In some of these more eastern areas, it is probably introduced and this iscertainly also the case in all of N & S America (Hultén & Fries 1986, Map 1374).
M. spicatum is a serious aquatic alien nuisance weed in many regions of the world where it hasbeen deliberately or accidentally introduced. This is especially the case in NE North America where itposes a danger to native species in established vegetation (Aiken et al. 1979; Nichols & Shaw1986). Reed (1977) described M. spicatum as, "an economically important and ecologicallydangerous waterweed". Spiked Water-milfoil has been present in N America since 1848 and it hasspread from the E coast to the W coast of both the US and S Canada. Having said this, there have beenproblems in the past distinguishing M. spicatum from a native American species referred to asM. exalbescens Fern., although Reed (1977) is clear that both species occur and that it was M.spicatum that became a problem in large water bodies in N America in the late 1950s and 1960s.
M. spicatum was present in Canadian waters for around 50 years before it was recognised as asignificant aquatic weed. The population there then exploded in the late 1950s, probably in part due tothe occurrence of dramatic weather events (Nichols & Shaw 1986). Some long distance dispersal ofM. spicatum in N America is known to be related to the aquarium and aquatic nursery trade (Reed1977). Shorter distance dispersal is attributed both to transport of plant fragments on boats andtrailers moving from lake to lake and also to natural water movements (Nichols & Shaw (1986).
Threats
None. This species copes well with highly eutrophic conditions and, indeed, it is often a good indicatorof them.
Native, occasional. Suboceanic boreo-temperate, but also in N America.
1882; Stewart, S.A.; Carrick Lough, Dresternan Td.
March to November.
Growth form and preferred habitats
This submerged, aquatic perennial with whorls of three or four much dissected leaves, grows in a widerange of still to swift-flowing waters, usually in the more acidic, soft-water, upland parts of thecounty although it can also occur in waters of high pH and low and high concentrations ofHCO3- (Spence 1967). It is a rhizomatous species with no specialised overwinteringturion buds, but is perfectly capable of vegetative regeneration and colonisation involving shortlengths of fragmented shoot. Since it prefers less productive, base-poor waters, in Fermanagh thiswater-milfoil species is much less common around both parts of Lough Erne than is M. spicatum(Spiked Water-milfoil). Compensating for this, however, it is very widely scattered around thecounty, particularly in fairly remote lakes and pools on the western plateau uplands.
As its species epithet indicates, M. alterniflorum bears its upper flowers alternately oropposite, not in whorls like M. spicatum does. The mid-stem leaves are also more divided than inM. spicatum, having 13-38 segments, rather than 6-18 in the latter (Crawley 2005).
The evidence of its widely scattered local distribution clearly illustrates that M. alterniflorumis more capable than M. spicatum of dispersing itself and colonising new water-bodes around itsexisting sites and at higher elevations above them. It has, for instance, been recorded from a pool inthe quarry beside Keenaghan Lough and also from a distinctly peaty pool in a cut-over bog in theWhitehouse Cave area. The latter example indicates the extremely wide ecological range of this species,which also tolerates highly calcareous waters and quite often accompanies M. spicatum in lesseutrophic situations (Preston & Croft 1997).
M. alterniflorum typically grows in water 30-300 cm deep, generally somewhat shallower than M.spicatum, although there is considerable overlap between their habitat requirements and they canoccur together.
Flowering reproduction and genetic variation
Alternate Water-milfoil flowers profusely in shallow water in mid-summer, producing wind-pollinatedflowers on a short, 1-2 cm, emergent spike that is female at the bottom and male at the top. The tip ofthe spike is drooping when in bud. Very occasionally, hermaphrodite (ie bisexual, perfect) flowers maybe produced between the unisexual flowers. The fruit is a four-lobed, 1.5-2.0 mm schizocarp, splittingwhen mature into four single-seeded mericarps (Sell & Murrell 2009); the seeds are viable. A studyof enzyme variation found M. alterniflorum displayed a large degree of variability within andbetween populations (Cook 1968; Harris et al. 1992).
Evidence from the enzyme study showed sexual reproduction via random mating was common in the M.alterniflorum populations and that therefore outbreeding is frequent. Sexual reproduction doesnot necessarily lead to genetic diversity, but the extent of outbreeding will also be important. Thefact that M. alterniflorum grows in sites with a great range of water chemistry, helps explainthe high degree of genetic variation found (Spence 1967). "For such a species, genetic diversitymay provide one means of exploiting the different environmental conditions." (Harris et al.1992).
The species is native, fossil pollen evidence from sediments in Scottish lochs indicating its presencefrom immediately after the last glaciation around 12,000 years BP (Pennington et al. 1972).Populations at different sites have therefore had a long time to accumulate genetic diversity and thereare a very large number of freshwater sites where the species occurs, meaning there is a substantialpool of genetic variation. If gene flow can occur between sites by transfer of vegetative shoots (eg bybirds), or by pollen transfer on the wind, this leads to increased genetic variation within a site(Sculthorpe 1967; Harris et al. 1992). The latter authors concluded, "Although a largeamount of the morphological response and the biochemical and physiological response to the environmentby freshwater macrophytes is phenotypic, the results presented here suggest that there is also thepotential for genotypic differences, and hence for the existence of ecotypes within a given speciesadapted to different conditions."
An alien American variety
Slender, worm-like, prostrate plants of M. alterniflorum with very short leaves (only 3-8 mm ascompared to normal 8-26 mm length), and with equally short internodes, only 2-8 mm rather than 8-17 mm,were first noticed growing in shallow water on sandy substrates in clear water in Lough Beg (H40) andLough Neagh (H36-H40), and again in Lough Ree along the R Shannon, just N of Athlone (H23, H24 &H25) by S.A. Stewart back in 1867. This unusual dwarf variety grew in these Irish sites in the apparentabsence of typical M. alterniflorum (Praeger 1938b). At the same date (and in the same Journalof Botany issue), Pugsley drew attention to the fact that these dwarf plants were very similarto the normal form of the species in N America, so he then named them var. americanum Pugsley.The same or a similar form was recorded elsewhere in Ireland on the larger lakes on the R Shannon, onsmaller lakes in W Donegal (H35) and in the Scottish Hebrides (VCs 103, 110 & 111) (Harron 1986;Preston & Croft 1997; C.D. Preston, in: Rich & Jermy 1998). However, due to subsequent habitatchanges, including eutrophication, some of these populations appear to have latterly declined ordisappeared. There is a definite need for further investigation of the true nature of the American formof the species (Preston & Croft 1997).
Fermanagh occurrence
M. alterniflorum is thinly and widely scattered in Fermanagh, but is more frequent in the west ofthe county at all altitudes.
There are records of it from 61 Fermanagh tetrads, 11.6% of those in the VC, but as the distribution mapindicates, eight of them are pre-1975 only, suggesting a possible slight decline of the species over theperiod from the mid-1940s. The local habitats range across lakes, tarns, rivers, drains and pools inquarries and on cut-over peat bogs.
Fossil occurrence
Fossil pollen of M. alterniflorum has been found in a full-glacial freshwater deposit of MiddleMidlandian age found at Derryvree, near Maguiresbridge in Fermanagh, and radiocarbon dated to 30,500 BP(Colhoun et al. 1972). The flora and fauna of the deposit indicate that open tundra vegetationand a periglacial climate prevailed at that period. Godwin (1975) reports similar evidence of longpersistence of this aquatic in Britain, since at least the middle of the last glacial period andpossibly a lot longer.
British and Irish occurrence
The New Atlas hectad map shows M. alterniflorum is frequent and widespread in both acidicand calcareous waters throughout N & W parts of B & I across the whole range of latitude.However, in common with other calcifuge species, in S & E England it is very much more scarce,scattered or absent and is confined to acidic, mesotrophic or oligotrophic waters. Suitably acidic, lowproductive habitats are infrequent in these parts of England since many sites have been destroyed in thelast century or so through drainage and agricultural intensification operations (C.D. Preston, in:Preston et al. 2002).
European and world occurrence
M. alterniflorum is mainly restricted to boreal and temperate zones in N & W Europe and mid-Eand mid-C parts of N America, making it an amphi-Atlantic species (Hultén 1958; Hultén & Fries 1986,Map 1375). As mentioned above, the American form of M. alterniflorum is very dwarf in comparisonwith the European plant.
Threats
None.
Introduction, neophyte, a very rare garden escape.
1 April 1989; Northridge, R.H.; streamside, Clonelly, NW of Kesh.
April to May.
Growth form, identification and preferred habitats
G. tinctoria is one of the most popular, decorative, architectural-scale garden plants for largegardens and demesnes with appropriately spacious water features to display it. It was introduced fromits native Chile and Argentina in the mid-19th century and was promoted in Victorian and Edwardianhorticulture for planting around ponds and in damp areas. This huge, herbaceous, slightly tender, coarseperennial with thick, surface-spreading stems that are entirely rhizomatous, has enormous rhubarb-likeleaves ± 2.5 m high and 1.5 m across, palmately 5-9-lobed, rounded in outline with sharply pointed,jagged-toothed lobes, cordate at the base, borne on 2 m tall, stout petioles, densely studded withshort, conic, prickly spines that give the leaf stalk a reddish-brown appearance (Sell & Murrell2009). The huge leaves and the up to 100 cm tall inflorescence at fruiting are deciduous, wilting anddecaying in the autumn to produce a very heavy leaf litter that protects the surface-exposed horizontalrhizome from frost. Massive, winter-resting, perennating buds, up to 25 cm long are formed on therhizome (Webb et al. 1988).
Throughout B & I, there has been a history of confusion between G. tinctoria and G.manicata Linden ex André from Brazil, that were introduced to European gardens at much the sametime, but the latter does not set viable seed in B & I. Other differences that distinguish G.manicata include its leaves that are even larger (2 m across) than those of G. tinctoriaand are pedately lobed (ie five leaflets arising from a single point on the petiole) and although thepetiole is covered with numerous small spines, each with a red tip, the overall appearance of the leafstalk remains green. G. manicata has a large, erect, conical inflorescence, while that of G.tinctoria is somewhat more cylindrical in shape and its numerous individual side branches orcatkins are shorter and less open than those ofG. manicata. Also, G. manicata isvery rarely cultivated in Ireland and there is only one reliable record of its naturalisation in thecountry, in W Galway (H16) (Clement & Foster 1994; Scannell & Jebb 2000).
In B & I overall, G. tinctoria is a fairly frequent escapee 'over the garden fence' and hasbecome naturalised and persistent in a variety of damp to wet and sometimes already shaded habitats,such as rough grassland on riverbanks, stream-sides, ditches and damp hollows, especially in lowlandareas near the coast in Ireland (Webb & Scannell 1983; Jarvis 2011). Remarkably, considering theenormous size of the plant, it sometimes establishes itself on sheltered sea cliffs and on old quarryrock faces. In addition to self-sown seed, or in place of it, some extra-garden populations mustoriginate from material discarded by desperate gardeners unable to accommodate the gigantic scale towhich the plant grows when given suitably damp, nutrient-leached, boggy ground (Clement & Foster1994; T.D. Dines, in: Preston et al. 2002).
The South American native range of G. tinctoria falls within the temperate climatic zone with apredominantly Mediterranean climate and a tropical moist climate subzone where annual rainfall is high(>200 cm) and mean annual temperatures are cool (10°-14°C) (Williams et al. 2005). Beyond itsnative range, the species grows and thrives in regions where winter temperatures are mild, any frostsare not severe, and both precipitation and humidity are typically high (Gioria & Osborne, 2013).
In garden growing experiments in Dublin, four days without water caused permanent wilting and desiccationof G. tinctoria leaves, with no sign of recovery of the outer edges of the leaves afterre-watering (Hennessy 2009). Young seedlings were found to be unable to survive being kept twoconsecutive days under waterlogged conditions (Gioria 2007). Adults are rarely found in areas where theentire rhizome is permanently under water (Campbell 1994). G. tinctoria grows on a range of soiltypes, but in Ireland is typically found on mineral soils and relatively acidic, wet soils and is notcommon on organic/peat substrates (Gioria & Osborne 2013).
G. tinctoria is commonly associated with a range of ruderal or
competitor-ruderal species, including Apium nodiflorum (Fool's-water-cress), Galium aparine(Cleavers), G. palustre (Marsh-bedstraw), Persicaria maculosa (Redshank), Stachyssylvatica (Hedge Woundwort) and Urtica dioica (Common or Stinging Nettle).
In coastal areas, it is associated with cliff faces dominated by Armeria
maritima (Sea Pink or Thrift), Festuca rubra (Red Fescue) and Plantago species(Plantains) (Gioria & Osborne 2010, 2013). Its competitive ability, said to be only realised in wetand/or humid habitats (Campbell, 1994), particularly along water courses, coastal cliffs and in wetmeadows, derives from a range of traits including its very large stature, perennial nature, capacity forfixing nitrogen through a unique intracellular symbiosis involving cyanobacteria (Nostoc), highrelative growth rate, early season growth, dense leaf canopy, abundant litter and the persistence of itsseeds and rhizomes (Gioria & Osborne 2013).
Gunnera tinctora is considered competitively superior to most associated species, although arecent study reported the displacement of long term G. tinctoria stands by Japanese Knotweed(Fallopia japonica) (Gioria et al. 2011). Any competitive potential of Japanese Knotweedfor control of G. tinctoria is of no benefit, as it also is a highly invasive alien species.
Flowering reproduction
The massive, erect, cylindrical inflorescence of G. tinctoria is a spike-like panicle up to 100 cmin height that arises directly from the rhizome. It consists of a central rachis or stalk, denselycovered with a very large number of slender, catkin-like branches 8-11 cm long, bearing very small,male, female and bisexual, petal-less flowers, each with parts in pairs. The flowers self-pollinate andproduce masses of small, red-orange fleshy, drupe fruits. It seeds freely and prolifically in the mild,wet, growing conditions of W Ireland and the Channel Isles, where frosts are infrequent and not verypenetrating. Each inflorescence can produce 8,000 seeds and an individual clone could muster 200,000 ormore in a season. The edible fruits attract birds that feed on them and disperse the seed and wind andwater are also likely to help dispersal to some extent. Webb & Scannell (1983) suggested that in theW of Ireland, seed might also be dispersed by sheep (presumably attached to their woolly coats).
Fermanagh occurrence
It is perhaps fortunate that G. tinctoria has only been recorded three times in Fermanagh, inwidely spaced streamside sites in which it does not yet seem to be actively spreading. In addition tothe first record above, the details of the other two sites are: Tattinweer Bridge, Tempo River, May1989, RHN; and on the Crom side of Inisherk Bridge, Upper Lough Erne, February 2010, RHN & HJN.
Irish occurrence
In Ireland, G. tinctoria is more commonly found along roadside banks and ditches in the milder,damper W & SW than elsewhere. It is extensively established and naturalised, forming large dominant,± single species stands in numerous parts (both coastal and inland) of W Mayo (H27) and W Galway (H16)(Connemara), and particularly frequent around Achill Island and Curraun. It is unquestionably invasivein these two latter VCs at least (Webb & Scannell 1983; Hickey & Osborne 1998; Reynolds 2002).It is distinctly alarming to see this huge, vigorous, nitrogen-fixing, invasive alien, thriving, seedingfreely and expanding its territory year by year as an abundant, fully naturalised species in the W ofIreland. It is doing so along the roadside bank overlooking the coast at Leenane in W Galway (H16), avery much visited tourist area of the country, and a site where it was first reported as naturalised andplentiful by Praeger as long ago as 1939 (Praeger 1939). In a few areas of coastal Mayo, dense thicketsof G. tintoria up to 3 m high are out of control and cover patches up to 0.5 ha in extent ofupland well-drained ground, in which no other species can compete (Jarvis 2011). Sylvia Reynolds (2002),in her Catalogue of Alien Plants in Ireland, lists records from twelve Irish VCs, not includingFermanagh (H33).
Under experimental garden cultivation, G. tinctoria was capable of very similar growth andproductivity in the more frost-affected Dublin area when compared with W Ireland, provided only that thewater supplied matched the higher rainfall of the west (Campbell & Osborne 1993).
British occurrence
G. tinctoria is an occasional, widely scattered garden escapee and discard in western England,Wales, the Isle of Man and Scotland and, to lesser extent, in parts of C & S England and the ChannelIsles. A high proportion of occurrences appear associated with or near the west coast, which fits withthe species' restricted temperature limits. Inland sites must be sheltered from severe frost in someother way that allows the plant to survive. Confusion with or misidentification of the Brazilianspecies, G. manicata, may make the New Atlas hectad map inaccurate to anunknown extent, as for instance the latter species is regarded as the more frequent of the two in Devon(VCs 3, 4) (T.D. Dines, in: Preston et al. 2002).
European and world occurrence
G. tinctoria is native to South America, predominantly in Chile. It is also considered tobe native to parts of Argentina, and in the Andean region of Colombia, Venezuela, Peru and Ecuador(Skeffington & Hall 2011; Gioria & Osborne 2013). Having been widely introduced to gardens, itnow has a current global distribution spanning both the northern and southern hemispheres. In thenorthern hemisphere, it is found in England, Scotland and Wales, the Isle of Man, Isles of Scilly, theChannel Islands, France (Osborne et al. 1991), Spain (Sanz Elorza et al. 2001), São MiguelIsland in the Azores (Schäfer 2002) and California (Howell 1970). In the southern hemisphere it is foundin Tasmania (Duretto 2013), New Zealand (Webb et al. 1988) and the Chatham Islands (de Langeet al. 2011).
Threats
G. tinctoria is too infrequent an escapee to be a problem in Fermanagh, but it is invasive, veryconspicuous and now extremely difficult or even impossible with current technology to control in manyparts of W and SW Ireland.
Despite their long-known potential to escape and become naturalised and/or invasive, G. tinctoriaand G. manicata are still advertised for sale on the internet as giant, tropical species suitablefor planting around lakes and ponds in gardens. G. tinctoria received the Award of Garden Meritin 2006 from the Royal Horticultural Society. Responsible horticultural traders should not stock G.tinctoria at all, but its seed remains readily available via the internet and garden centres mayinadvertently or in ignorance continue to trade the species. Gunnera tinctoria is listed on Schedule 9 of the Wildlife and Countryside Act inEngland and Wales, and therefore it is an offence to plant or otherwise to cause it to growthere in the wild.
Native, common and widespread. Eurasian temperate, but naturalised in N America and now circumpolar.
1881-2; Barrington, R.M.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
The numerous, bright magenta or pink-purple spikes of this 50-200 cm tall, wet ground perennial are avery spectacular feature of the Upper Lough Erne shoreline during the summer from July to September,especially when mixed among Filipendula ulmaria (Meadowsweet) and Senecio aquaticus (MarshRagwort) as is often the case. Purple-loosestrife is a common, prominent and sometimes abundant plant ofthe water-fringing, tall-herb, marsh and fen vegetation, including everything from fen-carr andreed-swamp to margins of acid Sphagnum bogs and under-managed, wet, marshy grasslands that areintermittently or seasonally flooded, and that are neither excessively grazed nor trampled. It is alsofrequent on floodplains beside muddy, slow-moving drains, streams and rivers and even occasionally alongdamp roadsides.
L. salicaria produces a strongly developed tap-root system that persists throughout the life ofthe plant without any adventitious roots developing (Shamsi & Whitehead 1974). The species grows ona wide range of soil textures, but is usually found in mud, silt, clay, or fen peat. While it cantolerate poor mineral nutrition, it most often appears under reasonably fertile, moderately acid toneutral, lime- or base-rich conditions (Sinker et al. 1985; Mal et al. 1992). It toleratessoils in the pH range 4.0 and above (Shami & Whitehead 1974).
Being a tall plant, in some sites it prefers shelter and can tolerate shading to about 50% of full sunfrom reeds, willows, alder and ash. On the other hand it is most vigorously competitive, abundant andconspicuous in sunny, marshy grassland and in linear, bank-side situations. The established strategy ofthe species is categorised as a C/CR (Competitor/Competitive Ruderal) by Grime et al. (1988,2007), reflecting successful germination of its buried seed after habitat disturbance, rapid growthrate, remarkable fecundity and dispersal ability, and vigorous colonising ability under a wide range ofsoil nutrient conditions. The principal limiting factors are temperature and moisture (it beingessentially a wetland species), so it is unsurprising that it performs best and can become dominant inthe mild, wet, W of Ireland and in warm, coastal parts of S Europe (Shami & Whitehead 1977).
Experimental evidence on reaction to nutrient levels suggests that L. salicaria may have spreadand increased its presence during the last 50 years or more assisted by nutrient enrichment of drainagewaters following increased use of agricultural fertilizers (Shamsi & Whitehead 1977).
Growth rate and vegetative reproduction
The plant has a persistent taproot from which it develops a horizontally spreading rootstock orwide-topped crown. From this perennial plate-like base arise 30 or more, crowded, erect, annual stems,some often reaching 150 cm or, occasionally, 200 cm in height. The growth rate of the species is so highthat despite its considerable vegetative plant size, L. salicaria seed can germinate in the earlysummer whenever temperature rises to 20°C, establish, flower and set seed, all within a single growingseason (Shami & Whitehead 1977). For this reason it is sometimes misleadingly referred to as a'facultative annual', although in reality it is both perennial and polycarpic, only the aerial partsdying back in late autumn (Shipley & Parent 1991). The lifespan of individuals is unknown and it isclearly a topic for further study.
The rootstock is the perennating organ and it produces fresh shoots each year from spreading root budsaround the crown, a localised form of vegetative reproduction and clonal development, resulting intightly tufted 'phalanx' plants that can represent an almost completely dominant monoculture on suitableground (Mal et al. 1992). Wide vegetative spread is unlikely with this form of growth, but theplant can persist in a particular spot for many years using this property (Shamsi & Whitehead 1974).In B & I and across its native territory, these monospecific stands usually do not persist long, buttend to be replaced quickly, active colonisation by other species commencing in the first year aftertheir establishment. L. salicaria also occurs as single individuals in a wide range of otherwetland plant community types (Shamsi & Whitehead 1974).
Flowering reproduction
L. salicaria flowering takes place from early July to September or occasionally October. Thespecies is capable of rapid spread in water systems by means of its lightweight seed, produced in vastnumbers despite a complex tri-stylous, self-incompatible (but not absolute), out-breeding system that isunique in the flora of B & I (Darwin 1865). The insect pollination requirement involved with thesethree flower forms is satisfied by bees and hoverflies attracted by nectar secreted at the base of thehypanthium or 'calyx tube' and copious pollen (Hickey & King 1981). Maximum seed production requiresthe presence of all three flower morphs (Mal et al. 1992).
The ovary is formed from two fused carpels each containing numerous ovules and the fruit is anoblong-ovoid bi-valved capsule, 3-4 mm long, enclosed in the calyx. It splits into two valves along theseptum (wall) between the two locules of the ovary (ie septicidally) to release the small, light seeds(Hickey & King 1981). In a study by Charles Darwin, the capsules contained means of 93, 130 and 83.5seeds in long-, mid- and short-styled flower forms respectively (Darwin 1877). The lowest capsules onthe inflorescence are ripe and seed is dispersed whilst the plant is still green and leafy. A normalhealthy plant produces about 900 capsules per year (Shami & Whitehead 1974).
Seed is very mobile, dispersal involving wind transport, water flotation and adhesion in mud to birds andother animals, plus on boats, boots and vehicle tyres (Shamsi & Whitehead 1974; Thompson etal. 1987). Seed viability in soil is also ecologically favourable, survival of at least severalyears duration, and sometimes exceeding five years, having been recorded (Thompson et al. 1997).It is important to note that germination and seedling emergence is negligible from seeds buried at 2 cm.Thus, irrespective of the large scale of the L. salicaria seed bank, small disturbances in thesoil surface layers are an absolute germination requirement, enabling further recruitment to the aboveground population from the dormant seed bank (Welling & Becker 1990).
Fermanagh occurrence
It is represented in 207 Fermanagh tetrads, 39.2% of those in the VC. In terms of frequency, in theFermanagh Flora Database, it ranks joint 25th with Crataegus monogyna (Hawthorn), although it isnothing like as widespread as the latter. As the distribution map indicates, it is predominantly foundin the basins of Lough Erne and Lough Melvin and along the River Finn and other major feeders, whileelsewhere in Fermanagh it is much more thinly scattered and local.
Irish occurrence
Although Purple-loosestrife is widespread in most lowland parts of Ireland and is particularly abundantin the milder west, it is much less frequent, local or absent in large areas of the NE of the island.Here it is more or less confined to the Lough Neagh and River Bann basin (FNEI 3). In hisFlora of Lough Neagh, Harron (1986) commented that L. salicaria has a very fragmenteddistribution around the shore of that very large lake, usually appearing only in small quantity. Heattributed the apparent population depletion of L. salicaria to past drainage operations,something The current author (RSF) considers rather unlikely, since this species very readilyre-colonises suitable damp ground. McNeill (2010) commented upon the unusual distribution of L.salicaria in Co Tyrone (H36). It is entirely absent from most of the county, yet is quiteabundant in three discrete areas: Lough Neagh shore extending up the River Blackwater; the Fairy Waterbasin; and along the rivers Finn and Foyle on the margins of the county.
British occurrence
Purple-loosestrife is locally abundant in lowland England and Wales. The New Atlas hectad mapshows that the southern and western distribution tendency noticed in Ireland is even more pronounced inBritain, since here, even more so than in Ireland, the species avoids areas of predominantly acidic peatsoils and critically colder early spring conditions that limit its germination and growth. Thus L.salicaria is excluded not only from upland regions of England and Wales, but also from most ofScotland except the milder SW (Preston et al. 2002).
European and world occurrence
L. salicaria is almost cosmopolitan in moist or wet, low-lying and coastal areas of Europe &Asia, except in extremely cold and arctic regions. It is native throughout Europe from Britain to CRussia, except in the extreme north. It is also indigenous around the Mediterranean basin including theislands, plus in Syria, Iran and Lebanon. It is also native in N Africa from N Morocco to Ethiopia andin W & N Asia. It is absent from NW Finland, Denmark and Iceland (Shami & Whitehead 1974).
L. salicaria is thoroughly naturalised in mild temperate areas of N America where, in the early1800s, it was introduced to New England, most likely as either a garden ornamental, a medicinal herb, oraccidently carried as seed in sailing ships' ballast (Mal et al. 1992). In the S Hemisphere, ithas been introduced to Australia, Tasmania and New Zealand and has become Circumpolar (Hultén &Fries 1986, Map 1348).
A serious invasive weed in N America
Although it had gradually spread throughout NE United States and SE Canada by 1900, L. salicariaonly became recognised as an aggressive wetland weed there in the 1930s. The initial spread was probablyassisted by habitat disturbance at the time caused by agricultural settlement, military activities andthe construction of canals, highways and railways. The species then spread explosively in N Americasometime after 1944, and by 1987 was already a major, out of control, alien weed and had become aconservation nightmare. It is now known there as 'The Purple Plague' or 'The Purple Peril', since itchokes wetlands, clogs drains and infills shallow water bays used by fish for spawning. It makesconditions inhospitable to native plants, out-competing them and forming near monoculture stands thatsupport far fewer plant and animal species than the communities they replace (Mal et al. 1992).Eradication appears impossible with present-day technology, but a great deal of research is currentlytackling the problem.
Medicinal and other uses
Apparently not used at present, there are old reports of L. salicariabeing used in herbal medicine in Ireland. Its astringent properties were used to treatwounds, diarrhoea and even dysentery (Allen & Hatfield 2004). Grieve (1931) says it was "highlyesteemed by many herbalists" and "well established in chronic diarrhoea and dysentery, and isused in leucorrhoea and blood-spitting". It has also been employed for fevers, liver diseases,constipation and cholera infantum and for outward application to wounds and sores. It has been stated tobe superior to Eyebright (Euphrasia spp.) for preserving the sight and curing sore eyes (Grieve1931).
Pharmacologists have taken an interest in the plant as it has been discovered that stem and flowerextracts produce significant hypoglycemia in hyper- and normo-glycemic rats, the extracts reducing bloodsugar by increasing insulin levels. This has potential implications for weight control medication andtreatment of diabetes (Lamela et al. 1985, 1986).
The species has also been regarded as a useful honey plant by beekeepers and at least ten cultivars havebeen selected for and improved for decorative garden use (Mal et al. 1992; Griffiths 1994).
Threats
None.
Native, very rare, but very possibly under-recorded. European temperate.
1902; Abraham, J.T. & McCullagh, F.R.; Castle Caldwell estate.
June to August.
Growth form and preferred habitats
While it is never found in Fermanagh in quantity, this dwarf, more-or-less prostrate, creeping, branched,reddish, quadrangular-stemmed annual, with opposite, fleshy, simple leaves and very inconspicuous, 2 mmdiameter purplish or pink flowers, solitary in the leaf angles, occurs on various types of bare muddy,gravelly or stony ground that floods in winter, but which dries out in summer. Alternatively it featuresin very shallow water on the margins of lakes, pools, streams, ditches and puddles. While it normallygrows in shallow water or on damp ground, L. portula can occasionally be found in water up to 1 mdeep (Preston & Croft 1997).
Due to its fairly small size and very inconspicuous flowers, this distinctly calcifuge, lime-avoidingannual, which is often very local in its occurrence, can be, as here, either difficult to detect, easilyoverlooked, or mistaken for something else equally diminutive.
Apart from the receding margins of all kinds of water bodies, L. portula should be looked for indamp depressions in sand and gravel pits and in temporarily wet ground near flushes, and in ruttedtracks and rides. While it is restricted to acidic, base-poor, mineral soils, Water-purslane avoidsstrongly acid, nutrient-impoverished conditions and it only rarely grows in open areas on organic peat(Preston & Croft 1997). The occurrence of a large but local population of the species on limestoneat Coole Lough in the Burren, Co Clare (H9) is very exceptional, but this is just another example ofanomalous behaviour by normally calcifuge species that takes place in this ecologically andbiogeographically highly unusual coastal region of W Ireland (Webb & Scannell 1983).
Variation
Two subspecies have been distinguished, somewhat arbitrarily on the basis of their mature fruits and aratio cline in the length of their outer calyx teeth. They are also partly distinguished by theirseparate geographical distributions, subsp. portula being widespread in Europe and extendingsouthwards to S Italy, Sardinia and Corsica, while subsp. longidentata, which has epicalyxsegments 1.5-2.0 mm, four times larger than those of subsp. portula, is more western in itsdistribution, being recorded from France, Portugal, Spain, Algeria, the Azores, and is the predominantform in at least western parts of Britain and Ireland (Allen 1954b). However, intermediates occur in thearea between the two subspecies where they overlap (eg on the Isle of Man) and they do not appear todiffer in their ecology, rendering them in the opinion of the current author (RSF) of only doubtfulutility and certainly better considered as varieties (Allen 1984; Preston & Croft 1997; Sell &Murrell 2009).
Flowering reproduction
The tiny pinkish flowers are produced from June to October. They have an epicalyx of six segments,0.5-2.0 mm, linear-lanceolate and with a long acute apex. The calyx has six lobes, each with a smallappendage. The six pink petals, when present, are very thin (they are described as 'fugacious' or'caducous', meaning they fall off early). The stamens are either six or twelve and are inserted beneaththe petals. The single style and stigma are very short and solitary and the superior ovary sits in ashallow, bell- or cup-shaped hypanthium.
The flowers are usually self-pollinated and produce a small, sub-globose, flattened seed capsule, about1.5 mm in diameter containing many seeds. Submerged flowers do not open and are automatically pollinatedin the bud (ie they are cleistogamous) (Melderis & Bangerter 1955). Plants growing in standing waterfruit less readily than those in terrestrial conditions (Allen 1954b).
The seeds are small (0.7 × 0.5 mm), ovoid and grey and, despite the apparent lack of a dispersalmechanism, they somehow, mysteriously, manage to reach suitable isolated habitats, such as sand quarriesand gravel pits (Preston & Croft 1997). Buried dormant seed survives for at least five and, perhaps,many years (Croft 1994; Thompson et al. 1997).
The established ecological strategy as determined by Grime et al. (1988, 2007) is given as R/SR,meaning it is intermediate between Ruderal and Stress-tolerant Ruderal. This designation, together withthe fact that seed germination requires light, suggests that some degree of soil or substratedisturbance is essential to bring dormant seeds to the surface and stimulate their germination (Preston& Croft 1997).
Vegetative reproduction
The usually creeping stems root at their nodes and, since they are rather brittle, fragments may dispersein water or mud and help propagate the plant (Preston & Croft 1997).
Fermanagh occurrence
In Fermanagh, Water-purslane has been recorded just 16 times in a total of 13 tetrads, 2.5% of those inthe VC. Eleven of the tetrads have post-1975 records and, as the tetrad map indicates, they are locatedmainly in the NW of the county.
British and Irish occurrence
While L. portula may very well be under-recorded in Fermanagh, the New Atlas hectad mapindicates that in B & I its distribution is very much better known now than was the case in theearlier BSBI Atlas (Walters & Perring 1962) and it is more frequent and wider in itsoccurrence than previously known. This is especially the case in all western parts of Britain and in theScottish midlands, between Glasgow and Edinburgh, where the species is now seen to be most prevalent. InIreland, L. portula has a strong representation in the SW, but is widely scattered throughout theisland.
Having said this, while the hectad map of B & I shows L. portula is frequent and widelyscattered, there is clear evidence of a species decline since the 1950s, especially in SE England andthe English Midlands. The losses are most readily explained by the typical loss of habitats associatedwith seasonally flooded ground, such as infilling, overgrowing, or draining of ponds, and the repair ofwet ruts in resurfaced tracks (Preston & Croft 1997; R. Wilson, in: Preston et al. 2002).
European and world occurrence
L. portula is essentially a lowland, European temperate species extending from S Scandinavia to SEurope and to scattered sites in N Africa and W Asia. It has occasionally spread within and somewhatbeyond its original range, for instance into isolated localities in more northern areas of Fennoscandia(Hultén & Fries 1986). It has also been introduced to California, C & S America, New Zealandand, possibly, also to N China (Hultén & Fries 1986, Map 1349; Sell & Murrell 2009).
Threats
None.
Introduction, neophyte, either a very rare garden escape or deliberately planted.
3 March 1996; Northridge, R.H.; Flaxfield Wood, Castle Coole estate.
Growth form, English status and preferred habitats
This glossy, bright green, lanceolate, leathery-leaved, evergreen, low-growing shrub up to 100 cm tall isa native of sub-Mediterranean and sub-Atlantic regions. Some botanists suppose it is native also incopses and woods in parts of C & S England – despite the total absence of fossil evidence and itswidespread cultivation from at least the 14th century onwards (Godwin 1975; Kent 1975; Harvey 1981).Even the first English record of the species in print refers to it being, "plētuously inhedges" (Turner 1548)!
Beyond the garden, Spurge-laurel typically, but often only locally, grows in deciduous woodland, on dry,near-neutral, lime or base-rich soils (hence its alternative common names 'Wood-laurel' and'Copse-laurel'). The name 'laurel' is misleading as it is unrelated to other laurels of the genusPrunus in the Rosaceae. D. laureola is very shade tolerant and capable of spreading bylayering, but it does not usually seed much, due to an absolute requirement for insect cross-pollinationand an early vernal flowering season. On account of these properties and its limited invasive potential,very occasionally it is deliberately planted as pheasant cover on Irish landed estates, as appears to bethe case in the deciduous woodland at Castle Coole, the solitary Fermanagh situation where this specieshas been found to date.
D. laureola occurs along with, or in similar situations to, other pheasant-rearing cover, eg theequally rarely found Ruscus aculeatus (Butcher's-broom) and the very common Symphoricarposalbus (Snowberry). Plants of D. laureola and R. aculeatus occasionally survive asscattered, isolated individuals, or more rarely as fairly large clonal patches, long after theirintroduction, when everyone who planted them is dead, and their human origins forgotten. It is easy toimagine this happening in England too, so that the so-called 'native' distribution has now become welland truly obscured by human, mammal- and bird-sown plantings (A.J. Richards, in: Preston et al.2002).
Flowering reproduction
The flowers are produced from January to April and are sweetly scented for a period only. They areyellowish green in colour and hang in axillary raceme clusters of 5-10 flowers below terminal rosettesof glossy leaves. The calyx is 4-lobed, on top of a tubular hypanthium; petals absent; stamens eight;style solitary and short. Nectar is secreted and concealed at the base of the hypanthium tube and isaccessible to bees, butterflies and moths (Melderis & Bangerter 1955; Hutchinson 1972; Sell &Murrell 2009).
The fruit is green at first, then bluish, and finally becomes a black, oval drupe up to 12 mm indiameter. Although the fleshy fruit is extremely poisonous to humans, livestock (including horses) and,indeed, possibly to all large mammals, it is taken by birds, including the Robin, and thus the seed canbe bird-sown (Snow & Snow 1988, p. 94). Despite its toxicity (see below), the fruit may also beeaten by bank voles and mice, which therefore can also act as seed vectors (G.H. Knight, in: Lang 1987,p. 121).
Irish occurrence
Apart from Fermanagh, in other parts of Ireland D. laureola crops up as a rare or infrequentgarden escape or discard, chiefly north of the political border, apart from on pheasant rearing estates.The plant usually occurs as isolated, presumably bird-sown individuals, in hedges, woods and in rockyplaces, or even rarely on old walls. Reynolds (2002) in her Cat Alien Pl Ir lists records ofthese types from nine Irish VCs, not including Fermanagh, and in some rare cases the shrub may becomequite thoroughly naturalised.
British occurrence
D. laureola is locally frequent in England north to Cumberland (VC 70) and Co Durham (VC 66) andis present in N Wales in Pembrokshire (VC 45) and Glamorganshire (VC 41). It also is present in theChannel Islands. It is introduced in Scotland and the Isle of Man (New Atlas; Sell & Murrell2009).
European and world occurrence
D. laureola is a submediterranean-subatlantic species that occurs from South-central and SouthEurope northwards into Hungary. It is also present in SW Asia and is rare in N Africa and the Azores(Sell & Murrell 2009).
Toxicity
Daphne species contain an acrid irritant sap in all parts of the plant, but particularly in thebark and the seeds within the berry-like drupes. Chemically, the toxins are described as tricyclicditerpenes with a daphnane carbon skeleton, daphnetoxin and its ester mezerein being the principalirritant compounds. Poisoning of stock animals is rare, as the plant is usually avoided because of itsunpalatability. Only really starving animals would be tempted to try eating it (Cooper & Johnson1998).
Human poisoning usually involves children who mistake the black berry-like fruits for currants. As thefruit is so very acrid, normally only a very few are eaten and the symptoms, which include a burningmouth sensation, nausea, abdominal pain, vomiting and diarrhoea, tend to be relatively mild andtransient, although obviously frightening at the time for all concerned (Cooper & Johnson 1998).
Uses
D. laureola is less acrid than D. mezereum (Mezereon), and its leaves were formerly used anemmenagogue (ie to provoke menstruation), but they may cause vomiting and purging. Both leaves and barkhave been used to induce abortion (Grieve 1931). It has been described as a brisk and rather severepurgative and was used as a cottage garden laxative. It was mixed with mistletoe and given for epilepsy.The crushed leaves were also reputed to have been used as a horse worming medicine although this wasfound to be dangerous and was discontinued due to fatal poisoning taken place (Cooper & Johnson1998). The roots were also once used as a supposed cure for venereal disease and for treating bothbenign and malignant cancers. The latter use has scientific merit since it continues to be a subject ofresearch (Allen & Hatfield 2004).
Threats
None.
Native, common and locally abundant. Eurasian southern-temperate, but widely naturalised in bothhemispheres.
1866-72; Smith, T.O.; Upper Lough Erne.
Throughout the year.
Growth form and preferred habitats
A common and widespread, up to 2 m tall, stoloniferous and rhizomatous, lowland, wetland perennial, theleaves of E. hirsutum are usually softly felted and are especially hairy on the veins. The robuststems are also clothed with a mixture of long, spreading, simple eglandular hairs and dense, shorter,glandular ones. As with other willowherbs, however, hairiness is extremely variable and rare almosthairless forms of E. hirsutum do exist (Clapham et al. 1962). When vegetative the plant isrecognised by its height (up to 200 cm) and its opposite, unstalked, oblong-lanceolate leaves, 3-12 ×1-3 cm, the bases of which run a short way down the stem forming wings on it (Sell & Murrell 2009).
'Great Hairy Willowherb', to give the plant its most descriptive English common name, often forms tight,vigorously growing, tall, clonal patches which can become locally dominant in lowland, sheltered, dampto wet, moderately acid to lime-rich ground of reasonable fertility (Kitchener 1992a). In the Sheffieldarea, Grime et al. (1988, 2007) described its ecological established strategy as a definite'Competitor' and found that it avoided strongly acid conditions (pH below 4.0), preferring those aboveabout pH 6.0 to neutral, which fits well the observed behaviour in Fermanagh. Much less frequently, itoccurs in shallow standing water of mesotrophic to eutrophic productivity and equivalent nutrientstatus.
In parallel with its smaller, less aggressively competitive relative E. parviflorum (HoaryWillowherb), E. hirsutum also grows and persists on relatively dry roadside banks andoccasionally in waste ground, including in old quarries and sand- and gravel-pits. However, it inhabitsthese drier habitats much more rarely than it appears in wetland situations, such as in ditches, onstreamsides and lake shoreline tall-herb fens and marshes. While definitely a wetland species, E.hirsutum performs less well in permanently wet fens and marshes than on periodically floodedwaterside ground (Grime et al. 1988, 2007).
E. hirsutum is intolerant of grazing or mowing and is quickly replaced by lower growing speciesunder such circ*mstances (Grime et al. 1988, 2007).
Fermanagh occurrence
E. hirsutum has been recorded in 260 Fermanagh tetrads, 49.2% of those in the VC. As the tetradmap indicates, it is very common and widespread around both parts of Lough Erne and is widely scatteredin many other suitable lowland, wet to damp ground sites throughout the county. As such E.hirsutum is considerably more frequent and widespread than E. parviflorum and it justbeats the very much smaller, calcifuge wetland species, E. palustre, in terms of both frequencyand distribution statistics.
Great Willowherb tolerates semi-shaded conditions in both fen-carr and on the margins of damp deciduouswoods. There are numerous records of the latter in Fermanagh, including, for instance around many of thewooded isles and along the shores of Lough Erne, plus in the upland Correl Glen NR mixed deciduous woodson the Western Plateau, Marlbank Wood, above Florencecourt and in fairly steep broken limestone rockyground in the steeply sloping Hanging Rock NR.
Reproduction
Since it dies down completely each autumn, regrowth of Great Willowherb begins in early spring. Being alarge, robust, stand-forming plant, a prolonged vegetative period is required before the very numerous,large, soft purplish-pink, insect-pollinated flowers can be produced. They are borne on long, branched,corymbose, inflorescences from late July into August, and are terminal on main stems and branches. Whiteflowered plants are very occasionally encountered and have been given the name var. hirsutumforma albiflorum Hausskn. in Sell & Murrell (2009).
The flowers are protandrous and pollination is carried out by bees and hoverflies. Flowering reproductionis extremely effective, vast quantities of plumed seed being released on the wind by the splitting ofthe 5-8 cm long, linear fruit capsules. The seed is long-persistent in the soil.
Additional vegetative reproduction takes place by the spreading growth of fleshy, white undergroundrhizomes or leafy surface stolons in the autumn and winter (Grime et al. 1988, 2007). Thus by oneof these means, the species can readily colonise freshly available, reasonably undisturbed, ungrazedsites, eg in quarries (Shamsi & Whitehead 1974, 1977).
Hybridization
In B & I, E. hirsutum rarely or very rarely forms hybrids with six other willowherb species,none of which have ever been found in Fermanagh (Sell & Murrell 2009).
British and Irish occurrence
In Ireland, E. hirsutum is common, widespread and sometimes dominant except in the more acidic bogsoils of the far west. It is also common and widespread in Britain, except in Scotland where it becomescoastal northwards. These facts suggest that low winter temperatures and acidic soils are factorslimiting its distribution (Preston et al. 2002).
European and world occurrence
A native of southern-temperate Eurasia, E. hirsutum has naturalised widely elsewhere, including ineastern N America and C & S Africa (Hulten & Fries 1986, Map 1357).
Threats
None.
Native, common and widespread. European temperate.
1881; Stewart, S.A.; Co Fermanagh.
April to November.
Growth form and preferred habitats
E. parviflorum is one of four erect Willowherb perennial species with four-lobed stigmas presentin the flora of B & I. The combination of dense, soft, short, spreading hairs on the stems, plusleaves softly felted on both surfaces, produces the usual 'hoary' appearance of this distinctiveperennial (Clapham et al. 1987). Unfortunately, the hairs are not always present in quantity anda gradation of forms exist which approach hairlessness (Kitchener 1992a). The latter near-glabrouscondition is very rarely found, however, and generally the appearance very well fits the English commonname. This is a smaller plant (up to 60 cm) and usually it is more narrowly upright than E.hirsutum (Great Willowherb). Its four stigma lobes in the smaller purplish rose-pink flowers arevariably compact and never curved back upon themselves as they do in the larger species (Kitchener1992a). The leaves are very variable but are usually 3-7 × 1-1.5 cm, the lower ones opposite and thoseabove the mid-stem alternate. The leaf shape is oblong-lanceolate and the base rounded, but neitherclasping the stem, nor decurrent at the sessile base (Clapham et al. 1987).
E. parviflorum is occasional to common in damp, marshy or swampy tall-herb fen conditions over avariety of lowland waterside habitats. Unusually and rather surprisingly, E. parviflorum alsofrequents very much drier, disturbed ground, as a pioneer colonising ruderal species (see below). Itsestablished strategy is therefore categorised by Grime et al. (1988, 2007) as C-S-R, meaning itis intermediate between a Competitor, Stress-tolerant and Ruderal species, and displaying features ofeach ecological approach to survival.
Fermanagh occurrence
In Fermanagh, Hoary Willowherb most often occurs on damp, calcium-enriched soils, including organic fenpeats around Upper Lough Erne and the SE part of Lower Lough Erne. In these wetland situations, it isespecially frequent where there is a recent history of soil disturbance, such as light trampling bycattle, ie animals, however heavy in weight, paying visits of brief duration, thus providing sporadic,occasional grazing pressure and manuring that keeps the ground ecologically open and primed forcolonisation.
Hoary Willowherb is seldom or rarely recorded in unproductive soils more acidic than around pH 5.5 (Grimeet al. 1988, 2007). Interestingly, E. parviflorum, like E. hirsutum (GreatWillowherb), also occurs widely, but less frequently and abundantly as a pioneer colonist of dry orwell-drained disturbed ground (Kitchener 1992a). Often, but not always, this involves stony or gravellysoils, including limestone talus slopes (eg below Carrickbeg cliffs), old quarries (it is present in atleast eight of these in Fermanagh), alongside gravel or sandy track-sides and in waste ground, includingthe margins of car parks and other similar, moderately to slightly disturbed urban sites. The ruderalpopulation remains associated, albeit to a minor degree, with the old disused railway lines and derelictstation areas which were abandoned in Fermanagh in 1942 and 1957.
Taking these two very different types of habitat moisture populations together, E. parviflorum isfound in 192 Fermanagh tetrads, 36.4% of those in the VC. The fact that it belongs to a genus equippedwith plumed, air-borne seed undoubtedly assists E. parviflorum in behaving both as a widespreadruderal colonist of disturbed ground, while holding on to its much larger, probably original, wetlandterritory, by means of longer-lived, repeatedly reproducing, species-sustaining populations.
Flowering reproduction
Overwinter perennation is by vegetative buds and short, leafy stolons and, of course, by seed. The leafybuds on stolons are produced in autumn and winter, but they are generally formed near the base of theplant and essentially they maintain existing individual plants and are neither an efficient means ofvegetative spread nor of species reproduction.
Flowers are produced in July and August and are borne in a ± corymbose terminal raceme. They are 6-13 mmin diameter (parviflorum means small-flowered), of a pale purplish-rose colour (rarely white)with deeply notched petals; the style is white, tinged mauve, club-shaped when young but the stigmaopens out into four fairly upright lobes. The style and stigma together about equal the length of thestamens (Sell & Murrell 2009). The ovary, 1.2-4.0 cm long, is inferior and resembles a thickenedflower stalk. The flowers are hom*ogamous (the two sexes maturing simultaneously) and are occasionallyvisited by hive bees and other Hemiptera, although they often self-pollinate (Clapham et al.1987; Fitter 1987). The fruit capsule is elongated, 30-70 mm, splitting into four narrow recurvedvalves, releasing numerous tiny seeds each with a tuft of silky hairs attached at the top that enablesready wind dispersal (Hutchinson 1972; Clapham et al. 1987).
Persistence of dormant seed in the soil greatly facilitates the rapid, opportunistic occupation ofvegetation gaps created by occasional disturbance. However, the survey of NW European soil seed banksfound that five of the six records that exist for E. parviflorum, described the seed as merelytransitory, while the other record suggested short-term persistence only (Thompson et al. 1997).In this context, the persistence of the small populations on long-abandoned railway land is veryinteresting, indicating the success the species is somehow experiencing in this portion of its habitatrange, given our decidedly damp, humid, often overcast weather conditions where drought conditions arerare.
Obviously other environmental factors assist survival by minimising competition from potentially dominantspecies that might otherwise oust E. parviflorum. It is not known for certain what theseecological conditions might be although low soil fertility and an intermediate level of vegetationdisturbance are probably involved (Grime et al. 1988). Further study of this topic is definitelyrequired.
Hybridization
Thus while it regularly consorts with its larger cousin, E. hirsutum (which unlike the majority ofwillowherbs, very rarely hybridizes with its relatives), when it is operating as a ruderal, E.parviflorum frequently overlaps and shares ground with another perennial member of the genus,E. montanum (Broad-leaved Willowherb), although it manages to avoid much contact with E.obscurum (Short-fruited Willowherb), again due to their differing soil pH preferences.Hybridization with E. montanum can and does occur at least occasionally, but to date this isrecorded only rarely in B & I.
Fermanagh has one of the four Irish records of this hybrid displayed in the New Atlas map. RHN andthe current author (RSF) tend to agree with G.D. Kitchener (in: Preston et al. 2002), that plantrecorders must all be overlooking this hybrid to some unknown extent.
British and Irish occurrence
E. parviflorum is common and widespread throughout most of lowland England, Wales andIreland, but becomes scarce and local in Scotland, apart from the urban Central Belt and coastal areas.However, the species does reach as far north as Orkney, but is absent from Shetland (New Atlas).
European and world occurrence
E. parviflorum belongs to the European temperate biogeographic element andis widespread throughout Europe northwards to S. Sweden. It is also present in N Africa, WAsia to India and in the Atlantic Islands (Sell & Murrell 2009). It is probably only casual in CFennoscandia and other outlying localities in both Europe and Asia (Hultén & Fries 1986, Map 1358).It is introduced to New Zealand, but was not collected there until 1967 and is thus probably a recentarrival. It may now be widespread in moist, fertile farming country around Hawke's Bay and Poverty Bayon North Island (Webb et al. 1988).
Threats
None.
Very rare, but probably seriously under-recorded.
1934-8; Praeger, R.Ll.; lakeshore, Carrickreagh Bay, Lower Lough Erne.
Stace, in his New Flora of the BI 1991, regarded this as one of the most common perennial hybridwillowherbs in the flora of B & I, and with just one Praeger record in Fermanagh dating from 1930s,it must certainly be under-recorded in this area. The same applies in the rest of Ireland, since in themap in Stace et al. (2015), apart from the solitary Fermanagh record, there are only five otherhectads with records scattered across the whole of the island.
The New Atlas hectad map of Britain also appeared so very sparsely peppered with records (56hectads with data), that the same comment might well apply throughout Britain too, although to be fairit indicated a slightly greater frequency of this hybrid in SE England, in particular in West Kent (VC16) and in Surrey (VC 17). The more recent Hybrid Flora of the British Isles hectad map, however,displays a very much greater frequency of records of E. × limosum in Britain,incorporating data from a total of 136 hectads (Stace et al. 2015). The distribution of thehybrid is now much more widespread, for instance, reaching the NE coast of England in NE Yorkshire (VC62) and Co Durham (VC 66), and it shows many more records scattered across Wales, the Midlands and SEngland. However, some of the mapped records have not been verified by specialists, so a margin of errormust be borne in mind when interpreting the map (Stace et al. 2015). The current author (RSF)felt that the New Atlas hectad map of this hybrid reflected the distribution of Epilobiumspecialists and their travels around the year 2000, rather than the real pattern of the plant'soccurrence. The most recent Hybrid Flora of the British Isles map, however, appears to show amuch more reliable picture of the distribution of this taxon.
Both parent species are common and widespread throughout B & I and the hybrid can arise at any pointwhere they meet. This generally happens in various forms of disturbed ground, such as for instance inrecently felled or coppiced woodland, quarries, chalk- and brick-pits, gardens and waste ground. It hasalso been recorded on woodland rides and margins, hedgerow banks, roadsides and railway sidings.
Wild hybrids in Britain and Ireland exhibit limited seed set and while they typically occur as solitaryplants, larger populations are known to occur and second generation backcrossed plants can, but onlyextremely rarely do arise (Hybridization; G.D. Kitchener, in: Preston et al. 2002; Staceet al. 2015).
Very rare, but probably overlooked and under-recorded.
1947; MCM & D; shore of Annashanco Lough, Annaghmartin Td.
This hybrid, for which the above is the solitary Fermanagh record, is almost certainly under-recorded asboth parents are widespread and common species.
As expected, the hybrid is intermediate in many of its characters, the stigma being either of a confuseddegree of lobing, or thick and entire. Middle and upper stem hairs are most like E. parviflorum,but are shorter and somewhat crisped. The pubescent leaves are sessile, linear-lanceolate and theirmargins are ± revolute like those of E. palustre, with the marginal teeth tucked underneath.Filiform stolons may be produced, like in E. palustre, but they are more leafy in the hybrid thanin the species (Stace et al. 2015).
The hybrid is recorded from wet marginal and disturbed habitats including semi-open areas in lowlandmarshes, fens and around ponds and only rarely does it appear on waste ground habitats (Stace etal. 2015). In his Flora of Cardiganshire, Chater (2010) described E. × rivulare as frequent there in tall herb fens and bogs, usuallygrowing amongst Juncus acutiflorus (Sharp-flowered Rush) or Molinia caerulea (PurpleMoor-grass) communities, where it sometimes was more abundant than E. palustre and often occurredin the immediate absence of E. parviflorum. The hybrid frequently resembled tall, much-branchedE. palustre (Chater 2010).
E. × rivulare is mostly sterile, although some wild material fromBritain shows more seed-set than found in experimental hybrids, and it may represent the F2generation or backcrosses. Whenever the hybrid is found in quantity, some of the plant spread andabundance may result from stoloniferous growth, rather than seed production (Stace et al. 2015).
The only three other Irish VCs with records of this hybrid in the Cen Cat Fl Ir 2 are Mid-Cork(H4), Antrim (H39) and Londonderry (H40). The latter two NI records are also Praeger finds dating from1937 and 1935 respectively, and the FNEI 3, which details them, adds another solitary record forCo Down (H38), made by the late Miss Doreen Lambert near Portaferry in 1971.
Stace (1997) states that E. × rivulare occurs scattered in Britain andNI, but it is not mapped in the New Atlas. The hectad map of this hybrid in the HybridFlora of the British Isles plots only two records from Ireland, one from Fermanagh and the otherfrom Mid-Cork (H4). The other three records from NI listed here are not represented. The map alsoindicates that E. × rivulare is rare and very thinly scattered inBritain (a total of 32 hectads), with a concentration of records in S Wales (VCs 44-46). Thedistribution stretches northwards, reaching coastal NW Scotland at Westerness (VC 97) (Stace etal. 2015). The hybrid is described as widespread in temperate Europe (Stace et al. 2015).
Native, common and locally abundant. European temperate, but also in C & E Asia.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
The most frequently occurring willowherb in Fermanagh, this common, stoloniferous 20-70 cm tall perennialhas a four-lobed stigma and no long spreading hairs, but short curled ones instead. The leaves areshort-stalked and toothed along their margin, including the rounded base. Most leaves are opposite, butthis does not apply to the leaf-like bracts above. Occasional plants have leaves arranged in threesrather than in pairs (Kitchener 1992a). After germination, a complex of primary and adventitious lateralroots soon develops, resulting in a fibrous root system similar to that of many grasses (Myerscough& Whitehead 1966).
Perennating overwinter as a leafy rosette with its own adventitious root system, in the spring and earlysummer Broad-leaved Willowherb becomes a pioneer colonist in a wide range of fully-lit to medium shadedand/or disturbed, open natural habitats, including woods, hedgerows, lakeshores, cliffs and screes. Itis most characteristic of lightly shaded conditions and only becomes abundant as a colonist of open,moist habitats (Grime et al. 1988, 2007). The species also displays opportunistic, weedycolonising behaviour in gardens, roadsides, walls and waste places, often on shallow, stony, skeletalsoils (ie rankers or lithosoils). E. montanum is wide-ranging in respect of water regime,occurring in habitats from damp woods to dry walls. Like Chamaerion angustifolium (RosebayWillowherb), it appears to have high mineral nutrient requirements, but it is also tolerant of hot, drygrowing conditions (Myerscough & Whitehead 1966).
Around Sheffield, Grime et al. (1988, 2007) found E. montanum preferred calcareous soilsand, indeed, in Fermanagh it is most prevalent in these too. However, most soils, habitats and plantcommunities (including not just pioneer but also more stable, mature vegetation) can and do accommodateE. montanum to some extent. In general, E. montanum prefers moderately acid to alkalineconditions and is increasingly uncommon in soils below pH 5.0. For these reasons, Myerscough &Whitehead (1966) aptly described it as having a wide range of tolerance, yet within its range it isseldom a very 'aggressive' competitor. Grime et al. (1988, 2007) categorize its establishmentstrategy as intermediate between Ruderal and C-S-R. As a garden or wayside weed E. montanum isseldom noxious to the extent of being of prime economic importance (Myerscough & Whitehead 1966).
Fermanagh occurrence
The wide ecological amplitude of the species results in it being represented in 359 Fermanagh tetrads,68% of those in the VC. In Fermanagh, E. montanum is remarkably widely and evenly distributed andit grows at almost all altitudes, except on the wet, blanket-bog peat of Cuilcagh, the highest mountain.It frequently occurs on the drier portions of lakeshores, including on many of the wood-fringed islandsin Lough Erne, but it completely avoids permanently waterlogged or truly aquatic situations.
Flowering and vegetative reproduction
Plants flower from June to August; as soon as an individual does so, it also begins to develop tight,vegetative buds in the axils of the lowermost leaves, which eventually become the over-wintering leafyrosettes. In normal circ*mstances, E. montanum is not considered well adapted to vegetativespread and dispersal, but in the autumn and early winter these overwintering buds and rosettes areeasily detached from the remains of the parent plant, and burrowing animals (eg rabbits) or thedisruption of horticultural practices (eg cultivation) helps promote their wider dispersal (Myerscough& Whitehead 1966). The possession of overwintering green leaf rosettes potentially allows thespecies to have a longer growing season in favourable years (Myerscough & Whitehead 1967).
The inflorescence consists of solitary axillary flowers forming a loose, terminal, leafy raceme, often ofless than 20 flowers (Grime et al. 1988, 2007). Flowers are 6-9 mm in diameter, ± drooping inyoung bud. Floral parts are in fours; the petals are pale at first, turning pink, deeply 2-lobed at thetip and narrowed at the base into a short claw. Stamens are four long and four short, the anthers creamor yellow. The style is white, solitary; the stigma has four short, non-revolute lobes, the wholeovertopped by the four longer stamens.
The flowers are hom*ogamous and attract relatively few insects (bees and flies), so most often they areself-pollinated (Fitter 1987). A slight possibility of crossing remains, since the flower opens beforethe receptive surfaces of the four-cleft stigma become exposed. Later in development, the stigma comesin contact with the anthers and selfing appears to be normal in nature. Experimental investigationsunder controlled or uniform conditions show there is very little variation, as might be expected with ahabitually inbreeding species (Myerscough & Whitehead 1966). However, at the same time it displaysgreat phenotypic plasticity with respect to the wide range of environments in which it occurs(Myerscough & Whitehead 1967).
Seed is set from July to September and the long slender fruit capsule, 40-80 × 1-2 mm, is four-angled andsplits along its length to release numerous small, light plumed seeds that are easily carried by wind(Grime et al. 1988, 2007). As with many other weedy species, seed is long persistent, the typicalplant producing around 2,300 (Salisbury 1964, p. 287). While the small seed size greatly facilitates itswide dispersal to suitably open sites, for colonisation, successful establishment requires nutrientresources the seed does not carry with it. Thus sites need to provide the minimum nutrient and moistureresources necessary for seedling growth, in order for effective colonisation to take place (Myerscroft& Whitehead 1967). The association of seedling colonisation with moist, shaded conditions is sharedby many common ferns such as Asplenium ruta-muraria (Wall-rue), Cystopteris fragilis(Brittle Bladder-fern) and Dryopteris felix-mas (Male-fern) which also exploit walls and steep,rocky habitats (Grime et al. 1988, 2007).
A limited degree of stoloniferous vegetative spread also helps maintain populations, particularly in moreshaded, wood or scrub environments (Grime et al. 1988, 2007).
Hybrids
Common and widespread throughout B & I, E. montanum forms hybrids with five otherEpilobium species that occur in this area (New Flora of the BI 1997). To date, only thehybrid formed with E. parviflorum (Hoary Willowherb) has been recorded (just the once) in theFermanagh Flora Database. Currently in Britain as a whole, the most common Epilobium hybrid isthat between this species and E. ciliatum (American Willowherb), namely E. × interjectum Smejkal (Kitchener 1992b). The Hybrid Flora of theBritish Isles map displays a total of 356 hectads for this particular hybrid, widely scatteredthroughout most of B & I, although scarce or absent from S & SW Ireland and northwards intoScotland (Stace et al. 2015).
Both Wurzell (1986) and Kitchener (1991) have pointed out that many field recorders are wary ofattempting identification of hybrids in this genus, since the species themselves are often rather trickyto determine. Thus there is scope for an unknown, but considerable degree of under-recording takingplace.
British and Irish occurrence
E. montanum is common and widespread throughout B & I although becoming somewhat more scarcetowards the N & W of both Scotland and Ireland and also on higher ground (New Atlas). Thereis no evidence of any change in the species distribution since the previous BSBI Atlas of 1962(G.D. Kitchener, in: Preston et al. 2002).
European and world occurrence
E. montanum extends from Arctic Norway and Russia southwards to S Europe and eastwards to middlelatitude Asia, including the Himalaya, Syria, Lebanon, Siberia and Japan (Hultén & Fries 1986, Map1359; Sell & Murrell 2009). The distribution in Asia is incompletely known.
Threats
None.
Introduction, extremely rare and very probably a mis-identification. Eurosiberian temperate.
1990; Montgomery, J. & Foster, S.; Castle Caldwell FNR, in a swamp or fen-carr scrub.
There is only one Fermanagh record for this species which is essentially southern in B & I and whichStace, in his New Flora of the BI (1991, 1997), states is very scattered in Ireland. E.tetragonum is one of the willowherb species with club-shaped stigmas, which are much moredifficult to identify than those with their stigmas split into a four-lobed cross. A total of five ofthese club-shaped stigma species are found in Ireland, although one of them, E. alsinifolium(Chickweed Willowherb), has only been found in one site in Co Leitrim (H29). Apart from E.tetragonum, the others are E. ciliatum (American Willowherb), E. obscurum(Short-fruited Willowherb) and E. palustre (Marsh Willowherb), all of which, in the opinion ofRHN and the current author (RSF), are much more likely to be found in Fermanagh than E.tetragonum.
Of the three Epilobium species in the Fermanagh area with club-shaped stigmas and ridged stems(the others being E. obscurum and E. ciliatum), E. tetragonum has no glandularhairs on the stem or the capsules, which are the longest of any B & I Epilobium species,ranging from 7 to 9 cm. The stem hairs of E. tetragonum are short, white and tightly appressed,looking like silk under the lens (Kitchener 1992b).
Stace's view of the distribution and rarity of E. tetragonum in Ireland is confirmed by the NewAtlas hectad map that shows it as being native and widespread in the southern half of Englandand all of Wales, but completely absent from Scotland and a very scarce alien in Ireland.
The wet, swampy habitat listed for the plant at Castle Caldwell does not completely rule theidentification out, since like many other willowherbs,
E. tetragonum tolerates both damp and dry situations. Consistent with its southern distribution inBritain, however, in general it prefers sheltered, warmer ground than that offered by fens (Sinker etal. 1985). The more usual habitats of this rather uncompetitive species are damp or moist,disturbed ground, including gardens, waste ground, urban pavements, or the base of walls. Alternatively,it may occupy ditches, streamsides, woodland margins and rides, rather than wet, swampy, fen-carr (G.D.Kitchener, in: Preston et al. 2002).
Clearly, the Fermanagh record needs to be verified and, being a first county record, it requires avoucher specimen to be acceptable. The New Atlas map for Ireland plots just 23 additional hectadsfor E. tetragonum and they are all on or close to the E & S coasts. This is furthercirc*mstantial evidence that renders the Fermanagh record even more obviously a remote outlier from therest of the species Irish range and, therefore, even more suspect. The solitary record is possibly anerror for either E. obscurum, or perhaps the recently spreading American species E.ciliatum.
Threats
None.
Native, frequent and widespread. European temperate.
1884; Barrington, R.M.; Co Fermanagh.
May to November.
Growth form, identification and preferred habitats
E. obscurum is an erect perennial, 30-80 cm tall, which overwinters by means of leafy stolons.Stem hairs are ± appressed, not spreading and the stem has two or four raised lines running down fromthe sessile, opposite leaf bases. Together with the closely related E. ciliatum (AmericanWillowherb) and E. tetragonum (Square-stalked Willowherb), E. obscurum has raisedlines or ridges running down the stems from the leaf bases and all three also have club-shaped stigmas.E. tetragonum differs from E. obscurum in having a square stem, a total absence ofglandular hairs and a longer fruit capsule (7-10 cm long). E. ciliatum is distinguished from theother two species by having the upper part of its stem covered with numerous spreading, glistening,glandular hairs. E. obscurum may have a few glandular hairs ± confined to the calyx tube (ie atthe top of what will become the fruit capsule), but it does not have glandular hairs lower down thestem. E. ciliatum also differs from E. tetragonum in having less prominent stem ridges anda shorter capsule (4-6 cm) (Garrard & Streeter 1983; Kitchener 1992b).
E. obscurum thrives in a wide range of damp ground habitats of moderate to low fertility, but likeother perennial willowherbs, it can also tolerate rather drier situations and can colonise waste ground,roadsides and walls, especially when the latter are periodically damp with rainwater or near waterbodies. This includes the parapets of bridges, which represent a typical site for this species. E.obscurum is absent from highly calcareous soils, but it occurs on bog peat, even when this isflushed with lime-rich groundwater as is the case in Fermanagh at Mullaghmore, Black River, the onlypossibly native Irish site of Erica vagans (Cornish Heath).
In terms of its establishment strategy, Grime et al. (1988, 2007) categorized E. obscurumas intermediate between Ruderal and C-S-R. It can only tolerate a moderate degree of shade, competitionfrom taller or more aggressive species, or environmental disturbance such as a widely fluctuating watertable or any prolonged exposure to trampling and grazing pressure (Grime et al. 1988, 2007).
Since its habitats very frequently ecologically overlap with E. palustre (Marsh Willowherb), it isworth knowing that whenever the two occur together, E. obscurum can be distinguished by itsleaves being less narrow, less shiny and its flower buds are held erect, not drooping (Kitchener 1992b).
Fermanagh occurrence
E. obscurum is a rather frequent willowherb in Fermanagh having been recorded from 122 tetrads,almost 23.1% of those in the VC. It is most frequent in marshy, base-poor grassland situations,especially on the shores of both parts of Lough Erne. It is also widely scattered on damp ground inhedgebanks and on woodland margins, along woodland tracks and in clearings. It is again common besideditches, streams and rivers, including in peat bogs and in ground that seasonally floods or iswinter-wet. This habitat range stretches to include damp hollows in at least one local quarry at Ederny.
E. obscurum is sometimes also quite difficult to distinguish from the American introduction, E.ciliatum, which has been spreading very rapidly in recent years in both B & I. It is knownthat this alien species arrived in Fermanagh, probably sometime in the early 1980s (the first record isdated 1983). It is therefore possible that identification errors may have been made overlooking the newarrival, resulting in an over-recording of E. obscurum. Both for and against this scenario arethe facts that to date E. ciliatum has only been recorded twice in Fermanagh, by expert visitingbotanical recorders Daniel Kelly and Ian & David McNeill
Hybrids
Since E. obscurum ecology overlaps with that of several other willowherb species, it formshybrids, particularly with E. ciliatum and with E. palustre, which further complicates itsidentification. In Ireland, none of the hybrids involving E. obscurum are common and some arevery rare (Stace 1975; Kitchener 1992b; Stace et al. 2015).
Flowering and vegetative reproduction
Reproduction is both vegetative, by means of long-running stolons (produced above or below ground) andplant fragmentation and, much more effectively, by large numbers of long-persistent, wind-dispersed,plumed seeds that permit colonisation of new environmentally suitable sites (Grime et al. 1988,2007). The spreading stolons are produced in late summer from the base of the erect plant and they bearleaf pairs at nodes along their length, but no terminal leaf rosettes (Sell & Murrell 2009). Aerialstems may also become broken when ground becomes flooded and fragments may then be transporteddownstream to colonise fresh sites (Grime et al. 1988, 2007).
Flowering takes place in July and August; the inflorescence is a branched raceme of around 20 small, deeppink flowers, each 7-9 mm in diameter. Flower buds are erect and acute in shape and the open flowerpetals are 5-6 mm, shortly 2-lobed. The flowers are hom*ogamous and self-pollinated and numerous, plumedseeds are released when the 4-valved fruit capsule splits (Sell & Murrell 2009).
British and Irish occurrence
E. obscurum is widespread in the N & S of Ireland, but it is less prevalent in C, W & Eparts of the island. It is also widespread throughout most of Britain, but either absent or apparentlydeclining in recent years in Highland areas, W Scotland and E & SE England. Reasons for thesedeclines are unclear, but they might possibly have arisen due to previous mis-identification andover-recording involving the arrival from N America and spread of E. ciliatum (G.D. Kitchener,in: Preston et al. 2002).
European and world occurrence
E. obscurum is widespread in W & C Europe, becoming less frequent to both northwardsinto Scandinavia and southwards to the Mediterranean basin, although it does reach N Africa, Madeira andthe Azores. It just reaches as far SE as Turkey and the Caucasus (Hultén & Fries 1986, Map 1363;Sell & Murrell 2009). It is introduced in New Zealand (Webb et al. 1988).
Threats
Eutrophication and soil nutrient enrichment may intensify competition beyond the survival ability of thisspecies.
Introduction, neophyte, very rare, but invasive, difficult to identify and very probably under-recordedin Fermanagh in disturbed ground near habitation.
August 1983; Kelly, Dr D.L.; disturbed ground, Enniskillen Town.
July and August.
There are just two records for this weedy alien willowherb in the Fermanagh Flora Database, the firstabove by Dr Kelly has vouchers in DBN and TCD, and the second, was made on 21 July 1997 byIan and David McNeill at Clabby Church.
This N American, rosette-forming perennial was first recorded in Ireland at the small E coast port ofArklow, Co Wicklow (H20) in 1958. The second discovery was in 1971 on an island in Lough Neagh. After adelay of approximately 20 years it appears that E. ciliatum then began to spread in Ireland inthe 'explosive' manner typical of many successful introduced species, although in reality it was fairlycertainly overlooked for an unknown period while it was actively spreading in the Dublin area prior to1980. The reason for this involves confusion with E. obscurum (Short-fruited Willowherb), fromwhich it can be very difficult to distinguish (Doogue et al. 1985; Cat Alien Pl Ir).
The 20 year 'delay' after an initial arrival phase for an introduced species, conforms to a patternfamiliar from other such cases and, indeed, the early history of E. ciliatum in Britain isremarkably similar to the Irish experience, the dates being much earlier, however. It was first found inEngland at a reservoir in Leicestershire (VC 55) as long ago as 1891, although it was not recognised andcorrectly named until 1934. By then, it was well established at least in Surrey (VC 17), having spreadrapidly especially around the 1930-34 period (Preston 1989).
The current widespread distribution of E. ciliatum throughout B & I revealed in the NewAtlas and the frequency with which it forms hybrids with other common members of the genus(particularly with the very common E. montanum (Broad-leaved Willowherb)) indicates that E.ciliatum ought to be looked out for in Fermanagh in all kinds of disturbed, mainly urbanhabitats. It certainly remains under-recorded, most probably being mistaken for E. montanum andE. obscurum.
Threats
None as yet, although it is invasive and is spreading rapidly elsewhere, especially in urban sites.
Native, frequent. Circumpolar boreo-temperate.
1900; Praeger, R.Ll.; Co Fermanagh.
May to November.
Growth form, identification and preferred habitats
E. palustre is a stoloniferous perennial herb, 15-60 cm tall, with leafy, terete (cylindrical),pale green stems rising erect from a curved base (Sell & Murrell 2009). E. palustre caneasily be confused with several other Epilobium species and perhaps most easily with E.obscurum (Short-fruited Willowherb). However, E. palustre is distinguished from all otherwillowherbs that possess a club-shaped stigma, by the absence of raised ridges on the stem, althoughthere may be two lines of hairs (Kitchener 1992b). For other identification points separating members ofthis willowherb group, see the E. obscurum account above.
This calcifuge willowherb occurs at all elevations, almost right up to the summit of Cuilcagh, thehighest mountain in Fermanagh. It can grow on a wide range of usually base-poor, but occasionallyflushed, damp to wet, acid to neutral soils. Marsh Willowherb can also occasionally be found inlimestone areas, but it then appears confined to pockets of leached, acid soil, or in ground wherepatches of glacial drift overlie and bury calcareous bedrock.
In common with other primarily wetland willowherbs, the range of habitats occupied by Marsh Willowherb isextremely wide. It includes damp peat bogs, either cut-over, flushed, or neither of these, plus swampyand marshy lakeshore grassland, muddy bare ground or short sedge fen. However, E. palustre is notan aquatic species and it shuns both exposure on the margins of open water and shading from tall-herbfen or dense fen-carr environments. It can, however, tolerate semi-shade on woodland margins or inclearings and it very often features as a pioneer species in open, fully-illuminated sites on tracksides, ditch margins and stream sides of all dimensions.
As with other willowherbs, E. palustre can also colonise rather drier situations than it normallyfrequents, occupying open disturbed ground on wasteland, roadsides, disused quarries or sand- andgravel-pits. In upland areas, Marsh Willowherb can occur, in addition, on cliff ledges and on wet rockoutcrops. In every situation and on all occasions, it avoids competition from more vigorous, taller,aggressively crowding plant species. This is why the established strategy of E. palustre isdescribed as intermediate between C-S-R and Stress tolerator by Grime et al. (1988, 2007).However, it is important to bear in mind that this ecological determination is based to a considerableextent on the species' ability to garner nutrients from the environments it occupies, and E.palustre tolerates an unusually wide range of habitats and demonstrates a range of strategies tocope with different ecological niches (Craine 2009).
While Marsh Willowherb cannot survive heavy or regular disturbance in the form of grazing, cutting ortrampling, it is favoured by moderate levels of these and other growth limiting factors which helpprevent potential dominant species ousting it. At the same time, it tends to occupy soils ofintermediate fertility and moderate pH, rather than the extremes (Sinker et al. 1985; Grime etal. 1988, 2007).
E. palustre overwinters both as dormant seed and as small, tight, terminal buds on the ends ofthread-like underground stolons emanating from the base of the old plant. This is an importantdistinguishing character from the recently arrived and now widespread American alien, E. ciliatum(American Willowherb).
Fermanagh occurrence
In terms of its frequency and distribution statistics in Fermanagh, E. palustre trails E.montanum (Broad-leaved Willowherb) by a wide margin, but it more closely matches figures for thevery much larger species, E. hirsutum (Great Willowherb). Nevertheless, E. palustre isstill the third most frequent and widespread willowherb in Fermanagh, being present in 254 tetrads,48.1% of those in the VC. While it is well scattered throughout Fermanagh, E. palustre issomewhat more prevalent around Upper Lough Erne and on the Western Plateau.
Flowering reproduction
E. palustre flowers in July and August, the inflorescence consisting of a loose raceme of justaround ten small, mauvish-pink flowers, 4-6 mm in diameter. The flower buds are initially erectbut soon droop, so that the whole raceme hangs over to one side and the flowers are held almosthorizontally. The flowers are perfect (ie hermaphrodite), hom*ogamous (male and female ripensimultaneously) and they usually self-pollinate. Seed is set from July to September – the fruit capsulebeing green, flushed crimson, 5-8 cm long and splitting to release numerous, small, lightweight, plumedseeds that are wind-dispersed (Ross-Craig 1948-1973, Part XI Plate 27; Grime et al. 1988, 2007;Sell & Murrell 2009).
Vegetative reproduction
In summer, filiform (thread-like), often but not always subterranean stolons bearing distant pairs ofyellowish scale-leaves and terminating in autumn in swollen, bulbil-like buds with fleshy scales (ieturions), develop from the base of the aerial plant, providing the over-wintering mechanism of E.palustre. These bulbils or turions give rise to new plants in the springtime, often at up to 10cm from the original parent plant. Detached turions and portions of stem may also play a part invegetative spread of the species in some of its more disturbed habitats, for instance being carried awayin flood waters (Grime et al. 1988 & 2007; G.D. Kitchener, in: Preston et al. 2002).
Seed longevity
Thirteen studies examined in the survey of NW European soil seed banks gave mixed results and no clearindication of buried seed longevity (Thompson et al. 1997). Six of the studies found seedpersisted less than one year, three reported that they survived between one and five years, and oneconsidered them long-term persistent (at least five years). Three studies could not assign them to anyof these categories.
Most members of this genus have a proportion of their seed capable of surviving for more than one yearalthough others, like E. hirsutum and E. obscurum, have long-persistent seed buried in thesoil that can survive five or more years.
British and Irish occurrence
E. palustre is widespread and locally common throughout B & I, but has significantly declinedin SE England, mainly since 1950, most probably due to drainage and eutrophication associated withagricultural intensification (G.D. Kitchener, in: Preston et al. 2002).
European and world occurrence
E. palustre is widespread in Europe northwards to Iceland and Lapland. It becomes morescattered and scarce towards the Mediterranean. It is also widely distributed in Asia, N America andGreenland, making it Circumpolar Boreo-temperate (Hultén & Fries 1986, Map 1365).
Hybrids
It is capable of hybridising with numerous other willowherb species (eg E. alsinifolium (ChickweedWillowherb), E. brunnescens (New Zealand Willowherb), E. montanum (Broad-leavedWillowherb), E. obscurum (Short-fruited Willowherb), E. parviflorum (Hoary Willowherb),E. roseum (Pale Willowherb) and E. tetragonum (Square-stalked Willowherb)), but they areall of rare occurrence in Britain, and either extremely rare or absent in Ireland (Stace et al.2015). Also put in alpha
Threats
Eutrophication of water bodies and soil nutrient enrichment generally, could well stimulate competitionand oust this species.
Epilobium brunnescens (co*ckayne) P.H. Raven & Engelhorn, New Zealand Willowherb
Introduction, neophyte, a frequent garden escape now well naturalised.
1940; Faris, R.C.; Benaughlin.
Throughout the year.
Growth form and introduction
A low-creeping, small-leaved, prostrate, mat-forming perennial with often almost unopened, insignificantlooking, solitary pinkish-white flowers produced on relatively long stalks, E. brunnescens wasfirst recorded in the British Isles as a garden weed in Edinburgh in 1904, and as a naturalised plant ina disturbed, non-garden situation in Wiltshire in 1908 (Davey 1953). It made its first publishednaturalised appearance in Wales as late as 1930 (Harrison 1968), and did not appear in Ireland until1932 (Brunker 1950), even though Nelson (1994) has uncovered evidence that Irish nurserymen were sellingit to gardeners as a rockery plant in the early 1900s. Apart from this deliberate introduction, E.brunnescens very probably also arrived in B & I accidentally. Seed could easily have beentransported in soil on the roots of imported New Zealand trees, shrubs, alpines and other garden plants,many of which are popular and grow very successfully here.
In common with the pattern of other introduced species, after an initial delay, E.brunnescens, which seeds itself freely, entered an 'explosive phase',spreading very rapidly from numerous localised focal points. As a result it is now widely distributedand frequent to locally abundant in northern and western parts of B & I at elevations from sea-levelto high mountain summits, preferring the higher ground (Davey 1961; New Atlas).
Preferred habitats and ecology
The plant has a shallow fibrous root system penetrating only 6 cm at most, a limitation which makes itrequire either an almost permanently damp, although well-drained gritty, sandy or stony soil, or anotherwise drought-protected substratum. Thus although the plant is a wintergreen perennial, spreadingand forming low, clonal mats by means of frequent branching and rooting at leaf-nodes, individuals aresusceptible to drought and may be killed if it is severe or prolonged (Davey 1961). This undoubtedlyhelps explain why the species is scarce or absent from most of the E, SE and Midland counties of Englandwhich regularly experience high summer temperatures and droughts that would be lethal to this species(Preston et al. 2002). This explanation does not cover the similar scarcity of E.brunnescens in C Ireland and, at present, the current author (RSF) is unable to supply a logicalexplanation.
E. brunnescens occurs on various types of soil, both acidic and lime- or base-rich, alwaysprovided they are sufficiently damp, free-drained and the habitat open to seed colonisation (Davey 1953,1961; Harrison 1968). The habitat openness requirement for establishment frequently involves a modestdegree of substrate and vegetation disturbance, either through natural instability of the site, or humaninvolvement, eg trampling or grazing by stock. While typically found in fully illuminated conditions,the species can tolerate some degree of shade, for instance along woodland paths or in ditches, or fromolder ericaceous shrubs (Davey 1961). The plant is regularly found in close association with othersimilar low-growing or creeping species, eg Lysimachia tenella (Bog Pimpernel),Chrysosplenium oppositifolium (Opposite-leaved Golden-saxifrage) and Sagina procumbens(Procumbent Pearlwort) and various mosses including Sphagnum species. Care is then needed in itsidentification, since non-flowering specimens of the species might easily be confused with Lysimachiatenella.
Reproduction
Although limited vegetative spread does occur (ie local diffusion as a result of horizontal growth),drought sensitivity makes the species very dependent on seed for both population maintenance anddispersal. The numerous solitary flowers, produced over a prolonged period from April or May through toNovember, are almost always self-pollinated (but see below), often in bud or in the unopened flower.They are self-fertile and the abundant tiny seed (c 120 per capsule) thus produced are equipped forefficient wind dispersal with the tufted plumed pappus typical of the seeds in this genus (Davey 1961).Germination can occur after only a fortnight, but late autumn seedlings seldom survive while thoseproduced in early spring have the capacity to flower later the same season (Davey 1961). The currentauthor (RSF) has not found any information regarding seed bank longevity, except that Davey (1961)reported no germination in seed after 17 months storage.
Fermanagh occurrence
E. brunnescens is frequent, widespread and is probably still actively spreading in Fermanagh,having been recorded from 87 tetrads to date, 16.5% of those in the VC. It has even reached the summitof Cuilcagh, the highest mountain in Fermanagh – a decidedly remote situation that is not all thatfrequently visited.
Hybrids
Hybrids between E. brunnescens and no less than five other species of willowherb have beenrecognised from various parts of B & I, proving that cross-pollination in the wild clearly occurs.Irish hybrid material involved originates from Cos Antrim (H39) and Tyrone (H36), and comprises ofcrosses with alien E. ciliatum (American Willowherb) and native E. obscurum (Short-fruitedWillowherb) (Kitchener & McKean 1998).
Name
The Latin specific epithet 'brunnescens', means 'approaching brown', ie tending to become brown orbrownish, but implying not attaining full development of that colour; a reference to the leaf colour(Stearn 1973).
Threats
There are no threats to it and while E. brunnescens is a frequent and widespread relatively recentintroduction, especially in upland sites, it appears to co-exist with native species rather than oustthem.
Chamaenerion angustifolium (L.) Scop. (= Epilobiumangustifolium L., Chamerion angustifolium (L.) Holub), RosebayWillowherb
Native, frequent and widely scattered throughout. Circumpolar boreo-temperate.
1904; Praeger, R.Ll.; western end of the Cliffs of Magho.
Throughout the year.
Growth form and preferred habitats
A very tall, leafy herbaceous perennial, 1-3 m in height, with a combination of long, horizontallyspreading and deeply penetrating roots. Stems can be either simple or branched and are glabrous belowand hairy above. Leaves are alternate, spirally arranged, 5-20 × 1.0-3.5 cm, oblong-lanceolate in shapewith margins entire or with small, distinct horny teeth. The leaf margin is also often wavy and the baseis narrowed and either sessile or obscurely petiolate. Shoots die back completely in autumn, buds on theold stem base overwintering and giving rise to new aerial shoots in spring. A strongly competitivespecies with high mineral nutrient requirements, it often forms large, dominant clonal patches under awide range of usually ± fertile (ie nutrient-rich, well-aerated and favouring nitrification) disturbedconditions (Myerscough & Whitehead 1966; Grime et al. 1988 & 2007; Broderick 1990). It israrely found in long-established plant communities, being more characteristic of disturbed, neglectedderelict land (Myerscough 1980).
The history of its spread
In Praeger's prime recording days (the late 1880s to the early decades of the 20th century), this specieswas recognised primarily as a plant of mountain cliff ledges and rocky gullies, chiefly confined to thenorth of both B & I. By the mid-1940s, however, it effectively modified its tolerances and hadalready colonised a variety of once-disturbed lowland habitats. While this tall-growing speciesis now a pioneer, rapidly invading recently disturbed ground, it cannot survive regular, continueddisruption in the form of trampling, grazing, cutting or burning (Myerscough 1980). Suitable fresh sitesoften involve major works of vegetation clearance and an extensive exposure of bare soil or burntground, eg in clear-felled or wind-thrown forestry plantations, major roadworks, building or bomb sites,or where land has been recently reclaimed from scrub invasion.
Having successfully colonised and established numerous new, sometimes dense, dominant populations in thisrange of disturbed lowland sites, C. angustifolium then used these foci to colonise a furtherseries of often much smaller patches of disturbed, burnt or droughted ground, in a very wide variety ofsoil types. The soils range from wet mud to rather dry sand and from acid to fairly alkaline (Myerscough& Whitehead 1966; Myerscough 1980; Broderick 1990). This later more 'weedy' colonisation was muchless aggressive and more gradual in Ireland than was the case in Britain, so that Webb (1972) was onlythen reporting the plant's recent arrival in parts of central Dublin.
Reproduction
In unshaded conditions, C. angustifolium flowers from July to September. The inflorescence is along, spike-like raceme, often bearing up to 200 bisexual, perfect, purple flowers, 20-30 mm indiameter. The inflorescence size generally indicates the reproductive investment the plant can afford.The flowers are protandrous and can self-pollinate, although they do also attract insect visitors(mainly bees) for pollination (Schmid-Hempel & Speiser 1988). Nectar is secreted by the epigynousdisk at the base of the style (Clapham et al. 1987). Thanks to the height of the plant (0.5-3 m),species mobility is conferred by the considerable output of well-plumed, airborne seed, released fromlong capsules frequently 200-250 cm above the ground.
It has been shown, however, that once established in a site and having developed a ± dense population,C. angustifolium plants direct a very high proportion of their photosynthetic resources intovegetative growth and horizontal root or rhizome spread, rather than towards flower and seed production(van Andel & Rozema 1974; van Andel & Vera 1977).
Having a very extensive underground network of rhizome and mycorrhizal fibrous roots, the plant is aneffective accumulator of mineral nutrients after forest fires and has a conservative system of internalmineral recycling within the root and shoot (van Andel & Nelissen 1979). The extensive annual shootlitter is also rapidly recycled, while the root system of existing stands appears to inhibit its ownseed germination (van Andel & Rozema 1974). These properties enable dense colonies to persist for 30or more years and individual root segments can survive for up to 47 years, and perhaps much longer (vanAndel 1975; Broderick 1990).
Species native status
The suggestion that the C. angustifolium population that spread so noticeably in the 20th centurywas alien rather than native was not supported by morphological, genetic and cytological studies(Myerscough 1980). Molecular genetic tests have yet to pronounce a verdict on this question.
Fermanagh occurrence
Consequent to all this, as the tetrad map shows, C. angustifolium is now widely scattered across162 Fermanagh tetrads, 30.7% of those in the VC. It occupies a range of weedy, ruderal habitats,including quarries, roadside banks, urban waste ground, the unmanaged rough ground margins of bogs andcar-parks, and it can even grow and survive to flower on damp walls.
British and Irish occurrence
C. angustifolium is common, widespread and often abundant over most of B & I, except for thefar west of Ireland and in patchy areas of NW Scotland including the islands (Preston et al.2002).
European and world occurrence
A completely circumpolar species (Circumpolar Boreo-temperate element), C. angustifolium occursacross Eurasia and N America from 25oN to within the Arctic Circle. It is a more recentintroduction in some parts of its range and, for instance, has now been recorded in N Africa. In Europe,it is widespread and generally common in the north, but is more scattered and rare in the south(Myerscough 1980; Hultén & Fries 1986, Map 1355; Sell & Murrell 2009). In the more southern partof the species distribution, Hultén & Fries (1986) separate off subsp. circumvagum Mosquin,mainly in N America, and Myerscough (1980) follows this decision.
Threats
None.
Introduction, neophyte, a rare garden escape.
July 1982; Northridge, R.H.; roadside waste ground on the Tempo road near Enniskillen.
Oenothera is a large, originally American genus of mainly annuals and biennials, (rarelyperennial) many of them taxonomically complex and critical, the species limits being still a matter ofopinion amongst taxonomists (Stace 1997; Stace & Crawley 2015). Some species have large,spectacular, yellow flowers, making them popular with gardeners. At least four species ofEvening-Primrose (so called because of their evening-opening primrose-yellow petals), plus severalhybrids, have 'jumped the garden wall' and become quite commonly semi-naturalised 'in the wild' inlowland B & I, chiefly in the southern half of England and in Wales, while remaining only rare andthinly scattered in Ireland. Around 18 additional species and hybrids of the genus are also reported inBritain as rare or very rare casuals (Clement & Foster 1994).
The biennial O. glazioviana is the most frequent and widespread of the four species that regularlyappear as B & I garden escapes and it displays a definite preference for sandy soils, both at thecoast and inland (C.A. Stace, in: Preston et al. 2002). It flowers from June to September andbears its stigmas above the anthers and, therefore, is more likely than other species to outbreed, andhence to form hybrids (Stace & Crawley 2015).
It has long been argued that this species arose in Europe in recent centuries, where it is establishedlocally in northern and central regions, rather than it being a native of America like the rest of thegenus (Clapham et al. 1962). It could be a neonative, evolved in Europe by mutation andhybridisation; if this were the case, it would be a second British case of hom*oploid hybrid speciation,in addition to that of Oxford Ragwort (Senecio squalidus) (Stace & Crawley 2015).
There are just three records of this large flowered species in the Fermanagh Flora Database, all found inand around Enniskillen town. In addition to the first record listed above, the other two records are:ten plants on waste ground behind the old town jail, 19 July 2006, RHN; and Silverhill, Enniskillen, 6October 2006, RHN.
O. glazioviana (= O. erythrosepala Borbás) is a tall plant, approaching 2 m in height, withnumerous large pale yellow flowers which seed profusely. Unlike Epilobium species (willowherbs),the seed is not plumed. The seed output of this and related species is so huge however, that it is nowbeing exploited commercially, both as an oil-seed crop and for its beneficial effects in medicine (GwynnEllis 1993).
O. glazioviana typically colonises open vegetation in waste places, car parks, roadsides and otherforms of disturbed ground (including railway ground, where this survives). At the coast, it invades sanddunes, the equivalent natural vegetation. While large populations can rapidly develop in freshly opened,disturbed ground, these usually decline equally quickly as the vegetation closes, so that the speciesmay be regarded as a non-persistent pioneer colonist or casual.
The genus Oenothera seems to have become more popular and fashionable with Irish gardeners duringthe 1970s, since the majority of records on the island post-date this period and they have increasedfrom rarity to occasional in status. This species is still only reported from seven Irish VCs, four ofthem in NI (ie H33 & H38-H40) (Cen Cat Fl Ir 2; Cat Alien Pl Ir). We expect to seemore of this attractive alien in Fermanagh in the future.
Threats
None.
Introduction, neophyte, both deliberately planted and naturalised, frequent.
1953; MCM & D; Letterbailey Td.
March to November.
Growth form, origin and introduction
This familiar, bright red and purple pendulous flowered, deciduous shrub grows up to c 3 m. It haspapery, peeling brown bark and opposite or 3- or 4-whorled, ovate to elliptic leaves with toothed toentire margins. Popular with gardeners as it flowers from June to October, F. magellanica is anative of Argentina and Chile and was thought to have been introduced to horticulture in the BritishIsles in 1788, although this may really have happened as late as the 1820s (Sell & Murrell 2009). Itwas first recorded in the wild in Britain in 1857 (D.A. Pearman, in: Preston et al. 2002).
Fermanagh occurrence
Fuchsia is never anything like as abundant in Fermanagh's hedgerows as it is in parts of W Donegal (H35)and in other Irish seaboard situations stretching anticlockwise from Antrim (H39) to E Cork (H5).Despite this, it is still quite frequently found in Fermanagh, having been recorded in 65 tetrads, 12.3%of those in the VC. Locally, Fuchsia is especially associated with old hedges near small upland orbogland homesteads in N & W Fermanagh, but it also occurs in a wide variety of other habitatsincluding cliffs, scarps and by waterfalls. Its habitats range from cut-over bogs to hedges beside lakesand along waterways and waysides.
Sexuality and sterility: an important limiting factor: Observations suggest that the greatmajority of Fuchsia bushes in Ireland were originally deliberately planted from around the 1850s onwardfor roadside or field hedging, or along woodland paths. Over a century later the shrub had becomenaturalised in more wild, semi-natural situations, but only occasionally so, and to a surprisinglylimited extent. In their Flora of Connemara and the Burren, Webb & Scannell (1983) reckonedthat in those regions of W Ireland, "less than one plant in a thousand grows anywhere but where itwas planted". In Fermanagh, RHN and the current author (RSF) equally roughly estimate this ratio ismore likely one in 50,000! The reason for this is that nearly all the hedgerow Fuchsia bushes belong toa cultivar of hybrid garden origin, possibly triploid, and despite forming the occasional fleshy fruit,they are either completely sterile or almost so (Valentine 1978). However, there are reports ofgynodioecy in some species of Fuchsia (ie where females and hermaphrodites coexist), suggesting that sexexpression may possibly be unstable (see Richards 1997a, pp. 318-9).
Other workers have found that in two American Fuchsias, F. thymifolia and F. microphylla,90% of the apparently hermaphrodite flowers are female sterile (although still bearing fully-formedfemale organs) and the plants are therefore functionally male (Arroyo & Raven 1975). Thus, thesexuality of Fuchsia species may well be more complex than appearances first suggest and thereason(s) for sterility is equally opaque. Probably in most parts of Ireland only a minority of hedgerowplants belong to the true species, or to a form of it which is capable of sexual reproduction.
In the FNEI 3, Hackney et al. also comment that, "some stands fruit abundantly, othershardly so". According to a short note on some Kerry plants (H1 & H2) by Donaldson et al.(1976), until recently Fuchsia seed was sold in agricultural seed merchants in that county, "forsowing along banks where it served as feed for both cattle and sheep". The current author (RSF) hasnever heard of stock eating Fuchsia anywhere else, nor observed any evidence of it happening, and istherefore sceptical of this aspect of the report. However, without him distinguishing the two forms thatoccur in W Ireland, both of which he was perfectly aware of, Nelson (1994) stated clearly that,"Fuchsia is a useful shrub for hedging fields, because cattle do not like eating the foliage andshoots." This is not to deny that when grass and other forage is scarce or rare and animals arenear starving, they will eat almost anything to survive.
Reproduction and dispersal
In its native S American habitats, the bright red flowers of F. magellanica and its relatives arepollinated by humming-birds feeding on the copious nectar. In the absence of humming birds in B & I(for the moment!), the flowers are visited by hive bees and other insects (Valentine 1978, p. 121). Whenit is formed, the four-celled fruit ripens between August and the onset of the first frosts. It is aplump, burgundy-coloured fleshy capsule or berry containing numerous seeds (Hickey & King 1981).Presumably the fruit is edible and attractive to birds, but the current author (RSF) has not uncoveredany mention of this in the literature, although Ridley (1930) details examples of various foreign birdsand of Opossum dispersing fruit of the New Zealand species, F. excorticata, a plant which is onlyrarely grown in Irish gardens (Forrest 1985).
In B & I, we may be thankful that F. magellanica seldom spreads here by seed, since on theisland of La Réunion, in the Indian Ocean, it is a much more aggressive invader, penetrating previouslyundisturbed native montane forest and forming dense stands therein (Macdonald et al. 1991). Thereis evidence to suggest that in W Ireland there is a tendency for self-incompatibility in Fuchsia, andthe close presence of a second clone appears to be essential for pollination and fruit production(Nelson 1994). This requirement appears to be seldom met and, consequently, fruit set appears to beoccasional in Ireland beyond the garden setting, the majority of Fuchsia shrubs planted in hedgerowstending to increase and diffuse slightly, and over many decades, by a minor degree of suckering to formclonal scrub stands.
However, since Fuchsia wood is brittle, and the slender, young branchlets are easily detached, eg beingremoved by passing human and animal traffic, a much more rare and sporadic form of vegetativereproduction occurs through the rooting of transported shoot fragments. This might enable a somewhatwider dispersal, but obviously it must chiefly occur along narrow linear habitats, such as paths andstreams and be very occasional in its occurrence.
Whenever Fuchsia bushes do manage to produce seed, it is fertile and seedlings are occasionallyreported (Nelson 1994). Seedlings are frost-sensitive and can only survive overwinter in sheltered sitesin very mild areas of W Ireland. Thus, isolated Fuchsia shrubs are only rarely found, likely derivedfrom occasionally formed, self-sown seed, almost certainly involving fruit eaten, transported and voidedby birds. The bushes found locally in Fermanagh on the cliffs in Bolusty More townland, by Pollophouca(or Poulaphouca) waterfall, and on cut-over bogs near Knockennis and Farncassidy, are concrete examplesof these very probably self-sown plants.
Cultivar 'Riccartonii'
By far the most common hardy Fuchsia in Ireland and Scotland is cv. 'Riccartonii', which is believed tohave been raised in Riccarton garden near Edinburgh by a gardener called Young, sometime around 1830.The parentage of cv. 'Riccartonii' is still clouded by uncertainty, but it is said by some to be aseedling of a garden hybrid called cv. 'Globosa', which was raised sometime before 1832 by Bunney ofStratford. Cv. 'Globosa' is considered to be a cross involving F. magellanica var. conicafrom Chile, with the Brazilian species F. coccinea being the most likely other parent. Othersfeel that cv. 'Riccartonii' is merely a variety of the species, F. magellanica (eg M. Rix,in: Phillips & Rix 1989).
The hardiness of cv. 'Riccartonii', compared with other garden Fuchsias, was not appreciated at first,but when this property of the plant was discovered around the late 1830s it ensured its wider garden andlandscape use (Bean 1973, Vol. II, p. 246). 'Riccartonii' can be recognised by its short tube,only about 8 mm long and its stiff 18-22 mm long spreading sepals (Phillips & Rix 1989). Its calyxis a richer crimson and its unopened flower buds are much fatter and globose than those of the truespecies, but less so than those of cv. 'Globosa' (Webb & Scannell 1983; Krussmann 1985). Nelson& Walsh pointed out that the fat flower buds of cv. 'Ricccartonii' are so bulbous, and the calyx sofirm in texture, that they can be audibly 'popped' by squeezing them with one's fingers, a featimpossible with the other form of the plant (Nelson & Walsh 1991, p. 247).
When growing in hedgerows, F. magellanica cv. 'Riccartonii' prefers moist, peaty soils and eitherfull sun or partial shade. When established and protected by its neighbours it can tolerate temperaturesdown to -15oC.
British occurrence
In Britain, F. magellanica in its various hedgerow forms occurs from the Channel Isles toShetland. While it is frequent in W Scotland and along the S coast of England, elsewhere it is very muchmore scarce, scattered and coastal in comparison with its distribution in Ireland (Preston et al.2002).
European and world occurrence
Forms of F. magellanica are also naturalised in the wild from planted hedges in the Azores (Tutinet al. 1968) and in New Zealand (Webb et al. 1988), where cv 'Riccartonii' is avery common relict of cultivation in old or long-abandoned settlements all over the country. Asmentioned above, F. magellanica is a serious woody weed of native forests on La Réunion.
Names
The genus name 'Fuchsia' is named for Leonhart Fuchs (1501-1566), a German physician and herbal writerwho, of course, pre-dated its introduction to Europe and never witnessed the plant. The Latin specificepithet 'magellanica', refers to the area of the Straits of Magellan in S America, from near which theplant was first imported to Europe (Stearn 1992).
Uses
As children at play in Londonderry and Donegal, RSF remembers sucking the nectar from the flowers, andVickery (1995) reports the same from Cos Antrim (H39) and Down (H38), where the common name given theplant was 'Honeysuckle'. In Co Donegal (H34 & H35), the flowers were boiled to give a dark red dye,and on Merseyside there was a tradition that it was unlucky to bring the flowers indoors (Vickery 1995).
Threats
None. Although naturalised to a limited extent and very persistent irrespective of origin, Fuchsiamagellanica is mainly planted and not at all invasive.
Native, common. European temperate, but also in Asia and N America.
1882; Stewart, S.A.; Co Fermanagh.
March to December.
Growth form and preferred habitats
A clonal, potentially patch-forming, rhizomatous, polycarpic, herbaceous perennial with erect 20-60 cmstems, swollen at nodes, ± densely pubescent with some glandular hairs. Leaves 4-10 cm, ovate andtruncate or slightly cordate at base, sinuate-toothed or distantly denticulate, sub-glabrous or withmargins and underside of veins hairy. Inflorescence without bracts, consisting or about 50 well-spaced,open white flowers with parts in twos (Clapham et al. 1987; Sell & Murrell 2009).
While C. lutetiana is most frequently found in damp, shady woods, scrub and hedges, often nearwater, it also appears more rarely in shaded areas on screes and cliffs. It appears more or lessconfined to base-rich, moderately fertile, damp soils, and Grime et al. (1988) reckoned thespecies is intolerant of acidic conditions below pH 4.5. C. lutetiana is especially successfulcolonising moist, shaded sites with a relatively high proportion of exposed soil or litter and itsestablished strategy is described as Competitive-Ruderal (Grime et al. 1988, 2007).
Enchanter's-nightshade possesses spreading rhizomes and their rapid growth, even in shaded situations,enables the species to vegetatively colonise moist, relatively productive, somewhat disturbed siteswithin woodland, such as along tracks and rides, in hedge banks and on shaded riverbanks. Other obviousreasons why the plant may be most prevalent or most obvious in linear sites such as these could be theresult of better lighting there than under full woodland canopy and the fact that the fruit is a one ortwo-seeded burr. From August onwards each year, these burrs adhere to the coats of passing woodlandanimals, or to the lower parts of clothing including the socks of human traffic (Knight 1960). Isolatedpopulations of C. lutetiana are often found in woods a considerable distance from their nearestneighbours, providing circ*mstantial evidence of effective fruit dispersal by this means.
Fermanagh occurrence
A common and locally abundant plant of shady, damp places throughout lowland Fermanagh,Enchanter's-nightshade is mainly found in the Lough Erne basin. It is recorded in 173 tetrads, 32.8% ofthose in the VC. Many authors mention C. lutetiana occurring as a garden weed, perhaps colonisingshrubbery or other dark corners from nearby hedges, but as far as RHN and the current author (RSF) know,it is not recorded in any Fermanagh garden.
Flowering and vegetative reproduction
For the truly woodland shade plant that this species is, C. lutetiana flowers rather late in theseason, from June through into September. The attractive, pure white flowers each have parts in twos,with two caducous sepals (ie soon shed), two deeply bi-lobed petals, two stamens and one style, that isusually also deeply two-lobed. The flowers are self-compatible but they contain nectar and thereforealso attract insect pollinators, mainly small flies (Clapham et al. 1987). The 3-4 mm diameterfruit is pendulous and burr-like, being densely covered with stiff hook-tipped white bristles. Passinganimals, including man, pick up the burrs and transport the one or two contained seed (Clapham etal. 1987).
Seed is transient in the soil, persisting less than a year (Thompson et al. 1997). The leafy stemdies down after fruiting, but below ground at the base of each one, 2-6 horizontal, fleshy whiterhizomatous, overwintering shoots are produced. Although slender and somewhat brittle, the rhizomes canspread up to 30 cm annually through suitably loose soil or surface litter. When the parental axis diesin winter, the underground shoots are disconnected and the following spring the shoot tips break surfaceand commence growth as new clonal individuals surrounding the position of their forerunner (Salisbury1964, p. 348).
Despite its lack of evergreen aerial perennation and being noticeably late in developing spring vernalgrowth, C. lutetiana manages to persist in woods in competition with typical woodland floorspecies that display these adaptive advantages. This most likely reflects Enchanter's-nightshade'ssuperior colonising ability, conferred by its vigorous rhizome growth (Grime et al. 1988, 2007;Sell & Murrell 2009).
British and Irish occurrence
Enchanter's-nightshade is widespread and common in suitable shady, lowland habitats throughout Ireland,although less frequent in the far west. In Britain, it is similarly widespread except in upland Scotlandand in the more acidic terrain north of the Great Glen (Preston et al. 2002).
European and world occurrence
C. lutetiana belongs to the European temperate biogeographical element and is widespread in mostof Europe except the NE. It also stretches eastwards into C Asia and southwards to N Africa (Hultén& Fries 1986, Map 1351; Sell & Murrell 2009).
Threats
None.
Native, occasional, perhaps in decline. European temperate, but also in Asia and N America.
1839; Ball, J.; shore of Lower Lough Erne near Enniskillen.
June to August.
Growth form and preferred habitats
Upland Enchanter's-nightshade is a delicate, sterile hybrid herb with a more slender perennating rhizomethan its more common parent, C. lutetiana (Enchanter's-nightshade). It also produces over-groundspreading stolons from the lower leaf axils of its aerial stem that are usually absent in C.lutetiana. It occurs either with or without its parent species in moist woods and other shadyand rocky places, including streamsides and block scree in mountain gorges, on river shingle, lakesideboulders and in hedgerows, gardens and waste ground. Like its more common parent, and despite itsEnglish common name, Upland Enchanter's-nightshade is most frequent in lowland situations and it againprefers to grow in moist, mildly basic, fertile soils. C. ×intermedia has also inherited the rather weedy, colonising tendencies of C. lutetianaand, although it is totally sterile, producing no seed, it can spread into sufficiently damp and shadydisturbed wayside habitats by means of the vegetative growth of its rhizomes and stolons and theirtransport (Stace et al. 2015).
Identification
This inter-specific cross between C. lutetiana (Enchanter's-nightshade) and the misleadingly namedC. alpina (Alpine Enchanter's-nightshade) is distinctly intermediate in appearance between itsparent species and is easily recognised. Since C. alpina is extinct in Ireland, botanists herehave only to distinguish the hybrid from the common woodland Enchanter's-nightshade, C.lutetiana, with which it ecologically overlaps in damp, shaded, disturbed habitats, often nearwater. While the hybrid often occurs along with C. lutetiana, it can also appear on its own.
The hybrid has a less robust habit than C. lutetiana, its leaves are thinner and more definitelytoothed, more cordate at the base and more shiny (less hairy) than those of C. lutetiana. Thehybrid inflorescence is also much shorter and fewer flowered than C. lutetiana, although it stillbears the flowers rather remote from one another. The hybrid flowers are absolutely sterile; to quotefrom Raven (1963), "The anthers of C. × intermedia often fall undehisced, and I have neverseen a single well-filled, morphologically normal, pollen grain in a plant of this taxon. Likewise,plants of C. × intermedia fail to produce mature fruit."
Fermanagh occurrence
C. × intermedia has been occasionally, almost quite frequently recorded in 31 Fermanaghtetrads, 5.9% of those in the VC. As the tetrad map indicates, all but two of the twelve Fermanaghtetrads that have post-1974 records are from the woods and islands of Lower Lough Erne. In view of the14 widely scattered older records that were made by Meikle and his co-workers, RHN and the currentauthor (RSF) reckon the current distribution of C. × intermedia inthe rest of Fermanagh requires further investigation before it can be determined if it really hasdeclined locally or not.
Irish and British occurrences
This hybrid is quite scarce, widely scattered, yet locally abundant in the northern counties of Ireland(FNEI 3; NI Vascular Plant Database; New Atlas). In comparison, it has just three southernoutliers in Cos Wicklow (H20) and Co Dublin (H21) (Brunker 1950; Flora of Co Dublin). The recordsfrom both these VCs are pre-1975 – the Wicklow record, discovered by Praeger, dates from 1894 and thetwo Dublin ones from 1967 and 1973, and they all are or were associated with garden cultivation ordisturbed ground.
The occurrence of C. × intermedia is much better recorded nowthan in the early 1960s when Raven was studying it, yet it still displays a frequent andwide-ranging distribution in the W & N of Britain, becoming rarer eastwards and southwards, where itreaches the mouth of the River Severn in S Wales (Raven 1963; Preston et al. 2002).
Population maintenance and vegetative spread
While easily recognised, C. × intermedia is still variable in manycharacters, a fact which strongly suggests it arose locally on many occasions in the past when C.alpina was more widespread than at present, and the parent species much more frequentlyoverlapped. The alternative explanation is that the hybrid consists of a single or a few genets, whichperhaps arose in a somewhat drier than the present day post-glacial period 5,000 years ago, and whichthen gradually spread to the current B & I distribution through the vigorous vegetative reproductionthis hybrid possesses (Raven 1963).
However, this latter hypothesis fails to explain the observed variability of the hybrid and, in the viewof the current author (RSF), it greatly overestimates the dispersal ability of the sterile hybrid intonew ground. While the vegetative reproductive capacity of the hybrid allows it to maintain populationsin existing sites, it fails to offer a feasible mechanism of jump dispersal permitting anything morethan a very rare or occasional fortuitous colonisation of fresh habitat at any appreciable distance.However, it is necessary to consider the possible role of man in the dispersal of the hybrid, and thereis no doubt that C. × intermedia can and does at least match C.lutetiana in its ability to reproduce vegetatively. Prof. Webb pointed out to Peter Raven thatthe Dublin and Wickow records of the hybrid represented garden weeds, and these occurrences are or wereso remote from the other Irish records in the north of the island, that their origin very probablyderived from horticultural activity. Thus it is a sensible assumption that these remote southern Irishoutliers were introductions, perhaps the result of rhizome fragments that might have been transferredwith soil on the roots of imported garden trees or shrubs (Raven 1963).
Nowadays the C. alpina L. (Alpine Enchanter's-nightshade) parent of this hybrid is totally absentfrom Ireland and is a rare or scarce species in N & W Britain, relatively common only in the EnglishLake District and on the Scottish isle of Arran. While the scattered populations of C. alpina maywell be stable at present, there can be little or no doubt it is a relict species of cooler climaticconditions, representing a truncated, probably previously much wider distribution across both B & Iin the cooler, early post-glacial period.
C. × intermedia thus appears to be both a relict native and a gardenweed and this is very probably a situation not just confined to its Irish occurrence, since Raven (1963)noted, "Also in Merioneth [in Wales], for example, it [the hybrid] is mainly restricted todisturbed areas and man-made habitats, particularly in gardens and along roadsides, and is probablyextending its range in connection with cultivation (Benoit & Richards 1961)."
Threats
None.
Probably native, but possibly some sites planted or representing escapes from cultivation, rather rare.European temperate.
1872; Smith, T.O.; Colebrooke River (unspecified portion).
April to October.
Growth form, status and preferred habitats
One of the most distinctive identification features of the genus Cornus is the very obvious curvedleaf veins (in this species in 3-5 pairs), that are impressed on the upper surface, and stand outprominently beneath. The autumn leaf colour is a spectacular burgundy or claret reed and the bare redtwigs provide additional winter colour. A distinctive, often blood red-twigged, freely suckering, smalltree or shrub up to 6 m with a rounded crown and opposite, simple, deciduous leaves, Dogwood is regardedas native in Ireland although it is also frequently planted in hedges and for cover on estates. C.sanguinea is widely scattered in rocky places and thickets on lime or base-rich soils. In relictpatches of woodland and in scrub and hedges in Fermanagh and elsewhere across Ireland, C.sanguinea often grows associated with Crataegus monogyna (Hawthorn), Prunusspinosa (Blackthorn) and Rhamnus cathartica (Buckthorn), plus Euonymus europaeus(Spindle) and Viburnum opulus (Guelder-rose).
Although scattered and relatively rare elsewhere on the island, C. sanguinea is possibly alsoindigenous around the Lough Erne basin in Fermanagh (An Irish Flora 1996). The records at theCrom and Belleisle landed demesnes strongly suggest the possibility of deliberate planting there, andthe related American species C. sericea (Red-osier Dogwood) has definitely been introduced at theCrom estate (Kelly 1990). There might even be a suspicion that mis-identification of C. sericeafor C. sanguinea might be involved in one or two of the Fermanagh cases. Nevertheless, recordsshow these two dogwood species are present in woodland sites around Lough Erne, although both arerelatively rare.
Fermanagh occurrence
The Fermanagh Flora Database contains records of C. sanguinea from a total of 14 tetrads, nine ofthem with post-1974 dates. It is found on riverbanks and the margins of moist woods and thickets onbase-rich lakeshores and on limestone rock outcrops like Knockninny Hill. It is chiefly distributedaround both parts of Lough Erne and on adjacent ground.
These two Dogwood species can occur in similar habitats and, perhaps, even sometimes grow near oneanother. Only C. sericea, however, is a definite introduction in Fermanagh. Generally the latteroccurs in sites with easy access that are clearly and unambiguously planted, eg along tracksides inconifer plantation at Crom and in a shrubbery below Crom Castle itself. C. sanguinea is moresporadically scattered than C. sericea, sometimes in rather remote sites a considerable distanceoff road, eg on the limestone outcrop of Knockninny Hill and also near Lisnarrick, Crockhaver, Enaghanand at Scottsborough lakelet.
Reproduction
C. sanguinea reproduces both sexually by seed and vegetatively by vigorous suckering. Theinflorescence is a many-flowered, ± flat, umbel-like corymbose cyme that develops before the leavesfully open. The white flowers open in June and July and have their parts in fours with a 2-celled ovary.They contain nectar and give off a faint, but pleasant scent that attracts pollinating bees (Edlin1964). The fruit, which ripens in small clusters in September, is a shiny, purplish-black, pea-likefleshy drupe, with red-staining juice and a bitter taste (Lang 1987). Each drupe contains a single,hard, two-celled stone which is dispersed as a unit over the next few months by a range of common birdsof the thrush and crow families. The Starling generally is the most active seed vector (Snow & Snow1988).
British and Irish occurrence
In Britain, C. sanguinea is widespread and locally common on limestone, chalk and base-rich claysin lowland S England and E Wales, which means it has a rather definite south-eastern distribution,becoming rare or absent both northwards and westwards. In Ireland, as a native (or presumed native), itis thinly scattered, mainly in the W & C of the island. Both to the N & S of this zone, it israre and is regarded as a planted introduction (New Atlas).
In recent years, Dogwood has become commonly used in amenity landscape planting along roadside andmotorway embankments and in similar groundcover planting in towns and cities in both B & I. As aresult, Dogwood has spread sporadically beyond its native range and the boundaries between native andplanted are now rather blurred and more work is required to clarify the status in individual sites(Hackney et al. 1992; G.T.D. Wilmore, in: Preston et al. 2002).
European and world occurrence
C. sanguinea is restricted to temperate parts of Europe, absent from the NE and extreme N, andthinning southwards and becoming rare in the Iberian peninsula and in SE Asia and Asia Minor (Turkey,Iraq and Iran) (Hultén & Fries 1986, Map 1378; Sell & Murrell 2009).
Names
The genus name 'Cornus' is derived from the Latin 'corneus' or 'crann' meaning 'horny', an allusion tothe wood's hard texture. The Latin specific epithet 'sanguinea' means 'blood-red', referring to the twigcolour (Gledhill 1985). The English common name 'Dogwood' is also believed to derive from the hard whitewood that the branches offer, and which previously were used (often by travellers) to make good skewers,since the term 'dog' is or was used by timber-men to mean a sharp spike (Edlin 1964). The hard timberwas also used for millcogs, pestles, spikes and wedges. It also made arrows, ramrods, toothpicks andpipe stems and its charcoal was used to make gunpowder (Grigson 1987). Sixteen alternative Englishcommon names listed by Grigson include 'Cat-wood' and 'Gatter-bush', contemptuous references to cats andgoats associated with the sometimes bad or off-putting smell of the bush and dislike of its inedibleblack fruits (Grigson 1987).
Threats
None.
Introduction, neophyte, deliberately planted, rare and thinly scattered.
1950; MCM & D; Inishroosk, Upper Lough Erne.
June to August.
A native deciduous shrub of N America introduced to B & I ornamental gardens in the late 17thcentury, C. sericea has become rather commonly used in landscaping and amenity plantations forthe decorative effect of its many, bare, reddish-brown or blood-red winter twigs. It is also used forground- and game-cover, particularly in damp woods, hedges and other ground near water and in areassubject to occasional flooding during the last 60 or so years. Given eutrophic wetland conditions, C.sericea readily spreads by suckering and it can form large, dense clonal thickets up to 3 m tallthat are capable of suppressing virtually all pre-existing herbaceous vegetation (Kelly 1990).
C. sericea can be distinguished from C. sanguinea by its larger leaves usually having atleast six pairs of curved side veins as compared to at most five pairs in the latter. The young twigs ofC. sericea are a brighter red in spring and summer (although they turn brownish-red in autumn)and they are often covered in prominent dots. The flowers are also smaller (6-7 mm across) and theberry-like fruit is white rather than purplish-black (Sell & Murrell 2009; Parnell & Curtis2012).
In Fermanagh, C. sericea was recorded from five sites by Meikle and his co-workers between 1950-3and it has been found at a further eight stations since then, meaning that it is now thinly scatteredacross a total of just 13 tetrads, 2.5% of those in the VC. Fortunately, Red-osier Dogwood does not setseed and thus control of this potentially invasive, woody weed ought to be perfectly feasible. In acounty with as much damp and wet ground as Fermanagh, C. sericea has the capacity to become areal threat to biodiversity through suckering and layering and therefore planting of it should bestrongly and actively discouraged (Kelly 1990).
The New Atlas hectad map shows C. sericea is now widely distributed throughout Ireland andis particularly well represented in the north of the island. It has also spread rapidly in most oflowland Britain as far N as Glasgow and Edinburgh, becoming rarer and more coastal N & E ofInverness.
Threats
Potentially an invasive thicket-forming species in damp ground habitats.
Native, widespread, locally frequent. European temperate, but also widely naturalised.
1884; Barrington, R.M.; Ely Lodge Forest.
February to December.
Growth form, preferred habitats and Fermanagh occurrence
In many situations, inconspicuous for ten months of the year, Spindle is an occasional to frequent shrub(very rarely of tree-like dimensions), with sage-green, straight, slender twigs and leaves and greenbuds in opposite pairs. The pale green leaves have 6-8 pairs of veins, ± impressed above and prominentbeneath. The opposite buds and leaves lead the plant to produce almost square sectioned twigs throughthe development of four corky ribs on the somewhat older shoots. Further along the branch, rough, palebrown bark is eventually formed. Beneath the bark lies the firm, smooth, whitish wood that gives theplant its name. Spindle wood is strong, smooth and does not splinter and, although the diameter isalways small, as its English name declares, it was used for making spindles to spin wool into threadbefore the invention of the spinning wheel (Edlin 1964, p. 63; Sell & Murrell 2009).
E. europaeus strongly prefers and is almost confined to well-drained, base- or lime-rich soils. InFermanagh, it is found in the more open parts of woods and in hedgerows, especially locally around theshores and on the islands of both parts of Lough Erne. Although more frequently found in the lowlands,it also appears in shaded upland glens, again almost always in limestone districts, eg above Doagh Loughand at Hanging Rock NR. E. europaeus has been recorded in 129 Fermanagh tetrads, 24.4% of thosein the VC.
Flowering reproduction
Spindle is said to be polygamous, the flowers varying from bisexual to those with separate sexes,sometimes on different plants (ie it can be dioecious). The flowers are small, greenish-white andinconspicuous. They are borne in stalked cymose clusters in the upper leaf axils and open in June.Flower parts are in fours and they provide nectar and attract small insects as pollinators (Edlin 1964).Self-pollination can occur if there are no insect visitors (Sell & Murrell 2009).
The shrub is conspicuous only in the autumn (September-October) when in addition to the leaves turning adeep red, the colourful fruit is apparent. This is a capsule apparently consisting of a quartet of palepink berries dangling from a stalk, which splits to emit the bright orange seed – certainly providingone of the most startling colour contrasts in Nature! The fleshy bright orange colour is in fact anedible pulpy outgrowth of the seed coat, called an 'aril' (Edlin 1964).
Seed dispersal and plant toxicity
Despite their attractive appearance these 'berries' and almost all other parts of the plant are decidedlypoisonous to both domestic animals and man, although not so to birds (particularly Blackbirds),which eat the aril and disperse the seed (Lang 1987; Nelson & Walsh 1993). All or most parts of theplant contain a co*cktail of toxic substances, some of which are specific to the plant, eg evobioside andeuonymin. The poisoning symptoms suggest that some of the toxins are cardiac glycosides, while otherssuch as evonine are definite alkaloids. Poisoning of sheep and goats eating spindle twigs and leaves hasbeen reported and two horses died of digestive paralysis after browsing. About twelve hours after eatingthe fruits, human victims suffer vomiting, diarrhoea, stimulation of the heart and, in some severecases, hallucinations, convulsions and loss of consciousness. Fortunately, recent reports of humanpoisoning have only involved relatively mild symptoms (Cooper & Johnston 1998).
British and Irish occurrence
E. europaeus is native, frequent and widespread in England, Wales and Ireland, particularlyin more southern and eastern regions. However, it becomes scarce to rare northwards into N England andScotland, where its status also becomes questionable (T.D. Dines, in: Preston et al. 2002).
European and world occurrence
E. europaeus is widespread across most of temperate Europe, but becomes scarce or absent bothtowards the hotter Mediterranean basin and northwards into S Scandinavia. It is probably almost entirelyintroduced or naturalised in most of Scandinavia and this is definitely the case in its quite wide NAmerican occurrence where it is alien (Hultén & Fries 1986, Map 1296; Sell & Murrell 2009).
Uses
Despite its toxic nature, Euonymus has been used for a long time in herbal medicine as a purgativeand a diuretic, or externally as an unction for abscesses, chilblains, acne and wounds (Launert etal. 1981). The purgative fruits were also baked, powdered and rubbed into hair as a cure forhead-lice. Apart from its previously mentioned use in spinning, the wood was also frequently fashionedinto clothes pegs and sold mainly by members of the travelling community, a practice that gave rise tothe English common name 'Pegwood'. The wood was also used for making small items such as skewers,toothpicks and knitting needles (Sell & Murrell 2009).
The interesting autumn leaf colours and unusual fruit make E. europaeus to some extent gardenworthy and in parts of B & I a proportion of the shrubs recorded may represent naturalised escapesfrom cultivation (T.D. Dines, in: Preston et al. 2002).
Spindle eradication
Both Rackham (1986) and Sell & Murrell (2009) mention a programme of hedgerow eradication of E.europaeus that took place in the UK during the Second World War since the plant is considered asecondary host of the Peach-Potato Aphid, Myzus persicae. This sap-sucking insect is an importantpest on a wide range of crops, including many crucifers, potato and its relatives, peach and othergreenhouse crops including ornamental plants. The aphid spreads viruses such as Potato Leaf Roll Virusand Potato-Y virus. However, the aphid has many other secondary hosts, mainly amongst annual herbs, sothe removal of Euonymus from hedges provided no respite from crop infestation by this parasiteand its accompanying pathogen.
Threats
None.
Native, common and widespread. Suboceanic southern-temperate, often cultivated beyond its native range.
1882; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Fermanagh occurrence
Abundant and recorded in 441 tetrads, 83.5% of those in the VC, Holly is the fourth most common tree orshrub in Fermanagh in terms of record numbers and the fifth most widespread woody plant in tetrad terms.
It is common and widespread everywhere except on the most extremely exposed or the wettest ground. Itappears in woods, scrub, hedgerows, riverbanks and loughshores, plus a lesser presence on cliffs,gullies on rock outcrops, limestone scree and in crevices in limestone pavement. Holly is also veryfrequently planted around habitation (see below under Fodder).
Growth form and preferred habitats
A suckering, much branched, evergreen small tree or shrub up to 15 m tall, with alternate leathery, oftenprickly leaves, while its blackish-green, foliage casts a heavy shadow, Holly itself is extremely shadetolerant and it is most typically found as a secondary canopy understorey shrub in woods. Like otherevergreen-leaved species, the leaf lifespan is merely extended beyond a single year, the leaf eventuallyfading a greyish brown and dropping off the branch after about two years (Edlin 1964). Leaves drop atall times of year, but the main period of shedding is in mid-summer (Peterken & Lloyd 1967).
Thanks to suckering growth from its roots and occasional layering of pendulous lower branches that rootadventitiously, and depending upon the degree of animal disturbance it faces, I. aquifolium cansometimes form ± dense clonal groves in mixed woodland along with Oaks, Ash, Rowan, Hazel, Hawthorn andDowny Birch (and previously Elm). Vegetative spread may be important for filling gaps in stands, but itdoes not appear to be an effective means of species dispersal and spread (Peterken & Lloyd 1967).
However, Holly also commonly occurs as isolated individuals, sometimes of more tree-like dimensions, inhedgerows, or along the banks of rivers or streams mixed among other woody species. More or less stuntedHolly bushes are also frequent in crevices on cliffs, rock outcrops, screes or even in shady grykes, iethe deep, often dark, narrow fissures that occur in limestone pavement.
Holly grows in such a very wide variety of soils, acid to lime-rich that it appears almost indifferent tothe normal range of soil pH and nutrient status it encounters. While generally it avoids very wet, coldsoils and the extremely acidic nutrient-starved conditions of peat bogs, Holly can survive on the drierparts of fen alder-carr scrub around lakeshores and, in some areas of Britain, it manages to occupysoils that are waterlogged even in summer to within a few cms of the soil surface (Peterken & Lloyd1967).
Soil improvement
Like birches and, indeed, all other trees, Holly has the ability to improve podsol conditions by theproduction of a deep leaf litter which slowly decays to form mull humus and thus the tree graduallygenerates a more fertile brown earth soil beneath it. However, the growth rate of birch is very muchfaster than Holly, so that any comparison of the two in terms of soil improvement could only be made inthe extremely long term. In regions of high rainfall like Fermanagh and other western oceanic parts of B& I, leaching tends to override the mull-forming abilities of Holly litter, so that unfortunatelythe podsols persist and must thus limit the tree's growth (Peterken & Lloyd 1967).
Temperature limits
Holly is sensitive to prolonged winter frost and the natural distribution of the species in Europe fallswithin the -0.5oC winter isotherm for the coldest month, a feature which chiefly affects itsnorthern and eastern boundaries (Iversen 1944, pp. 474-6). The species also has a threshold minimumrequirement for summer temperature and this means that in B & I Holly is not found naturally inareas where the July mean does not exceed 12.8oC (= 55oF) (Peterken & Lloyd1967). It is very likely that reasonably high summer temperatures are necessary for the successfulformation of fruits on the female tree. Holly trees are susceptible to fire on account of theirremarkably thin bark, a property which is probably also involved in the damage they suffer in frost(Iversen 1944; Peterken & Lloyd 1967).
Seedling and sapling growth and first flowering
Given good sunlight, young Holly bushes or trees first flower when they reach a height of between 1.5-3.0m, generally reached when they are 8-15 years old. Bushes flower very poorly or not at all when growingin shade (Richards 1988). Seedling growth is exceedingly slow to begin with, achieving onlyapproximately 1.0 cm per year for the first four or five years after germination. This is followed by averitable growth spurt, when vigorous individuals can for a few years (obviously the duration variablewith circ*mstances) put on more than 50 cm per year. Shaded, understorey plants, however, cease activeheight growth somewhere between 5-10 m, depending upon the dominant tree species and its level of shade;17 m appears to be the absolute upper height limit achieved in woodland conditions. After the activegrowth spurt, height increase drops back to around 2.0 cm per year in trees over 30 years of age(Peterken & Lloyd 1967). The greatest recorded age of an individual Holly tree appears to be 254years, but as this particular specimen was healthy and not conspicuously large, greater ages areconsidered certainly attainable (Peterken & Lloyd 1967). The tree expert, W. Dallimore (1908)reckoned that without doubt, Hollies could live for 250-300 years.
Flowering reproduction
Mature Hollies, in good light, flower reasonably freely most years, but berry production variesenormously, probably following a masting cycle similar to that in Beech and Oak (see the Beech speciesaccount on this website for a description of the controlling factors and possible masting rationale).The trees and flowers are dioecious, although there have been very rare reports of perfect flowers (iehermaphrodite, bisexual – containing both sexes), which Richards (1988) suggests may really belong to apopular hybrid cultivar, I. × altaclerensis 'Pyramidalis'.
The sex ratio of trees is either a straightforward 1:1 or, in some populations, may be male predominantto some degree, although in semi-natural woodland shade a high proportion of non-flowering plants canobscure the picture in this respect (Richards 1988). One large Holly tree in Cologne was observed tochange sex from female to male between 1910 and 1916 (Hegi 1924-7, 5), but the true identity ofthis specimen is unknown.
Flower buds develop in late summer on the current year's shoot growth, and often are produced only highon the crown of the bush or tree. The flowers typically open in May and June of the following season (ieon second-year wood) but, occasionally, they bloom prematurely out of season between September andmid-December, presumably due to a combination of inherent variation and local environmental conditions,although artificial lighting may well play a part through mimicking the day-length flowering trigger(Taylor 1992; Stace 1993; Wheeldon 1993; Nicolle 1995).
Small, white flowers, 6 mm in diameter, are borne in few-flowered axillary cymes with their parts infours. In female flowers, nectar is secreted from tissue at the base of the ovary and bees and flies areattracted and serve as pollinators. The familiar bright red Holly 'berries' are in fact globose drupes,1 cm in diameter, each weighing 130-150 mg and sometimes up to 60 may be tightly packed on a singletwig. Towards December, the fleshy fruits ripen to crimson and attract birds such as Thrushes,Blackbirds and Finches that eat the pulp and void the small, hard, black seeds. Heavy berry crops arethe result, not of hard winters as many of the public imagine, but of a sunny summer 18 months previouswhen the flower buds were laid down (Edlin 1964).
Each 'berry' (ie drupe) contains up to four seeds and each female twig on a tree may bear up to 50berries. Potentially, even a small tree standing alone, 4.7 m tall and 3.7 m in diameter at the crown,might produce 120,000 seeds (Peterken & Lloyd 1967).
In common with other bird-dispersed stony fruits, germination is delayed and normally takes place in thesecond or third spring after formation. However, passage through the gut of a bird may speed up theweathering process, allowing the seed to germinate sooner (Hyde 1961).
Fossil history
I. aquifolium fossils are easy to identify both macroscopically by its wood, charcoal, seeds andleaves, and microscopically by its tricolpate pollen grains with distinctive sculpturing of the heavyexine which is unmistakable. The tree or shrub being entomophilous and having very much lower pollenproductivity than wind-pollinated species, its pollen grains are unlikely to be subject to distanttransport, giving the species high climatic indicator value.
Fossils from the glacial stages are absent except for scattered pollen grains, but there are plentifulinterglacial records, macroscopic and pollen alike, from the Hoxnian onwards. The return of Holly in theFlandrian interglacial was delayed until the Boreal period (zone VI), when it first appeared in Irelandand S Wales. Afterwards, it slowly spread east into England, and Holly sites became very frequent in thewetter Atlantic period (zone VIIa) and still more abundant from then on in the Sub-Boreal (increasedsummer warmth) and Sub-Atlantic (milder winters). Fossil pollen frequencies in B & I in the earlystages of this post-glacial have been highest of all in Co Kerry, SW Ireland. In the later historicstages of the Flandrian, pollen curves of I. aquifolium parallel those of weed and cereal pollen,marking the positive response of Holly to forest clearances and exploitation for fodder and other uses.However, this responsiveness to clearances and cutting tends to confuse any explanation of changingpollen abundance in climatic terms alone (Peterken & Lloyd 1967; Godwin 1975, p. 173-5).
In Ireland, there are no longer any pure stands of Holly such as are rarely encountered in Britain, eg inEpping Forest, or the unique ancient stand on shingle at Dungeness (Peterken & Hubbard 1972). TheseEnglish stands are considered relicts of medieval planted Holly woods which are believed to have oncebeen widespread in these islands, and particularly so in the west and north of Britain (Rackham 1980).
There is evidence that in past ages, Holly used to grow much larger trunks than it does at present; it isnow rarely seen larger than a shrub. In Ireland, it once formed pure stands from which valuable timberwas cut (Fitzpatrick 1933). At Inniskallen, Lough Leane, at Killarney, Co Kerry, in 1897 a 'celebrated'Holly tree was measured with a girth of 15 ft 4 in [4.67 m] (at 4 ft [1.22 m]) (Nelson & Walsh1993). There are historical records of large Holly trees being cut down on river banks in Scotland andin Northumberland in the 1830s and 1850s, although no measurements are given (Peterken & Lloyd1967).
British and Irish occurrence
Occurs throughout most of lowland Britain and Ireland and reaches 520 m in Argyll (VC 98) and 550 m in co*kerry (H1 & H2). The distribution becomes more scattered northwards and westwards into Scotland andsimilarly, but to a lesser extent, it thins towards the W coast of Ireland. In both cases, this mostlikely reflects the very open country, the absence of woodland, unsuitable soil conditions, excessivealtitude and exposure (Godwin 1975; New Atlas).
European and world occurrence
I. aquifolium has an oceanic and suboceanic distribution in Europe and adjacent parts of Asia. Itoccurs throughout NE, C & S Europe, SW Asia and N Africa but is often cultivated beyond itsnative area. It reaches 64oN in Norway where it is confined to the coast and the southernlimit is at 34oN in Algeria and Tunisia. It is absent in the Mediterranean from Cyprus andCrete. An eastern outpost is said to occur in Heupeh province, C China (Peterken & Lloyd 1967;Hultén & Fries 1986, Map 1295; Sell & Murrell 2009). Holly is also introduced in N America andNew Zealand.
Holly woods and use as fodder
Despite its prickly leathery foliage, Holly once provided important winter fodder for sheep and cattle(Radley 1961; Kelly 1997). For this reason, Holly was planted and encouraged in hedgerows near farmhabitation, making it readily available for cutting whenever winter fodder was required. In the 8thcentury, in Irish Laws of the Neighbourhood, I. aquifolium was one of the seven 'Nobles ofthe wood', the high ranking undoubtedly due to its fodder value, but possibly also on account of itsvery hard, heavy, ivory white timber which was used for 'chariot shafts' and for cooking spits (Peterken& Lloyd 1967; Nelson & Walsh 1993; Kelly 1997, p. 382).
It is important in this context to realise that in Ireland hay-making was rare before the coming of theNormans. The Old Irish law-texts, which provide extraordinarily detailed information on farming routine,contain no reference whatsoever to hay-making and no term for 'hay' has been identified in the languageof this period (Kelly 1997, p. 47). As a response to stock browsing, prickly leaves are concentrated onthe lower part of Holly bushes or trees, and the relatively spine-free upper branches werepreferentially cut for feeding to cattle and sheep. A 14th century Irish source also recounts how Hollyleaves were roasted a little to soften their prickles before feeding them to cattle (Kelly 1997, p. 46).One might imagine that boiling the leathery foliage would even better serve this function.
Holly hedges withstand pruning or clipping extremely well provided the tool used is sharp enough, in thecurrent author's (RSF) experience, and apart from animals being allowed to browse Holly repeatedly inwoodland, there is evidence that some Holly stands were deliberately planted and actively managed ascoppice or pollard to supply cut winter fodder (Rackham 1980, 1986, p. 120; Spray 1981).
The previous occurrence and distribution of these long gone managed Holly woods is hinted at chieflythrough the survival of place-names. The Old Irish name for Holly is 'cuillen' or 'cuileann', while'cullenagh' is 'a place producing Holly', so that many places still have 'cullen' or 'cullion'(anglicised from 'cuillionn'), as a word element, for example, 'Cullen Hill' near Ely Lodge inFermanagh. Irish townland names, either called 'cullion', or with it as a word element, are very commonthroughout the whole island (Joyce 1968; Milner 1992).
Names, uses and folklore
No less than 16 English common names are listed by Grigson (1987), reflecting both the variety of uses(timber, quick-lime also known as bird-lime, fodder, medicinal and Christmas seasonal decoration),magical folklore beliefs (lucky and unlucky) and physical properties of this familiar woody species,especially its prickles. Medicinal uses included treatment for chilblains, arthritis, rheumatism andwhooping cough. In Ireland, it was used for treating burns and for curing a stiff neck (Vickery 1995;Allen & Hatfield 2004).
Threats
None.
Introduction, planted or naturalised, very rare. Sub-Mediterranean-Sub-Atlantic, but widely naturalised.
11 June 1988; RHN & RSF; County Bridge, south of Garrison.
April to August.
Growth form, native status in England and preferred habitats
Although still considered native in up to ten famous, long-known sites on chalk and limestone in SEngland, in the vast majority of its other very numerous, widely scattered sites across B & I, thisdistinctive, slow-growing, evergreen shrub or small tree with opposite, dark green, smooth, leatheryleaves, 1.2-2.5 cm in length, is now recognised as being a naturalised introduction of garden origin(Wigginton 1999).
Biogeographically, B. sempervirens is regarded as a Sub-Mediterranean orSub-Mediterranean-subAtlantic species, and apart from the fact that Box appears as an element in manyplace names in S England, the reasons for doubting its native status in England was based on threeadditional pieces of evidence: the quite definite lack of a fossil (pollen or macrofossil) from theearly part of the current Flandrian interglacial in B & I (although it had been recorded from thetwo previous interglacials – the Hoxnian and the Ipswichian) (Godwin 1975); the supposed (casuallyassumed) absence of Box from adjacent areas of N France, except where it had obviously been planted forgame cover; and because the hard, fine-grained wood has always been considered a very valuablecommodity, and therefore B. sempervirens might be one of the plants introduced to Britain by theRomans on account of its usefulness (Tansley 1939).
Native status is very important and significant within the framework of biological conservation efforts,often attracting Government and private funding for preservation and enhancement of existing sites.Research by Pigott & Walters (1953) showed that B. sempervirens remains present and appearsperfectly native in numerous sites in N France that in geography and habitat conditions closely mirrorexisting Box sites in S England, and therefore no discontinuity whatsoever exists between English standsand those on the near continent. It is noted that several French local Floras have treated B.sempervirens as native in places around Paris and E, W & S Normandy (Pigott & Walters1953). On account of the value of the wood, planting of Box-groves over several centuries has certainlyaugmented native stands in both France and England, but the evidence of Anglo-Saxon place namesinvolving the element 'box' from the 8th century onwards indicates impressive antiquity to the stands ofthe plant.
After glacial ice melted, 10,000 BP, plants migrated back into B & I. Pigott & Walters (1954)suggest that, even with forest cover increasing (at first Betula and Pinus, laterQuercus and Ulmus) chalk and limestone scarp slopes in southern Britain remainedfree of trees for a longer period into the post-glacial, due to removal by solifluction of pre-glacialsoils, than did the neighbouring clay and plateau areas. These sites would have favoured thecolonization by Buxus therefore and enabled the plant to spread over southern Britain on thechalk and northwards on the oolite, reaching the Carboniferous Limestone of the Lake District by theclimatic optimum, 3,000 years after entering Britain. With an increasing forest cover, the Buxusdistribution would have become fragmented, and the deterioration of the climate in the Sub-AtlanticPeriod restricted the range of the species until, in Neolithic to Roman times, it was confined tosouthern England, where it still occurs today (Staples 1970).
Box might therefore be native in at least a small number of southern localities, such as Box Hill inSurrey (VC 17), on unstable limestone landslip at Noar Hill, Hampshire (VC 12) (Brewis et al.1996), and along ancient trackways, for instance the Fleam and Devil's Dyke in the Chilterns andCotswolds, and this proposition is widely accepted today. Part of the argument hinges on archaeologicalfinds of probable Box charcoal in association with Neolithic camps on the South Downs 2,000 years beforethe Roman invasion, suggesting (since Stone Age farmers are not considered to have been planters orimporters), that B. sempervirens grew wild in S England at that time (Marren 1992).
B. sempervirens usually grows as an understorey tree or dense shrub in woodland on limestone(usually with Beech or Yew), or as a major component in woodland canopy, on chalk in S England or onoolitic limestone in Gloucestershire. Nowadays, it is often most prominent on steep slopes where theroots of larger forest trees cannot maintain those species on the shallow soils (Staples 1970; Wigginton1999).
Box has a reputation for long persistence after plantation (deliberate or bird-sown) in woods and slopethickets, estate pleasure grounds and gardens and it may also occur in hedges or in other shady sitesnear habitation after 'jumping the garden wall'. Box can also survive and establish as a discard fromcultivation, often dumped in less appropriate waste or derelict ground habitats, but also surviving inlong abandoned gardens or in other neglected, unmanaged ground around ruins. Since Roman times, Box hasbeen a traditional hedging and topiary subject and, in a dwarf form, it remains a very popular edgingplant for beds in both decorative and kitchen gardens.
Flowering reproduction
Although it will flower when not in deep shade and does so early in the year in April or May, the blossomis small, yellowish green and insignificant in appearance, attracting little attention. Each smallcluster in a leaf axil contains both male and female flowers, usually 5-8 males and a solitary female.Pollination involves insect visitors and the subsequent fruit does not ripen until October. The fruit isan ovoid, white, papery capsule or pod containing up to six shiny, hard, black seeds that are releasedexplosively when it eventually splits (Edlin 1964; Sell & Murrell 2009). Each seed possesses anelaiosome or nutritive oil body in the form of a caruncle, developed from the lips of the ovulemicropyle. The food body attracts ants which assist local dispersal of the seed, although this is onlyactive and positive over a few metres at most (Ridley 1930, p. 524). Only vigorous bushes in the moreopen parts of woods or gardens produce viable seed (Pigott & Walters 1953).
British and Irish occurrence
As mentioned above, B. sempervirens is considered native in just ten sites scattered across SEngland. For instance, in parts of Berkshire (VC 22), Box stands appear to be well grown and are said tooften give rise to self-sown individuals (Crawley 2005). However, seed production in Ireland at least ispoor, so that self-sown seedlings are rare or very rare, with only eight sites from four VCs listed inCat Alien Pl Ir, in Cos Waterford (H6), Cavan (H30), Tyrone (H37) and Armagh (H38). The NewAtlas hectad map, in comparison, plots records of the most recent date class (1987-99) from 14Irish VCs. Three of the four Co Tyrone records are associated with major landed estates in the VC wherethe plant is undoubtedly planted (McNeill 2010) and all eight records from Co Wexford are of plantedsites, the majority from around ruins of past habitation (Green 2008).
Fermanagh occurrence
There are just six widely scattered Fermanagh stations for B. sempervirens. In addition to thefirst record above, the details are: lane approaching cliffs, Tiranagher Beg Td, 24 April 1994, RHN& RSF; Druminiskill, 26 April 1995, RHN; Old Crom Castle, 3 July 1995, RHN & RMHN; Bigwood Td,Lower Lough Erne, 13 August 1996, RHN, RSF & Don Cotton; Kilcoo Crossroads, 4.5 km SE of Garrison, 8September 2010, RHN & HJN.
European and world occurrence
B. sempervirens is regarded as a member of the Sub-Mediterranean-Sub-Atlantic biogeographicalelement and is considered native in SW & WC Europe and in the mountains of NW Africa (Sell &Murrell 2009). It is widely cultivated and naturalised beyond its native range (T.D. Dines, In: Prestonet al. 2002).
Toxicity
All parts of the plant are EXTREMELY POISONOUS, containing a complex group of steroidal alkaloids. Theplant gives off a disagreeable smell and has a bitter taste, so fortunately animals avoid it. Horses andother large stock animals have died of respiratory failure after browsing it or eating clippings (aslittle as 750 g of consumed foliage is sufficient to kill a horse) (Cooper & Johnson 1998).
Threats
None.
Introduced, archaeophyte, occasional. Eurasian southern-temperate, but naturalised in N America and nowcircumpolar.
1882; Stewart, S.A.; Co Fermanagh.
April to November.
Growth form and preferred habitats
A taller (20-45 cm), hairless and somewhat more robust, weedy spurge than either E. peplus (PettySpurge) or the locally extinct E. exigua (Dwarf Spurge), the oval, yellow-green leaves, that arefinely toothed in their upper part, make Sun Spurge an easy species to identify. Plants on poor drysoils may be very dwarf, but still manage to flower. E. helioscopia frequently has an erect,unbranched stem below, but above it terminates in regular side branches that give the entire plant acandelabra-like appearance (Salisbury 1964).
Like the other two spurges mentioned, Sun Spurge is a quick growing summer-annual, ruderal of openhabitats and appears in and on the margins of cultivated and disturbed ground in gardens and nearhabitation, plus in arable root and leaf crops, waste ground, soil heaps, roadsides, or in gravel orpavement cracks (Grime et al. 1988, 2007). E. helioscopia is also similar to its tworelatives in terms of its preferred ecology, being essentially a plant of dry or well-drained,near-neutral, calcareous or base-rich soils. As its current English common name 'Sun Spurge' suggests,it grows best in sheltered, lowland, sun-warmed sites. The Latin specific epithet 'helioscopia' means'turning towards (or following) the sun'.
Fermanagh occurrence
In Fermanagh, E. helioscopia usually occurs as isolated individuals or in rather smallpopulations, never appearing in any great quantity. It is thinly but widely scattered throughout thelowlands, particularly in the SE of the county where the better agricultural land lies. In terms offrequency, E. helioscopia is very similar to E. peplus, being found in 29 Fermanaghtetrads, 5.5% of those in the VC. Ten tetrads have pre-1975 records only, a fact which RHN and thecurrent author (RSF) believe reflects the casual nature of the occurrences of this weedy species.
Flowering reproduction
Both E. helioscopia and E. peplus flower from May to October or November. The inflorescenceis an umbel, usually with five main rays and yellowish-green bracts similar in shape to the leaves. Thesmall groups of flowers that make up each inflorescence consist of a single female and a few maleflowers, all without perianth parts, held in a cup-like axillary structure called a cyanthium. This has4-5 small teeth alternating with conspicuous green glands at the top. The female flowers have threestyles, each with bifid stigmas. The ovary is 3-celled, with a single ovule in each cell, and it iscarried on a pedicel (stalk) that elongates in fruit. The flowers are pollinated by small flies and theresultant fruit is a 3-valved, globose, smooth capsule, c 2.5 cm in diameter (Sell & Murrell 2009).
The seed number per plant varies from 600-700 (Guyot et al. 1962) and the mean seed number is 257per plant (Pawlowski et al. 1970). The seeds of Sun Spurge are very much larger than those ofE. peplus, weighing on average 0.00248 g, five times the mean weight of the latter, and they areof the order of size approaching that of typical woodland perennials, ie plants of closed, shadedvegetation (Salisbury 1942, pp. 23-4). Compared with smaller, lighter propagules, the large seed ofE. helioscopia may create dispersal difficulties associated with fewer numbers and less readytransport. However, this could be more than offset by greater competitive ability at the extremelycritical stage of seedling establishment.
In common with other spurges, E. helioscopia seeds are released by the sudden splitting of the3-lobed fruit capsule which throws the three contained seeds a short distance from the parent plant. Theseed possesses an elaiosome nutritive oil body (sometimes called a caruncle), that attracts ants, and ithelps scatter the seed, minimising possible seed predation (Salisbury 1964). Again, as with otherruderal spurges, a small proportion of E. helioscopa seed can survive dormant in the soil formany years (more than five) and some estimates reckon this could be for over 20 years (Thompson etal. 1997).
British and Irish status and occurrence
Although Stace in the New Flora of the BI (1997) still regarded E. helioscopia as native inB & I, it is now considered an archaeophyte. Its date of introduction is unknown, but believed to beprior to 1500 AD (Preston et al. 2004). Sun Spurge is widespread and common throughout most oflowland B & I, becoming less frequent and more coastal northwards in Scotland and westwards inIreland. There has been no change in the overall distribution of E. helioscopia during the last60 years, except for a slight decline at the margins of its range (J.H.S. Cox, in: Preston et al.2002).
European and world occurrence
E. helioscopia probably originated in the Mediterranean area and in W and possibly E Asia. It hasspread as a common agricultural weed of root and leaf crops across most parts of the world, including B& I, N Europe, S Africa, N & S America, Hong Kong, Australia and New Zealand (Hultén & Fries1986, Map 1282). It is a more casual weed in the extreme north of Europe and has become circumpolar(Clapham et al. 1987).
Toxicity
All spurge tissues contain a poisonous milky white latex sap that exudes when the plant is cut orcrushed. The toxic principles still require further elucidation, but a resin, an alkaloid (euphorbin,euphorbine or euphorbane), a glycoside, dihydroxycoumarin and other complex compounds such as aditerpene ester called ingenol mebutate have been reported in various Euphorbia species. Thepoisonous action of the toxins is not affected by drying and storage, so that feeding dried fodder cropscontaining spurges could still poison animals. Having said this, very few cases have been reported inBritain (Cooper & Johnson 1998).
E. helioscopia caused severe swelling and inflammation of the mouth, salivation and some diarrhoeain sheep allowed to graze a field of poorly grown kale that was infested with Sun Spurge. The animalsrecovered fully when transferred to good pasture (Cooper & Johnson 1998).
The milky latex is very caustic and quickly burns and irritates skin with which it comes in contact. Evenhandling broken stems (eg after weeding) and then touching the face, especially around the eyes, is avery bad idea, resulting in painful regret.
Uses
The corrosive latex has a universal reputation for removing warts, as has that of the even more commongarden weed, E. peplus, and the much rarer E. hyberna (Irish Spurge) endemic to S & WIreland. In NE Scotland, E. helioscopia was used against ringworm, while in SE England it wasrecommended as a poultice for adder bites and other venomous wounds. An unusual and rather dangerous usein Northumberland was as an infusion of the plant, drunk twice daily, to relieve the pain of chronicrheumatism (Allen & Hatfield 2004), perhaps by fatally poisoning the patient! The internal use ofthe latex has been abandoned by herbalists owing to the severity of its action (Grieve 1931, p. 764).
Names and folklore
Grigson (1987) lists a total of 25 English common names from around B & I for E. helioscopiaand Vickery (2019) tops this with 29 names, five of which refer to the famous wart cure, 'Wart-grass','Wart-gerse', 'Wart-weed', 'Wartwort' and 'Wret-weed' (another form of 'Wart-weed'). Gerard (1597, 1633)is quoted as saying that the juice or milk, "cureth all roughness of the skinne, mangines, leprie,scurffe, and running scabs, and the white scruf of the head. It taketh awaie all maner of wartes, knobs,and the hard callouses of Fistulaes, hot swellings and Carbuncles". An excellent name (highlyappropriate) listed by Vickery (2019) is 'Little good', or variants such as 'Little giddie', 'Littlegweedie', 'Little guid' and 'Littlegude, the Devil'.
Threats
None.
Introduction, archaeophyte, a very rare casual, now locally extinct. European southern-temperate.
1952; MCM & D; waste ground by railway at Belcoo.
Fermanagh and Irish occurrence and decline
There is just one 1952 record for this small, 5-20(-30) cm, erect, ± glaucous, ruderal annual spurge inFermanagh. Meikle and co-workers in their 1975 Revised Typescript Flora commented that it ishardly more than a casual in the VC. Principally a weed of lowland arable farming with fibrous roots anda slender taproot and flowering nearly all year round, it prefers light, dry, base-rich soils and opensunny situations; E. exigua was always quite a rare plant in NI where such specific growingconditions were fairly uncommon (Hutchinson 1972). Dwarf Spurge does not stray far from arable fieldsand only rarely features on other forms of disturbed ground including gardens and bare patches in wasteground. Nowadays, it is all but extinct in NI. The New Atlas map plots a solitary hectad with apost-1987 record in Co Armagh (H37).
Elsewhere in Ireland, according to the 1987 Cen Cat Fl Ir 2, Dwarf Spurge was previously recordedin all but four of the 40 VCs (S Kerry (H1), W Mayo (H27), Sligo (H28) and Fermanagh (H33) and waschiefly found in the east and centre of the island, but with the major decline in arable tillage and theexpansion of intensive farming practices involving changes in crop management, increasing pasturage,reseeding, the use of broad-spectrum herbicides, increased fertilizer application and the development ofcompetitive crop varieties, this weedy little spurge naturally and unnaturally declined everywhere. Inmost of Ireland, the major decline of E. exigua took place around the 1930s and 1940s, thuspreceding this major agricultural shift and intensification, but it is now a rare and definitelydwindling species presence that is categorised as 'Near threatened' in the Ireland Red List No. 10Vascular Plants (Parnell & Curtis 2012; Wyse Jackson et al. 2016).
Good examples of the historic contraction of E. exigua are found in Cos Wicklow (H20) and Dublin(H21): in his Flora of the County Wicklow, Brunker (1950) regarded the species as rare and helisted all known VC records from 1866 to 1932, a total of just eight localities. In adjacent Co Dublin,immediately to the north, Colgan (1904) regarded E. exigua too common a cornfield weed to listlocalities. By 1961, when a Supplement to Colgan's Flora was published (infuriatingly without anindex to either genera or species – but on p. 60), Dwarf or Cornfield Spurge is listed as 'occasional',although only one 1947 station, on the banks of the River Dodder, is actually listed. The understatedaccompanying comment was, "Apparently not so common as in Colgan's time." (Brunker etal. 1961). In comparison, the most recent (1998) Flora of Co Dublin listed a total of sixstations for E. exigua located during the 1984-93 recording period (Doogue et al. 1998).
While the peak level of this casual weed species presence and the timing and rate of decline must havevaried locally to a considerable but now unknown extent, this did not stop Praeger & Megaw (1938)from claiming in the 2nd edition of the Flora of the NE of Ireland that Dwarf Spurge was a,"frequent and locally abundant colonist" among crops on light soils in all three VCs covered(Down (H38), Antrim (H39) and Londonderry (H40)). Fifty years earlier, the first edition of this localFlora by Stewart & Corry (1888) was more circ*mspect in its appraisal, describing E. exiguaas, "rare but locally abundant", and in Co Londonderry (H40) as, "common in cornfields,and in sandy or gravelly land".
Status in Britain and Ireland
While always regarded as introduced or probably introduced in Ireland, E. exigua has previouslyand traditionally been assumed native in most Flora accounts in Britain prior to 2012. Stace (1997), forinstance, was unusual in considering it, "probably native". However, Webb (1985) listed itamong 41 species he believed were probably introduced to Britain by man. Subsequently, Preston etal. (2002 & 2004) carried out the necessary analysis of the available circ*mstantialevidence and they have declared it an archaeophyte, vindicating Webb's opinion.
British occurrence
E. exigua remains fairly common and widespread in SE & C England, but is much less common inthe N & W of Britain and in Wales where it has always been more or less coastal. Losses in the N& W of Britain were occurring before 1930 and, elsewhere, continue into the present day due tointensification of farming (J.H.S. Cox, in: Preston et al. 2002).
European and world occurrence
The species belongs to the European Southern-temperate element and probably originated in theMediterranean region. It remains widespread in W Europe stretching north into mid-Norway (around65oN), N Sweden and S Finland, Atlantic Isles (Azores), N Africa, eastwards to Palestine andN Iran. It was introduced and naturalised in some of these northern areas, probably as an agriculturalseed contaminant and was also introduced into N America, SE Australia and New Zealand (Clapham etal. 1987; Hultén & Fries 1986, Map 1283).
Threats
None.
Introduction, archaeophyte, occasional. European southern-temperate, but widely naturalised and nowcircumpolar.
1882; Stewart, S.A.; Co Fermanagh.
April to December.
Petty Spurge is a little, bushy, ruderal summer-annual, 15-30 cm tall, with a relatively large, diffuseinflorescence of small flowers in umbels, usually with three main branches. The glands in the flowersare crescent-shaped with long, slender points. The fruit capsule is slightly keeled or winged, havingtwo dorsal ridges on each of the three carpel valves (Sell & Murrell 2009; Parnell & Curtis2012). Petty Spurge is a plant of warm, well-drained, fertile, disturbed or cultivated soils, includinggravel, shingle, stony ground and pavement cracks. Although it can survive in full sun if watered orrained upon at intervals, as it is shallow-rooted, plants generally perform best in semi-shaded siteswhere the substrate is less likely to dry out entirely.
Like the smaller E. exigua, whose fruits are smooth and not keeled, it was included in the list of41 species previously assumed to be native in Britain that Webb (1985) considered were probableintroductions by man, and whose traditional status required further study. In response to this clarioncall, the New Atlas editors and Preston et al. (2004) reviewed its status and nowrecognise E. peplus as an ancient or pre-1500 introduction.
In Fermanagh, Petty Spurge is only an occasional, widely scattered lowland species of gardens and urbandisturbed ground. It has been recorded in a total of 29 tetrads, 5.5% of those in the VC. In view of itsdiminutive, useless and weedy nature, this is obviously an accidentally imported species, most likelyarriving from Mediterranean Europe either as a crop seed contaminant, or with soil on the roots of othermore desirable imports, or carried on the feet or on the coats of transported animals, or with othercargo. E. peplus has also been transported very much further afield, to S Africa, Japan, N and SAmerica, Australia and New Zealand, so that nowadays the species has a discontinuous circumpolardistribution in the N Hemisphere and in wider temperate and sub-tropical regions of the world (Hultén& Fries 1986, Map 1284). In areas outside its native range, E. peplus becomes naturalised andis often invasive, although on account of its small size, it is rarely or never a serious weed.
In B & I, the distribution of E. peplus, although widespread and common, has a southern andeastern preponderance, becoming more coastal in the N & W of both islands. This suggests a winterlow-temperature limitation exists, a feature consistent with the supposed Mediterranean centre of originof the species. The B & I distribution appears stable, with no noticeable change in presence duringthe 40 years between the two BSBI Atlases (Perring & Walters 1962; Preston et al. 2002).
Petty Spurge can germinate and grow all year round provided water is available. Plants grow rapidly andreach maturity in 12-14 weeks. Although it normally develops only a small, bushy individual about 20 cmtall, the seed production of E. peplus plants is high with a mean value around 260 per plant.Pollination of the flowers involves small flies, although selfing may also be possible (Fitter 1987).Seeds possess a small attached edible elaiosome oil-body that attracts ants which help them to disperseand avoid seed predation (Ross-Craig, Part XXVI, Plate 41: 1967-70). Since seed can survive for over 20years (perhaps 50-100 years) in the soil, the species is very well protected from local extinction(Thompson et al. 1997). Seedlings appear between May and August and flowering occurs from July toNovember (Salisbury 1964, p. 315).
Toxicity and uses
Like other larger spurges, E. peplus contains milky sap which is toxic, corrosive and can causecontact dermatitis. Nowadays, like other Euphorbia species it is sometimes called 'Cancer Weed'or 'Radium Weed' because components of its sap are being studied for the treatment of skin cancer,leukaemia, warts and sunspots (https://plants.ces.ncsu.edu/plants/euphorbia-peplus/ accessed 23August 2021). E. peplus has a long and wide traditional medicinal use for treating sunburn,corns, waxy growths, asthma, catarrh, reducing blood pressure and as a purgative. If RSF may be alloweda personal comment, "Not such a petty species after all!"
(See also the E. helioscopia account on this website for other spurge herbal medicinal uses.)
Threats
None.
Native, locally frequent. Eurosiberian temperate, but widely naturalised.
1825; Mackay, J.T.; islands in Lough Erne, near Enniskillen.
May to October.
Growth form and preferred habitats
A dioecious, deciduous, rather variable small tree or shrub (usually 5-10 m in height), with branches inopposite or sub-opposite pairs and small greenish flowers in axillary clusters. The prominent,impressed, curved veins on the opposite, finely-toothed, oval leaves of 'Purging Buckthorn', togetherwith the spiny branches and, in autumn, small clusters of shiny black 'berries' on short stalkspersisting after the leaves fall, make this an easy species to identify. It is the only tree (or shrubin wild habitats) in B & I that has both paired buds and thorns. A closer examination showsthat R. cathartica has three sorts of twigs: long shoots that extend growth; short shoots thatbear leaves and flowers but remain short; and shoots modified into sharp woody thorns (Edlin 1964).
R. cathartica demands base-rich or calcareous soils. On alkaline, lime-rich, lakeshore fen peat,R. cathartica is an undershrub of the more open areas of damp to wet, seasonally flooded, woodyfen-carr scrub dominated by ash and/or alder and accompanied by various willows. Young bushes are veryshade tolerant (Godwin 1943), but R. cathartica is not usually a woodland plant in B & I,being much more confined to scrub and hedgerows (Rackham 1980). When it does appear in woodland, it isgenerally in more open areas of Beech and Oak. Buckthorn often associates with Euonymus europaeus(Spindle), Prunus spinosa (Blackthorn) and, occasionally, Ligustrum vulgare (Wild Privet),although the latter is generally planted.
History and status
The fact that prehistoric R. cathartica fossils or sub-fossils (eg charcoal remains) first becomerecognised in the current Flandrian (or Irish, Littletonian) interglacial, during the Atlantic period(7500-5200 BP), and always from archaeological sites, is probably because the species is insectpollinated (see below). As Neolithic farmers are not known to have been actively involved in importingplants apart from cereal crop seeds and root vegetables, the late date of recognised fossil materialdoes not necessarily suggest this woody species was introduced, although this could yet be the case(Godwin 1975; Rackham 1980). All B & I Floras assume R. cathartica is a native species inEngland, Wales and Ireland (eg Sell & Murrell 2009) and, since it is bird dispersed, this appearsreasonable.
Flowering reproduction
Reproduction is entirely by seed and trees and bushes flower early for a woody species, certainly by theage of eleven (Godwin 1943). Tiny male and female flowers (3-7 mm across) appear from May to Juneon separate trees or bushes. The sex ratio is highly skewed towards fruit-bearing female bushes; theratio can be as high as 1:6 or 1:7 (Godwin 1943). Flowers are borne either solitary, or in denseclusters of 2-5 on short shoots. Flower parts are in fours: the yellow-green sepals are the mostconspicuous element, while the greenish-white petals are so tiny (1 mm) they are often consideredabsent. The calyx tube is adnate (fused) to the base of the ovary and its inner surface forms a nectary.Depending on gender, the flowers have either four stamens, or a pistil with a solitary style, a4-branched stigma, and a 4-celled ovary (Sell & Murrell 2009). Sepals and petals are shorter andnarrower in female flowers than in the male (Godwin 1943). The flowers are, as Edlin (1964) comments,"seldom noticed", but nevertheless they are strongly honey-scented and are clearlyinsect-pollinated, mainly by Honey Bees. Although insect visitors are said to be abundant, they arelittle recorded (Godwin 1943; Lang 1987).
The sexual process is efficient as most female bushes fruit heavily from September and the species ismost prominent when it is festooned with shiny ovoid black berries, each c 6 × 8 mm, with a smalldepressed central scar at the apex. Ripe fruits often persist on the tree or bush well into December oreven later. Bushes usually fruit heavily, for instance one bush 2.1 m high at Wicken Fen inCambridgeshire produced 1,455 'berries' (Godwin 1936). Botanically, the fruit is a drupe, since each'berry' contains three or four hard-layered stones or pyrenes (Godwin 1943).
The strong association of R. cathartica with hedgerows very probably reflects the frequency ofbird dispersal of the seed. Blackbirds, other thrushes, Starlings, Robins and Great tit* are the mainfruit consumers and agents of seed transport, while the Bullfinch is a seed predator (Godwin 1943; Snow& Snow 1988, p. 80). The observation that fruits remain available on bushes for several months afterthey ripen suggests birds prefer other items of diet and really only consume Buckthorn drupes when theyrun out of other hedge fruit, or bad weather induces near starvation (Godwin 1943). Seed can survive insoil for at least two years, but germination usually takes place the spring after seed is released(Godwin 1943).
Variation
The species is rather variable and Sell & Murrell (2009) list no less than six varieties recorded inB & I, one of which, var. prostrata Druce, is a prostrate plant! Var. cathartica andvar. ambigua J. Murr appear to be widespread within the range of the species; var.hydrensis (Hacq.) DC. is in scattered localities and var. schroeteri DC is in the north ofEngland. The distribution and ecology of these four varieties is described as, "notunderstood". Var. longipetiolata Grubov is an introduction planted by motorways and probablyelsewhere and seems to have been imported from E Europe and Russia (Sell & Murrell 2009). No hybidsare known (Godwin 1943).
Irish occurrence
R. cathartica has a rather scattered distribution in Ireland. The strongly expressed calcicolousecological requirements result in it being restricted to a wide belt across the central plain where itcan be frequent, but it is absent from southern counties and rather rare or absent in NI. R.cathartica is close to the northern limit of its natural range in these islands and, elsewherein NI, it is a rare, sparse and decreasing species. In fact, beyond Fermanagh it is confined to justthree hectads around Lough Neagh and the adjacent Lough Beg (Flora of Lough Neagh), plus oneisolated hectad in north Co Down (H38) (NI Vascular Plant Database).
Fermanagh occurrence
In Fermanagh, R. cathartica is locally frequent in damp wood copses, scrub and bushy hedgerowsaround both parts of Lough Erne and, as the map indicates, is seldom seen beyond the Lough. There arerecords in the Fermanagh Flora Database from 50 tetrads, representing 9.5% of those in the VC. The twomost isolated sites shown on the tetrad map are Druminiskill Td, 2 km SE of Arney Bridge, where it wasrecorded in July 1992 by an RSPB survey, and in a wood just N of Summerhill Lough, where it was found inAugust 1995 by RHN and HJN.
In addition to its more usual damp to wet ground habitats listed above, Buckthorn is also found inFermanagh to a lesser extent in dry, sunny, sheltered areas of scrub on rock outcrops and in hedgerowsof limestone districts.
Although R. catharctica has declined in other parts of its NI distribution over the lasthalf-century or so (Hackney et al. 1992), it seems perfectly stable in Fermanagh. Buckthorn isthe obligatory larvae food plant of the Brimstone butterfly caterpillar (Gonepteryx rhamni), aconspicuous bright yellow adult butterfly species. The insect was first recorded in Fermanagh in 1918and has been seen sporadically since then. The population's best years in the VC were possibly1983-1985, from which the Brimstone has declined markedly to extinction as a resident species in NI(Nash et al. 2012). The fact that the shrub population of R. catharctica has not declinedin this period in Fermanagh, makes the insect's fluctuation, and the reasons for its apparent localdecline, something of a mystery.
British occurrence
In England, it is frequent to locally common in the Midlands and S & SE wherecalcareous rock outcrops and soils occur. It is absent from Cornwall, N Devon andNorthumberland, rare and not native anywhere in Scotland; it is also absent from most of Wales exceptparts of the S & NW (New Atlas). It has been planted along motorways and elsewhere.
European and world occurrence
Originally restricted mainly to Europe and W Asia, it belongs to the Eurosiberian temperate element.R. cathartica is widespread across most of Europe to 62oN in Norway & Sweden andeastwards across Russia into W Asia, Afghanistan and Turkestan. In S Europe, it stretches to Mid & ESpain and east to Italy, Sicily and Macedonia, but it is absent from the Balearics, Corsica, Sardiniaand Greece. In N Africa, it is present on high mountains in Morocco and Algeria. See also the section onVariation above.
It was introduced to N America and is quite widespread in E and Midland regions (Hultén & Fries 1986,Map 1298).
Names and uses
The purging 'catharsis' in the name derives from the herbal and veterinary use of the pea-sized fleshyfruits. These are very poisonous when both ripe and unripe, so that the juice of ripe drupes (describedas acrid, nauseous and bitter) must be extracted, sweetened, aromatics added (eg aniseed, cinnamon andnutmeg) and the resultant syrup diluted six-fold before its use as a very strong laxative for severeconstipation (Grieve 1931; Lang 1987). Perhaps on account of this powerful medicinal effect and the'cross-like' (crucifix) shape of the twig arrangement, the branches were also wielded as a folk charmagainst witches!
'Buckthorn' was well known to the Anglo-Saxon herbalists and was named in their medicinal writings as'Hart's-thorn'. Both of these names are thought to refer to the wide-spreading branching of the shrubtwigs, that is like a Deer's anthers. Another common name is 'Way-thorn', ie highway-thorn, translatedfrom the German 'wegedorn' (Prior 1879). Other English names include 'Rainberry Thorn' and 'Rhine-berry'– the latter a translation of the German name on account of, "there being much of them alongst theRiver Rhene" (Lyte 1578, quoted by Prior 1879). It is also possible that the derivation of'Rainberry Thorn' comes from 'rain', meaning 'a boundary', the usual place of growth of the shrub (Prior1879; Britten & Holland 1886). Gerard (1597) referred to it as 'Laxative Ram', suggesting the fiercestrength of the drug. It remained official in the British Pharmacopœia until 1867 but was discontinueddue to its severity, being considered more suitable for veterinary practice treating dogs and cows(Grieve 1931; Allen & Hatfield 2004).
Grigson (1955 & 1987) and Grieve (1931) list another use in dyeing. Dried 'berries' provide a rangeof rich but fugitive colours. The fruit used to be sold under the name 'French berries' and importedalong with those of R. infectorius from the Middle-east (Britten & Holland 1886). If gatheredgreen and unripe, the berries give a saffron yellow dye that was fixed with alum and used to tint paperand also leather, especially for making gloves (Grigson 1955 & 1987). Ripe berries mixed withgum-arabic and limewater formed the pigment called 'Sap- or Bladder-green', said to be well known towater-colourists. The bark of the plant also provides a yellow dye (Grieve 1931).
Threats
None.
Introduction, neophyte, previously deliberately planted, now very rare and casual.
21 August 1989; Northridge, R.H.; Farnaght, SE of Tamlaght.
July to October.
One of the three recent sporadic records in Fermanagh of this tall and conspicuous blue-flowered annualwas a plant found growing in a reseeded field. The owner, Mr Carrothers, informed RHN that his fatherhad grown flax for fibre in the field during the World War I over 60 years previously. However, thecurrent author (RSF) presumes that the plant RHN found was a recently introduced grass-clover seedcontaminant. Experimental work and vast local experience of the plant throughout NI, which was a majorflax fibre growing region in the 19th and early 20th centuries, indicates that L. usitatissimum(a species that probably arose in cultivation from the Mediterranean-Atlantic species, L. bienne(Pale Flax)) is only transient in the soil seed bank surviving for less than one year (Toole & Brown1946; Thompson et al. 1997). RHN and the current author (RSF) very much doubt that anyrecent occurrences of the plant in Ireland are relicts of this earlier period of widespread cultivation,despite speculation to the contrary reported in Cat Alien Pl Ir.
Details of the other two Fermanagh records are as follows: plentiful in disturbed ground in severalfields of a farm E of Melly's Rocks, 14 July 2001, RHN; and one plant at the end of the jetty,Knockninny Quay, Corraslee Point, Upper Lough Erne, 5 October 2002, RHN.
The New Atlas map plots 24 post-1986 hectads in Ireland with records of L. usitatissimumscattered fairly randomly across the island. The Cat Alien Pl Ir lists ten Irish VCs with recentrecords, not including Fermanagh. It appears that white-flowered forms frequently occur in bird-seedmixtures and several other Linum species may also have been introduced in this way (Clement &Foster 1994).
At an agricultural research station near Belfast in the 1980s, there was a failed attempt to develop achemical means of 'scutching' flax plants to extract the fibre. The project may have grown somevarieties for a time, although the small surviving local Irish linen industry imports all of its flaxfrom Egypt. The rare and sporadic occurrences of L. usitatissimum in disturbed ground inFermanagh and elsewhere around Ireland are most probably derived either as seed contaminants ofcommercial grass and clover mixtures, or are dispersed from wild bird seed-mixtures on bird-tables,carried on the wind or the wing.
Short, branched cultivated varieties grown for linseed oil were reintroduced to Britain, mainly in thesouth of England, during the early 1990s (Clement & Foster 1994), but neither RHN nor RSF on theirtravels have yet seen these grown anywhere in Ireland. The resultant explosion of recent records of theplant throughout S England is very apparent in the New Atlas hectad map but, as G.T.D. Wilmorecomments, these escapes from cultivation usually represent only transient casuals (G.T.D. Wilmore, in:Preston et al. 2002).
Threats
None.
Native, common and widespread. European temperate, but also in N America.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
The unbranched, slender, wiry stems, 5-30 cm tall, of this little biennial (or rarely annual) ruderalherb, with its small opposite, sessile, well-spaced, wintergreen leaves (5-12 mm long), remain attachedto its tap-root and shallow fibrous roots all winter, so that the species is identifiable throughout theyear. Although individuals are short-lived, the species is a constant and sometimes abundant plant ofspecies-rich, short turf in unimproved, infertile, limestone grassland growing on sunny, dry orwinter-wet, rocky or stony, calcareous or base-rich soils.
As the words 'purging' and 'catharsis' associated with its English and botanical names suggest, anddespite its small size, L. catharticum contains significant toxins (eg linamarin, a cyanogenicglycoside), so that stock animals know to avoid it (Cooper & Johnson 1998). It therefore toleratesgrazing very well and is prevalent on upland rocky pastures.
Fairy Flax is also found in a range of other habitats including short-sedge dominated flushed ground onmoderately acidic heaths, in winter-wet marshy grasslands and in the drier parts of lime-rich fens,common enough around Upper Lough Erne. Both in the uplands and in the wetter ground, L.catharticum is particularly associated with sites that are either steep, rocky, swampy orotherwise inaccessible to farm machinery. This type of ground means the vegetation cannot be sprayedwith agrochemicals, and where the ground is too rocky, or the soil too shallow, it is also impossible toplough and reseed, so the species manages to avoid major agricultural disruption and can persist.
Variation
Two varieties have been named: the standard form, var. catharticum has solitary stems 5-30 cm,usually branched only in the inflorescence and leaves, oblong, 5-12 mm long; the alternative form, var.dunense (Druce) Druce, has stems only 6 cm tall, densely branched from the base and bears short,2-5 mm, elliptical leaves. Var. dunense is a coastal sand dune form, probably widespread aroundthe shores of B & I, but not yet widely recorded (Sell & Murrell 2009).
Fermanagh occurrence
L. catharticum is frequent to common in Fermanagh, having been recorded in 194 tetrads, 36.7% ofthose in the VC. As the tetrad map shows, the species is widely but very unequally distributed inFermanagh. Fairy Flax is very much more often found on the limestone pastures, rocks, cliffs andquarries lying to the W of Lough Erne, where it may be locally abundant in soils that do not completelydry out in summer.
Ecology, established strategy and phenology
Being a very small biennial plant, often flowering when stems are only a few cm tall and seldom exceeding15 cm in height, L. catharticum must avoid competition in order to survive amongst other plants.It has adopted, therefore, the life strategy of a rather slow-growing, but extremely stress-tolerantruderal (Grime et al. 1988, 2007). Following spring germination, the plant grows in gaps in turfwhere it is supported and protected by taller vegetation. At the end of the growing season itoverwinters as a very small wintergreen individual. In its second spring, the plant pokes up an erectwiry stem through the surrounding vegetation, which then branches and flowers that summer. Like otherbiennials, after flowering the exhausted plant dies.
Flowering reproduction
Small white flowers, 5-6 mm in diameter, are produced from June to September in an open, well branchedcymose inflorescence of c 20 stalked blossoms. Floral parts are in fives. The flowers are pollinated byvarious insects, or may self-pollinate and fruit capsules are formed from July onwards (Grime etal. 1988, 2007; Sell & Murrell 2009).
Kelly (1984) showed that very small, relatively few-flowered and therefore habitually 'depauperate'species, like L. catharticum and other similar species of infertile soils (eg Erophilaverna (Common Whitlowgrass) and Gentianella amarella (Autumn Gentian)) can manage toregulate their seed production in relation to the available resources in a particular season in just thesame way that larger plants, including perennials, do. All the species studied that produced smallplants managed to adjust the number of their flowers that set seed and the number of seeds produced perfruit to the resources available in the particular environment. The more fruits they formed, the greaterthe number of seeds per fruit were produced (Kelly 1984).
The Linum fruit capsule can develop up to ten internal segments, each containing a solitary seed.Seeds are shed from July to October. The seeds are small (0.15-0.19 mg) and they have no structures toenhance dispersal by wind or involving animal transport. However, they can persist and accumulate in thesoil seed bank, since they possess an annual dormancy cycle controlled by soil temperature, plus a lightrequirement for germination. Soil disturbance is therefore required to bring buried seed up to the lightat the soil surface. Thus seeds almost exclusively germinate in the spring after a cold winter breaksboth primary and secondary dormancy (the latter created by high temperatures in early summer). Any formof soil disturbance may assist the germination process in spring or early autumn when soil temperaturesare moderate (Milberg 1994).
Fossil history
Fossil seeds and pollen of L. catharticum have been recorded in the British Pleistocene and, asGodwin (1975) points out, apart from one exceptional occurrence in the late Hoxnian (at Gort inIreland), all the other 24 records come either from glacial stages, or from late Flandrian (IrishLittletonian) sites associated with cultivation. The seed and pollen records from across B & I agreeclosely with one another and indicate, "the plant has a strong preference for fresh soils and openconditions, together with tolerance of climatic conditions of at least the early and late phases ofglacial periods" (Godwin 1975).
British and Irish occurrence
Thanks to its tolerance of a wide range of poor soil habitats, Fairy Flax or Purging Flax is common andvery widespread throughout B & I. Population losses due to intensification of agriculture since 1950are ameliorated to some extent by colonisation of a range of infertile, artificial, open habitats suchas quarries and embankments alongside railways and roadsides (Grime et al. 1988, 2007; G.T.D.Wilmore, in: Preston et al. 2002).
European and world occurrence
L. catharticum is widespread in temperate regions of Europe, stretching northwards toIceland and to 69oN in Fennoscandia. It also reaches eastwards into W Asia. Insouthern parts of Europe, it is mainly confined to the mountains (Hultén & Fries 1986, Map 1277;Sell & Murrell 2009). It is introduced in eastern N America and New Zealand.
Names and uses: 'Fairy Flax', 'Purging Flax', 'Dwarf Flax' and 'Mountain Flax' are just four ofthe numerous English common names applied to Linum catharticum, each telling us somethingabout the species, although not always clearly! It is a small plant (fairies are usually imagined asbeing tiny, if not dwarf) and it regularly grows in mountains and it contains toxic compounds that havestrong purgative and emetic medicinal properties (Grieve 1931; Grigson 1955, 1987; Allen & Hatfield2004). The whole plant, small, wiry and sparsely leaved as it is, was collected in July when in flower,and used fresh or dried (Grieve 1931). In the second and greatly improved edition of Gerard'sHerball (1633), Thomas Johnson described how this plant, also known to him as 'Mil-mountain' or'Mill Mountain' in Hampshire and Somerset, was taken for its somewhat fierce purging effect to cureconstipation and was prepared by bruising it and gently cooking it in a pipkin of white wine. It isbitter tasting and had a powerful purgative action described by Quincy (1718) as, "evacuatingviscid and watery humours from the most remote lodgements"; it was apparently also recommended tocommon people for rheumatism and led Quincy to advise its use, "only for very robust strongconstitutions" (Allen & Hatfield 2004). Grieve (1931) says, "it is generally takencombined with a carminative [ie a drug that relieves flatulence], such as peppermint". Fairy Flaxhas also been prescribed by herbalists for catarrhal infections, liver complaints and jaundice and was,"employed with benefit" (Grieve 1931).
Threats
None.
Native, frequent. European temperate.
1882; Stewart, S.A.; Co Fermanagh.
March to January.
Growth form and preferred habitats
A small perennial with a short, tufted, somewhat woody basal rootstock that sends up many branches, P.vulgaris is a rather variable, low-growing, ± hairless species with stems 7-35 cm, ascending toerect but very often sprawling, branched and leafy. The stalk-less, leathery leaves are all alternate onthe stem, measure up to 35 mm long (the uppermost the largest), lanceolate-elliptic in shape and haveentire margins. The irregular flowers of complex structure are borne in slender racemes, the largest ofwhich contain 10-30 flowers. The flowers also vary greatly in colour, ranging from purplish-blue, lilac,pink to white, but never the clear dark blue that is so typical of P. serpyllifolia (McCallumWebster 1978; Sell & Murrell 2009; Parnell & Curtis 2012).
P. vulgaris is a species of short, dry, sunny, semi-natural, calcareous grassland. Typicalhabitats include pastures, cliffs and rock outcrops (some with a covering of open scrub), plus limestonepavement, stabilized scree, drier parts of lakeshores, alkaline to neutral fens, wayside verges and,much more rarely, on waste ground. The range of places where it can occur suggest it has some degree ofshade tolerance. The requirements of the species are met chiefly on somewhat leached, unproductive,calcareous soils, while in base-rich regions in other parts of B & I the plant tolerates a pH rangebetween 5.0 and 8.0, ie mildly acid to alkaline, avoiding the most acidic soils (Grime et al.1988, 2007).
Flowering reproduction
Although P. vulgaris often produces numerous radiating prostrate branches from a central woodyrootstock, these do not root, so that species reproduction is entirely by seed.
P. vulgaris flowers from May to September, with the main flush occurringin the first three months. The inflorescence is a ± dense, slender, terminal raceme, usually only 10 cmtall, bearing 10-40 bisexual (perfect, hermaphrodite) very irregular (zygomorphic) flowers. Theinflorescence elongates considerably when it reaches the fruiting stage. Floral parts vary in number:three of the five sepals are small and green, while the other two are large and petal-like, forming'wings', although later they may persist around the fruit capsule and turn greenish; petals are three innumber, 4-10 mm long, fused to form a tube with three lobes, of which the middle one is fringed at thetip; stamens are eight in number, the filaments fused below for more than half their length into an opentube adnate to the petals, separate above forming two sets of four, the anthers distinct and opening bypores; the ovary is superior, formed of two united carpels and the curving style is 1.0-1.5 times thelength of the stigma, which is 2-lobed and spoon-shaped. Nectar is secreted.
Pollination
Not only is the flower structure complex, pollination is also a very intricate mechanism. Insects, mainlybees, are attracted to the flowers by the two large, petaloid sepals and they alight on the median,fimbriated petal that presents itself at the front of the flower as a landing platform. Folds on theupper side of this petal enclose the anthers and the spoon-shaped end of the style. Just behind this'spoon' is the sticky, hook-like stigma; the anthers are exactly arranged so that when they dehisce thepollen falls into the terminal spoon. When an insect visitor probes for the nectar secreted at the baseof the flower, it first touches the pollen in the spoon and then the stigma. However, it is not untilthe insect's proboscis has become stickily smeared from touching the stigma on the way in that any ofthe pollen will adhere to it. This means the pollen is only collected as the insect leaves the flowerafter receiving the nectar reward, this arrangement thus definitely favouring cross-pollination (Hickey& King 1981).
Self-pollination can also occur, however, at the early stages of anthesis (flowering), when the amount ofpollen shed into the spoon of the style is so large that some of it may get pushed back onto the stickystigma as an insect’s proboscis enters the flower. It may also take place in the absence of insectvisitors as a fail-safe mechanism, as the stigma is able to bend down and touch against the pollen lyingin the spoon (Hutchinson 1945, 1972; Hickey & King 1981).
Fruit and seed
The fruit capsule is 4.0-8.5 mm, ovate and laterally compressed and narrower than the inner sepals whichpartly conceal it. When ripe, the capsule bursts along its margins to release the two contained 3.0 ×1.0 mm, downy, ovoid seeds (Melderis & Bangerter 1955; Butcher 1961; Sell & Murrell 2009;Parnell & Curtis 2012). Each seed bears a relatively large, 2-lobed elaiosome oil food body (orcaruncle), that attracts ants that help disperse them locally, thus minimising seed predation (Hickey& King 1981). The survey of NW Europe soil seed banks failed to discover seed persisting more than asingle year (Thompson et al. 1997). Seed germinates in the spring (Grime et al. 1988,2007).
Variation
P. vulgaris is the most common and ecologically wide-ranging species of the genus Polygalain Europe and, genetically, it is very variable. Five varieties of P. vulgaris are recognised bySell & Murrell (2009) in their critical Flora of B & I: var. caespitosa Pers. is agrassland plant; var. intermedia Chodat is also a plant of grassland as well as of coastalsituations; var. dunensis (Dumort.) Buchenau is a coastal sand-dune plant; var. balliiNyman ex A. Benn. is a rare endemic form known only from the limestones of Ben Bulbin, Co Sligo (H28);and var. vulgaris is the most widespread variant on grassland, fens and river-banks, often, butnot always, on chalk or limestone. All five of these B & I varieties belong to subsp.vulgaris (Sell & Murrell 2009).
Hybrids
Sterile, intermediate hybrids are known to occur extremely rarely in the genus Polygala and onlythree botanists have contributed all of the British records. There are just two hybrids featured inStace et al. (2015): P. vulgaris × P. calcarata F.W. Schultz,for which twelve hectads in SE England have records in sites shared by both parent species, and evenrarer, P. vulgaris × P. amarella Crantz, where there is a solitaryBritish record from near Wye in E Kent (VC 15). No Polygala hybrid has ever been recordedanywhere in Ireland.
Fermanagh occurrence
In Fermanagh, this is a frequent and locally common, but never abundant, low-growing, often sprawlingperennial of short, dry, calcareous grassland. P. vulgaris is recorded in 112 Fermanagh tetrads,representing 21.2% of those in the VC.
The very specific soil requirements and tolerances of P. vulgaris, together with the fact thatall the leaves on the plant are alternate, including those on the lowermost portion of the stem,permits the safe distinction of this species from the closely related P. serpyllifolia (HeathMilkwort). Having said that, one other habitat situation where Common Milkwort can occur very locally ison wet acidic heaths where base-rich ground water springs flush the surface. This situation arisesrather frequently in one particular area of Fermanagh – that around Monawilkin and the Correl Glen NR onthe Western Plateau. In this rather unusual situation, the otherwise quite definite ecological barrierbetween the two most common Polygala species becomes lowered and here they may grow adjacent toone another.
British and Irish occurrence
Throughout the whole of B & I, P. vulgaris is still a locally frequent and widespread species,although it has decreased as a result of habitat losses during the last half century or more. It reallycan no longer be described as 'common' in B & I in comparison with previous years (M.J.Y. Foley, in:Preston et al. 2002). Many of the old, traditionally managed, relatively undisturbed, infertile,low-productivity pastures which provided a suitable habitat niche for this and other species of lowcompetitive ability have been destroyed by post-1945 intensification of agriculture involvingbroad-spectrum herbicides and fertilizer sprays, or 'field improvements' that require drainage andploughing of pastures and meadows, followed by reseeding. The several previously common forms ofspecies-rich, calcareous grassland vegetation that supported P. vulgaris now only persist whereground is inaccessible for modern agricultural machinery which itself has greatly grown in scale inrecent decades as manufacturers have produced larger, heavier and more powerful tractors. Examples ofspecies-rich calcareous grassland vegetation are now rare and are confined to lake islands, or to smallparcels of ground hemmed in by cliffs or water – only protected by being either too small, awkward orhazardous, or just too uneconomic for the farmer to work with.
Consequently, as with Linum catarcticum (Fairy Flax or Purging Flax) and other ecologicallysimilar stress-tolerant species (Grime et al. 1988, 2007), P. vulgaris has suffered adecline in the available area of suitable habitat. Populations have been pushed to survive in rocky,steep, shallow or otherwise too difficult terrain for the farmer to manage. Alternative sites for thesespecies also exists in artificial habitats such as derelict quarries and any neglected, under-managedground that is subject to sufficient disturbance or other growth-limiting factors that reducecompetition from taller, more vigorous species. Locally these less than ideal habitats include urban andvillage waste ground, churchyards and wayside verges of suitable base-status.
European and world occurrence
In phytogeographical terms, P. vulgaris belongs to the European temperate elementand is widespread across most of Europe and also in W Asia and N Africa. The species map inHultén & Fries (1986), Map 1288, shows it well represented throughout the Iberian & Italianpeninsulas and eastwards into Greece. It is also present in Corsica, Sardinia and Sicily, but not in anyother Mediterranean islands, although it is shown as occurring in the Azores.
Names and uses
The Latin genus name 'Polygala' is the name of an unknown plant taken to be the 'polygala' of Pliny andthe 'polugalon' of the Roman medic Dioscorides from a combination of the Greek 'polus' and 'gala',meaning 'much milk', so called because it was supposed to increase the secretion of milk(Gilbert-Carter 1964). There is considerable doubt surrounding the application of the name to plants ofthe Polygala genus we know today, and whether infusion of the plant that improved milk flow wasfor human mothers or for cows in the field, since apparently Dioscorides did not make this clear.Herbalists took it to be for nursing mothers and prescribed it accordingly (Grigson 1955 & 1987).Allen & Hatfield (2004) cast considerable doubt on the whole hopeful identification of the ClassicalPolygala as this genus and any recognition of its milk inducing benefits, regarding the folklorecredentials as highly questionable.
Another English common name for Polygala spp. from Waterford in S Ireland is 'Four sisters', anallusion to the four colours – white, pink, blue and purple – of flowers on different plants (Vickery2019).
Eleven alternative English plant names, including 'Cross-flower', 'Rogation-flower', 'Procession-flower'and 'Gang-flower', are listed by Grigson (1955 & 1987), mostly so-called book names, associated withRogation week processions that were once a regular feature of Christian church worship, when the plantwas picked, made into garlands and carried around the parish when the bounds were beaten and cropsblessed. In the procession, the parish Cross was carried and bells were rung. In England, the Mayblossoming of the plant must have usually been a little late for Rogation.
In Wales, P. vulgaris was named 'Llysiau Crist' ('Christ's herb'), no doubt from the samecontinental tradition of the Milkwort garlands and the associated junketing before Ascension Day. In CoDonegal, the plant was called 'Fairy Soap' from a local belief that fairies make a lather from the rootand leaves (Grigson 1955, 1987).
On Guernsey, P. vulgaris was known as 'Herbe de paralysie' and was allegedly used to prevent orcure paralysis or stroke. The same local name was also applied to Potentilla erecta (Tormentil)(Vickery 1985).
Threats
The expansion of agricultural set-aside and other environment-friendly government funded farm subsidyschemes may allow species such as this to recolonize lost ground. Post-Brexit it remains to be seen whatkind of farm support package(s) will be available and whether it will enable plant species like this oneto retain and regain ground in suitable habitat.
Native, frequent. Suboceanic temperate, but also present in Greenland.
1881-2; Barrington, R.M.; Co Fermanagh.
April to January.
Growth form and preferred habitats
This is a variable, low-growing, 6-25 cm, decumbent to ascending, slender stemmed, few-flowered (3-10),perennial of wet to dry, acidic, pastures and often more peaty habitats in upland grassland, heaths,bogs and moorland. Its predominant occurrence on higher altitude, acidic habitats compared to thosefavoured by the similar P. vulgaris (Common Milkwort), together with its lack of a woody base andthe opposite or slightly sub-opposite leaves on the lower stem of this plant, readily allow P.serpyllifolia to be distinguished from P. vulgaris which has all of its leavesalternate (Sell & Murrell 2009). However, since quite often a number of the significant lower leavesdrop off early in the growing season, in order to absolutely confirm the identification of this species,it then becomes necessary to examine the relative positions of the leaf-scars on the nodes, or toexamine other more critical characters (see R. FitzGerald et al., in: Rich & Jermy 1998).
Apart from the distinctions regarding lime-tolerance (calcicole) or lime-avoidance (calcifuge) and apreference for base-poor, acid conditions in the case of P. serpyllifolia, the ecology andbiology of these two Milkwort species is really remarkably similar. Both species frequent andcharacterise the species-rich vegetation of old, grazed, but otherwise relatively undisturbed,nutrient-impoverished grasslands. Both Milkwort species are stress-tolerant competition avoiders.Conducive circ*mstances are more often located on higher ground in the case of P. serpyllifolia,for instance on sheep-grazed, hilly moors, heaths and blanket bogland, but it can also occur ondamp hummocks in lowland raised and valley bogs, so it is not confined to the uplands.
Since P. serpyllifolia is a mainly upland species, reaching almost to the summits of the highesthills, its habitats are less severely affected by agricultural management grassland 'improvements' thanthose of P. vulgaris (see RSF's species account on this website).
Flowering reproduction
Both these Milkwort species demonstrate a similar complete dependence on seed for their reproduction.They flower from May to August or September and produce a continuous spectrum of flower colour rangingfrom the typical and most common deep purple-blue, through pale blue to almost white, to pink and astrong purple. Heath Milkwort regularly demonstrates this colour range on a single hill slope, with thefrequency emphasis on the deeper colours rather than among the paler hues. The raceme inflorescence ofP. serpyllifolia has significantly fewer flowers (3-10) than that of P. vulgaris (10-40).The individual flowers are also slightly shorter, the petaloid sepals (wings) being 4.5-5.5 mm long,whereas those of P. vulgaris measure 4.0-8.5 mm. The flowers of both species are cross-pollinatedby bees rewarded by nectar, but they may also be selfed (Hickey & King 1981) (See RSF's P.vulgaris account on this website for a fuller description of Polygala pollination).
Seed production is not sizeable in both Milkworts, each flower producing just two relatively large seeds.The fact that P. serpyllifolia has many fewer flowers than P. vulgaris normally does,obviously affects its relative level of seed production. The fruit capsule splits to release the seeds,each of which possess a nutritive oil body or elaiosome attractive to ants, which help to disperse themlocally, thus minimising seed predation (Ridley 1930). There does not appear to be any mechanism of longdistance dispersal, but since these two Polygala species are widespread, it must remain to bediscovered. Seed of P. serpyllifolia can persist in the soil for more than a year and perhaps forup to five (Thompson et al. 1997).
While these two Polygala species can very occasionally occur near one another on flushed heathsand moors (see RSF's P. vulgaris species account on this website), no hybridizationresults (Stace 1975; Stace et al. 2015).
Variation
In B & I, three varieties are recognised and named in their critical Flora by Sell &Murrell (2009): var. vincoides (Chidat ex Davy) P.D. Sell, with leaves on the upper stem ±opposite, elliptical and shortly pointed; var. serpyllifolia, upper stem leaves obviouslyalternate, lanceolate, rounded to pointed; and var. decora C.E. Salmon, with upper leavesobviously alternate, oblong-elliptical, pointed at apex and flowers usually larger than the other twovarieties.
Fermanagh occurrence
In Fermanagh, these two milkwort species are almost equally common in terms of both numbers of recordsand tetrads. Heath Milkwort has just a slight edge on the lime-tolerant P. vulgaris with recordsin 133 tetrads (25.2% of the total), compared with 112 tetrads for P. vulgaris. However, incommon with other local Flora writers, RHN and the current author (RSF) believe P. vulgaris isprobably somewhat over-recorded, some field workers having missed the opposite basal leaves whichdistinguish P. serpyllifolia.
There are large areas of strongly acidic, wet, peaty and podsolized ground in Fermanagh, especially onhigher ground, but also in specific areas of bogland and, more generally, anywhere a layer of glacialclays impedes drainage. Even within the limestone areas in the VC, leached peaty soils sufficientlyacidic to exclude P. vulgaris do occur. This situation led RHN and the current author (RSF) toexpect Heath Milkwort to be the commoner of the two species by a greater margin than the existingFermanagh data demonstrates.
British and Irish occurrence
The New Atlas map shows that P. serpyllifolia has a definite, quite marked, northern andwestern tendency in its B & I distribution, somewhat less pronounced in Ireland than in Britain, butpresent nevertheless. There has been a substantial loss of suitable habitat in S England and inspectionof the record database shows this to have occurred from 1950 onwards (M.J.Y. Foley, in: Preston etal. 2002). Of the three varieties mentioned above, var. serpyllifolia is the widespreadplant, var. vincoides occurs in Cornwall (VCs 1 & 2) and Staffordshire (VC 39), and var.decora is recorded from mountainous areas of all three countries of Britain, plus Ireland (Sell& Murrell 2009).
European and world occurrence
P. serpyllifolia belongs to the Suboceanic temperate phytogeographic element and is mainlyconfined to W Europe, although also present in C Europe to a much lesser extent. It is poorlyrepresented in the Mediterranean basin but is present in Corsica. It is reported from SE Greenland(Böcher et al. 1968) and the Faeroes, where it is said to be, "common, especially on thelowlands [ie heaths]" (Ostenfeld & Gröntved 1934), although it is completely absent fromIceland (Hultén & Fries 1986, Map 1289). It is considered native on Greenland by Hultén (1958) and,therefore (if this position is agreed), it belongs to the Amphi-Atlantic plants that bridge the AtlanticOcean.
Threats
Since P. serpyllifolia is a mainly upland species, reaching almost to the summits of the highesthills, its habitats are less severely affected by agricultural management grassland 'improvements' thanthose of P. vulgaris.
Introduction, neophyte, deliberately planted, frequent.
1976; Dawson, Miss N.; Necarne Forest.
Throughout the year.
Growth form and Fermanagh occurrence
As in most lowland parts of these islands, in Fermanagh Horse-chestnuts, 25(-40)m tall, deciduous,broad-crowned specimen trees, are very commonly planted in town and country parks, landed estates,larger gardens, school playgrounds and along roadside hedgerows. From these sites the tree readily seedsitself into adjacent areas, including the shade of semi-native deciduous woodland and scrub, or in moreopen conditions on waste ground or rough grassland. The Fermanagh flora survey found that the tree iswidespread in lowland areas that are neither permanently too wet nor subject to seasonal flooding and ithas been recorded across 156 tetrads, 29.6% of those in the VC total.
British and Irish occurrence, status and introduction
Despite its undoubted widespread, common and familiar nature and presence throughout lowland B & I,apart that is from peaty areas in the far north of Scotland, the Scottish isles and down the W coast ofIreland, where it is either rare or absent (New Atlas), Horse-chestnut is an introduced alien andit is rarely completely naturalised anywhere on these islands. Its present distribution and speciespopulation size definitely appears to rely on man's assistance – both active and passive. Furtherevidence of the alien nature of the tree is provided by the fact that in Britain it supports a total ofjust nine species of leaf-eating insects and mites, compared, for instance to the 98 supported by thenative Beech (fa*gus sylvatica) (Kennedy & Southwood 1984).
In the wild, as a native tree, A. hippocastanum is confined to three small mountain populations onthe converging political boundaries of Greece, the former Yugoslavia and Albania. Botanists did notdiscover it until 1557, when a Flemish physician in Constantinople sent a specimen to the Italianherbalist Pietro Mattioli, indicating at the same time its use by Turks as a medicine for horses (Nelson& Walsh 1993, p. 131). Another, differing account, has it that seeds were sent from Constantinopleto the Dutch botanist Charles de L'Ecluse (Clusius) who was working in Vienna, and he forwarded samplesto England and to France early in the 17th century (Sell & Murrell 2009).
Irrespective of who first received Horse Chestnut seed in W Europe, samples were first dispatched toEngland around 1612 or 1615 and, despite its southern origin, the species has proved sufficientlyvariable and perfectly adaptable to the range of environmental conditions throughout these islands thatspecimens very often grow to large proportions and seed themselves abundantly.
Fuelled by the unusually large starting capital of the very substantial seed, A. hippocastanumsaplings grow extremely rapidly in their first few decades. They first fruit when around 20 years oldand reach their optimum seed-bearing stage by the time they are only 30 (Thomas 2000, p. 183). The seedmust germinate soon after it is produced, as it is non-persistent, quickly loses viability and rots indamp conditions either on or under the soil surface.
Toxicity
The glossy, mahogany-coloured chestnut seed, like the bark, young leaves and flowers, is very bitter anddistasteful to horses and other stock animals, although starving beasts obviously will eat anything.Cattle are reported to have been killed in the USA by a related Aesculus species, while dogs havealso suffered and sometimes died from eating Horse-chestnuts. The poisonous principle is generallyagreed to be a saponic glycoside named esculin (or aesculin), although it has been suggested that thetree may also contain alkaloids (Cooper & Johnson 1998). Soaking Horse-chestnuts in lime-water, orsoaking and then boiling and grinding them to flour, makes them more palatable (and safer) fodder.
Seed dispersal
Since the familiar fruit, the 'conker', is relatively large and decidedly bitter and toxic, it is noteasy to explain how the tree achieves natural dispersal. Ridley (1930, p.374) mentions rats nibbling andtransporting chestnuts without killing them and rooks have at least once been observed taking andburying them, as also have squirrels in London, although they were the introduced American Grey Squirrel(Ridley 1930, p. 451; Nelson & Walsh 1993, p. 133).
A very quirky correspondence began in BSBI News in 1989 and continued over four issues of thenewsletter, being triggered by a query from a schoolgirl to Mary Briggs, the late secretary of theBotanical Society of the British Isles (Briggs 1989). The original topic was conker dispersal, whichgave rise to a number of imaginative suggested vectors, including extinct dwarf elephants andhippopotami – featuring rapid passage of large quantities of vegetable matter through the gut – thusallowing the possibility of some chestnuts surviving an incomplete digestion process. Alternativetransport mechanisms considered were wild boar, bears and mountain torrents (Akeroyd 1990a, b; Whitehead1990).
An unusual use for conkers
Additional chestnut topics, aired in BSBI News during late 1989-1990, included the widespreadharvesting of the fruit in Britain during both World Wars, when Chaim Weizmann, a Jewish chemistryprofessor, discovered that the solvent acetone required for the production of cordite (the smokelesspowder propellant for bullets and shells, superior to traditional gunpowder), could be manufactured bybacterial fermentation of starch (Ounsted 1990; Stearn 1990; Akeroyd 1990b).
Maize transported from America was used at first in place of the original source, wood, but when theGerman blockade of the Atlantic convoys threatened the maize supply, an appeal went out forHorse-chestnuts, which were collected across the country by school children. While a small quantitywere, indeed, used by the acetone factory, it transpired that the vast bulk of the harvested conkersrotted away in railway sidings across the country (see the related Internet pages of the-tree.org.uk andthe Imperial War Museum (http://collections.iwm.org.uk/server/show/ConWebDoc.1267),accessed 2004.
Timber and tree longevity
Horse-chestnut timber is soft, weak and fractures smoothly, so that large branches have a tendency tobreak off when deluged with heavy rain during thunderstorms. This makes it unsafe to shelter under thetree during such weather conditions (Mitchell 1996, p. 193). Largely on account of the weak timber, itis difficult or impossible to estimate the age of large, old trees, but many specimens in parts ofEngland at least were reckoned by the famous British tree expert, the late Alan Mitchell, to be around300 years, and he gives examples of some of these in his posthumously published book Alan Mitchell'sTrees of Britain (Mitchell 1996).
Names
The genus name 'Aesculus' was a Latin name originally given to a variety of Oak (possibly even Quercuspetraea), but it was applied by Carl Linnaeus to the Horse-chestnut instead. The Latinizedspecific epithet is a combination of two Greek words 'hippo' meaning 'horse' and 'kastanos' meaning'chestnut'. The specific epithet (and by translation, the English common name) was coined by the Frenchbotanist, Tournefort, possibly to suggest chestnuts unfit for human consumption (Gilbert-Carter 1964),the fruits of the tree closely resembling, but only in appearance, the edible fruits of the SweetChestnut, Castanea sativa. 'Horse' and 'Dog' applied as elements to common names are usuallyintended to be derogatory.
Apart from this, the tree has two other horse connections: according to early medical lore,Horse-chestnuts were fed by Turks to their horses as a stimulant and to make their coats shine; and thelarge shield-shaped leaf-scars on the twigs are curved rather like a horseshoe, the vascular tracesrepresenting the nail holes!
Threats
Being an introduced species, A. hippocastanum does not qualify in terms of conservation threatassessment, but since it is so widely planted, doubtless most people would nevertheless greatly regretany dwindling of its development. Horse-chestnuts are prone to several pest and pathogen problems,undoubtedly intensified by the species limited genetic base (the founder effect). At present, thespecies is under severe, potentially lethal attack by the combination of a leaf mining micro-moth(Cameraria ohridella) and a bacterium (Pseudomonas syringae pathovar aesculi), thattogether cause defoliation and an ugly bleeding canker on the trunk and major branches. This attack onthe species is spreading rapidly across both B & I and is all too easily observed now in NI. Whetherit kills the tree or not remains to be seen.
Introduction, neophyte, deliberately planted, occasional, under-recorded. Eurosiberian temperate, butabsent as a native from much of W Europe.
October 1998; Northridge, R.H.; Castle Coole NT estate.
Growth form and preferred growing conditions
Cultivars of this familiar, variable, large-leaved, deciduous maple are commonly planted in largergardens, estates, amenity landscaping and along roadside embankments. It tolerates a wide range of soilsand planting situations and recommends itself to tree-planters of civil amenity and a private nature onaccount of its healthy, vigorous growth and two annual seasons of prominent beauty. It does, however,also seed itself rather freely irrespective of soil and site and can, therefore, spread and naturaliseitself sometimes more widely than is welcome in rough grass, scrub and neglected ground near where it isplanted (Mitchell 1996).
In terms of beauty, the tree blossoms in early spring (March-April), usually before mid-April, producingbright acid-yellow flowers, 8-10 mm in diameter, in 30-40 flowered, erect terminal corymbs that openbefore the leaves unfurl, and last until the leaves come out (Sell & Murrell 2009); in the autumn(mid-October) the leaves turn a bright butter-yellow, then turning orange-brown if they persist. In sometrees, the outer leaves turn scarlet, making a real statement in the landscape (Mitchell 1996).
Species introduction
A widespread native of Europe from Norway to the Crimea, A. platanoides was introduced to Britainsome time before 1683, when the first unambiguous written reference to it was made by George Sutherlandin a list of trees that he moved from the botanic garden at Holyrood Palace to a new garden elsewhere inEdinburgh. Nowadays, there are no trees of known planting date much over 100 years old, although severalmuch larger trees are known than the recorded specimens and will be considerably older. However, A.platanoides does not seem to be a very long-lived species and probably no individual tree datesback to before 1800 (Mitchell 1996). It has been known 'in the wild' in Britain since at least 1905(T.D. Dines, in: Preston et al. 2002).
Variation
A great range of cultivars exist, varying for instance in tree habit, size, number of leaf lobes, leafcolour, surface texture, degree of dissection and leaf margin (eg entire to revolute or crisped).Griffiths (1994) lists no less than 33 named cultivars in the Royal Horticultural Society Index ofGarden Plants.
Fermanagh occurrence
There are only four records of this tree present in the Fermanagh Flora Database, all made by RHN.Details of the other records are: Riversdale Forest, on the banks of the Ballinamallard River near LowerLough Erne, July 2000; planted widely apart near the old castle ruin, Castle Caldwell, 11 November 2006;and Derrychara Playing Fields, planted between pitches and the lake, 16 May 2008. None of these stationscould be assumed to contain naturalised specimens, but rather all the trees have been deliberatelyplanted.
This is another example of a species almost entirely ignored in the Fermanagh survey carried outpost-1975 by RHN and the current author (RSF), essentially because it is a planted introduction.
British and Irish occurrence
In Ireland, A. platanoides plantations occasionally produce seedlings, but self-sown trees rarelyoccur anywhere except in or near planted stock (Cat Alien Pl Ir). The New Atlas mapcontains Irish records of all date classes in a total of just 47 hectads widely scattered across theisland. This contrasts with a total of 1,420 hectads with records in Britain, where A.platanoides is not only more commonly recorded, but is much more evenly distributed fromPlymouth to Inverness.
European and world occurrence
Originally a European and W Asian species, A. platanoides is native from SScandinavia to the Pyrenees and E France, and stretching eastwards to Switzerland, Italy and the Balkansand parts of Turkey as far as S Kazakhstan and the southern shores of the Caspian Sea. Further north, itis native from S Norway into Russia. Strangely, as a native, it is absent from much of W Europe, but iscultivated well beyond its native range, including in B & I, W France and E & C areas of NorthAmerica, where it is a very common street tree in towns and cities (Elias 1980; Hultén & Fries 1986,Map 1291; Sell & Murrell 2009).
Names
Strangely, both the scientific and English common names of this tree are rather inappropriate andunhelpful. The broad native range of the species stretches across Europe from the Caucasus mountainswestwards to the Spanish Pyrenees, but only minimally reaches the SW tip of Norway, yet the tree getscalled 'Norway Maple'. Even more surprisingly, there does not appear to be any alternative Englishcommon name. Likewise, when in 1758 the Swedish botanist, Carl von Linné was looking for a name to giveto this species, the leaf was until then the only one on a Maple that resembled the London Planefoliage. Linnaeus was unaware that the Scots were shortly afterwards to begin to refer to the Sycamore(A. pseudoplatanus) as the 'plane', and the North Americans were to call their EasternPlane tree (Platanus occidentalis L.), American Sycamore (Elias 1980; Mitchell 1996).
Introduction, neophyte, deliberately planted, rare or very occasional, probably somewhat under-recorded.European temperate, but widely planted and naturalised.
1975-85; Faulkner, Dr J.S.; Marble Arch/Cladagh River Glen NR.
April to October.
Growth form and preferred habitats
Field Maple is a modest, small-leaved, deciduous tree with no conspicuous property to make itoutstanding. Even its normal autumn leaf colour is a rather unexciting yellow, although occasionally itcan muster a more impressive bright gold turning orange, or a dull crimson fading towards purple. Itprefers lowland habitats on moist, ± fertile, light, base-rich, chalk or limestone soils, but it canalso occur on wet clay soils. It reaches its best development on the Downs of SE England (Sell &Murrell 2009). Trees can reach 20 m, or a little more in height, but often is very much smaller thanthis, especially when growing in hedges or on poorer soils. When growing in hedges that are regularlycut back, the twigs develop corky wings that are an unusual feature in a maple (Hadfield 1957; Mitchell1996).
In S & E England, it occurs in woodland, scrub and old hedgerows and, in addition, is regularlyplanted in public amenity areas, on farmland as hedges and along roadsides (T.D. Dines, in: Prestonet al. 2002). It is not a gregarious tree and does not colonise waste ground (Hadfield 1957).
Variation
The critical Flora of B & I lists two subspecies, one containing three varieties (Sell & Murrell2009). Subsp. campestre has hairy fruits (samaras), while the recently introduced subsp.leiocarpum (Opiz) Pax has samaras glabrous, or nearly so. With regard to the three varieties ofA. campestre subsp. campestre, only var. campestre iswidespread within the range of the species. Like subsp. leiocarpum, the other two varieties ofsubsp. campestre are recent introductions, probably having arrived in Britain within the last 30years or so (Sell & Murrell 2009). The two introduced varieties of subsp. campestre are var.lobatum Pax which has slightly larger 5-lobed leaves than var. campestre (leaves up to 8cm) and has a central leaf lobe often wider than long, and a more rounded leaf shape than the other twoforms. Var. oxytomum Borbás has the largest leaves of the three varieties (up to 8 × 10 cm), thecentral lobe usually longer than wide and pointed (Sell & Murrell 2009).
Fossil record and status in Britain
A. campestre is the only Maple species considered native in Britain and then mainly in S & Eparts of England and the Welsh borders, although it does also reach northwards to Durham and Cumbria.However, it has been for a long time frequently planted elsewhere to the N & W of England and intoScotland, so its supposed indigenous range has become considerably obscured (T.D. Dines, in: Prestonet al. 2002). Even so, despite its native designation, the fossil record does not provideconvincing proof of its British status. Most of the fossil records of A. campestre are based onwood or charcoal from archaeological sites and the very few fossil pollen records (the tree is insectpollinated, making fossil pollen exceedingly rare) are really only capable of being identified to genuslevel.
The bald fact is that no British Flandrian post-glacial record for A. campestre exists prior tothe Neolithic period (zone VIIb) when the first farmers arrived on the land. Records from the BronzeAge, Iron Age and Roman period are all of similar frequencies to one another (Godwin 1975). A singlepollen diagram from a bog in Somerset (Decoy Pool Wood) is the only one where there is anything like acontinuous Acer pollen curve and there it seems probable that the Maple increased in the earlypart of zone VIII as a consequence of forest clearing or thinning by Neolithic people. The pollen curvesin this particular case for Ilex, Fraxinus and Acer appear to show some tendency(no more than that) to follow the pattern of the weed and cereal pollen curves and this may indicatethat these three woody species were favoured by woodland clearance, opening up the vegetation andallowing them to increase and spread (Godwin 1975, Fig. 168).
In his major work on ancient woodland in S England, Rackham (1980), referring to A. campestre,states his view clearly, "The record for Maple is very unsatisfactory." An explanation for thepoor or under-represented fossil record is offered in terms of geography – Maple is rare or absent in N& W regions of Britain from which much of the pollen evidence comes. In addition, it is pointed outthat Maple is insect-pollinated and is a poor pollen producer, "though rated no worse than Ash orLime". It also grows on clay soils well away from pollen-collecting basins (Andersen 1970; Rackham1980). "Nevertheless, it is remarkable that even in East Anglia, the present stronghold of Maple,it should have so little prehistoric record." (Rackham 1980).
Rackham concludes, "The pollen record of Maple, taken at face value, indicates that the tree hasgreatly increased, or could even have been introduced, since the development of agriculture."Neolithic people are not known to have intentionally introduced plants other than cereals and otheredible species. Perhaps a few medicinal herbs might also have been imported, but no woody plants arebelieved to have been deliberately introduced by early farmers. Rackham goes on to say, "This isdifficult to reconcile with its [Maple's] modern ecology: Maple is a non-aggressive tree which does noteasily form secondary woodland. As a coloniser of clearings in woodland or of abandoned farmland it isless effective than Ash or even Hazel. The question [of native status] must be left unanswered; the lackof evidence tells us less about the prehistory of Maple than about the limitations of palaeobotanicaltechniques." (Rackham 1980).
In phytogeographical terms, Matthews (1955) placed Field Maple in the Continental-southern element of theBritish flora, and the strongly southern restriction of its range in England and Wales agrees well withits recorded late appearance in the fossil (or sub-fossil) record. A. campestre appears to beunaffected by frost in England and it is notably resistant to both shading and coppicing. Nowadays, itcan become a very substantial woodland tree, although far more frequently it occurs as an undershrub orhedgerow plant (Godwin 1975).
Status in Ireland
A. campestre has always been regarded as an introduced, planted species in Ireland (Mackay 1825).
Fermanagh occurrence
Very occasionally, this species is planted in hedges and as specimen trees in demesne parkland or woods,or in landscaping schemes around the grounds of larger public or private buildings, eg the KillyhevlinHotel, just outside Enniskillen. Very rarely, as at Crom Castle, it produces a large tree, but much morefrequently it is only recently planted and is still of shrubby proportions. The tree may eventually seeditself into adjacent ground, but as yet RHN and the current author (RSF) have no record of thishappening anywhere in Fermanagh in a wild, or even in a semi-natural situation.
Elsewhere, in England, when seedlings do occur they are known to be very hardy and tolerant of heavyshade, but under even good conditions they grow slowly. Self-sown seedlings of A. campestre aremuch less common than those of Norway Maple (A. platanoides) and Sycamore (A.pseudoplatanus), though they are more in evidence after seasons of heavy fruiting, which areirregular and sometimes only occur at long intervals (Hadfield 1957).
There are records in the Fermanagh Flora Database from a total of just 17 tetrads (3.2%), so it iscertainly under-recorded. As the tetrad map indicates, it is widely but rather rarely and very sparselyplanted in the lowlands.
European and world occurrence
On the European continent, A. campestre just manages to reach S Sweden (where it is rare) andextends southwards from Denmark, Poland and S Russia to C Spain, Corsica, Sicily and N Greece. It thenextends through Asia Minor and the Caucasus to N Iran and Turkestan. It is also found, though rarely, inAlgeria (Hadfield 1957).
Names
The English common name 'Maple' is derived from Old English 'mapultreow', 'maple-tree'. Place namesincluding this and other early names as an element are found in the midlands and south where the speciesis most commonly found and considered native. Examples are 'Maperton' (Somerset), 'Maplebeck'(Nottinghamshire), 'Maple-durham' (Hampshire and Oxfordshire) and 'Mappowder' (Dorsetshire) (Hadfield1957).
Uses
Nowadays, the wood of A. campestre, being mainly sourced from hedgerows, is usuallysmall-scale and only suitable for minor items. However, in the past, or when larger trees do becomeavailable, the timber is whiter, harder, finer grained, often handsomely figured and far superior tothat of Sycamore or even Beech, belonging indeed to the pre-20thcentury age of craftsmen whoused it for turning dishes, cups and trays. In the past it was also used for inlays and for making harps(Grigson 1955 & 1987). While the wood is hard, tough and strong, it is not durable in the open whensubjected to rain and sun (Hadfield 1957).
Introduction, neophyte, commonly naturalised. European temperate.
1934; Praeger, R.Ll.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Sycamore can form a large, or very large deciduous tree when it is then at its most handsome, although itis very much more frequently of modest proportion or as saplings. Irrespective of size, Sycamore trunksalways stand erect even in windswept situations, perhaps because its young shoots are surprisingly stout(Hadfield 1957). The buds are green and their time of opening varies greatly from tree to tree; theearliest trees break buds around mid-March in NI. In other places, in B & I, buds can break as earlyas January. The familiar palmate 5-7-lobed leaves, 10-18 × 11-25 cm, are very often blotched with blackspots by the fungus Rhytisma acerinia (Pers.) Fr. Autumn leaf colour is a disappointing drab,rather dingy yellow, turning brown (Sell & Murrell 2009).
Despite the disfiguring leaf-spot fungal attacks, Sycamore suffers little harm from pests and diseases,although Grey Squirrels and other rodents may cause extensive damage by peeling the bark (Edlin 1964).
A. pseudoplatanus will grow on all types of soil chemistry, although it performs best on moist,fertile, base-rich substrates. Young plants are extremely hardy and tolerant of most environmentalfactors, except the most acidic and waterlogged soils. However, they prefer woodland conditions, or atleast some shade. It avoids, or does poorly in badly drained sites where there is a degree ofwaterlogging or inundation that could lead to root-rot. This includes wet podsols and heavy clay soilswhere gleying comes close to the surface. Having said this, it occasionally strays onto peat bogsurfaces where there is a little drainage, although it is unlikely to grow to tree dimensions. Sycamoreis also generally absent in overly hot, dry situations where soils are subject to drought. It iscommonly abundant on limestone, although where this is very pure and gives rise to very shallow lightsoils, the tree does poorly, even when planted. Sycamore naturalises itself best where humus decay israpid and nitrification is active (free). Jones (1945) regards it as, "to some extent anitrophilous species", and "its presence usually indicates a fertile soil".
Sycamore trees withstand exposure to wind and to salt spray better than almost any other large deciduousspecies and hence it has been commonly planted around exposed farms in hilly areas and on or near seacoasts either as copses or shelter-belts, which despite wind-pruning, provide excellent weather cover(Jones 1945).
A. pseudoplatanus is often associated with Ash (Fraxinus excelsior), the habitat ranges ofthe two species largely overlapping, although Ash is usually on somewhat heavier, wetter soils thanSycamore, and pure stands of the latter are rare. In Ash or Beech woodlands, Sycamore generally occursas scattered individual trees, rather than in pure stands. It often seeds into gaps in coniferplantations, anywhere there is bare ground and along the margins of such stands (Jones 1945).
In many older woodland stands, A. pseudoplatanus represents an undesirable woody weed as it cangerminate and establish in moderately deep deciduous woodland shade where there is at most a thindiscontinuous carpet of herbaceous species and little litter, with plenty of mineral soil exposed. A.pseudoplatanus can form an under-storey under the canopy of Oak, Ash, Birch or Larch woods, butSycamore's own leaf litter is plentiful, very slow to decompose, and where it accumulates it suppresseswoodland ground and herb flora layers, limiting local species diversity. Again, it is noticed thatSycamore seedlings appear to be somewhat nitrophilous, their distribution often coinciding withMercurialis annua (Annual Mercury), M. perennis (Dog's Mercury), Sambucus nigra(Elder), Silene dioicum (Red Campion) and Urtica dioica (Stinging Nettle) (Jones 1945).
A. pseudoplatanus now occurs in B & I from sea-level to at least the upper limits ofcultivation and habitation. The potential upper limit of the tree is determined more by the presence ofsuitable soil and regeneration conditions rather than by levels of exposure or temperature (Jones 1945).
Flowering reproduction
Flowers open from late April and they continue being produced for at least a month. Yellowish-green, theyare borne in narrow pendulous racemes up to 20 cm long containing 6-100 flowers in stalked clusters 4-6mm in diameter. It has been noted that often up to 60% of flowers in a raceme are male and that maleflowers open first. The proportion of male to hermaphrodite (perfect, bisexual) flowers varies greatlyfrom tree to tree and it is possible that some trees are entirely male, although if this is the case,they are very rare. Trees with few hermaphrodite flowers are not uncommon (Jones 1945).
The flowers are scented and produce nectar, attracting bees (including honey-bees) and flies aspollinators. The samara fruit are formed very quickly, the earliest being of appreciable size beforeflowering on the tree has finished. Trees in open situations can produce fertile seed when only 20 yearsold and good seed crops are produced practically every year (Hadfield 1957). Samara are released fromthe tree gradually, beginning about the last week in October and continuing through into the first fewmonths of the following year. Wind dispersal usually carries the fruit only a short distance (ie up toaround 80 m), but high winds can occasionally disperse it for up to 4 km. Seed does not germinate, evenunder favourable temperature conditions, before the mid-January after production. No hybrids have beenreported (Jones 1945; Stace et al. 2015).
Sycamore readily seeds itself in pre-existing woodlands and in hedgerows. Well-established trees areextremely difficult to extirpate, the root and stump very stubbornly regenerating the plant. Since windtransport of the winged samara fruit is very effective, small seedlings often appear in the spring aslawn weeds, in gravel paths and as pioneer colonists of neglected ground in parks, gardens and roadsideverges. The lawnmower, however, quickly dispatches them. In wilder situations, A. pseudoplatanusis generally kept in check by competition from more vigorous, taller herbaceous species and also by thebrowsing of grazing animals (Jones 1945).
Status and introduction to Britain
This is such a common and often a well-grown, large tree in B & I that many people assume it isnative, while in reality it is certainly a neophyte, ie a post-1500 AD deliberate human introduction.A. pseudoplatanus is a native tree of upland parts of C & S Europe and the oldest trees inthese islands are in Scotland, possibly planted around 1550 AD or a little earlier (Mitchell 1996, p.183; see also Jones 1945, p. 236). Subsequently, the tree became very widely planted all across B &I for its excellent timber and it quickly naturalised itself from the 18th century onwards in older,native or semi-native woods and other more open habitats, especially hedges (Rackham 1980, p. 58). InIreland, there is reliable evidence that Sycamore was being planted near Derry (Londonderry) around1610, during the plantation of Ireland by Scottish settlers in the reign of James I (Nelson &Walsh1993).
The same level of plantation occurring on the continental mainland means it is difficult or nearimpossible to determine its original centre of origin and natural distribution (Hadfield 1957).
Variation
A. pseudoplatanus is sufficiently variable that as many as 34 distinct cultivars are listed in theRoyal Horticultural Society Index of Garden Plants (Griffiths 1994). In the critical Flora of B& I, two varieties are recognised. Var. pseudoplatanus, with samara wings 20-40 mm, containsfour formae : forma pseudoplatanus; forma variegatum (Weston) Rehder; formapurpureum (Loudon) Rehder; and forma erythrocarpum (Carrière) Pax, the latter threecorresponding to cultivars. The other variety is var. macrocarpum Spach, which has samara withlarger wings (50-60 mm) (Sell & Murrell 2009).
Fermanagh occurrence
Sycamore is now an extremely common and widespread self-sown tree in woods and hedges throughoutFermanagh, as is the case almost everywhere in B & I. In Fermanagh, it has been recorded in 403tetrads, 76.3% of those in the VC, making it the 9th most widespread woody plant in the post-1975 surveyby RHN & the current author (RSF), although it still lies well behind Fraxinus excelsior(Ash), Alnus glutinosa (Alder), Ilex aquifolium (Holly) and Corylus avellana(Hazel) in terms of tetrad frequency.
Seedlings or saplings are frequent just about everywhere in Fermanagh, except in the wettest, mostacidic, most exposed or very heavily disturbed situations.
European and world occurrence
Some authorities claim that A. pseudoplatanus reaches SE Denmark and SW Sweden in the north, butothers believe it spreads southwards only from NW France, S Germany and Poland to N Spain, Corsica,Sicily and C Greece and thence into Asia Minor and the Caucasus (Jones 1945). It grows mixed with othertrees, particularly Beech and Ash and is seldom found growing in pure stands (Jones 1945; Hadfield1957).
Uses
Sycamore wood is very pale-brown to yellowish-white in colour, darkening with age. It is fairly hard,moderately heavy and strong, but it is not durable outdoors, being unsuited to prolonged exposure toweather (Edlin 1964). When polished it is one of the most beautiful timbers available to the cabinetmaker and wood turner. Some forms have colour flecks or wavy patterned grain, when it is called'fiddle-back' sycamore. It is also suitable for steam bending and can be used for making musicalinstruments including violins and harps. Sycamore wood has also been used for veneers, carving,furniture making (including for tables). It is smooth enough for making good dance floors and, as it canwithstand wear and scrubbing without the grain picking up, staining or tainting food, it is perfect forthe manufacture of smaller everyday kitchen items such as bowls, plates, ladles, spoons and for rollersfor printing machines and clothes' mangles. It also makes excellent firewood, but its greatest use sofar has been as a shelter tree for buildings (Edlin 1964; Nelson & Walsh 1993; Mitchell 1996; Sell& Murrell 2009).
Names: 'Sycamore' properly is the common name of a species of Fig, Ficus sycomorus L.(Mulberry-leaved Fig), native in W Asia and tropical Africa, but the name 'Sycamore' was more looselyapplied in 14th century England, and somehow here became attached to Acer pseudoplatanus. Gerard(1597) wrongly called it 'Sycomore', but he preferred to refer to it as 'Great Maple' or 'Giant Maple'.To add to the confusion, in N America the name 'American Sycamore' is applied to Platanusoccidentalis L., their species of Eastern Plane and, in Scotland, the tree is often called the'Plane' (Elias 1980; Mitchell 1996). Grigson (1955, 1987) lists twelve additional English common names,several of them linked with children using the twigs to make whistles.
Threats
The presence of Sycamore might well downgrade the conservation status of old woodland. The recentlyarrived, rapidly spreading fungal pathogen, Phytophthora ramosum, reportedly attacks A.pseudoplatanus in southern England.
Introduction, neophyte, invasive garden escape, so far very rare.
August 1997; Northridge, R.H.; Enniskillen Town.
Growth form, variation and Irish occurrence
This yellow-flowered, small, creeping Oxalis sp., 15(-20) cm tall, is so very variable,phenotypically and genetically, that local forms have often been described as new species (Lovett Doustet al. 1985). It has only twice been recorded in Fermanagh, once as var. atropurpurea (seebelow) and once in this guise as the green-leaved form. Almost all forms of this species, however,demonstrate some degree of purplish tinge in the leaflets and, again, this character varies in responseto the prevailing light levels. Most B & I introduced Oxalis species originate in warmerclimes, they are not very competitive and typically they only occupy very open, often disturbedhabitats, sometimes ± confined to greenhouses (Louvett Doust et al. 1985).
The more-or-less green-leaved form of O. corniculata was found naturalised in Enniskillen town asa persistent (perhaps pernicious) garden weed, rather than in a truly 'wild' situation. The Cat AlienPl Ir lists records from seven other Irish VCs. The records from NI (Cos Tyrone (H36), Down(H38) and Antrim (H39)) all report this species as being confined to gardens, greenhouses (where it caneasily run riot and become a real nuisance weed) or nurseries (FNEI 3). The survey of urbanBelfast found it in seven 1-km squares, mainly as a weed of flowerbeds (Urban Flora of Belfast).
The status of introduced plants
It is often difficult to decide whether or not introduced plants like Oxalis in gardens shouldever be considered 'wild', or, if so, in what sense they are wild or feral? The ability of aplanted garden subject to perpetuate itself without the continuing assistance of human maintenance isinsufficient by itself. As D.P. Young (1958, p. 53) pointed out in an article specific to Oxalisgrowing in B & I, the real criterion hinges on the plant's capacity to withstand all attempts toeradicate it!
In areas with mild winters and at least indoors, O. corniculata is a freely seeding, short-livedperennial with a tough taproot. In cooler conditions, it behaves as an annual, overwintering exclusivelyby seed. The plant produces procumbent, creeping stems or stolons that root at the nodes and it canprove invasive and persistent, but it is not quite as pernicious as those species that also producebulbils (Louvett Doust et al. 1985).
Introduction and spread in Britain and Ireland
O. corniculata is believed by Eiten (1959) to have originated somewhere in tropical Asia,Malaysia, Australasia, or in high ground on islands in the W Pacific. In other words, it is an 'OldWorld' species and has been introduced to northern temperate areas of the world in historic times(Louvett Doust et al. 1985). It was introduced to gardens in Britain prior to 1585 when it wasfirst reported as a garden weed in Somerset (Raven 1953; Stace & Crawley 2015). It was firstrecorded beyond the garden confines in 1770.
O. corniculata did not begin to spread much until the 20th century, but since the 1960s has becomeincreasingly invasive; having consolidated in the S & W of England, it is now spreading north and inaddition to gardens, is colonising paths, walls and waste ground (Sell & Murrell 2009). The NewAtlas hectad map shows O. corniculata has reached Inverness, although the occurrence isonly rather scattered north of a line between Hull and Liverpool (M.F. Watson, in: Preston et al.2002).
In the rather warmer climatic conditions of S Ireland (eg Mid-Cork (H4) and Cos Waterford (H6), Dublin(H21) and Louth (H31)), a solitary plant has been found at a seaside bay in Mid-Cork, while the otherVCs reported habitats 'beyond the garden wall' included streamside and roadside banks and on wasteground.
Reproduction and dispersal
The small yellow flowers, petals 4-7 mm, are borne in 1-7 flowered umbels. They successfullyself-pollinate if insect visitors fail to oblige and are completely self-fertile (Louvett Doust etal. 1985). Although the plant can spread vegetatively to a limited, local extent, dispersal ismainly by seed which is explosively shed from the capsules, travelling a distance of approximately 2 mfrom the parent plant. The seeds are also sticky and may be spread further afield by rodents. Since thespecies has shown itself capable of spreading quite rapidly in Britain, and has gradually become apernicious weed, quite often it may also be transported with waste as a garden discard. Thanks to thecharacteristic generosity of gardeners sharing plants with their friends, it very probably is alsoaccidentally passed in pots from garden to garden, as well as through the horticultural trade (Young1958).
Threats
Colonising behaviour, spread and persistence by this small species is not really a problem in Ireland asyet, but it might eventually become one if it follows the long-delayed invasive pattern it has displayedin Britain.
Introduction, neophyte, invasive garden escape, so far very rare.
July 1993; McNeill, I.; roadside waste ground, Ballycassidy.
A rather insignificant-looking, short-lived perennial, often selfing and behaving as a small ruderalannual, this species can regularly produce more than one generation in a season (Young 1958). Despitethe minor scale of the plant, the contrast between the clusters of little yellow flowers and, in thisvariety, the bronze-brown trifoliate leaves, makes some folk regard it as a desirable garden subject,although perhaps it is most suitable for troughs and other entirely confined, miniature gardens. Whileit usually remains a garden, greenhouse and conservatory weed, frequently infesting bare soil around thebase of plants in pots, as listed above O. corniculata var. atropurpurea has escaped intothe wild in Fermanagh at least once. RHN and the current author (RSF) suspect that the species wastransported and dumped among garden waste in roadside waste ground. Fly-tipping of this nature isunfortunately an all too common practice in Fermanagh.
The New Flora of the BI (1997) describes this species as being a pernicious weed of gardens,paths, walls and waste ground, which has it just right! Although the short, wiry procumbent stems rootat their nodes, by far the most effective means of spread is by the copiously produced, lightweightseed, apart that is from deliberate human transport in the back of a vehicle to a quiet, concealed,rural spot, where garden waste is all too often deposited.
Threats
It has the potential to become a common ruderal weed of open habitats and especially of disturbed groundnear habitation.
Introduction, neophyte, garden escape, very rare.
1995; McNeill, I.; waste ground near a house, Moynaghan North Td, south of Drumblane House.
Another small, yellow-flowered annual or short-lived perennial garden escape belonging to this genus,O. exilis is very often mistaken for a tighter, neater, even more prostrate, mat-forming,greener-leaved form of the very variable and more common garden subject, O. corniculata(Procumbent Yellow-sorrel) (Young 1958). O. exilis can be distinguished from O.corniculata by its always solitary flowers having just five fertile stamens and developing asmaller fruit capsule, 5-8 mm, about three times as long as wide and abruptly narrowed at the tip (Reid1975). As a result of the identity confusion between these two Oxalis species, the truedistribution and frequency of O. exilis in B & I is not really known at present. M.F. Watsonhas provided a helpful key to O. corniculata and its allies in the BSBI Plant Crib1998.
There are just two records of O. exilis in Fermanagh, both found by Ian McNeill, who has recordedthe species on seven occasions in Co Tyrone (H36) since 1987 (McNeill 2010). His second find of theplant in Fermanagh was at Irvinestown in July 2009.
In Ireland, O. exilis has appeared almost exclusively as a garden weed, or more rarely a ruderalin disturbed ground, eg around gateways or along roadsides, usually in the vicinity of gardens. TheCat Alien Pl Ir lists rare occurrences from 1958 onwards in four additional Irish VCs (CosWaterford (H6), Wicklow (H20), Down (H38) and Antrim (H39)). In Britain, O. exilis is anestablished garden escape and a persistent weed in gardens, becoming especially frequent in the warmersouth of the country (Clement & Foster 1994; New Atlas).
Like O. corniculata, most populations are probably only casual in both shaded and open, disturbedground situations where they can avoid competition. Occasionally, however, they may be rather morepersistent, surviving for many years (Young 1958). The species is believed to be of Australasian originand it was first recorded in the wild in Britain in 1926 (M.F. Watson, in: Preston et al. 2002).
Threats
Too rare and casual to be a problem in Ireland.
Native, common and widespread throughout. Eurasian boreo-temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This attractive, variable, low-growing, white-flowered, wintergreen, patch-forming, perennial with aslender creeping rhizome occurs in moist, shady situations and, locally, appears even in the smallestfragment of woodland cover. Wood-sorrel has beautiful, light- to dark-yellowish green, trefoil leaves,the colour varying with light levels. The leaflets, borne on long petioles up to 10 cm, fold down twicea day, carrying out 'nastic movements' in response to changing turgor pressure in the hinge-like pulvinicells at the leaf base. The rhizomes of O. acetosella run along the soil surface and the rootsystem penetrates only the top few cm, thus although it can cope with shallow soils, it is very muchrestricted to permanently moist habitats (Packham 1978; Grime et al. 1988, 2007).
In some parts of S Britain, O. acetosella is confined to primary (ie ancient), relativelyundisturbed woodland, or has a strong affinity for old woods, making it a useful indicator of thesesituations deserving conservation action (Rackham 1980, p. 54). This more restricted occurrence isclosely related to species-limiting levels of moisture- stress which the plant experiences in the morecontinental climate of S & SE England in a way that is quite foreign to other wetter, cooler milderareas of B & I like Fermanagh. In SE England, the species lies closer to the southern margins of itsdistribution and, as is also the case elsewhere that enjoys a drier, more continental climate, O.acetosella does not grow well in dry soils (Packham 1978). In Fermanagh and other wetter partsof B & I, numerous other types of permanently damp, shaded situations which to some extent mimicwoodland conditions, allow Wood-sorrel to occur much more widely in non-woodland situations and,occasionally, to absolutely thrive in them. This is especially the case in well-drained, yet moistureretaining organic soils, often with an insulating depth of leaf litter. Being a low-growing herb,however, excessive amounts of leaf litter inhibits the plant's growth. The latter situation confinesO. acetosella to localised sites where less material accumulates, for instance on even slightlyraised spots or mounds on the ground, on slopes and at the base of tree trunks.
Plants cannot tolerate any degree of prolonged trampling, but are little affected by grazing as theycontain toxins that are sharp and distasteful to animals (see below).
The range of Wood-sorrel habitats additional to old, established woodland, include darker, damper areasin plantations, scrub thickets, hedgerows and along river and streamside banks. O. acetosella isone of the very few vascular plants which can penetrate conifer plantations, but even then it is onlycommonly found in marginal situations, or where the evergreen canopy density is partially opened orthinned by some form of damage (Packham 1978; Grime et al. 1988, 2007).
Locally, in Fermanagh, O. acetosella also occurs in damp shade on cliff and quarry ledges andunder and among rocks in block scree or limestone grykes (± deep, shady crevices). It also grows inrough montane grassland and under bracken, gorse and ericaceous heath at relatively high altitudes.
The established growth strategy is described as intermediate between stress-tolerator and stress-tolerantruderal (Grime et al. 1988, 2007). The rather low growth rate and small stature of O.acetosella creates stress for the individual plant when faced with shade and competition forresources from taller, shade-tolerant, woodland floor species.
While it occurs across a wide range of soil reaction and parent materials (pH 3.5 to >8.0), O.acetosella is most frequent in the acid end of the range, below pH 5.0, and it is predominantlyfound on infertile, undisturbed, rather deeply shaded situations, where the ground is permanently dampand never subject to drought (Packham 1978; Grime et al. 1988, 2007).
Toxicity
All Oxalis species contain oxalic acid and oxylates that can cause poisoning. However, it isunlikely that sufficient of the plant would be eaten in B & I to cause actual poisoning. A totaloxylate content of 7% has been estimated in air-dried plants and sheep appear to be most at risk ofpoisoning, as little as 600 g of the plant leading to severe calcium deficiency that can lead to kidneyfailure and death. Stock animals appear aware of the toxins from their unpleasant, lemon-sharp, sour,sorrel-like (Rumex acetosa) taste, or their odour, and they quickly learn to avoid the species(Cooper & Johnson 1998).
Flowering reproduction
Beautiful, solitary, bisexual, pendulous bell-like, white flowers with parts in fives are produced on5-15 cm long peduncles in April and May. On close inspection, the petals are observed to be exquisitelyveined with fine, lilac to purple stripes, making it one of the most delicate and subtly decoratedwidespread, native wildflowers in B & I. Pink, lilac and purple petal colour variants do exist, butthey are very rare and local: the current author (RSF) has never come across them anywhere. The springflowers, which are so welcome every year, appear to attract few insects and they set little seed.However, later in the summer, an abundant crop of petal-less flowers are produced that self-pollinateand self-fertilize while still in the bud (ie they are cleistogamous). These later flowers produce mostof the seed for the season in 3-4 mm ovoid capsules (Clapham et al. 1987). Up to ten seeds areproduced per fruit capsule; their dispersal is explosive, the capsule rupturing and scattering the seedsup to a metre from the parent plant. Wet seeds are sticky and may occasionally enjoy secondary dispersalon leaves or other wind-blown litter (Ridley 1930, p. 22).
The survey of soil seed banks in NW Europe concluded that seeds are transient to short-term persistent inthe soil seed-bank, ie their survival ranges from less than one year up to several years, but less thanfive years (Thompson et al. 1997)
Vegetative reproduction
O. acetosella tends to grow in patches, many of which represent a single clonal mat of growth,achieved by spread and branching of its horizontal surface-creeping rhizomes over the individual'sseveral years lifetime. In darker areas of woodland in particular, clonal patches may measure up to 50cm across, and they can meet and mingle with adjacent clones to create a larger mat or sheet of thespecies, potentially dominating the ground layer. In this way, it can, in some cases, oust otherfamiliar woodland species such as Hyacinthoides non-scripta (Bluebell), by spreading amongst theadult plants and shading out their seedlings (Packham 1978). Reproduction by seed allows invasion offresh habitat, but the extensive vegetative spread shown by Wood-sorrel is important in maintainingestablished communities.
Fermanagh occurrence
As a result of its wide ecological tolerances, in NW Ireland O. acetosella is recorded from noless than 392 Fermanagh tetrads, 74.2% of those in the VC. Locally, in addition to damp, shadysituations in woods, scrub, hedge- and river-banks, cave-mouths, screes, rock ledges in quarries and inlimestone crevices, O. acetosella can occur in upland grassland under bracken, gorse andericaceous heath on boggy peat at relatively high altitudes. This accounts for the fact that it has beenrecorded from all the tetrads which cover the long summit ridge of Cuilcagh, the highest mountain in theVC, although the species does not actually reach the more exposed areas of the ridge itself.
British and Irish occurrence
In Ireland, O. acetosella is widespread and common in most of the country and, while recorded inevery VC (Scannell & Synnott 1987), it is less prevalent and more scattered inConnemara and in the Irish Midlands. In Britain, it is even more widespread, common and occasionallyabundant, except in the drier parts of the country with the lowest rainfall in S & SE of England,and especially around the Wash and the Humber estuary where the mean annual rainfall is less than 635 mm(Packham 1978; Preston et al. 2002).
European and world occurrence
O. acetosella s.l. occurs from Iceland, Faeroes and N Scandinavia, S & SE-wards across most ofEurope to the mountains of C Spain, Italy and Greece. It occurs at over 2000 m in European alpine heathand Krummholz communities just beyond the timberline (Packham 1978). It becomes rarer towards theMediterranean coast and Corsica is the only Mediterranean island it reaches.
O. acetosella is also native in large areas of N & C Asia, although further east there are anumber of closely related taxa that can be confused with it, and a similar situation involving alliedspecies exists in N America. Previously some of these taxa were considered subspecies of O.acetosella and the latter was then reckoned to be a single species, circumpolar in distribution(Hultén & Fries 1986, Map 1260). In mainland Asia, the species is typically subalpine and alpine inhabitat at altitudes from 2130-4110 m (Packham 1978).
O. acetosella is a member of the Eurasian boreo-temperate phytogeographic element (Sell &Murrell 2009).
Uses
Despite their sharp, sorrel-like flavour and oxalate toxin, some people still consider Wood-sorrel adesirable salad vegetable or suitable for addition to cream-cheese sandwiches (Mabey 1996)! The 'SourDocks' or Sorrels cultivated for centuries as pot-herbs were Rumex acetosa (Common Sorrel) andR. scutatus (French Sorrel or Rubble Dock), as well as the smaller R. acetosella (Sheep'sSorrel) and the totally unrelated Oxalis acetosella (Wood Sorrel) on account of the 'gratefulacidity' of their herbage due to the bioxalate of potash, also present in Rhubarb. The foliage of O.acetosella has been eaten as a spring salad "from time immemorial, their sharpness takingthe place of vinegar" (Grieve 1931).
In herbal medicine, O. acetosella was used as a diuretic, an antiscorbutic and for its'refrigerant action'. It was used to treat high fever, to quench thirst and allay the fever. As Allen& Hatfield (2004) point out, the medicinal benefits of this species have been often confused withthose of Rumex acetosa, due to the similarity between their vernacular names, and therefore caremust be taken in applying their use. There are always suspicions voiced whenever a medicinal use isconfined to a particular region of B & I, for instance the application of O. acetosella forhealing bruises only in Devon, or for making a poultice for scrofula in Colonsay in the Inner Hebrides.In Ireland, Wood-sorrel has been used to treat diarrhoea in Cos Mayo and Wicklow, as a blood tonic inCavan, as a heart tonic in Wicklow and to treat palsy in Limerick. It was also prescribed for stomachcancer in an unknown part of Ulster (Allen & Hatfield 2004).
While used by herbalists to treat haemorrhages and urinary disorders, on account of its oxalic saltscontent, Grieve (1931) cautions against its general use for all constitutions, "especially those ofa gouty and rheumatic tendency".
Names
Both botanical names, 'Oxalis' and 'acetosella' refer to the acidity of the foliage, the genus name beingderived from the Greek 'oxys' meaning 'sour' or 'acid' and the Latin specific epithet 'acetosella'meaning 'vinegar salts'.
O. acetosella must be one of the species most frequently given English common names, since Grigson(1987) lists no less than 58 and Vickery (2019) lists almost the same number, although there are verymany minor variants along the same lines. Many of the English names refer to the fact that people(especially children), consume the leaves either as a salad, or as a sharp tasty snack, eg 'Egg andcheese', 'Bread and cheese', or 'Cuckoo bread and cheese', the latter one of several that give anindication of springtime, when young leaves are at their most mild flavour (and oxalate content), andtherefore safest to eat. Warnings are given, however, since too many leaves could lead to poisoning!
Other animals are mentioned, including 'Fox's meat', 'Rabbit meat', 'Hare's meat' and 'Sheep's sorrel'.References to 'lady' or 'woman', such as 'Lady cakes', 'Lady's meat' and 'Woman's nightcap' bringreligion into the subject as they mean Mary or 'Our lady'. Easter is another topic, with names such as'Alleluiah' and 'Hallelujah' referring to the period of the year when the flowers first appear and thecuckoo is about. The word 'sour' is often a name element also, such as 'Sour sab' and 'Sookie Sourach'(Grigson 1987).
Threats
None.
Introduction, neophyte, an extremely rare garden escape.
1946; MCM & D; roadside bank near habitation, Farnaght, SE of Tamlaght.
A garden perennial with attractive, trumpet-shaped, white and strongly net-veined magenta-pink flowersand wintergreen leaves, this species is a native of C & S Italy, Sicily and the S Balkan Peninsula.It was introduced to cultivation in Britain as early as 1629, making it one of the first four hardygeraniums in decorative cultivation in this part of the world (Yeo 1985).
From the early decades of the 19th century onwards it has become a frequent garden escape or discard,widespread but thinly scattered throughout the length and breadth of England and Wales, although itremains extremely rare and sparsely scattered north of a line between Lancaster and Scarborough (NewAtlas). Since it possesses a sturdy compact rhizome or rootstock, it readily establishes itselfand can become thoroughly naturalised and long persistent in places such as grassy banks, hedgerows andsemi-shade in woods (Clement & Foster 1994). The New Atlas map shows that G.versicolor is very much more frequent and persistent in the southern half of Britain and it isespecially frequent in the extreme SW of England.
This species has only been recorded once in Fermanagh and that was as long ago as 1946. While it is knownto persist in other parts of Ireland, RHN and the current author (RSF) have no knowledge of it doing sohere. It first appeared in the wild near Lisburn, Co Antrim (H39) in 1837 but, in comparison withBritain, it very much more rarely 'jumps the garden wall' on this side of the Irish Sea. The few Irishrecords that have accumulated show it is largely clustered around Belfast and Dublin, although in factit has been recorded at least once in twelve Irish VCs, ie the eleven listed in Cat Alien Pl Ir,plus Fermanagh.
The Latin specific epithet 'versicolor' is a combination of 'verto' meaning 'change' and 'color' meaning'colour' and translates either as, 'changing colour' (Gilbert-Carter 1964), or 'variously coloured'(Stearn 1992). This name probably refers to the fact that the close net of fine magenta-pink veins onthe white (or pale pink) petals, fades as the flowers age (Yeo 1985, p. 69).
The English common name 'Pencilled Crane's-bill' clearly is a book name, and the reference to the magentaveins is an obvious one.
Threats
None.
Introduction, archaeophyte, occasional. European southern-temperate and widely naturalised.
1860; Smith, T.O.; Ardunshin.
February to November.
Growth form and preferred habitats
This species is distinctive and, when flowering, quite unmistakable; it is 10-60 cm tall, straggling,tap-rooted, with sub-rounded to kidney-shaped (reniform) leaves, lobed almost to the base and smallflowers with bright purple petals 4.5-6.0 mm, notched at the tip (Sell & Murrell 2009; Parnell &Curtis 2012). A usually overwintering biennial or annual of disturbed lowland, it is found in sunny tosemi-shaded sites, on moderately moist, fertile, calcareous to weakly acidic soils. The 1987-2004Change in the British Flora monitoring survey found that it can survive competition and increaseand thrive in arable crops, even if field nitrogen levels are high (Braithwaite et al. 2006). Thesame survey found evidence of a slight expansion of its range in the north, the areas involvedsuggesting that set-aside ground may have been partially responsible for the increase. G.dissectum is decidedly weedy in its ecology, its established strategy being categorised as R/CRby Grime et al. (1988, 2007), meaning it is intermediate between Ruderal and Competitive Ruderal.The Change survey gives good examples of this in action across Britain (Braithwaite et al.2006).
Typical habitats of G. dissectum include grasslands (frequently in rather disturbed areas, likearound gateways), hedgebanks, wayside grass, stony waste ground and in old quarries, but it is alsofrequent as a weed of arable cultivation, including winter wheat fields and garden situations. Itcontinues to be introduced with temporary grass leys or clover crops as a seed impurity. A study of theweed flora of southern England between 1960 and 1997 suggested that Cut-leaved Crane's-bill wasincreasing in frequency (Marshall et al. 2003). In grassland grazed by horses, Cut-leavedCrane’s-bill is often associated with latrine areas (Gibson 1997). The grasslands it occupies inFermanagh include riverbanks, lakeshores and parkland.
Flowering reproduction
From May to August, the plant produces tightly clustered cymose inflorescences of usually deep pink orbright purplish flowers, each with petals slightly notched and shorter than the hairy sepals. The rathersmall flowers mature in pairs, but even so they generally appear on the plant only a few at a time sothat the species is quite inconspicuous and far from 'garden worthy'. Although the flower is reported tobe protogynous (Hutchinson 1972), and is also said to attract at least saw-flies (Proctor & Yeo1973, p. 134), in common with other weedy, small-flowered species in the genus, it is habituallyself-pollinated and regularly sets seed (Garrard & Streeter 1983; Proctor et al. 1996, p.332). When the fruits ripen, each of the five seeds is ejected explosively and thrown in a sling-likemanner for a considerable distance (up to 6 m) out of its mericarp container, leaving the familiar andcharacteristic candelabra-like appearance of the discharged crane's-bill fruit (Yeo 1985, p. 33).
The seed is relatively large, oval, its surface pitted and it is heavy in comparison with many otheragricultural weeds, weighing 0.002-0.003 g (Salisbury 1964). The seed production per plant is veryvariable, often ranging from 50-200 (Guyot et al.1962), but a large plant may produce 10,000 to20,000 seeds (Salisbury 1964). Germination of seed shed in the summer begins in July and extends in anintermittent manner throughout the autumn into November in mild years. By the end of the year, between18-59% of seeds have germinated and appreciable numbers of seeds continue to germinate intermittentlyduring the following year, mainly from May-September, peaking in mid-June. They continue to germinateover the following four or more years. The timing of emergence within this period is largely determinedby soil moisture and to a lesser extent by cultivation disturbance in early summer (Salisbury 1964, Fig.28 & p. 320; Roberts & Boddrell 1985).
Fermanagh occurrence
G. dissectum is widely but rather thinly scattered throughout the whole VC, being represented in84 tetrads, 15.9% of those in the VC. Nine of these tetrads contain only pre-1975 records, whichsuggests a definite decline in suitable habitats, or perhaps cleaner sown seed mixtures.
British and Irish occurrence
G. dissectum is widespread and common throughout the whole of B & I, except in a few areas ofC & W Ireland and C & N Scotland, where it becomes scarce or absent. The ability of the speciesto compete and thrive in nutrient-rich disturbed habitats makes it likely that it is increasing itsrange northwards and becoming more abundant in many areas of B & I (S.J. Leach, in: Preston et al. 2002).
At the hectad scale in the New Atlas maps, G. dissectum appears as common and ubiquitous asG. robertianum (Herb-Robert) in NI. However, this certainly is not the case; Cut-leavedCrane's-bill is very much more occasional or even casual in many of its cultivated or more transitorywayside and waste ground habitats than is G. robertianum. The comparative post-1975 statisticsfor Fermanagh are: G. robertianum 1156 records in 450 tetrads while for G. dissectum thefigures are 104 records in just 70 tetrads.
British status revised
Before the 20th century seed cleaning techniques were developed, G. dissectum frequently was alarge-scale seed contaminant of clover mixtures, particularly affecting seed of Trifoliumincarnatum (Crimson Clover), which was extensively cultivated in S England towards the middle ofthe 19th century. Up to 50% of the seed sown as clover could actually be Cut-leaved Crane's-bill.However, more normal contamination levels would have been closer to 5% (Salisbury 1964, pp. 136, 167).
The weedy behaviour and persistent seed bank of G. dissectum, together with the fact that the onlyfossil evidence of the species presence in B & I from the current Flandrian interglacial (=Littletonian in Ireland) is from two archaeological sites of the Roman period (100-400 AD) (Godwin 1975)has resulted in a recent revision of the plant's status. The New Atlas is the first floristicpublication to regard G. dissectum as an archaeophyte throughout B & I, ie an ancient orpre-1500 AD introduction. In the light of this recent modification, The current author (RSF) regards ita little surprising that the same change of status was not applied to G. molle (Dove's-footCrane's-bill) and possibly also to G. lucidum (Shining Crane's-bill), two further Crane's-billsof very similar biology, ecology and fossil history to G. dissectum.
European and world occurrence
Cut-leaved Crane's-bill is believed to have originated in Europe and W Asia and it is widespread in S, W& C Europe, along the Mediterranean coast of NW Africa and in Macaronesia. In a discontinuousmanner, it extends eastwards in southerly latitudes only reaching the Caspian Sea. Northwards thespecies stretches up the Atlantic and Baltic coasts of Scandinavia and crosses the Arctic Circle, butprobably only with the assistance of man. It is very widely dispersed across the globe as a naturalisedintroduction in regions including Iceland, the W Himalaya, N & S America, the Cape Province of SAfrica, New Guinea, Australia and New Zealand (Hultén & Fries 1986, Map 1272). It was first recordedin New Zealand in 1940, but recently has become a prominent weed of arable land, especially in N and ESouth Island (Webb et al. 1988).
Names
The Latin specific epithet 'dissectum' means 'deeply cut', referring very aptly to the upper leaves(Gilbert-Carter 1964). The most frequent English common name is simply a translation of the Latinspecies name, but there are two alternatives, 'Jagged Crane's-bill' (Watts 2000), which is not accurateand deserves to be ignored and a much more interesting name 'Pink Needle' (Britten & Holland 1886),which they suggest Lyte (1578, p. 48) applied to this species. Prior (1879), however, followed WilliamTurner's Herball (1551-68) and referred this name to Erodium moschatum (Musk Stork’s-bill)and E. cicutarium (Common Stork’s-bill) (see also Grigson's account of these two species in hisEnglishman's Flora (1955 & 1987)). Prior suggested the derivation was due to the longtapering fruit awn resembling the needle used in pinking, ie for making eyelet holes like pinks, inmuslin (Prior 1879, p. 185). 'Pink Needle' is a name sometimes also given to Scandixpecten-veneris (Shepherd's Needle).
Threats
None.
Native, or possibly an ancient introduction, occasional. European southern-temperate, also widelynaturalised in both hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
March to October.
Growth form and preferred habitats
A small, spreading, or if supported, quite large, 10-50 cm, climbing, softly hairy winter annual orbiennial, G. molle is not a common species in Fermanagh, nor indeed in the N & W of Irelandgenerally, although it is widespread and common throughout most of Britain and some parts of Ireland(see below). It grows in a wide array of open, lowland habitats and is probably most frequently found indry to droughted, often shallow calcareous or sandy, open, sunny, regularly disturbed wayside orgrassland sites (Grime et al. 1988, 2007). It also appears locally on sandy lakeshores. In otherregions of these islands, but not in Fermanagh, it is commonly found in arable fields and on sand dunes(arable farming is virtually extinct in Fermanagh). G. molle also regularly crops up in what canbe regarded as permanently open sites, eg in rock crevices, screes and on walls (Segal 1969; Rodwellet al. 2000, eg community types OV38 and OV39).
In all these different situations, ecologically it behaves as a stress-tolerant, ruderal weedy species,avoiding competition from taller and more vigorous perennials. It achieves this avoidance by colonisingconditions that are too dry, disturbed, or nutrient-impoverished to allow perennials to growaggressively and thrive (Grime et al. 1988, 2007). It is therefore quite possible that in the N& W of Ireland and, indeed, in similar, but usually somewhat more upland regions of Scotland, theclimate is consistently simply too wet and, perhaps, the soils excessively acidic (it prefers a pH above5.0) to allow G. molle to compete and be as frequent as it is elsewhere in these islands.
Like its even more common, but much more shade-tolerant relative, G. robertianum (Herb Robert),G. molle often overwinters as a small, green rosette of downy, dissected leaves on a short, erector decumbent stem attached to a quite long, penetrating taproot. Like Herb Robert, the bruised leavesare unpleasantly pungent, the odour somewhat reminiscent of mouse, a feature which, along with theirexcessive glandular hairiness, appears sufficient to deter the depredations of herbivores.
Variation
There are no subspecies described for G. molle, but four varieties have been distinguished andnamed in Sell & Murrell (2009). These are var. arenarium N. Terrace, a form that produces aplant lying flat to the ground in a circle, with leaves 0.5-3.0 cm in diameter, flowers 5-6 mm indiameter, and with ribbed mericarp fruits; var. molle, is an erect or ascending plant with leaves1.5-5.0 cm in diameter, flowers 5-8 mm diameter and mericarps ribbed; var. aequale Bab. is anerect plant with leaves 1-3 cm in diameter, flowers 5-6 mm in diameter and mericarps smooth and withoutribs; and var. grandiflorum Vis. is another erect plant with leaves 3-6 cm in diameter, muchlarger flowers 9-13 mm in diameter and with mericarps ribbed.
Var. molle is the widespread plant, while var. arenarium is an ecotype of sandy soils nearthe coast and sandy heaths inland, occasionally found elsewhere.
The distribution of var. aequale is unknown while var. grandiflorum occurs in scatteredlocalities across S Britain and in Continental Europe (Sell & Murrell 2009).
Fermanagh occurrence
G. molle has been recorded in 25 tetrads, 4.7% of those in the VC. It is only occasional andthinly scattered in the survey area, but as the tetrad map clearly shows, it is much more frequent inthe eastern half of the county, there only being two tetrads with records in all the ground to the W ofLough Erne. Other habitats where there are rare Fermanagh occurrences include disturbed upland pastures,roadside verges, quarries, bridges, urban walls and car parks.
Flowering reproduction
G. molle flowers from April onwards, continuing throughout the summer until September, the timingdependent upon the favourability – or lack of it – of the local environment. In shallow, lighter, driersoils, plants have already completed their flowering and fruiting cycle by the beginning of June, asituation which might sometimes allow more than one generation within a growing season as happens withother successful small weedy annuals such as Senecio vulgaris (Groundsel) and Capsellabursa-pastoris (Shepherd's-purse). Although the current author (RSF) does not know of anydetailed population study of G. molle, I certainly would expect that two or three 'cohorts' ofseedlings germinating in different periods of the growing season might complete their life-cycle duringa single year, as is known to be the case for G. robertianum (Falinska & Piroznikow 1983).
Habitual self-pollination
The paired, mauve or pale pink flowers with their distinctive deeply notched petals, barely longer thanthe sepals, must originally have evolved to attract insects to carry out cross-pollination. The degreeto which this strategy still occurs requires more study to elucidate, but nowadays it appears thatflowers of G. molle habitually self-pollinate. The selfing process is described as 'automatic',the small, self-compatible flowers having their anthers and stigmas borne on the receptacle in closeproximity and maturing simultaneously. The transition from reliance on an external agency of pollentransport to an automatic selfing breeding system is a common strategy adopted by many small, ephemeral,annuals, many of which live only a few weeks rather than the year which the term 'annual' suggests. Theyappear to have evolved this very reliable reproductive method to enhance the chances of their survivalin unpredictable, open, often harsh physical and chemical environments. There is a balance of advantagesand disadvantages to the various strategies of pollination and fertilisation and a continuum ofvariation exists between the two extreme types of sexual reproductive mechanism. Furthermore, as hasalso been seen to be the case with Poppies (Papaver spp.) and other weedy plants, a species mayonly require the occurrence of a very occasional cross-fertilisation to replenish population variationand to maintain its much longer-term genetic options (Proctor et al. 1996, pp. 330-3).
Seed size and natural dispersal
In comparison with many other common, more or less ephemeral weeds, G. molle seed is quite large,weighing on average 0.001539 g. This is five times heavier than seed of Senecio vulgaris andthree times that of Stellaria media (Common Chickweed), two species with which Dove's-foot oftenassociates (Salisbury 1942, Table 9; Rodwell et al. 2000, OV5, p. 345)). Apart from thecharacteristic Crane's-bill jerked, sling-shot mechanical seed release of the Geranium genus,viable seed samples of G. molle have been recovered from the dung of cows and Wood-pigeon, bothof which obviously must feed on the plant and transport it (Ridley 1930, pp. 361, 498).
Seed dispersal by man
In addition, in past years, G. molle was widely disseminated through being a frequent and,occasionally, a rather large-scale contaminant of commercial clover seed mixtures (Salisbury 1964).Another means of accidental long distance dispersal by man was in ships' ballast, or in the discardeddung of transported cattle. G. molle features in a long list of European species introduced andrecorded in Philadelphia on dumped ballast heaps of this nature, given by Ridley (1930, p. 646).
Seed germination and persistence in soil seed bank
After an experimental sowing made by Roberts & Boddrell (1985) at the end of July, 11% of seedgerminated in the first autumn, followed by appreciable numbers in each of the following four years, soa persistent soil seed bank clearly exists. In this particular study seedlings emerged in a consistentpattern, mainly from May to September, the timing largely determined by soil moisture, but with a mainflush following a programmed soil disturbance in mid-June simulating cultivation (Roberts & Boddrell1985).
Possible confusion with G. pusillum: G. molle is rather similar inappearance and ecology to G. pusillum (Small-flowered Crane's-bill – another frightful Englishcommon book name), but it has larger, more spreading and more deeply notched petals and hairless, ribbedmericarps (although the actual seeds are smooth and brown) (Butcher 1961; Yeo 1985). The two species aresaid to be capable of hybridizing (Stace 1975) and undoubtedly they are sometimes confused.
In NI, G. pusillum has been very rarely recorded in Cos Down (H36) and Tyrone (H38), where it isregarded as an introduction, while elsewhere in S & E Ireland and in Britain, where it is a muchmore frequent and widespread species, Small-flowered Crane's-bill is considered a native (Preston etal. 2002).
British and Irish occurrence
G. molle is native, common and widespread throughout lowland Britain ascending to around 550 m. Itbecomes less common and more scattered northwards in Scotland (New Atlas). In Ireland, it is alsoquite common and widespread, although most frequent in the E & S and increasingly scatteredwestwards, where it again becomes more obviously confined to coastal sites. Being so widely introducedelsewhere in the world, the current author (RSF) feels it worth considering that the patchy Irishdistribution of G. molle might represent the outcome of many repeated accidental introductions,stemming from a native range that, as the New Atlas map proves, is much more consolidated in thewarmer lowland areas of the southern half of Britain.
European and world occurrence
G. molle is indigenous and widespread in all of S Europe, eastwards to the Balkan Peninsula andnorthwards to S Sweden. Beyond this it is much more scattered and occasional, but stretches to the WHimalaya and, in Scandinavia, to the N Baltic. It is also present in N Africa and the Atlantic Isles.Beyond this, it has been widely spread as a weed of cultivation in parts of Scandinavia, C Russia,Iceland, The Faeroes, S Africa, N Japan, N & S America, Australia and New Zealand (Hultén &Fries 1986, Map 1269).
Names
The Latin specific epithet 'molle' means 'soft', an easily appreciated reference to the characteristicdense coat of soft hairs on the leaves. The English common names 'Dove's-foot' (16th century), or'Dove's-foot Crane's-bill' if we absolutely must (!) (20th century) and the older 'Culverfoot' (15thcentury), derive as direct translations of the medieval Latin herbalist's name 'pes columbae' or 'pescolumbinus', dove and culver being alternative names for pigeon. The shape of the leaf is erroneouslysupposed to resemble a pigeon's foot, but the soft hairiness of the whole plant is reminiscent of thebird's downy feathers. In other European countries, this name referred to G. columbinum, but in B& I, G. molle is the commoner species, so the name was transferred (Dony et al. 1974;Grigson 1974; Watts 2000).
Grigson (1955 & 1987) lists a few alternative English common names, all of which are interesting butdecidedly puzzling in their derivation, eg 'Dolly Soldiers', 'Jam Tarts', 'Mother of millions' and'Starlights'. The latter author tells us that the plant was used like Herb Robert as a vulnerary tostaunch bleeding. Nicholas Culpeper (1653), whose herbal ran to over a hundred editions, recommendedDove's-foot for a list of cures almost identical to those attributed to G. robertianum (see thespecies account on this webpage), and he memorably described it as, "a very gentle, though martialplant".
Threats
None.
Native, uncommon to occasional, but locally frequent. Submediterranean-subatlantic and native in NAfrica, SW & C Asia.
1864; Dickie, G.; Enniskillen Road, E of Belcoo.
Throughout the year.
Growth form and preferred habitats
A most distinctive weedy winter or summer annual, very variable in overall size with slender, succulent,branched stems, 10-50 cm, and characterised by almost hairless, slightly succulent, glossy, shallowlylobed, red-tinged leaves which go an even deeper waxy red towards the end of their life span. The plantis a quite markedly calcicole and is found in open, sunny to partially shaded, relatively dry sites oncalcareous rocks or in sandy soil where conditions are weakly acid, or near neutral, and moderatelyfertile, or in recent years, becoming nutrient-enriched (Hill et al. 1999). It thereforefrequents warm, dry rocky ground, stony banks, bare ground and walls, mainly in calcareous areas, butalso artificial habitats which meet the particular needs of the species.
Shining Crane's-bill colonises and can sometimes spread rapidly and invasively, becoming abundant incalcareous or basic, relatively open, artificial habitats that offer bare soil, typically nearhabitation. This includes sites on or at the base of old lime-mortared walls, churchyards, quarries, thegravel drives of houses, overlooked or neglected corners in cultivated beds, rock-work and crevices ingarden paving (Tatlow 1898).
As with G. molle (Dove's-foot Crane's-bill, the established strategy of G. lucidum iscategorized as R/SR (ie intermediate between Ruderal and Stress-tolerant Ruderal), and it appears to bepoorly equipped to compete with persistent perennials (Grime et al. 1988, 2007). It thereforetends to avoid perennial competition by colonising and being resident in less than ideal growingenvironments that the former avoid. This means G. lucidum is often confined to ground that issubject to intermittent disturbance or other forms of stress environments, including for instance thedwarfing effect of summer drought, or of the damaging effect of substrate movement, eg on screes, or ongravel paths subject to trampling. In these sorts of habitats it can be long persistent (Crawley 2005).
There appears to be some degree of tolerance, if not appreciation of this weedy annual amongst gardenersin B & I. Ingwersen's (1978) Manual of alpine plants, while recognising the true weedy natureof G. lucidum, suggests that on account of its colourful appearance, especially its rich redautumn leaf tints, "it may be allowed to colonise a cool wall or a shady nook".
Fermanagh occurrence
During the last quarter of the 20th century, G. lucidum has had a very similar frequency ofrecording to G. dissectum (Cut-leaved Crane's-bill), but it is more restricted in itsdistribution, being present in only 10.0% of the Fermanagh tetrads, compared to a figure of 15.4% forG. dissectum. Previous records from the VC indicate that except in the upland Western plateausouth of Lower Lough Erne (District 4 of Meikle and friends), G. lucidum was only very rarelyrecorded in Fermanagh. A total of just six records appear in their Revised Typescript Flora forthe other three VC districts during the period 1864-1953 (Meikle et al. 1975).
G. lucidum is nowadays considered occasional to locally frequent in Fermanagh, there being recordsfrom a total of 53 tetrads. The specific habitat conditions required are chiefly realised in uplandareas of Fermanagh, eg on stony ground, cliffs and scree slopes around Knockmore, Hanging Rock and aboveFlorencecourt. However, the species occurs less abundantly in more or less disturbed habitats at lowerlevels, including around the limestone lakeshores of Lough Erne and especially of the Lower Lough.
In other parts of B & I, G. lucidum is often associated with the ballast of railway lines andin Fermanagh it still persists in the vicinity of the old railway station at Maguiresbridge, whichbecame abandoned in 1957 when the transport system closed down.
Flowering reproduction
The paired, bight pink flowers, not more than 10 mm in diameter are profusely borne between May andAugust and can be either bisexual and protogynous (ie female when they first open), or wholly female, abreeding system which clearly encourages out-crossing. Nevertheless, the bisexual (hermaphrodite orperfect) flowers are capable of self-pollination should insect visitors fail to find and service themand thus seed set is assured (Hutchinson 1972).
As is generally the rule with winter annuals, overwintering leaf rosettes of this species arising fromseedlings emerging in the autumn are stronger and grow into larger plants than those produced by springand summer germinating seed (Yeo 1985, p. 158). In common with other Geranium species, the seedis large and heavy for a weedy annual (mean weight 0.0015 g) and it is mechanically dispersed distancesup to 6 m by the carpel projection sling arrangement typical of Crane's-bills (Salisbury 1964, p. 169;Yeo 1985, pp. 157-8).
Germination and seed longevity
Germination is both in the autumn and the following spring and summer. RSF has noticed seedlings, withtheir first true leaves, appearing in gravel at the beginning of August. Despite a search ofconsiderable library resources, RSF has discovered no information on secondary seed transport orpopulation behaviour. Evidence regarding seed longevity in soil has been hard to locate, but the revisededition of Comparative plant ecology contains a qualified suggestion that a small proportion ofG. lucidum seed can survive soil burial for up to five years (Grime et al. 2007, Table7.3). A report by Kent (1975, p. 234), also suggested that G. lucidum was introduced as a weed toa garden in Ealing in soil that had been brought from Wiltshire, indicating that a seed bank of someduration does exist. Even a minor study could add significantly to our knowledge of the populationdynamics and ecological behaviour of this rather colourful little annual.
Chromosome counts and hybrids
G. lucidum is the solitary species belonging to Section Lucida of the genus. It has theunusual chromosome diploid number of 20 (the most common numbers in the genus being 2n=26 and 2n=28),and additional counts of 40 and 60 have also been rarely made (Yeo 1985, p. 40). Although G.lucidum and G. robertianum (Herb-Robert) (usual diploid number, 2n=64) are quitefrequently found growing intermingled or close to one another in disturbed or rocky ground, a hybridbetween them has only ever been claimed (twice) by one knowledgeable but eccentric late-19th centuryEnglish naturalist (F.A. Lees). In view of the difference in chromosome counts, he almost certainly musthave been mistaken. No voucher specimens exist for his hybrid plants and the reports may be discounted(see D. McClintock, in: Stace 1975, p. 192; Yeo 1985, pp. 44, 46).
The question of native status
Traditionally, but in the absence of any fossil proof, G. lucidum is considered a native speciesthroughout most of B & I. In the Census Catalogue of the Flora of Ireland, it is regarded anintroduction in Co Wexford (H12) and Co Down (H38), but present and taken as native in all of theremaining 38 Irish VCs (Scannell & Synnott 1987).
British and Irish occurrence
G. lucidum is widespread across low ground at all latitudes across both B & I,appearing most prominent to the S & W of both islands, although this is less marked in Ireland(Preston et al. 2002).
The three editions of the Flora of the NE of Ireland appear to show that from the end of the 19thcentury onwards in the three NE counties of NI covered by that Flora (Down (H38), Antrim (H39) andLondonderry (H40)), Shining Crane's-bill was always very local and rather coastal in its distribution.It was not always persistent either, particularly in Co Down where it was not recorded until as late as1902 and it appeared to be extremely dependent upon man to provide suitable sites (Stewart & Corry1888; Praeger & Megaw 1938; Hackney et al. 1992).
Comparison of the two BSBI Atlas surveys of the 1960s and the late 1990s (Atlas, NewAtlas) showed a major increase of the known distribution of the species. The calculated changeindex has a value of +1.42, indicating an increase of 142% over the 40 year period. This appears a verysignificant potentially invasive expansion by the species – despite there being some reservations abouthow the index calculation was made for species covered by the New Atlas (Hodgson 2003; Prestonet al. 2003).
The invasive nature of the plant, its weediness, association with horticulture and with sites of humandisturbance, including the involvement of nutrient enrichment, may, when taken together, have offeredthe species conditions for rapid, perhaps short-term colonisation of fresh sites in recent years. It isjust a little puzzling, however, that this trend did not appear to register as significant during the1987-1988 Monitoring Scheme partial survey conducted by BSBI members (Rich & Woodruff 1990,2: Map and Table on p. 70).
European and world occurrence
Beyond the shores of B & I, G. lucidum is regarded as indigenous in W & S Europe andadjacent parts of NW Africa & W Asia. It is rather thinly scattered throughout C Europe and, indeed,there appears to be a gap in the distribution in N Central Europe. In Norway, G. lucidum becomesconfined to coastal areas. In SE Europe, it becomes scattered in Turkey and is even more sparseeastwards towards a few isolated stations in the Himalaya (Hultén & Fries 1986, Map 1273). Althoughthe latter reference does not show it, G. lucidum has spread to one or two states in Pacific NWAmerica, where currently it is considered quite an invasive alien weed.
Uses
One Internet Website suggests that G. lucidum has diuretic and astringent medicinal value (https://pfaf.org/user/Plant.aspx?LatinName=Geranium+lucidumAccessed 28/10/2021).
Names
The Latin specific epithet 'lucidum' from the verb 'luceo' means 'shining', an obvious reference to thealmost hairless leaves and stems (Gilbert-Carter 1964). The English common name 'Shining Crane's-bill'is therefore a simple, unoriginal translation 'book name'. Britten & Holland (1886) offer two localalternatives, 'Bachelor's Buttons' (from Preston, Lancshire) and 'Robin' (from Devon), both borrowedfrom the 111 names also applied to G. robertianum (see Grigson 1955, 1987 and RSF's speciesaccount on this webpage).
Threats
Under certain conditions an invasive weed, but possibly not very competitive and therefore may notpersist for long.
Native, common throughout, locally abundant. European temperate, also native in C & E Asia and widelynaturalised beyond its native range.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form
A very common and variable herb, its overwintering, wintergreen rosettes of heavily dissected leaves areperfectly frost hardy. In terms of life-form, it is a rather malodorous, slightly succulent annual orbiennial; it is almost always monocarpic, flowering once only and then dying. Very rarely it behaves asa short-lived perennial, living for three years, ie germinating in the autumn of the first year,reaching flowering size late in the second season of growth and flowering a second time in the followingspring. It only manages to do this if it flowered late in its first season of generative growth – whenit probably did not set much (if any) seed. These 'triennial' plants never exceed 5% of the speciespopulation and represent individuals whose development has been inhibited in the juvenile phase. In suchcases they live for three years in the form of poorly developed leaf-rosettes, and some of them diebefore attaining the generative phase. Great plasticity exists within the species, so that individualsof the same growth phase differ considerably in size. The greatest variation in this respect is seen inindividuals in the generative phase (Falińska & Pirožnikow 1983).
It is thus somewhat more than a 'facultative biennial', in the sense of Kelly (1985),ie a 'pauciennial', defined as a short-lived monocarpic perennial, rather than a 'strictbiennial', which always flowers only once during its second year of growth.The most familiar strict biennials in this part of the world are Linum catharticum (Purging- orFairy Flax) and Pedicularis sylvatica (Lousewort).
Biennial species
Most so-called biennial species growing wild in the field take more than two years to arrive at the sizeand accumulate the photosynthetic resources necessary for successful flowering and fruiting (Harper1977; Forbes 1989; Bertin 2001). The life-form of this type of plant is very much conditioned by thefertility (or rather, the degree of infertility) of the site they occupy, plus the competitionthey face and the extent and frequency of grazing and other deleterious pressures that they must endureto survive to reach the flowering stage.
The current author (RSF) regards G. robertianum as 'an extended annual' or 'a conditional annual',ie it is an annual, completing its life-cycle in twelve months, provided it is given mesicgrowing conditions. However, it regularly requires several years to achieve flowering in less favourableenvironmental situations and circ*mstances.
Preferred habitats
As might be expected with such a common and widespread species, G. robertianum has an extremelywide ecological amplitude, allowing it to grow and survive in a tremendous variety of habitats. It ismost frequent and abundant in limestone terrain, or in moist, shady, mesic woodland or scrublandsituations, ie moderate in terms of acidity, moisture availability and nutrient status. However, thisspecies is also surprisingly tolerant of open, sunny, dry conditions. In the latter circ*mstance, it isoften dwarfed and very strongly pigmented, the whole plant turning either red or purple-brown.
In terms of habitat, the species is really only excluded from pastureland and from extreme examples ofacidic, disturbed, wet or aquatic environments (Grime et al. 1988; Hill et al. 1999; Tofts2004). In coastal situations, a more succulent form, referred to by some as subsp. maritimum, mayoccur on non-saline shingle and, where railways survive, this plant form can also sometimes be found onthe ballast between or along the rails (Tofts 2004).
In quarries, arable farmland, or in other types of disturbed ground, Herb-Robert usually appears as acasual ruderal or as a pioneer annual in wayside situations or on skeletal soils. In these more openhabitats, it can colonise and often grow extremely rapidly. When it reaches the flowering stage, ittends to form diffuse, rounded plants, up to 60 cm or more in height. The open-habitat forms of thespecies are, however, generally unable to compete with other vigorous ruderals, either annual orperennial, such as Galium aparine (Cleavers), thistles, docks and hogweed, with the result thatsuch G. robertianum populations often quickly become overgrown and diminish or completelydisappear after their second year.
In very damp oceanic deciduous woodland conditions, Herb-Robert very occasionally is reported growing asan epiphyte on the trunks and lower branches of trees and shrubs, although we have no immediaterecollection of this happening in the Fermanagh area (Tansley 1968, p. 109).
The root/shoot ratio of the species is low and the root system is shallow, fibrous and may or may notcontain a vescicular-arbuscular mycorrhizal fungus. If the mycorrhizal association is present, itconfers a significantly greater ability on the plant to take up P and N (Boerner 1990).
Plant communities
While G. robertianum is predominantly a plant of woodland shade, its wide ecological tolerancesresult in it having been recorded in no less than 21 varied communities of the National VegetationClassification (Rodwell et al. 1991a, b, 1992 & 2000). In almost all cases, however, itoccurs at a frequency of 40% and, quite often, below 20 %. In the communities where it is represented,the 'constant species' most frequently associated with it is none other than the Rubus fruticosusagg. (Common Bramble) (Tofts 2004).
Variation
At present three subspecies and three varieties of one of them are recognised by thecritical Flora of Great Britain and Ireland (Sell & Murrell 2009). In coastalsituations, a more succulent biennial form of the species takes over, named subsp. maritimum(Bab.) H.G. Baker, of which three varieties are distinguished on their degree of hairiness: var.maritimum Bab. where the plant is glabrous or nearly so; var. intermedium Wilmott, wherethe plant is fairly hairy; and var. hispidum Druce, where the plant is densely hairy. Subsp.maritimum may occur on non-saline shingle and, where railways survive (which they no longer do inFermanagh), this variant can also sometimes be found on the ballast between or along railway lines. Itoccurs in widely scattered locations around the coasts of Britain, but mostly along the southern coastsof England and Wales (Tofts 2004).
Another variant, subsp. celticum Ostenf., is an annual and a characteristic plant of crevices incoastal limestone rocks and cliffs, having small, pale, almost inodorous leaves, reddish only at thenodes and petiole bases. It is said to have large pale pink or whitish flowers, the petals bearing awhite streak on their under-surfaces (Rich & Rich 1998, p. 211; Sell & Murrell 2009). Experiencein Fermanagh, however, leads RHN and the current author (RSF) to believe that this form is not verydistinct and it seems to intergrade with the typical plant subsp. robertianum (Meikle etal. 1975). Stace (2019) may well be correct in saying that subsp. celticum is confined tocoastal limestone crevices in central W Ireland and S Wales.
The third subspecies is the regular, widespread form of the plant subsp. robertianum, which isusually biennial, has stems ascending, flowers deep pink and schizocarps ± hairy.
There can be striking differences between different stocks of G. robertianum growing inquite close proximity, yet remaining distinct from one another due to the general tendency forself-pollination that exists in the species. The degree of hairiness is especially variable and can makeidentification of the subspecies/varieties problematic (Tofts 2004).
Similar species
A very similar species, of entirely annual life-form is G. purpureum (Little-Robin). It isvery much more frost-sensitive, more drought-resistant and hence it has a more southerly B &I and Mediterranean distribution and prevalence than G. robertianum. Physically, it isdistinguished by its very much narrower and smaller petals and by its yellow anthers. In Ireland, G.purpureum is confined to just four hectads along coastal areas of counties Cork (H3, H4 &H5) and Waterford (H6) and, while it is more frequent in SW England, it is also a scarce plant there(Preston et al. 2002).
Flowering reproduction
G. robertianum entirely relies on seed for its survival and dispersal, having no means ofvegetative reproduction. Indeed, the vegetative portion of the stem itself is extremely short, althoughprocumbent flowering plants often tend to give a different impression of both these matters. Themuch-branched flowering stems arise from the axils of the upper rosette leaves and, as alreadyindicated, they can be either erect, forming a rounded bushy plant, or decumbent, giving rise to asprawling, trailing or dangling mass (Tofts 2004).
Natural forest and experimental population studies in Poland involving garden cultivation found thatthree cohorts of seedling emergence could be distinguished: vernal, aestival (ie summer) and autumnal.The vernal cohort were the most important of the three for the population dynamics of the species, forthey demonstrated the greatest fecundity and survival rate (Falińska & Pirožnikow 1983). In B &I, flowering stretches from April onwards, with the occasional unseasonable flower appearing in anymonth of the year in mild years. These late or very unseasonable flowers probably never set any viableseed.
The main flowering period (April to September), coincides with the expanded canopy of deciduous trees andshrubs and, indeed, G. robertianum does flower surprisingly freely under moderately dense shade,eg in garden experiments it flowered at light levels above 12.5% of ambient illumination (Board2001; Tofts 2004). Flowering tends to peak in May-June, with a secondary burst of colour and sexualactivity in August.
Flowers are borne in pairs (ie a reduced cyme or cymule) but individual flowers sometimes abort. Floralparts are in fives, the mericarps each containing two ovules, although usually only one develops into aseed. Individual flowers open early in the day, but they last only one or two days at most. The blossomoffers both nectar and pollen and they are visited by various long-tongued bees, butterflies and flies.Thus outcrossing would normally be expected yet, on the other hand, the stability of observed distinctlocal population differences strongly suggests that levels of self-pollination are generally high inthis species (Baker 1956; Yeo 1985; Proctor et al. 1996). In N America, however, Bertin (2001)found uniformly high levels of seed set from self-pollinated (96%), cross-pollinated (96%) andunmanipulated experimental flowers (88%).
Seed production and dispersal
Under natural conditions, G. robertianum individuals typically produce between 10-40 fruitingheads or capsules, and although each flower contains ten ovules, the maximum seed number set is five percapsule. Tofts (2004) found the average seed (partial fruit or mericarp) production was 4.95 per flowerfor British material. The fruit splits when ripe and forcibly slings the seed a distance of up to 6.5 m.Obviously the distance travelled depends upon numerous factors, including eg the density of surroundingvegetation, the original height above ground and the prevailing weather conditions at the moment ofrelease.
Seed germination and soil-seed bank longevity
Most (but not absolutely all) G. robertianum seed requires an after-ripening requirement of two ormore months before germination can take place (Baker 1957) but, after this time, light or darknessappears to have little effect on emergence (Grime et al. 1988 & 2007).
As with flower production, germination and the appearance of new seedlings may continue throughout thewhole growing season, although it tends to peak strongly in May (giving rise to the vernal cohort ofseedlings) and rises again, but to a lesser extent, so that summer and autumn cohorts may also berecognised in July and September (Falińska & Pirožnikow 1983; Bertin 2001). As expected, inPoland the spring cohort performs best of the three in terms of growth, survival to flowering and seedoutput (80-90% attaining the generative stage in their first year of life), while by comparisonseedlings emerging in autumn have very low levels of overwintering survival (ie 5-6%). In the very muchmilder Atlantic winters in NW Ireland, survival rates of winter leaf-rosettes in the wild is undoubtedlymuch greater than this, although at present there are no figures to prove this suggestion. (Anotherproject awaiting someone.)
On the other hand, in experimental sowings made at the end of September, Roberts & Boddrell (1985)found some seedlings emerged that autumn, but the main emergence was not until the following year. Fewseedlings appeared after the second year, but some continued to appear for up to five years aftersowing. These workers could not discern any clear pattern of emergence, but flushes of seedlingsappeared throughout the period from March to October, not apparently associated with their threeregularly spaced and very thorough experimental soil disturbances, designed to simulate cultivation.Re-inspection of their results suggests to the current author (RSF), that emergence peaked in April,July and September (in close accord with the Polish work mentioned above), although the scale of thiseffect was not very predictable from year to year (Roberts & Boddrell 1985, Table 2, p. 233).
Herbivory and secondary plant substances
The stems of Herb-Robert are fragile, and when they are broken, or the leaves bruised, the plant givesoff a rather strong, unpleasant, mousey or musky, some say foxy smell (Genders 1971). Perhaps for thisreason, Mabberley (1997) describes all Geranium species as, "rabbit-proof herbs", yetwhile G. robertianum contains a long list of flavonoids, terpenoids, polyphenols, acids andtannins, it possesses no toxic alkaloids (Tofts 2004). Thus it may be that the plant is distasteful toherbivores, vertebrate and invertebrate, without actually being poisonous.
Medicinal uses
Of all Geranium species, Herb-Robert was formerly the most frequently used in folk and veterinaryherbal medicine. It was the standard remedy in Ireland for Red-water Fever, a very common disease ofcattle. It was also used to treat worms in horses and cattle, and for both constipation and diarrhoea.The very commonness of the species made it also a popular medicine for a range of human ailments heretoo, especially for kidney problems, stomach gripe, gravel and water retention. It was also an Irishremedy for coughs and sore throats and here, as in other areas of Britain and Europe, it was regarded asa vulnerary, ie, used for staunching bleeding. In England and Wales, it was used on the skin foreverything from a wash for the complexion, to the treatment of erysipelas and skin cancer (Allen &Hatfield 2004).
Fermanagh occurrence
In Fermanagh, it has been recorded in a total of 472 tetrads, 89.4% of those in the VC. This makes it the19th most widespread species in the county measured on this scale and, in terms of record frequency, itranks 37th in the Fermanagh Flora Database, so it is quite simply, a very common plant.
British and Irish occurrence
G. robertianum is found in every VC throughout B & I and, indeed, in very nearly 90% of theNew Atlas hectads in these islands. It is, however, very much rarer in the Outer Hebrides (onlypresent on South Uist) and it is very scattered on the Orkney and Shetland Islands (Tofts 2004).
European and world occurrence
G. robertianum is described as belonging to the European temperate element and is widespreadacross all of Europe from Ireland in the west to N Scandinavia, being absent only from the extremenorth; in S Europe, it is present throughout the Mediterranean basin from the Iberian to the BalkanPeninsulas, plus the western Mediterranean islands as far east as Crete. It is also indigenous across NAfrica (possibly as far south as Uganda), the Azores, Madeira, the Canaries and the Cape Verde Islands.In Asia, it stretches from SW Russia, the Caucasus, to the Himalaya, W Siberia, C Asia and SW China.
Beyond this it is widely naturalised so that it is difficult to distinguish native stations in E Asia andN America, although some authorities consider it naturalised in at least some areas of the PacificStates. It is possibly native but definitely expanding its range in eastern N America. It is introducedin Japan and SE China.
In the southern Hemisphere it occurs in temperate S America as far south as Chile, and it is alsonaturalised in New Zealand (Hultén 1971, Map 193; Hultén & Fries 1986, Map 1274; Tofts 2004). Ittherefore belongs to the discontinuous circumpolar plants (Hultén & Fries 1986, Map 1274).
Names
The genus name 'Geranium' comes from the Greek diminutive 'geranion' meaning 'a little Crane', an obviousreference to the narrow beak-like structure at the top of the fruit (Yeo 1985, p. 9). The specificepithet 'robertianum' and the most frequent English common name 'Herb-Robert' are both derived from themedieval Latin name 'herba Roberti', or 'herba Sancti Ruperti'. The English form, as 'Herbe Robert', wasalready in use by the 13th century (Grigson 1955, 1987). The origin of the name and the connection withthe Austrian saint may both lie in the distinctive redness of the typical plant, which could be acorruption or modification of 'herba rubra', this in turn leading through the colour to an associationwith blood, and with the eighth century St Rupert or Rudbert of Salzburg, who was invoked to healbleeding wounds, ulcers and erysipelas (Grigson 1974).
A list totalling 111 English common names is provided by Grigson (1955, 1987), who details numerous linkswith sacred and frightening phenomena, such as adders, cuckoos, goblins, sex and death. Twenty-four ofthe English common names begin with the letter 'R' and revolve around Robert, Robin (its diminutive) andthe colour Red. Grigson suggests that the name 'Herb Robert', and many of the illusions in thealternative folk names, may derive from the unpleasantly malevolent and resource-demanding house goblin,Robin Goodfellow. The German equivalent of this mischievous elf is Knecht Ruprecht, and theGerman plant name is 'Ruprechtskraut'. In support of this contention, 16th centuryreferences to Robin Goodfellow have described him as a hairy goblin, red-featured and wearing a redsuit.
Threats
None.
Introduction, neophyte, an occasional garden escape.
1900; West, W.; near the Silver Swallow Inn, Enniskillen.
April to September.
A tall or medium-sized, clump-forming, sub-alpine perennial of damp meadows and wood margins from themountains of C & S Europe, the native range of the species stretches from the Pyrenees acrossthrough the Alps and continues eastwards to reach upland areas of W Russia and its satellite states. Itis often naturalised outside this range, especially northwards (Yeo 1985, p. 165). G. phaeum hasbeen considered a very useful and easy garden subject for shade or semi-shade in Britain ever since itwas introduced sometime in the 17th century or even earlier (Yeo 1985, p. 12). It is appreciated for itsearly flowering (April onwards), in a range of mainly dark or dusky petal colours, some of which changehue as they age (Yeo 1985, pp. 163-6).
Possessing a stout rootstock and displaying a wintergreen leaf habit, the plant can readily establish*tself from dumped garden rubbish or as an escape from cultivation and it quite often becomes wellnaturalised and persistent. In Britain, it was first recorded in the wild as long ago as 1724 and isfrequently recorded in suitably moist, shaded sites, chiefly close to habitation on roadside grasslandsand hedgerows, churchyards, railway embankments and woodland margins (Clement & Foster 1994; S.J.Leach, in: Preston et al. 2002).
The thinly scattered Fermanagh locations are likewise almost exclusively on roadsides near habitation, orthey are obvious relicts of cultivation near ruined cottages, mainly in the lowland east of the county.There are local records from a total of 14 Fermanagh tetrads, nine of them with post-1975 dates. Onesite N of Tattykeeran Td is rather different from the remainder in being very much more remote. However,since the large clumps involved include plants of several distinct colour forms, it is highly likelythey all originate from fly-tipped excess garden material.
G. phaeum is widely scattered throughout Britain from Land's End to Inverness but, in Ireland, itis very much more frequently recorded in the north of the island than in the RoI. However, in thisrespect it may be important to realise that Reynolds (2002) lists 16 Irish VCs from which at least onerecord of G. phaeum has been made, five of which in the RoI are not represented in theNew Atlas map (ie VCs East Cork (H5), Meath (H22), Co Sligo (H28), Co Cavan (H30) & EastDonegal (H34)).
The very dark, sometimes nearly black flowered form of G. phaeum has been particularly fashionablein recent years, as have near-black varieties of other garden genera and one of the English common namesfor G. phaeum is 'Black Widow' (Griffiths 1994). The Latin specific epithet 'phaeum' is aLatinised Greek word meaning 'dusky', 'dun' or 'dusky brown' (Gilbert-Carter 1964).
Threats
None.
Introduction, neophyte, an invasive, well-naturalised garden escape, sometimes locally dominant.
1946; MCM & D; roadside at Ballagh Crossroads.
May to November.
Growth form and preferred habitats
Growing up to 2.5 m tall with thick, hollow, translucent, succulent, ribbed stems and with buttress rootsto keep it erect, I. glandulifera regularly forms dense pure stands in moist sites, often alongthe banks of waterways, but also in open areas of damp woods, especially where there has been moderatedisturbance, plus in flushes and bogs. It is the tallest annual in the British flora and is branchedabove, swollen at the nodes, with lanceolate to ovate, serrately toothed leaves, 10-20 cm long, heldopposite or in whorls of about three, borne on 8 cm petioles (Grime et al. 1988, 2007; Sell &Murrell 2009).
Introduced to these islands from India as a garden ornamental around 1839, I. glandulifera seedsabundantly and very soon escaped from cultivation. Valentine (1978) says it was first recorded in thewild in 1848, only nine years after its first garden introduction, although another report puts thefirst record in the wild as being 1855 in Middlesex (Kent 1975). It has since then become naturalised inmoist soils in many parts of lowland Britain and, to a lesser extent, throughout Ireland (Beerling &Perrins 1993; R.M. Burton, in: Preston et al. 2002). The spread was so rapid and the plant sodominant that it had been given weed status as early as 1898 (Perrins et al. 1993). This isremarkable considering that, unlike most serious weeds, it is neither a perennial nor a weed ofa*gricultural importance.
A summer annual therophyte with no powers of vegetative reproduction, the plant does however produceabundant seed. These require chilling to break their dormancy, germination being timed to occursynchronously in the spring. On suitably damp soils growth is very rapid and produces tall, dense standstypically of 20 or more plants per square metre. It is tolerant of a wide range of soil texture,structure and chemistry, ranging from free-draining mineral soils to maritime shingle and peat bogs(Beerling & Perrins 1993).
The 'competitive-ruderal', or perhaps better, 'ruderal dominant' growth strategy of I.glandulifera (Grime et al. 1988, 2007), allied at the end of the season with a heavyaccumulation of persistent litter, enables the species to shade out competitors, including previouslyestablished perennials. The accumulating litter then suppresses other seedlings. In this way, I.glandulifera holds its ground and greatly reduces plant species diversity in all forms ofinvaded habitat (Grime 1977; Grime et al. 1988, 2007).
Flowering reproduction
Plants flower continuously from June to October, a period of about 12 weeks. The large, bisexual flowersare borne in a diffuse cyme of 3-12 in the axils of upper branches. They are strongly zygomorphic(irregular) in shape and are pollinated by bumble and other types of bees that crawl right inside thelarge open corolla tube to reach the stout nectar spur. Wasps and moths also frequently visit theflowers and, presumably, they can carry out pollination (Beerling & Perrins 1993). It is unusual foran alien insect-pollinated species to be successful in B & I without having to resort toself-pollination, as most introductions do not manage to attract sufficient suitable insect visitors(Valentine 1978). Having said this, I. glandulifera is self-compatible and can set seed to alesser extent without insect involvement (Valentine 1971). Seed is set from mid-July onwards. Flowercolour varies from purple, through shades of pink, to white, indicating some degree of geneticvariation. After fertilisation, the fruit capsule begins to develop and the whole corolla,including the nectar spur, falls off the plant (Beerling & Perrins 1993).
Seed production and dispersal
Seed production per plant varies with the density of plants in a stand. Medium-sized plants growing at adensity of 20 per square metre produce between 700 and 800 seeds per plant (Salisbury 1964). I.glandulifera generates a peak seed rain of around 1,200 per square metre measured at 2 m fromthe parent plant. Short-distance dispersal is dependent on explosive dehiscence as the 15-35 mm longcapsule dries and ruptures along linear sutures (lines of weakness), firing the contained 1-6 seeds adistance of up to 5 m from the parent plant (Salisbury 1964; Sell & Murrell 2009). The dispersaldistance achieved depends on the height of the capsule above the ground, the direction and strength ofthe prevailing wind at the relevant moment, and the degree of crowding by adjacent stems. Waterside seedmay also spread rapidly with the flow. Upstream movement is slower but does occur; apart from the wind,it may involve seed catching by small rodents or human intervention.
Fermanagh occurrence
From its first recorded occurrence on a roadside in 1946, in recent years, I. glandulifera hasspread along the banks of the Colebrooke, Kesh and Ballinamallard rivers and around the eastern shoresof Lough Erne. It has also appeared along a few damp grassy roadsides, forest paths, in urban and ruralwaste ground, field corners and even into much drier conditions in a neglected area of a demesne yard.It has been recorded in a total of 37 Fermanagh tetrads (7%), 34 of them with post-1975 dates and it isdefinitely actively spreading in the lowlands E of Lough Erne. The fact that it is beginning to appearin drier ground, sometimes quite distant from water bodies, suggests that, like Heracleummantegazzianum (Giant Hogweed), it may be entering a phase of secondary colonisation, duringwhich it may spread into the general countryside.
Weed control
Man has sometimes created favourable conditions for the spread of the species through the destruction ofnatural riverbank communities and unwittingly has facilitated long-range dispersal by trade andtransport of seed-contaminated soil. Fortunately for present and future control of the species, there islittle or no persistent seed bank (Grime et al. 1988, 2007; Perrins et al. 1993). Likeanother eastern alien, Fallopia japonica (Japanese Knotweed), I. glandulifera is sensitiveto late spring and early autumn frosts and it cannot survive prolonged drought. Unlike JapaneseKnotweed, I. glandulifera can tolerate moderate shade, and it can flower and seed in openwoodland and amongst tall reeds (Grime et al. 1988, 2007). In Fermanagh, I. glanduliferahas near ideal conditions as the climate is hyper-oceanic, with typically mild temperatures and evenlyspread precipitation right throughout the growing season.
Weed control of I. glandulifera generally requires at least two applications of herbicide (2,4-D-amine), as the canopy of taller plants may protect stunted individuals. Spraying must be carried outprior to flowering and seeding, since plants sprayed while flowering can still set seed (Beerling &Perrins 1993).
Herbivory
While birds do not appear interested in the seed, sheep and cattle graze and trample all the aerial partsof the plant and a limited range of insects feed and lay eggs on the plants (Beerling & Dawah 1993).Otherwise, I. glandulifera appears immune to attack from both parasites and diseases.
British and Irish occurrence
Common and widespread throughout lowland Britain and most frequent on the banks of waterways. It is stillspreading and has reached more outlying areas including the Isles of Scilly, Orkney and Shetland. As theNew Atlas hectad map indicates, it becomes confined to coastal or near coastal sites furthernorth into Scotland (Beerling & Perrins 1993; New Atlas). In Ireland it is very much morethinly and widely scattered although it is quite a lot more frequently recorded in NI than in the RoI.Even here, the occurrence thins out westwards and southwards, although the mild, damp, temperate oceanicclimate should be very suitable for its growth (New Atlas).
European and world occurrence
I. glandulifera is widespread throughout 18 countries in temperate Europe between 30°N and 64°N.Large populations have been reported from S Sweden, especially along watercourses and it is widespreadand spreading in both Czechoslovakia and Switzerland. It is also widespread in the Pyrenees, the Alpsand around the Mediterranean basin. In the north, it is widespread throughout the Baltic and S & SWparts of Russia (Beerling & Perrins 1993). It has also been introduced and become naturalised onboth islands of New Zealand (Webb et al. 1988).
Threats
I. glandulifera is still spreading and poses a threat to native species in a variety of damphabitats.
Native, common, invasive and often abundant. European southern-temperate. H. helix s.l. has beenintroduced and partly naturalised in N America and New Zealand.
Pre-1739; Henry, Rev W.; Hanging Rock (now a NR).
Throughout the year.
Growth form and preferred habitats
Ivy is such a common and well known plant, its lobed, spear-shaped, evergreen leaf (although extremelyvariable) has become an ornamental emblem in western art and an ancient icon of the Christmas season,probably borrowed from paganism, while in classical times the Ivy leaf was a symbol of fidelity. Theevergreen habit allows Ivy to photosynthesise throughout the year, and the plant climbs vertically bymeans of a clinging 'adhesive' double row of short, unbranched rootlets that are intermittently producedon the stem. Alternatively, it scrambles and sprawls over the ground in woods, thickets and on shadybanks, its stems when growing horizontally, rooting at their nodes. Ivy is a woody-stemmed, perennialthat often surrounds and overtakes old or slow-growing trees in moderately open or less-shaded parts ofwoods of many kinds (Metcalfe 2005; Strelau et al. 2018).
The very wide range of woods and scrubland types in which Ivy plays a part in B & I can be glimpsedby scanning the National Vegetation Classification (NVC), where H. helix features in 21 of the 25defined communities of woody vegetation. It is recognised as a 'constant species' in four of them and isabsent only from a few highland and montane woodland types (Rodwell et al. 1991 Volume 1).
Ivy can survive in deeper shade than any other woodland plant, managing to hold on even under the canopyof well-established Gorse (Ulex europaeus) and Bramble (Rubus spp.) thickets. It is alsothe last species to be excluded by the shade under Rhododendron ponticum (Rhododendron) andTaxus baccata (Yew), and it will climb even these for a metre or two if they are by the edge oftracks and are relatively open at the base allowing sufficient light to penetrate. Ivy tends to be mostfrequent in soils that are mesic with respect to moisture and pH, while preferring medium to highnutrient status. It avoids wetland habitats, waterlogged, very dry or very acid soil (scarcely everbelow pH 4.0), exposed or heavily disturbed sites and, as mentioned above, higher, colder altitudes(Grime et al. 1988; Hill et al. 1999).
In Fermanagh, hedgerow Ivy really is omnipresent. Hedgerow trees and shrubs are more vulnerable to Ivyinfestation than those in the darker environment of closed canopy woodland, since at the first sign ofweakness in their growth, light penetrates the hedge immediately, giving the Ivy that crawls at theirbase the opportunity to launch itself upwards into competition for light and space. Hedgerow Fraxinusexcelsior (Ash) is particularly vulnerable to Ivy invasion since its pale green, compound leafcasts a relatively light shade and it probably bears leaves for fewer weeks than any other tree speciesin the British Isles (Thomas 2000, p. 232).
The widespread modern practice of mechanical hedge-trimming using a tractor with a spinning disk saw,flails and mauls trees and shrubs so savagely that it must severely check their growth for considerableperiods. This also facilitates the Ivy invasion of the hedgerow, initiating competition by the evergreenclimber, leading to its eventual dominance of hedges 'managed' in this brutal and abhorrent manner.
In addition to climbing, Ivy can often equally well cover the ground in shaded or partially-shadedsituations like a springy, evergreen blanket, 15–25 cm deep. Long-established Ivy can form clonalmonocultures, almost completely dominating the herb layer of woods and scrub thickets in this manner.Although as detailed below, Ivy exhibits prolific seed production, its vigorous vegetative reproductionis chiefly responsible for its success as a species (Strelau et al. 2018). Crawling or climbing,the leathery, more or less glossy leaves arrange themselves in a leaf-mosaic with minimum overlap, thuscapturing the maximum amount of sunlight (Melderis & Bangerter 1955).
Associated species
In woods and thickets the main competitors of H. helix are Brambles (Rubus spp.),Honeysuckle (Lonicera periclymenum), Gorse (Ulex europaeus), Bracken (Pteridiumaquilinum) and other ferns with large fronds. Other close associates of H. helix are thepre-vernal and vernal patch- or carpet-forming herbs, such as Bluebells (Hyacinthoidesnon-scripta), Ramsons (Allium ursinum) and, in demesnes at least, Snowdrops (Galanthusnivalis), all of which often grow up through the Ivy regardless of its vigour. In Britain, thislist would of course include Dog's Mercury (Mercurialis perennis), but not so in Ireland (seeRSF's comment in the Allium ursinum species account on this website). It appears that subsp.hibernica is a better competitor than subsp. helix under conditions of woodland shade(McAllister & Rutherford 1990).
Fermanagh occurrence
In winter, in Fermanagh, as indeed elsewhere in W Ireland, Ivy is often the most conspicuous plantspecies in the countryside, being completely hardy in this part of the NW Atlantic coastline. It hasbeen recorded in 483 Fermanagh tetrads, 91.5% of those in the VC. It is almost ubiquitous in woods,hedges, shaded cliffs and on neglected walls and buildings. It can become superabundant, burgeoningeverywhere, growing up and shrouding its natural and man-made supporting structures.
Variation
Recent studies have determined that two forms of Ivy exist, a diploid (2n=48), variously referred to asH. helix, H. helix subsp. helix, or H. helix var.helix with the common name Common Ivy, and a tetraploid (2n=96), called either H.hibernica (G. Kirch.) Bean, or H. helix subsp. hibernica(G. Kirch.) D.C. McClint. and given the common name 'Atlantic Ivy'. McAllister & Rutherford (1990),who have done much of the work to establish the existence of these two forms and define theirdifferences, give them species rank, but An Irish Flora (1996) reckons the distinction onlymerits recognition at the varietal level. The current author (RSF) prefers the compromise positionproposed by Lum & Maze (1989) after their analysis of trichomes in the taxa and consider them asseparate subspecies, a stance also preferred by Stace (New Flora of the BI 1997).
The subspecies are not always easy to separate, but a full account of their differences, written by A.Rutherford, appears in the Plant Crib (Rich & Jermy 1998). One of the differences is odour;Atlantic Ivy has a strong, sweet, resinous, pine-like smell, while Common Ivy has only a weak,disagreeable, usually rather acrid odour. Unfortunately, the current degree of clarity separating theIvy subspecies was not available when the bulk of the Fermanagh fieldwork was carried out and there arenot yet any records that distinguish them. The same situation pertains in the great majority of IrishVCs, and although Alison Rutherford (pers. comm., January, 2002) considers that the Atlantic Ivy is theprevalent form in Ireland and Stace (New Flora of the BI 1997) appears to agree with this notion.How the two taxa are distributed and their degree of overlap is unknown.
Effects of climbing Ivy
Since Ivy does not penetrate the living tissues of the plants it climbs or clambers over and it does notderive any water or nutrient from them, it is classed as an epiphyte rather than a parasite (Holmes1979). Nevertheless, a perennial debate arises as to whether or not Ivy harms other plants or buildingswith its vigorous growth and evergreen shading leaves.
Generally it does not become dominant on woodland trees until they become old and senescent, when theircanopy opens and allows light to penetrate to lower strata. Undoubtedly the evergreen climber indirectlyhastens the demise of trees it smothers, since they are much more likely to catch the wind in wintergales and be felled (Thomas 2000, p.232).
Direct harmful effects, such as branch breakage are sometimes caused on older trees by the physicalburden of the climber and there could also be some degree of restriction on growth of the supportingtrunk due to layers of the rope-like network of H. helix stems gradually accumulating on it.However, despite its specific epithet 'helix' (Greek, meaning 'anything that assumes a spiral shape')(Gilbert-Carter 1964), Ivy does not climb by winding itself around its support. Thus, while it is one ofthe few temperate representatives of a plant family otherwise exclusively tropical in its distribution,the behaviour of Ivy is not analogous to the strangler vines characteristic of tropical forests.
Climbing Ivy does not harm structurally sound walls and buildings but damage does occur if the ivy stemfinds a weakness or an opening it can grow through, since it will then act as a living wedge; then itwill also, given time, become very difficult and expensive to extricate.
On the other hand, ornamental cultivars of Ivy are valued as a means of covering unsightly buildings andit is reputed to be the only climber which does not make walls damp. The leathery evergreen leaves forma sheltering curtain, holding and absorbing rain and moisture, but do not transfer it to the underlyingsupporting structure (Grieve 1931).
Ivy in ancient and secondary woodlands
Although a plant of prehistoric woodland, Ivy is remarkably local in ancient woods (at least in SEEngland), being much more associated with secondary woods (ie woodland on ground that previouslysupported other vegetation or use), and with straightforward plantations or re-plantations (Rackham1980). In his detailed study of well documented, managed ancient woods, Rackham found Ivy had problemsof initial colonisation, but when it does manage to achieve establishment, it radiates inward from thewoodland edge and invades along rides, or into disturbed areas within the vegetation boundary, forinstance, where quarrying has taken place.
In secondary woods, Rackham suggests Ivy is associated with a Hawthorn dominated scrub stage in thesuccession towards woodland development (the NVC community W21, Crataegus monogyna-Hederahelix scrub of Rodwell et al. (1991)), or with more permanent Hawthorn scrub itself(Rackham 1980, p. 353). When Ivy arrives in ancient or well-established woodlands it can persist forcenturies, but Rackham felt that it has little power of dispersal into existing closed woodlandcommunities (Rackham 1980, p. 95) (see below on dispersal ability).
Fossil history
A Danish palaeobotanist has used the occurrence of fossil pollen to develop a technique to construct athermal correlation graph which describes the climatic range of certain species in NW Europe over thelast ten to twelve thousand years (Iversen 1944). He used Ivy and two other species of contrastingbiology, Viscum album (Mistletoe) and Ilex aquifolium (Holly), as his three indicatorspecies. It is perhaps a little surprising that Ivy, being an insect-pollinated species, produces anddisseminates sufficient pollen to be followed accurately in the fossil record preserved in peat and lakemud sediments along with that of wind-pollinated species, but evidently this is the case (Godwin 1975).
Iversen found that the distribution of Ivy is governed by a double form of temperature control; itrequires moderately high summer temperatures, together with winter temperatures not falling below -1.5˚Cas the average for the coldest month. Climbing H. helix plants suffer severe damage at lowertemperatures than this and seed production is also reduced, since growth of Ivy is then restricted tolevels where adequate snow-cover gives protection from the worst winter weather (Godwin 1975). Iversentherefore reclassified Ivy as a eu-oceanic plant (ie 'true-' or 'truly'-oceanic), rather thansub-oceanic (Iversen 1944) and, indeed, the present distribution within B & I does reflect thesefindings, Ivy being most prominent in the Atlantic fringe areas of Europe, where the climate is wetterand milder throughout the year.
H. helix is not only present as fossils throughout the current interglacial period, but is alsorecorded in two earlier ones, called the 'Hoxnian' and 'Ipswichian' in Britain, the former referred toas the 'Gortian' in Ireland. The pollen record for these three interglacial warm periods in B & Isites accords well with those from Denmark (Andersen 1966), especially in that Hedera tends tohave its maximum development early on in the interglacial cycle of rising and falling temperatures, incontrast with Ilex aquifolium. Andersen attributes this chiefly to the development of more acidicmor (ie peaty) soils in the latter part of each interglacial cycle, giving soil conditions that arebetter tolerated by Ilex than by Hedera. Ivy does extend south in Europe and Asia,however, reaching Asia Minor, Palestine and N Iran, and it is also naturalised in N America where it iscommonly known as 'English Ivy' (Clapham et al. 1962).
Leaf and stem morphology
Ivy is dimorphic, having juvenile and mature growth forms of differing leaf shape (ie it is also'heterophyllous') (Rose 1980). When creeping horizontally, it is always completely sterile and has'juvenile phase' deeply lobed leaves in a great range of size, shape and pigmentation. Climbing Ivy,which some describe as 'arborescent' or tree-like, has quite different 'mature phase' adult growth form,with unlobed, or much less lobed leaves (Whitehouse 1992). Given sufficient illumination, the correctday-length regime and a temperature below 16°C, this 'adult' form of the plant will initiate and developflowers in late autumn from September to November (Wallerstein & Hackett 1989). Since the length ofstem internodes changes between horizontal juvenile stems and vertical mature ones, the latter beingmuch shorter, hormonal control involving gibberelins in some way controls the change of growth phase,altering the species' response to light. The juvenile stem is negatively phototrophic, growing away fromlight into dark places, while the vertical stem is positive and grows towards a light source. At thesame time the plant hormone initiates the ability of the plant to flower (Hopkins 1995).
Flowering reproduction
Flowers are produced on well-lit, aerial, adult shoots from about 10 years old (Clark 1983). The small,bisexual, lime-yellow flowers are borne in September to November in compact, globose, terminal umbelsand have their parts in fives (although the five carpels are fused together and there is a solitarystigma). The five sepals are very small, so that the five greenish petals can be mistaken for the calyx.The flowers, which are strongly self-incompatible, can be either protandrous (ie male stage first), orhom*ogamous (ie sexes maturing at the same time). Either way, the terminal, rounded umbels of flowers arecross-pollinated by flies (including hoverflies), Small Tortoiseshell, Peaco*ck and other butterflies,plus moths, wasps and honey bees.
The abundant late-season pollen and plentiful, fully exposed nectar of Ivy flowers are of greatimportance to all these insect species, but especially so for honeybees which avidly collect and storesupplies in the hive comb in order to feed next season's early brood. In late October, other hibernatinginsects, such as queen wasps, also frequent Ivy flowers to collect food (Knight 1962). A succession offurther umbels may be produced below the terminal one from November onwards, and functioning flowers mayeven be found in December, but the chances of pollination decreases with advancing cold weather andconsequent fewer active insect visitors (Knight 1962).
Since flower induction requires a minimum light level, H. helix never blossoms and fruits wherethe plant occurs most abundantly, ie in the deeper shade of woods and thickets (Whitehouse 1992).Despite the popularity of the flowers with insect visitors, examination of umbels reveals the levels offertilisation achieved are far from ideal. Although the branch tips may bear 50 or more flowers, oftenonly 40-50 % of these develop fruit, even in a mild winter (Knight 1962).
The fruit is a 6-8 mm diameter, green, turning black or deep purple, urn-shaped berry, which does notfully ripen until the following spring. Early ripening fruit may wither and abort if exposed toexcessive cold (Snow & Snow 1988). The berry is smooth, succulent, thin-skinned and contains purplepulp and up to five rather soft seeds, although generally fewer than this are produced since some of theovules abort. The main berry crop ripens in March and April when the weather is becoming milder and foodsupplies for birds are already plentiful.
Seed dispersal
Since Ivy berries are among the most nutritious wild fruit available in B & I, having an especiallyhigh fat content, they are keenly sought, eaten and transported by the members of the Thrush family(Simms 1978, pp. 108-11), plus Robins, Blackcaps, Starlings and Woodpigeons. Where the latter arecommon, a very high proportion of Ivy seed, perhaps up to 75% of the total production, is destroyed inthe birds' stone-filled crop by being taken when the berries are unripe (Snow & Snow 1988, p. 33).
Since berries can contain more than one seeds, and birds often consume numerous fruits at a time, asingle bird dropping may deposit numerous seeds and give rise to clusters of seedlings. As it grows, Ivyoften continues to maintain its own company and many stems, often of different ages, may be observedtogether climbing trees, pillars and walls (Metcalfe 2005).
Toxicity
Although the bird species mentioned above eat the berry pulp without harm and Blackbirds feed the pulp totheir young nestlings (Ridley 1930, pp. 408-9), all parts of the Ivy plant contain saponins which arebroken down with loss of sugars to form toxic substances called 'hederins'. In sufficient quantity,these poisons are harmful to mammals including farm stock, producing vomiting, diarrhoea, muscularspasms, paralysis and even coma. The sap of the plant is also dangerous and can give rise to irritantand allergic dermatitis (Cooper & Johnson 1998).
Despite this, ivy has a history dating back to the ancient Romans of use as fodder (Grieve 1931;Troels-Smith 1960); it was collected in winter, when grazing became scarce, and fed to cattle inparticular, but sometimes also to horses and sheep (Kelly 1997). There are plenty of folklore recordsattesting to this fact and, indeed, Ivy was also used as a medicine for ewes that were poorly afterlambing (Vickery 1995). Quite apart from providing spring food for birds, climbing 'arboraceous' Ivyalso provides ideal homes for nesting birds, the dense evergreen foliage giving them and their youngdry, secure shelter.
Seed germination and seedling establishment
Ivy seeds are quite large, weighing on average 20.4 mg (more than twice the weight of Holly seed forexample) and they can germinate immediately on being sown in the spring, the soil seed bank being onlytransitory (Grime et al. 1988, 2007). Germination is inhibited somewhat by light (Grime etal. 1981) and establishment from seed in woodland shade is probably a rather slow, prolongedprocess (Lang 1987, p. 134). A study of the behaviour of Ivy seedlings and the factors influencingestablishment of the species from seed in existing woodland is sadly lacking and would be a veryworthwhile project. Although quantitative measurements do not appear to exist, subjective observationssuggest that establishment from seed is likely to be rare, and while bird populations do provide vectorsfor jump dispersal, the frequency of this happening is, as usual, completely unknown.
Vegetative reproduction
In comparison with establishment from seed, vegetative spread is very obviously significant once Ivy hasarrived at a site, and it remains to be shown what factors influence the balance between and theeffectiveness of sexual and asexual reproductive processes in Ivy. Ivy can invade garden hedges,including clipped evergreen privet. It does so from a previously established base on adjacent soils ofmore open or rocky ground, or sometimes from the base of adjacent walls. Grime et al. (1988,2007) point out that this ability to extend over soil-less habitats from a base rooted locally in soilis unique within the flora of B & I, but the current author (RSF) suggests this is a matter ofdegree, since brambles can also behave in much the same manner.
H. helix climbs straight up trees, poles, walls and rock without twining, adhering by means ofvery short adventitious roots on the shaded side of the stem, which attach the stems to bark or anyother surface irregularities by secreting a sticky substance; subsequently they keep their grip withgreat tenacity. The climbing stem obtains all its water and dissolved mineral nutrients in the normalmanner, through a taproot in the soil (Brimble 1962). Climbing Ivy stems can be immensely long-lived,with reports of them developing trunks 90 cm in circumference (Simpson1989), or greater 'than a fat man' (Rackham 1986, p. 117), and of being up to 400 years old(Rose 1980, p. 12). The weight of ivy removed from a single tree in Olympic Park, Washington wasestimated to be 953kg (Simon 2002). The typical climbing Ivy stem, however, branches veryfrequently to form a interlaced network of much narrower diameter than the giants just mentioned, thelower, older parts of the stem being clothed with a buff coloured 'fur' of adventitious rootlets, eachabout 2-3 mm long (Metcalfe 2005; Strelau et al. 2018).
Herbivory and disease
A number of insects, including a weevil and a moth, have Ivy as their principal host food-plant, but themost prominent feeder is the Holly Blue Butterfly, Celastrina argilus, which attacks the buds,flowers and fruit of Holly in the spring generation, and switches to those of Ivy in the summergeneration (Asher et al. 2001). One insect, the aphid, Aphis hederae, feeds exclusively onIvy leaves and stems and is, therefore, described as 'monophagous' (Peat 2002: Ecological FloraDatabase, York). Ivy leaves are also attacked by at least five species of microfungi which causespots, the most common species probably being Colletotrichum trichellum (Ellis & Ellis 1985).
In NI, very rarely, Ivy roots host the total parasite Orobanche hederae (Ivy Broomrape) itself aflowering species which is more frequent further south in both B & I, and is chiefly coastal in theformer (Perring & Walters 1976; Webb et al. 1996; Stace 1997; Preston et al. 2002). Ithas been suggested, perhaps somewhat tentatively since information is scarce, that Ivy Broomrape is morefrequently found associated with Atlantic Ivy, H. helix subsp.hibernica, than with Common Ivy, subsp. helix, since the distributions of the two plantsappear to be quite strongly correlated (Rutherford 1985).
British and Irish occurrence
H. helix is a widespread and abundant species throughout B & I, except in parts of theScottish Highlands since it is essentially a lowland plant, reaching a maximum altitude of around 610 min Isla and Atholl (Wilson 1949; Garrard & Streeter 1983). It reaches the same altitude in theMourne Mountains, Co Down (H38)(Hackney et al. 1992). H. helix is recorded as introducedin both Orkney and Shetland (Preston et al. 2002).
European and world occurrence
Although hardy and apparently completely so in western parts of the British Isles, the distribution ofH. helix is undoubtedly climatically limited in parts of Europe, where it is native in 30countries. It declines to rarity in the N & C parts of Europe which suffer cold winters and it isentirely absent, for example, from most of Russia, E Poland, Iceland, the Faeroes and Finland. In SEEurope it reaches Ukraine (Metcalfe 2005).
H. helix has been introduced in India and South Africa and become naturalised in Australia, NewZealand, Hawaii, Brazil, Canada and the United States (Larocque 1999, cited in Grivet & Petit 2002,quoted in Metcalfe 2005).
Medicinal uses
The value of Ivy is not much regarded by many working in traditional, orthodox medicine today, but in thepast it was used in cases of dysentery and jaundice, and a paste of Ivy boiled in butter was even use torelieve sunburn (Grieve 1931). More recent medicinal folklore suggests a poultice using boiled Ivyleaves is effective in treating corns, or again boiled in butter will reduce or remove burn scars. Inview of the fact that Ivy can cause contact dermatitis, it is interesting that it has been recommendedin herbal medicine for cooling eczema, and also for healing a scalp rash (Vickery 1995).
At present, Hedera species are being examined as a potential cure for the uncomfortable andsometimes fatal condition known as leishmaniasis (Ridoux et al. 2001; Abbasifar et al.2017). This condition occurs in tropical and sub-tropical regions around the world and affects over400,000 people annually. Ivy is most frequently used today in the hom*oeopathic treatment of a number ofcommon conditions and diseases, and extracts of the plant are also being examined as possibleingredients in cosmetic creams and as a possible tumour reducing agent (Rose 1996). Studies have shownIvy extracts to provide effective control against liver fluke (Julien et al. 1985). HederasaponinB extract from H. helix can also be used as a novel drug candidate with antiviral activityagainst subgenotypes of enterovirus 71 (EV71), the foremost source of hand, foot and mouth disease (Songet al. 2014).
Garden use
Ivy has a very long history as a garden plant. The Classical Roman writer Pliny the Elder reported thatTheophrastus, around 314 BC stated that Ivy did not grow in Asia Minor, but that Alexander the Great hadcome back from India wearing wreaths of Ivy because of its rarity (Rose 1980). Pliny also described thecultivated Ivies he knew, including the first mention of a variegated form and of a stiff, erect 'Treeivy', which stands without a support (Rose 1980). Eleven species of the genus Hedera are ingarden cultivation (Griffiths 1994) and Rose (1980) reckoned that about 200 cultivars exist, the greatmajority being forms of H. helix.
Names
The genus name 'Hedera' is an ancient classical Latin name for the plant (Gledhill 1985). In Greeklegend, a crown of Ivy intermingled with flowers was first used by the God of Wine, Dionysus (or hisRoman equivalent, Bacchus), such wreaths initially being reserved for the gods, but gradually their usespread to sacrificial animals, to priests and, finally, to the faithful worshippers themselves. In somecults, Dionysus bore the designation of 'kissoi' (Ivy), because legend said that as a baby he had beenwreathed in Ivy by the forest nymphs (Baumann 1993). The connection with alcohol continued intohistorical times when English taverns bore over their doors or on a pole the sign of an Ivy 'bush', toindicate the excellence of their liquor: hence the saying 'Good wine needs no bush.' (Grieve 1931).
Considering how common and conspicuous a plant Ivy is, and how long it has been associated with man hisanimals and his dwellings, it is rather surprising that it appears to have accrued so few English commonnames. Grigson (1955, 1987) lists only seven, while Britten & Holland (1886) manage ten, includingthe one with which we are most familiar 'Ivy', which is derived from the Old English 'ifig' (Grigson1974). There are several close variants of it among the English common names rounded up by Grigson,including 'Ivin', 'Ivery', 'Ivory', 'Eevy' and 'Hyven'.
An alternative suggestion for the origin of the name 'Ivy' is that it might be derived from the Latin'ibex' meaning 'climber' (Rose 1980, p. 18). Several other names refer to the woody character of theplant and the idea that Ivy binds and bends around the objects it climbs, for example, 'Bentwood',Bindwood', 'Woodbind' and 'Benewith Tree', names it often shares with Lonicera periclymenum(Honeysuckle) (Britten & Holland 1886).
Threats
None.
Native, common and widespread. Sub-oceanic southern-temperate.
1881-2; Barrington, R.M.; Co Fermanagh.
February to December.
Growth form and preferred habitats
One or more orbicular, crenate, peltate leaves, 8-35 mm in diameter, borne on sparsely hairy petioles upto 25 cm long, arising from slender, pale green stems, long-creeping in mud or floating in water androoting at the nodes, make Marsh Pennywort a readily recognised aquatic perennial. H. vulgarisgrows in usually sunny, open, moist or wetland situations and is extremely common and patch-forming in awide range of moderately acid, infertile muddy ground, or in short-sedge sward of Sphagnum orother species of moss-lawn, and in shallow, still aquatic habitats including ditches. It is onlyoccasionally found in half-shady situations, but it can form an understorey to taller species such asJuncus effusus (Soft-rush) and in Phragmites reed beds (Grime et al. 1988, 2007;Jonsell & Karlsson 2010).
Taxonomy
The genus Hydrocotyle has been moved about from family to family on numerous occasions. It hasbeen included in the Apiaceae (= Umbelliferae) (eg Tutin 1980; Stace 1997; Sell & Murrell 2009), orthe Araliaceae (Ivy family) (eg Jonsell & Karlsson 2010), or sometimes it is given its own family,the Hydrocotylaceae (Pennywort family). A morphological study of a few representatives of the twoclosely related families Apiaceae and Araliaceae indicated that neither is monophyletic (Judd etal. 1994, quoted in Jonsell & Karlsson 2010). Hydrocotyle was found to be in the sameclade as Araliaceae based on molecular studies (Chandler & Plunkett 2004, quoted in Jonsell &Karlsson 2010). A morphological character uniting Hydrocotyle with Araliaceae is the hardendocarp on the seed or mericarp (Jonsell & Karlsson 2010). However, most members of the Araliaceaeare woody, have either one or five styles, and the fruit is a berry (Sell & Murrell 2009). None ofthis fits Hydrocotyle which lies rather better in a subfamily within the Apiaceae, theHydrocotyloideae, although it differs from the rest of the family in possessing leaf stipules and thefruit does not have oil bodies. In truth, Hydrocotyle appears anomalous in either of the twoclosely related families and while Sell & Murrell (2009) preferred to keep the genus within theApiaceae, Stace (2010, 2019) decided to recognise the Hydrocotylaceae.
Fermanagh occurrence
Patches of H. vulgaris are a very familiar sight around Fermanagh since this truly amphibiousperennial has been recorded in 234 tetrads, 44.3% of those in the VC. It is particularly frequent aroundthe sheltered, grazed lowland shores of Upper Lough Erne, where the muddy-organic substrates, seasonallyflooded water-meadows, marshy pastures with temporary wet depressions, swampy fen and fen-carr, andshallow, eutrophic, peat-brown lakeshore waters must provide near-ideal growing conditions for thespecies.
In both lowland and more upland bog sites, such as for instance on the Western Plateau, it is commonlyscattered on the margins of pools and peaty ditches, as well as in more moderately nutrient-rich,flushed organic muds on grassy moorlands, blanket-bogs and heaths. Here it grows where taller, shadingand more vigorous species are limited or excluded by grazing, trampling, periodic flooding, or otherforms of disturbance or exposure, and also by the restricted availability of plant nutrients (Grimeet al. 1988, 2007).
Despite its leaning towards wet to moist, moderate to mildly acid, often infertile conditions, H.vulgaris does occur in some more calcareous situations too, eg around the shores of Lower LoughErne, in pools on the distinctly marly River Finn and around Rooskey turlough (ie a 'vanishing lake' inlimestone terrain, where drainage is vertical into a subterranean cave system). In Scandinavia, Jonsell& Karlsson (2010) consider H. vulgaris is indifferent to lime.
In the frequently wet, mild, oceanic climate of Fermanagh, where prolonged drought is very rare, MarshPennywort can also occur in a purely terrestrial mode on much drier, but still moist or constantly damppeaty banks, for instance in open areas in wet deciduous woods, eg along paths in the Correl Glen NR andthe Cladagh River Glen NR (also known as the Marble Arch NR).
Vegetative reproduction
The slender creeping horizontal shoots of H. vulgaris root at the nodes and form more or lessextensive clonal patches, allowing the plant to quickly colonise adjacent ground bared by any form ofdisturbance (Grime et al. 1988, 2007). Fragmentation of the stem as a result of grazing,trampling or flooding, also enables detached material to disperse by water and re-establish the plantvegetatively in fresh sites.
Flowering reproduction
Although the casual observer will always be unaware of it, the plant may produce its minute, extremelyreduced, insignificant, well-hidden flowers from June to August in very small inflorescences, onpeduncle stalks only about half the length of the leaf stalks (ie 3-18 cm high). The little axillaryumbel clusters are borne on these short leafless stalks springing from nodes on the horizontal stems.There are three to six almost sessile flowers in each very imperfect umbel, which is sometimes elongatedto form a secondary cluster. The 3 mm diameter flowers have very small or no calices, and pink orgreenish-white, entire petals. The tiny schizocarp fruit measures 1.5 × 2.5 × 0.7 mm and has two muchflattened, keeled, nearly circular carpels and four lateral ridges on the twin mericarps (Melderis &Bangerter 1955; Butcher 1961; Sell & Murrell 2009). In these very inconspicuous flowers,self-pollination is possible and probably obligatory. Although the anthers mature in succession, thestigmas mature before the last of the five anthers bursts to release its pollen, allowing selfing tooccur.
Irrespecitive of how pollination and fertilisation is achieved, seed regeneration in H. vulgarismay be less important than vegetative reproduction in many situations, the plant often being flowerlessdue to excessive competition (Grime et al. 1988, 2007). Seed is still significant in the longerterm, however, both to maintain the species genetic variation and for long distance dispersal betweenalready colonised and fresh or newly created habitats. No persistent soil seed bank has been detected.
Mericarp fruits float and, in addition, adhesion in mud on the feet or feathers of ducks or otherwaterfowl has been proposed as the external transport mechanism for short distances; seed recovered fromthe gut of wide-ranging, migrating ducks such as Gadwall, suggests that they may be the vector to muchmore isolated stations (Ridley 1930; Grime et al. 1988, 2007).
British and Irish occurrence
The subfamily to which H. vulgaris belongs, the Hydrocotyloideae (previously treated bytaxonomists as a separate family, the Hydrocotylaceae) is mainly distributed in the S hemisphere andH. vulgaris is the only native member in B & I. H. vulgaris is very common andwidespread in most of Ireland, but becomes significantly less frequent in the drier and more fertile SEpart of the country. It is also very widespread at all latitudes in Britain, but again there is a markedspecies predominance in the wetter and more acidic environs of the west and it is absent or rare inlarge areas of the English Midlands and N Scotland (Tutin 1980). Drainage and development since aboutthe 1960s has wiped out many former sites of the species, especially in SE England (M. Southam, in:Preston et al. 2002).
European and world occurrence
Beyond the shores of B & I, H. vulgaris is mainly confined to W, C & S Europe, thegreatest frequency being in W & C regions. Rather unusually for a plant quite widely distributed inEurope, the species does not extend appreciably into Asia (Sculthorpe 1967). It does occur, however, inW Iceland, S Scandinavia to around 60°N, in the remote Azores and previously in N Africa, although ithas not been seen there recently (Tutin 1980; Hultén & Fries 1986, Map 1382; Sell & Murrell2009). Previous reports of the species in New Zealand are rejected and discounted by Webb et al.(1988).
Names and folklore
The English common name 'Marsh Pennywort' is the most frequently one applied to H.vulgaris (an obvious reflection of its orbicular leaf shape), but there are at least ninealternative common names applied in different regions around B & I (Grigson 1955, 1987). Otherunderstandable common names include 'Fairy Tables', and several that refer to a once widespreaderroneous belief that sheep grazing the plant were liable to contract flukeworms causing liver rot,hence names such as 'Farthing Rot', 'Flowkwort', 'Sheep Rot', 'Shilling Rot' (another coin reference),'Water Rot' and 'White Rot' (Grieve 1931; Grigson 1955, 1987). Another plant that has been similarlyaccused is Ranunculus flammula (Lesser Spearwort), which does contain acrid toxic protoanemonin,although it really does not affect stock to any observable extent, as they avoid consuming it (Cooper& Johnson 1998).
There is a suggestion that H. vulgaris might be the plant used by herbalists in Co Limerick fordressing burns, although, again, that might be another mis-identification for the more likelyPotamogeton natans (Bog Pondweed) (Allen & Hatfield 2004)
Threats
Drainage and development, including the intensification of agriculture. Not greatly threatened at presentdue to the upland nature of the majority of its sites, but lowland habitats are particularly affected bycontinuing changes in land management.
Native, common. European temperate, but also native in C Asia.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This erect, evergreen, rosette-forming long-lived perennial herb has a stout rootstock or verticalrhizome and numerous basal leaves, 2-6 cm in diameter, palmately 5-lobed, on petioles 5-25 cm long. Thewhitish inflorescence consists of a number of simple umbels each about 5 mm across, often giving theappearance of a compound umbel (Sell & Murrell 2009). The plant produces isolated clumps or morecontinuous patches in constantly damp, shaded (occasionally very deeply shaded), woodland situations.Having said that, Sanicle is more often found in somewhat better-lit, shady conditions, eg on exposedbanks and beside roads, paths and tracks in woods and glens, rather than found hiding under the verydeepest shade beneath Yew trees! It is difficult to distinguish, however, whether the difference infrequency of the species is due to light, moisture, or both of these factors, since even given the heavyannual rainfall of Fermanagh, the soil under the dark, evergreen canopy of Taxus trees isgenerally too dry for most ground storey species to tolerate, apart that is from Ivy (Hederahelix). S. europaea is sometimes associated with slopes where there is ground waterflushing the soil and it is sensitive to grazing and other forms of disturbance (Jonsell & Karlsson2010).
Sanicle very much prefers moderately fertile deciduous woodland and hedgerow soils developed oncalcareous or base-rich parent rocks, the substrate typically having a pH above 6.0 (Inghe & Tamm1985). In woods, these soil conditions support either pure, or more commonly, mixed stands of ash,hazel, beech (planted) and oak. S. europaea can be found growing less frequently on somewhat moreacidic loam substrates, down to a minimum pH around 4.5, eg on brown earth soils under oak in moreupland glens in Fermanagh (Grime et al. 1988, 2007).
Fermanagh occurrence
S. europaea is common in suitable damp, shady sites and is widespread throughout Fermanagh, beingrecorded in 243 tetrads, 46% of those in the VC. Apart from riverbank woods and hedges, which itcommonly frequents, in Fermanagh S. europaea is chiefly absent from wetlands, the more acidicpeaty soils in the Western Plateau uplands, plus in the area of better farmland on fertile soils to theE of Lough Erne. In all these latter situations, even in shade, it cannot compete with more vigorousspecies.
Vegetative growth and individual persistence
Sanicle possesses a short, stout rootstock or rhizome that grows very slowly and displays sympodialgrowth, ie a shoot from a lateral bud takes over the vegetative extension growth after flowering. Thereis no overwintering bud and leaves are produced and develop sequentially, some of them remaining greenand overwintering. Possession of the rhizome enables established plants to persist for an unknown period– perhaps or, indeed, probably, indefinitely. A pioneering 43-year population study of S.europaea was made by Prof Carl Tamm in a Swedish spruce forest (Tamm 1956; Inghe & Tamm1985, 1988). This proved that the horizontal rhizome is extremely slow-growing, the older portion of itdying away after 5-20 years. This allows a very "slow-motion" form of vegetative reproductionto take place by bifurcation, ie growth involving branching and separation as older parts of theindividual plant die off. Since the estimated half-life of an individual in this species is between 74and 221 years, similar to or longer than the lifespan of the canopy trees above it, replacement of theindividual genetic ramet is not a high priority.
Flowering reproduction
Flowering takes place in B & I from May to August. Small, globose, simple umbels are arranged in anirregular cyme, in appearance resembling a compound umbel. Two forms of flowers are borne together ineach head, male ones on short pedicels and three-six sessile bisexual ones. In each flower, the fivesepals are conspicuous and longer than the five white or pinkish petals. The five stamens have whitefilaments and pink anthers and the paired styles are divergent or somewhat curved, the stigmas capitate,and the ovary has a ± flat disc at its apex. Pollination is carried out either by visiting small fliesand beetles, or by selfing (Sell & Murrell 2009). The fruit is also small, about 3 mm, ± rounded andcovered with rigid, forward-pointed, hooked bristles which help it to spread on animals as a burr (Tutin1980; Jonsell & Karlsson 2010).
Seed is the main method of S. europaea reproduction and it provides the only really effectivemeans of dispersal. However, flower, fruit and seed production in Sanicle depletes stored resources tothe extent that the plant often rests for a year or more before flowering again. Flowering ischaracterised therefore by being strikingly irregular (Tamm 1948, 1956). Despite their small sample ofspecies data and incomplete meteorological records, Inghe & Tamm (1985, 1988) showed that in theSwedish forest they studied, ramet mortality, especially of juveniles, is very sensitive to summerdrought, and it adversely affects the probability of the plant flowering the following year (Tamm 1948).
It was found that single ramets often have a pattern of every-second-year flowering out of phase witheach other and sometimes there are repeatedly longer periods of three or four years between flowering(Inghe & Tamm 1988). These Swedish workers concluded that this interacts with the "uneven"levels of flowering resources of the plants in a population in a manner akin to "masting" inwoody species like Beech and Bilberry (see the current author's accounts of these species on thiswebsite). In the case of S. europaea, however, the uneven flowering and seed production behaviourcannot be explained in terms of a seed predator avoidance mechanism. S. europaea has a highreproductive effort per flowering season, resulting in high costs of reproduction measured in reductionof subsequent flowering. If the individual ramet is not very strong, as is often the case in heavilyshaded woodland settings, and the environmental conditions are not unusually favourable, one or moreyears of recovery was needed before the Sanicle plant could muster a new flowering (Inghe & Tamm1988).
Seed persists in the soil and seedling mortality is, as usual, high, but growth of plants to maturity isalso very slow, most plants requiring, or having to wait in a suppressed condition for between eight and16 years before their first flowering (Tamm 1956). This makes S. europaea a classic example of anecological 'stress tolerator' in the model of Grime (1979), most of the stress apparently deriving fromshade and occasional drought periods.
British and Irish occurrence
The pattern and frequency noticed in Fermanagh is reflected throughout lowland B & I, the speciesbeing common and widespread in suitable moist, shady, neutral to mildly acid habitat conditions andabsent from wetlands and extreme acid or heavily disturbed ground, including farmland. However, althoughthere is no local evidence of it in Fermanagh, there has been some population decline of S.europaea across B & I since the 1960s (eg the New Atlas Change Index = -0.98).Presumably, this is due to losses of woodlands and of larger, older hedgerows (M. Southam, in: Prestonet al. 2002).
European and world occurrence
S. europaea belongs to the European temperate element and is restricted to Europeand adjacent parts of Africa and Asia. It is widespread in wooded regions of Europe, plus S, C & EAsia and N Africa. In the Mediterranean region and in the Caucasus, it is confined to the mountains.Also present in the mountains of Tropical Africa, and in S Africa (Hultén & Fries 1986, Map 1383;Sell & Murrell 2009).
Uses and folklore
Sanicle is described by Allen & Hatfield (2004) as, "an ancient panacea". S.europaea was praised so highly in the old herbals that its folk use raises the strong suspicionthat its many (almost universal) uses were entirely derived from the written tradition, being copied,embroidered and inflated from one herbal to another without any real world experience or evidence toback up the claims made. The main claim was that the plant could stop or staunch bleeding from wounds,ie it was considered a vulnerary. In the Scottish Highlands it was recommended for both bleeding woundsand ulcers, and in the Isle of Man for haemorrhages, dysentery, bruises and fractures. In Ireland, in CoLondonderry it was applied to cure bleeding piles, while in Co Donegal it was used to treat consumption(Allen & Hatfield 2004).
Lyte (1578), whose publication is a translation from Dutch of Rembert Dodoens' entitled A nieweHerball, or Historie of plantes, said that, "The iuyce of Sanicle drunken, doth makewhole and sound all inward, and outwarde woundes and hurtes." (Grigson 1974). With such a positivereputation it was sometimes called 'Self Heal', a name today applied to Prunella vulgaris of theLabiatae, a completely separate family of plants. Grieve (1931), who, in the view of the current author(RSF), gives the most complete modern account of herbalism, says, "As yet no analysis has been madeof this plant, but evidence of tannin in its several parts is afforded by the effects produced by theplant." She goes on to list uses of Sanicle as an astringent, 'alternative', such as it being givenin combination with other herbs (including Yarrow (Achillea millefolium) and Bugle (Ajugareptans)) "in the treatment of blood disorders". It is also recommended "for allchest and lung complaints, chronic coughs and catarrhal affections, inflammation of the bronchi,spitting of blood and all affections of the pulmonary organs".
Despite all this herbal acclamation, there is no entry for Sanicle in the B & I folklore compendiumsof Vickery (1995, 2019), nor for Ireland, of Mac Coitir (2015).
Names
The genus name 'Sanicula' is from the Latin 'sano' or 'sanus' meaning 'heal' or 'whole' andbest translates as 'healer' (Gilbert-Carter 1964; Gledhill 1985). On the other hand, Prior (1879, p.206) holds that to derive the genus name directly from the Latin 'sanare', 'heal' is incorrect andimpossible on the principles of etymology. Prior also points out that 'Sanicula' does not appear inClassical Latin writers and there is no such word as 'sanis' or 'sanicus' from which it could have beenformed. He goes on to query whether the name has its origin in Latin or German and he concludes the oddsare even. He also suggests the name may have arisen as a corruption of 'Saint Nicolas', called in German'Nickel', ie deriving 'Sanicula' from the German 'Sanet Nickel' or 'San Nicola' (Prior 1879).
The English common name 'Sanicle' is 15th century and is derived from the Old French 'sanicle', from themedieval Latin 'sanicula', a diminutive formed from Latin 'sanus' meaning 'whole' or 'sound', both namesbeing derived from the (most likely undeserved) past reputation the plant had as a wound herb (asdetailed above) (Grigson 1974). Apart from various forms of spelling of the name, eg 'Sanikle','Sinicle' and 'Sanikel' (Britten & Holland 1886), there appears to be one main English common namefor this species, although it does get supplementary labelled 'March Sanicle' and 'Wood Sanicle', butGrigson (1955, 1987) manages to dredge up 'Wood Elder' from Somerset.
Threats
Possible reduction in presence due to the destruction of woodlands and removal of hedgerows.
Introduced, neophyte, extremely rare and locally extinct. European temperate, very sparsely naturalisedin eastern N America and New Zealand.
1902; Abraham, J.T.; roadside at Lisgoole, Upper Lough Erne.
Growth form and preferred habitats
Although C. temulum is rather similar in appearance to Anthriscus sylvestris (Cow Parsley),Rough Chervil is distinguished from all other umbellifers by the purplish patches, or almost entirelypurple, rarely greenish, sometimes distinctly glaucous, hairy, solid (not hollow), angled stems that areslightly swollen below the nodes, and later by the mericarp fruits that are c 5 mm long, narrowingupwards, and dark brown with broad, rounded, lighter coloured ridges on them (Perring & Walters1989; Sell & Murrell 2009; Jonsell & Karlsson 2010). A robust, tall, up to 100 cm high, ratherhispid (hence 'rough') biennial, or rather, a monocarpic perennial (ie taking at least two, but usuallymore years to reach flowering capacity), C. temulum usually flowers later in the season thanA. sylvestris – in June and July, and before Torilis japonica (Upright Hedge-parsley), sothat these three umbellifers are not often confused (Grime et al. 1988, 2007).
Common and locally abundant in Britain, but rare in Ireland, C. temulum occupies the very samerange of habitats as the more widespread A. sylvestris, namely on open ground at the edge ofwoods and in hedgerows, along disturbed, roadside verges and in rough grassland or open scrub (thelatter particularly when near water). Also, it most often occurs on dry to damp, well-drained,moderately fertile and moderately acid to neutral or calcareous soils, in sun or half-shade (Garrard& Streeter 1983; Sinker et al. 1985). C. temulum has probably declined in Britain fromthe effects of eutrophication, which has led to a great increase in competition from species such asGalium aparine (Cleavers) and Urtica dioica (Stinging Nettle) (Braithwaite et al.2006). The established strategy of C. temulum is described as CR (meaning Competitive Ruderal) byGrime et al. (1988, 2007), which suggests it can readily enough cope with moderate levels ofcompetition in disturbed habitats, but nutrient enrichment greatly increases the level of stress itwould face from the likes of G. aparine and U. dioica.
Fermanagh occurrence
There are only two records of this species in the Fermanagh Flora Database: the one dating from 1902(listed above) and a second from another roadside at Cromaghy Lough in the 1945-53 period when Meikleand co-workers visited the county. The current author (RSF) and RHN firmly believe that in Ireland thisis always a very rare species of marginal, rough ground habitats and a noticeable 'follower of man',which has become locally extinct in Fermanagh since about the 1960s.
Flowering reproduction
C. temulum flower heads include both male and bisexual (hermaphrodite) flowers and, again, likeA. sylvestris, they attract as pollinators a range of unspecialised insects including beetles,flies and bees (Fitter 1987). Seed behaviour also follows the pattern of A. sylvestris, whilelongevity in the soil is described as transient or short-term persistent, ie survival is greater thanone year, but less than five (Thompson et al. 1997). Germination after chilling to breakdormancy, takes place chiefly in the spring.
Variation
Two varieties have been recognised in B & I by Sell & Murrell (2009). Var. temulum is ashort plant with a slender rootstock. The lower stem has spreading, long eglandular hairs with shortones in between them, but never forming a dense clothing tomentum. The inflorescence usually contains upto eight umbel rays, rarely up to 12. This contrasts with the much rarer, probably introduced var.canescens (Benitez ex Thell.) P.D. Sell, where the plant is often tall and has a stout rootstock.The lower stem differs in having a covering of dense, short, white eglandular hairs; less hairy upwards.The inflorescence in this variety consist of up to 17 umbel rays. In Britain, var. canescens isonly known from Cambridgeshire (VC 29) and appears to be an introduction from C Europe, probablytransported recently with horticultural material (Sell & Murrell 2009).
British and Irish occurrence compared
The complete contrast between Rough Chervil's behaviour and status in Ireland and in Britain is bothpuzzling and rather astonishing. In England and Wales, C. temulum is widely distributed and verycommon to local in lowland areas, but in Scotland it is more or less confined to the coast, especiallythe E coast, and to the vicinity of the two large north-of-the-border conurbations. In Ireland, on theother hand, C. temulum is very rare and extremely thinly scattered throughout, with a slightlygreater frequency towards the E, again possibly related to the marked anthropogenic character of thisumbellifer, ie it demonstrates a strong association with greater levels of human disturbance in areas ofdense urbanisation (New Atlas; Cat Alien Pl Ir).
There are parallel situations in numerous other species, including for instance in the same family,Clinopodium vulgare (Wild Basil). The most notable example of a major distribution difference inthese isles is possibly the perennial Mercurialis perennis (Dog's Mercury). In each case, acommon and widespread native British species is by comparison virtually absent or rare in Ireland and isconsidered an alien introduction. The current author (RSF) does not imagine that M. perennis isnative anywhere in Ireland, although others, including Parnell & Curtis (2012), join Scannell &Synnott (1987) in suggesting it may be so in "one or two places in the Burren", Co Clare (H9).
Species comparisons
C. temulum and Anthriscus sylvestris are not only related, alike and appear in the samerange of habitats, they also exhibit the same ecological tolerances and requirements as described bytheir Ellenberg Indicator Values (Hill et al. 1999). The fact that both of these umbellifers arechiefly found in linear, disturbed ground, roadside situations, preferably on basic, calcareous soils,makes the enormity of the difference in their Irish representation even more striking.
Species distribution differences like these are extremely difficult to comprehend, never mind explain!Surprisingly little is really known about plant species behaviour (ie the sum of their biology, ecologyand local history). It is very difficult to predict a species' competitive ability in newly invadedterritory, the consequent degree of success it may or may not have in maintaining a persistent colony,followed by effective dispersal from that spot to other available suitable sites, where it must again becapable of establishing viable breeding populations to achieve long term survival. Furthermore,insufficient data exist about the properties of this and many other wild flower species, in order toallow us to understand how they react to changes in the environment. Even a little more local naturalhistory observation might help clarify matters in many instances where information is scarce or totallylacking.
Status and decline in Ireland
The situation becomes even more baffling when one realises that even as a naturalised introduced species,C. temulum has a previous history of somewhat greater prevalence in Ireland prior to 1945. Colganat least recognised quite early on that Rough Chervil was probably a 'denizen', or only doubtfullynative (Cybele Hibernica 1898). This necessarily subjective assessment has been widely acceptedby the majority of Irish field botanists (eventually including even Praeger). Others reject thisopinion, despite it being endorsed by Scannell & Synnott in their listing in the Cen Cat Fl Ir2; other Irish local Floras, including for instance, Booth (1979) in Co Carlow (H13) andthe entire sequence of six publications on the three county FNEI dating from 1888-1992 (ieincluding the three supplements published in 1895, 1923 & 1972), plus the Flora of LoughNeagh and Beesley (2006), all continued to unquestioningly consider C. temulumindigenous.
More detail on the distribution and decline of C. temulum in Ireland is possible from an analysisof the species map in the New Atlas. Firstly, this shows records from just 19 or 20 Irish VCs andonly 11 Irish hectads have post-1986 records. A further two hectads have records from 1970-86 and 28more can claim the plant during the pre-1970 period. The current author (RSF) estimates that there arearound 1500 hectads on the New Atlas map of Ireland, making the present rarity and marked declineof C. temulum over a period of around 60 years very clear indeed (Preston et al. 2002).
The CEDaR Computer Database at the National Museums NI in Belfast holds a total of just 21 records of thespecies for the six VCs of NI, with dates ranging from 1837-1991. Sites in four VCs are listed fromaround Lough Neagh, but only the Co Antrim shore has a recent record of the plant.
Whatever its status, Rough Chervil has obviously been suffering a very slow, gradual decline towardsextinction on this side of the Irish Sea. The decline of the species has taken place despite seed veryprobably being sporadically and accidentally re-introduced from time to time, in mud and in cargo thatis transported on vehicles across the short sea crossing from Britain in vast quantities.
European and world occurrence
C. temulum belongs to the European temperate biogeographical element and its European distributionis similar to that of Aegopodium podagraria (Ground-elder), but it extends less far north, havingonly a few coastal localities north of 60°N in Sweden and Finland. It is regarded as only,"possibly indigenous in Denmark and S Sweden" by Jonsell & Karlsson (2010). It is absentfrom N Russia. However, C. temulum is present in most of temperate Europe, again becoming rare inthe Mediterranean region, although it is recorded on Sicily, Corsica and Sardinia. It is also present inSW Asia and NW Africa. As an introduction, C. temulum is very thinly represented in E USA and inNew Zealand (Hultén & Fries 1986, Map 1387; Sell & Murrell 2009).
Threats
None.
Native, common and very widespread, seasonally dominant along roadside verges. Eurasian boreo-temperate,perhaps introduced in C & S Africa.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
A very familiar, common and widespread, tall, wintergreen, tap-rooted, native, polycarpic perennial, CowParsley is a seasonally exceedingly conspicuous, characteristic, abundant and temporarily dominantumbellifer along wayside and riverside hedge banks, and the inner zone of roadside verge grasslands andgrassy waste places during the months of April to June (Dowdeswell 1987, p. 51). It is less common butstill very frequent around the unkempt margins of woodland and scrub and also appears in better-litareas within these shade habitats, eg in canopy gaps, and in more open areas beside paths. Cow Parsleyis also found in other lowland, marginal or moderately disturbed ground sites that similarly offer mesicor less extreme soil conditions in terms of moisture, shade, pH, base- and nutrient-content and which,therefore, support tall herb vegetation transitional between forest and managed grassland (Salisbury1964, p. 266). Finally, it also grows on damp cliff ledges of natural upland scarps and in theirartificial, man-made equivalent – quarries.
The ecologically wide soil preferences of A. sylvestris are also reflected in the overall B &I distribution at the hectad level, it being almost omnipresent throughout at this scale, exceptin the wet-peat boglands NW of the Great Glen in Scotland and along the western seaboard of Ireland(Preston et al. 2002). The marked presence on roadside verges which are not intensively managed,but are periodically mown two or three times during each growing season, indicates that A.sylvestris can cope with this level of human disturbance. However, the near absence from all butthe most neglected marginal areas of agricultural land, both in pastures or in cultivated fields,reveals the rather narrow limits of its tolerance of disturbance. The established strategy of thespecies is classed by Grime et al. (1988, 2007) as, "intermediate between competitor andcompetitive ruderal" (C/CR), indicating A. sylvestris can compete successfully andmaintain populations in tall, rough grassy vegetation and exploit a range of moist or shaded fertilehabitats.
A. sylvestris is a conspicuous component of all types of grassland that are not too heavily grazedand especially those that are mown first for hay or silage and grazed only towards the end of thegrowing season. Cow Parsley becomes most luxuriant where grassland is seasonally managed (ie cut,browsed and manured) in this manner (Clapham 1953).
Variation
Two varieties have been recognised in Britain since Clapham (1953) drew attention to them. In var.sylvestris, the whole plant is more open and sparsely hairy; the leaves are more rigid, ultimatesegments 10-25 mm wide, not overlapping; lobes linear or linear-lanceolate and gradually narrowed into aconspicuous mucro (ie a short abrupt point). The alternative from is var. latisecta Druce inwhich the whole plant is more dense and much hairier. Leaves are softer, ultimate segments 20-25 mmwide, overlapping; lobes lanceolate or ovate, rounded-obtuse-mucronate (Sell & Murrell 2009).
It appears that var. sylvestris is the native plant in Britain and is the only form in Scotlandand England south as far as Derbyshire (VC 57). It was probably the main plant of northern mainlandEurope also. In Wales, it is found on ancient trackways and on other grasslands.
Var. latisecta is the common form in England south of Derbyshire. Intermediate plants occur, butit is not known if an intermediate zone as such is formed (Sell & Murrell 2009). The situation inIreland is unknown. Clapham (1953) believed that the southern var. latisecta might be a quiterecent human introduction, and later work by Arthur Chater in Wales and towards London appears tosupport this notion (Sell & Murrell 2009; Chater 2010).
Vegetative and seed reproduction
Like many umbellifers, the A. sylvestris plant produces several perennating buds at the base ofeach fruiting stem as it dies off in summer. This provides overwintering semi-rosette plantlets and, atthe same time, it allows a measure of vegetative increase to take place – although effective dispersalof the buds would perhaps require an ultimately counter-productive, destructive level of habitatdisturbance of the site (Tutin 1980; Grime et al. 1988, 2007). The main reproductive thrust,however, is towards seed production, and although flowering occurs mainly and very conspicuously betweenApril and June, in fact it continues at a very low frequency throughout most of the year in the widerange of habitats that the species occupies. The flower-heads contain many male flowers arranged bothinternal and external to the perfect bisexual ones, the latter often only representing about a fifth ofthe total flower number in each inflorescence (Lovett-Doust 1980). Study by Lovett-Doust found between55% and 70% of bisexual flowers produced seed, and the presence of the additional male pollen flowersthat appear "cheaper to produce", attract more insects to the inflorescence than wouldotherwise be the case, thus improving outbreeding success (Lovett-Doust 1980; Lovett-Doust &Lovett-Doust 1982; Proctor et al. 1996, pp. 346-7). Incidentally, the flowers have a nasty tangof stale (human?) dung, a scent which short-tongued flies find attractive (Proctor & Yeo 1973, p.95).
Seed dispersal and germination
The 1.5 m tall fruiting stems dry out and die after July; the wind and rain wash the stems, scatteringtheir seed for a prolonged period, and do so particularly readily in the linear wayside habitats thatare so very characteristic of the species (Ridley 1930, pp. 31-2, 168).
The species is very definitely a 'follower of man' (ie it is anthropogenic in its occurrence), beingpromoted beyond its natural range of habitats and distribution by human disturbance and by the creationof artificial (man-made) linear habitats such as roads, railways and canals. Rapid movement of transportvehicles, the slipstream effect, also assists its dispersal.
Seed germinates in the spring following winter chilling, but it does not survive into the second year, sothere is no persistent soil seed bank (Roberts 1979; Thompson et al. 1997).
Fermanagh occurrence
A. sylvestris is recorded from 351 Fermanagh tetrads, 66.5% of those in the VC. It shows apreference for fertile lowland soils of near or above neutral reaction and it appears somewhat morefrequently in base-rich limestone districts than elsewhere in the Fermanagh flora survey. Having saidthat, a number of Fermanagh records in the Database are listed from sites described or named as bogs andloughs, but the plant never grows in wetlands, nor on extremely acidic soils and, therefore,these records must refer more loosely to the vicinity, rather than the actual wet or acid peat groundlisted for the particular location grid reference.
British and Irish occurrence
A. sylvestris is recorded in almost every hectad in England and Wales and is common and seasonallydominant in a wide range of suitable lowland habitats throughout both countries. In Scotland, it iscommon and widespread in the eastern lowlands and around the major conurbations, becoming more thinlyscattered in the wetter and more acidic terrain of the northern and western isles. It is absent from theHighlands for the same reasons.
In Ireland, A. sylvestris is common and locally abundant everywhere throughout the lowlands,except in the far west where unsuitable wet, strongly acidic, blanket boglands and dry, rocky limestonehabitats predominate (New Atlas). There has been little change in the hectad distribution in the40 years between the two BSBI Atlas surveys, and although the species has the ability to expand itspopulations and extend along linear habitats by seed, the detrimental effects of airborne pollution andnutrient enrichment on competitive ability appear to balance any tendency towards increase of thespecies.
European and world occurrence
A. sylvestris is widespread and abundant across Europe from Iceland to N Scandinavia (right to theshores of the Arctic Ocean), southwards to N Spain and eastwards to W Asia and SE Russia. However, itbecomes rarer in the south and is absent from all the Mediterranean isles except Corsica. It is alsoindigenous in mountain regions of N Africa. The population in NE Russia and Asia belongs to a separatetaxon, subspecies aemula Woron (Clapham 1953; Hultén & Fries 1986, Map 1388). A.sylvestris is also introduced in C & S Africa and it is even more obviously introduced in NAmerica (Jonsell & Karlsson 2010)
Herbivory, names and folk use
Although 'Cow Parsley' is one of the most familiar English common names of this well-known and much lovedearly summer harbinger, the plant does not survive much grazing pressure and trampling from cattle,sheep or horses. Horses in particular appear to seek it out. Deirdre Forbes, wife of the current author(RSF), describes having trouble when out riding dissuading her mare from stopping to pull it! Cattle andrabbits are also said to be partial to it.
Grigson (1955, 1987) lists a total of 59 local English common names and variants for the plant. Many ofthese names are derogatory, including those with 'cow', 'dog', 'sheep', 'pig', 'rabbit' (or 'coney'),'hare' and 'devil's' as elements. Several names were shared with the ultra-poisonous Hemlock, Coniummaculatum, such as 'Bad Man's Oatmeal' (ie 'Devil's oatmeal') and 'Lady's Lace', although theplant is only supposed or feared to be poisonous, rather than edible. The widely used name 'Queen Anne'sLace' is possibly transferred from the Virgin, as in 'My Lady's Lace', another name given to thisspecies. The reference to Queen Anne's Lace was probably not only to the lace-like flower-heads, butpossibly to the tragic child losses she suffered.
There is a substantial body of associated folklore (Vickery 1995), often regarding the plant asinauspicious, such as a prohibition on bringing the plant indoors, "because the snakes will followit". Picking the flowers and bringing them indoors was also feared to lead to the death of one'smother. Another of its names was 'Stepmother's Blessing', or 'Mother Die', all probably due to itssupposed similarity to the very poisonous Hemlock (Vickery 1995). In Ireland, another association waswith May as the month of Our Lady, and hence a saying, 'Lady's lace looks lovely on the altar'. Thehollow stem of the plant (often when dead) was frequently used as a pea-shooter in many parts of B &I (Vickery 2019).
Mabey (1972, p. 124) recommends A. sylvestris as a pot herb substitute for its close relativeGarden Chervil, A. cerefolium. In Gloucestershire, the Isle of Man and parts of Ireland, A.sylvestris was recommended as a cure for kidney or bladder stones or gravel. This might merelyhave been a carry over from the same property attributed to Garden Parsley (Petroselinum crispum)(Allen & Hatfield 2004).
Threats
None.
Casual introduction, archaeophyte, now extinct. Eurosiberian southern-temperate, very widely introducedin both hemispheres.
1904; West, W.; grounds of Portora School, Enniskillen.
Growth form and preferred habitats
Descriptions of this species vary so widely from book to book that the current author (RSF) has decidedto give a comprehensive compilation, based mainly on the outstanding, detailed description provided inthe excellent Flora of Cyprus (Meikle 1977).
A winter annual, therophyte herb with a tapering taproot and fibrous side-roots; stems 4-30(-50) cm tall,usually much branched, erect or spreading, glabrous or nearly so, becoming hollow when old, branched andleafy. Basal leaves numerous, narrowly deltoid, 1-10 cm long, 2-3-pinnate, the ultimate segments linearor lanceolate acute or obtuse at apex, on slender petioles often exceeding 10 cm. Stem leaves few andsmaller, with fine, almost linear segments. Flowering May-August (but mainly May & June),inflorescence composed of few (3-8)-flowered simple umbels, each with 1-3 rays, the terminal umbelbearing white, bisexual, 5 mm diameter flowers, the lateral umbels with varying proportions of male andbisexual flowers. Sepals absent or five, very small if present; petals five, very unequal, the outerones often radiate; stamens five, filaments 1 mm long, white, anthers cream; ovary linear-subulate, 3-4mm long, stylopodium flattened with an undulate margin, usually tinged purple, styles two, about 1 mmlong, erect, stigmas truncate (Meikle 1977; Sell & Murrell 2009).
Pollinated by unspecified insects (Fitter 1987). Schizocarp fruit, 3-7 cm long, ± cylindrical, with adistinct, flattened beak with rough ridges, 3-5 times as long as the seed-bearing portion. When ripe,the fruit splits in half with a violent jerk. Around the base of the umbel of flowers is a whorl ofsmall leaves which after fertilisation enlarge and become lobed (Salisbury 1964; Tutin 1980; Sell &Murrell 2009).
Fermanagh and Irish occurrence
Until around 1900, S. pecten-veneris was a quite common annual, arable weed in Ireland ondisturbed soils that dry out in summer, often on heavy, calcareous clays. Recorded only once inFermanagh at Portora School by one of the teachers, in the Revised Typescript Flora, Meikle etal. (1975) described S. pectin-veneris as, "hardly more than casual". Accordingto the Cen Cat Fl Ir 2 it was previously recorded from all Irish VCs except W Cork (H3) and WMayo (H27). It is now presumed extinct throughout Ireland, having last been seen in NI in 1972 on sanddunes at Murlough Bay, N Antrim (H39) (Curtis & McGough 1988; Beesley 2006).
British occurrence
Until around 1950, S. pectin-veneris was much more common and locally abundant in Britain in andespecially around chalky arable fields, to the extent that it could sometimes impede mechanicalharvesting. It was also known to occur in a number of semi-natural sites on coastal screes, perhapsrepresenting a form of refugium away from cultivated ground more akin to its natural habitat in warmerparts of S Europe including the Mediterranean basin (A. Smith, in: Stewart et al. 1994).
The main period of germination of this annual species is in the autumn, from October to early November,followed by a much smaller crop of spring seedlings. The autumn seedlings are often eradicated bypre-sowing cultivation associated with winter cereals, seasonal crops to which S. pectin-venerisoddly seemed to be almost entirely restricted. The spectacular decline of this arable weed since about1930, but accelerating from 1955 onwards, is thought to be largely due to its low level of seed dormancy(ie transient – persists for less than one year) (A. Smith, in: Stewart et al. 1994; Thompson etal. 1997), allied with important changes in agricultural practices, including greatly improved seedscreening and the widespread use of broad-spectrum selective herbicides (Stace & Crawley 2015).
In Britain, S. pecten-veneris has now largely retreated to an area SE of a line from the Humber tothe Severn with occasional plants elsewhere. The species remains occasionally locally abundant in partsof East Anglia (M. Southam, in: Preston et al. 2002; Sell & Murrell 2009). The Englishdecrease mirrors a similar decline of the species in both NW & E Europe.
Species status
Webb (1985) casts doubt on the previously presumed native status of this species in Britain, while inIreland the plant had long been accepted as a probable or definite accidental introduction, probablyarriving with cereal seed. The editors of the New Atlas now recognise it as an archaeophytethroughout B & I. Previously, Stace (1991, 1997) suggested S. pecten-veneris might possiblybe native in Britain, but in Stace (2010) he accepted it as an archaeophyte. Sell & Murrell (2009)have not yet accepted the archaeophyte status and continue to consider S. pectin-veneris"possibly native".
The fact that archaeophytes like S. pectin-veneris, that were once common cornfield weeds and wereconsidered thoroughly naturalised, have now very greatly diminished due to cleaner agricultural methods,strongly suggests that they were in fact always only casuals that owed their persistence to constantreintroduction with freshly imported crop seed (Stace & Crawley 2015).
European and world occurrence
The distribution of S. pecten-veneris in Europe is centred on the Mediterranean, extending northto Denmark, casual in Scandinavia, and westwards to B & I. Thus its presence in Ireland wasalways on the limits of its tolerances and geographical range, the oceanic influence of theAtlantic being much more extremely felt in Ireland than in the E & SE England in particular. BeyondEurope, S. pectin-veneris extends to N Africa, and from SE Asia to the borders of India. Inaddition to B & I, it is also introduced in S Africa, N & S America, Australia and New Zealand(Sell & Murrell 2009; Hultén & Fries 1986, Map 1390). On the Mediterranean isles, S.pectin-veneris is recorded in a much wider range of habitats than simply cultivated and fallowarable fields, being also associated with Olive groves, stony, bare hillsides, rocky places, roadsides,old walls and bare and disturbed ground (Haslam et al. 1977; Meikle 1977; Turland et al.1993; Press & Short 1994).
Names
The genus name 'Scandix' is an old Greek name for some edible plant. The Latin specific epithet'pecten-veneris' translates as 'Venus's comb'. In Britain, S. pecten-veneris was first mentionedby Turner (1548) as 'Crine Cheruel', a rather dull name, but the plant does have a resemblance toChervil before the formation of the needle fruits (Grigson 1955, 1987). The English common name'Shepherd's Needle' was first used by Lyte (1578), translating the herbalists' Latin 'Acus pastoris',although this and the English common name were both sometimes also applied to Erodium moschatum(Musk Stork's-bill), since that species also possesses long, beak-like fruits (Grigson 1974). Thanks tothe peculiar fruit, the plant is remarkable and has earned as many as 50 English common names, althoughGrigson (1955, 1987) lists only 26, many of which contain mention of needle or needles. Frequentfolklore associations are with goblins, the devil and witches (Grigson 1955, 1987). It is rather oddthat Vickery (1995, 2019) makes no mention of the species.
Uses
Herbal use is slight, but one report is of 'Adam's Needle', an alternative name for S.pectin-veneris, being used in Co Tipperary to treat toothache (Allen & Hatfield 2004).
Threats
None.
Introduction, neophyte, scarce and local.
1934; Praeger, R.Ll.; Castle Caldwell.
March to November.
Growth form and preferred habitats
A softly and rather sparingly hairy, 60-200 cm tall, 2- to 4-pinnate, fern-leaved perennial smellingstrongly and sweetly of aniseed when bruised, this ancient cultivated 'potable' (ie edible) andmedicinal herb tends to occur either as individual clumps or more extensive patches spreading in roughgrasslands along road- and path-side habitats (particularly in hedge and stream banks of more uplanddistricts) and in waste or neglected ground, usually near habitation. It prefers fairly dry, nutrient-and mull-rich bare soils, preferably in half-shade but also in full sun (Jonsell & Karlsson 2010).Additional Fermanagh habitats are described below. The leaves are pale beneath and they usually havevery characteristic, unique flecks or blotches of white on their upper surfaces (Tutin 1980).
Flowering reproduction
M. odorata flowers early in May and June, its inflorescence is a compound umbel of 4-10(-21)hairy rays, 1.5-3.0 cm long; its white flowers have radiating outer petals. The flowers are veryattractive to bees which act as pollinators. The schizocarp fruit is remarkably large, 15-25 mm long andit has a short beak. The tall, stout, hollow stems of the plant provide the necessary height to scatterthe seed efficiently in the late summer and autumn. Apart from a limited ability of the fruits to floatdownstream from established stands on waterside banks and potentially to recolonise suitable open soil,M. odorata is a poor colonist (Tutin 1980; Grime et al. 1988, 2007; Sell & Murrell2009).
The seeds germinate in the spring in any moderately fertile, well-drained soil, whether in sun or shade,provided it is neither too acid, too wet, or more than lightly disturbed or grazed (Grime et al.1988, 2007). By virtue of a stout, deep-penetrating taproot up to 35 mm thick, the species regularlybecomes established and long-persistent, as is proven in many sites around the country. While the plantis well equipped to survive and it often forms colonies that produce plenty of seed (it has the largestseed of any herbaceous wildflower in these islands – but it is really a dry fruit that splits into twomericarps), M. odorata seldom if ever travels very far from where it has been planted.
Fermanagh occurrence
In Fermanagh, M. odorata has been recorded from 22 thinly scattered tetrads (4.2%), 15 of whichhave post-1975 dates.
It is very often located in waste ground near houses or old ruined wall-steads, from the gardens orrefuse heaps of which it has escaped. Very occasionally it is also found in or near graveyards, as atMonea Roman Catholic chapel, where it persisted for at least 37 years, from 1951-88. Grigson (1955,1987) observed the churchyard association of Sweet Cicely in S Wales, where he thought it was cultivatednear headstones possibly as "a plant of memory and sweetness".
Status of the species
While in Ireland, M. odorata has always been recognised as a thoroughly naturalised garden escape(Cat Alien Pl Ir), in Britain it previously was regarded as at least 'possibly native' (egClapham et al. 1962). The third edition of the Flora of the British Isles declared thespecies an introduction (Clapham et al. 1987). However, the status of M. odorata remainedsufficiently questionable for Clement & Foster (1994) to accept it, "with reservations asnative" in their book Alien plants of the British Isles. The New Atlas recognised thespecies as a neophyte, ie introduced after 1500 AD, first recorded in the wild in 1777 (M.F. Watson, in:Preston et al. 2002).
British and Irish occurrence
The New Atlas indicates M. odorata is widespread throughout the whole length of Britain,but very much more prevalent north of Birmingham, becoming increasingly eastern in N Scotland. The Irishdistribution maintains the northern emphasis in the pattern, the plant being very nearly confined to thesix counties of NI and very much more frequent west of the River Bann and Lough Neagh. This distinctiveand rather odd pattern of naturalisation led Praeger (1915) to argue that M. odorata, along withother medicinal herbs of high repute such as Masterwort (Peucedanum ostruthium), Tansy(Tanacetum vulgare) and Elecampane (Inula helenium) was introduced by Scottish settlers atthe time of the Plantation of Ulster in the early 17th century. A glance at the distribution maps ofthese herbs will readily indicate that Praeger was probably correct in his supposition.
European and world occurrence
M. odorata is believed to have originated in SC Europe. Hultén & Fries (1986, Map 1391) plotthe native area as (presumably the foothills of) the Spanish Picos, Pyrenees, Alps, N Apennines, plusthe Dinaric Alps and mountains of Croatia, Hercegovina, Montenegro and Macedonia (ie the oldYugoslavia). It was introduced into cultivation northwards and eastwards of its origin and has becomewidely naturalised as far north as Trondheim in Norway and SW Iceland. It is said to be still spreadingin Scandinavian countries (Jonsell & Karlsson 2010). It has been widely introduced elsewhere,including in Chile (Hultén & Fries 1986).
Uses
Sweet Cicely (or 'Sweet Fern' – two among many English common names) was previously used as an additionto salads, or boiled as a vegetable when every part of the plant was eaten. The long, brown pod-likefruits are particularly full flavoured. As a medicine it was supposedly useful for both coughs andflatulence, and as a "gentle stimulant for debilitated stomachs" (Grieve 1931). The roots werealso regarded as antiseptic and were also used to treat the bites of vipers and mad dogs (Grieve 1931,p. 201).
Names
The genus name 'Myrrhis' is derived from the Greek 'myrrha' or 'murra' meaning 'perfume' or 'fragrant'(Johnson & Smith 1946), but there is also a connection with the ancient name of true myrrh from theBible, the gum resin from the small E African and Arabian shrub Commiphora myrrh (Grigson 1974;Gledhill 1985). The Latin specific epithet 'odorata' also translates as 'fragrant' (Johnson & Smith1946).
Fourteen English common names are listed from around B & I by Grigson (1955, 1987), of which mostrefer to the sweet aniseed aroma, eg 'Anise', 'Annaseed', 'Sweet Bracken', 'Sweet Cis', 'Sweet Humlick'(ie Hemlock). A Lancashire name listed by Grigson (1955, 1987) is 'Roman Plant', which probably refersto the growth of the plant as a potherb and medicine in Roman Catholic monastery gardens.
Threats
None.
Introduction, archaeophyte, rare.
1900; Praeger, R.Ll.; Co Fermanagh.
April to October.
Growth form and preferred habitats
In Ireland, this 50-150 cm tall, glabrous, robust, very conspicuous and fully naturalised biennial with afleshy, carrot-shaped, tuberous taproot 30-40 mm long in the first year of its growth, is mainly a plantof the E and S coasts, although it does also penetrate inland in a much more localised manner.Throughout B & I, it is locally abundant along roadside hedgebanks and wayside paths, often underthe dappled shade of tall trees. It also grows on sea cliffs and in waste ground and is especiallyfrequent and often abundant in open wayside habitats near the sea.
Fermanagh occurrence
In landlocked Co Fermanagh, NW Ireland it is only very occasional to rare, but it certainly is longpersistent in six of the seven known scattered sites it occupies in hedgerows and beside old dwellings.Alexanders is particularly abundant around Portora Castle in Enniskillen, where it could very well havebeen growing for several centuries. It has not been recorded at Newtownbutler since 1951. The local sitedetails are: roadside, Portora, 1951, Dr G. Gillespie (still there); hedgerow backing gardens,Newtownbutler, 1951, MCM & D; roadside, bridge and farmyard, Aghalane Td, Woodford River, SE UpperLough Erne, 1990 & 1996, RHN; on Wattle Bridge, 6 April 1996, RHN, still there 2005; roadside atGeaglum and Derrychree Tds, on the shores of Upper Lough Erne, 2 July 2002, RHN; large patch nearcrossroads at Black Lough near Glasmullagh House, 14 December 2003, RHN.
Introduction and uses
Archaeological evidence supports the idea that the plant, being a common edible native of S & SWEurope as far north as NW France, was originally introduced to Britain by the Romans. It is believedthat it was widely used blanched like Chicory as a culinary potherb (the young stems taste like Celery),until the latter replaced it in the 15th century, although in fact garden Celery did not become distinctfrom the wild form until the 17th century (Salisbury 1964, p. 272; Tutin 1980). It appears to have goneout of fashion as a useful vegetable, but is so well naturalised it has become no less characteristic ofmaritime habitats than many coastal natives (Stace & Crawley 2015).
The Romans possibly did not make it to Ireland (another subject of ongoing debate), but the plant wasprobably introduced along with the traditional monastery garden plants in early Christian times, if notbefore, since apart from its use as a potherb, the plant also enjoyed an ancient reputation as amedicinal herb. Among seafarers, for instance, it was believed to 'clear the blood' and to preventscurvy, sailors being known to put ashore specially to collect the plant (Allen & Hatfield 2004).
British and Irish occurrence
The New Atlas species account suggests that while the distribution of S. olusatrum is notmuch changed since the 1960s, it is increasing somewhat in both frequency and abundance along roadsidesin inland areas of B & I. Since the species is so completely naturalised and persistent everywherethat it occurs, there does not seem to be any simple explanation as to why it has such a predominantlycoastal distribution in these islands (M.F. Watson, in: Preston et al. 2002).
Threats
None.
Native, common and very widespread. Oceanic temperate and tightly restricted to W Europe.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This tuberous perennial, up to 8-70 cm tall, has glabrous, stems and leaves and displays a short, vernalseason of growth and reproduction (Grime et al. 1988, 2007). The basal leaves of the plant are2-4 times larger than the stem leaves and are borne on a petiole 4-15 cm long. The leaf blades of thebasal leaves are 2-pinnate, but they do not long survive, withering by late spring or early summer, sothat they often are completely absent when the plant flowers in May and June. The stem arises from anirregular, dark brown, nut-like subterranean tuber, 8-35 mm in diameter, which is deeply buried. Thetubers are edible, raw or cooked, and are described as having a pleasant, nutty flavour (Tutin 1980).Nowadays, for conservation reasons, it is forbidden to dig up wild plants, the only exception being thelandowner. The stem bears 2-4 leaves, each with a well developed greenish membranous sheath at its baseand a reduced petiole. The ultimate lobes of the stem leaves are elongated and linear and the terminallobe is much longer than the lateral lobes (Tutin 1980; Jonsell & Karlsson 2010).
C. majus is a very characteristic stress tolerant ruderal (SR) species of damp or shaded ground inwoods, scrub, heaths, shaded cliffs and in a wide variety of mesic agricultural grasslands (Grime etal. 1988, 2007). It is particularly frequent in relatively infertile soils derived fromlimestone, but occurs in both moderately acid and base-rich situations, most frequently in the pH range4.5-7.0. It also appears under similar growing conditions on relatively undisturbed, unproductive grassyroadsides and in waste ground. It is said to be scarce on chalk in England (Grime et al. 1988,2007).
Flowering reproduction
Despite the production of vegetative subterranean tubers, reproduction is entirely dependent onseed. The plant flowers early in the season, in May and June. The inflorescence is a compound umbel, 5-8cm wide, with 6-12 smooth rays, the peduncles longer than the rays. Flowers, 16-24 per 'umbellule' (ieultimate or secondary simple umbels), with a total flower count of >50 per complete inflorescence;petals white, outer ones not or scarcely radiating. As is normal in the Family Umbelliferae (orApiaceae), a proportion of the flowers are staminate (ie entirely male). In C. majus, staminateflowers form the majority, only one in five being hermaphrodite (ie bisexual or perfect) (Lloyd 1979unpublished; quoted in Lovett-Doust 1980). Nectar is well exposed and pollination is by insect visitorsincluding bees and wasps (Proctor & Yeo 1973). The crowded nature of the numerous flowers in umbelsfavours a mixture of self- and cross-pollination (Lovett-Doust 1980).
The fruit is c 4 mm long, ovate in outline, slightly flattened laterally and dark brown in colour and isformed and ripens in June and July. By the time seed (the twin mericarps) is produced, the aerial stemis already dead and it soon disappears, the plant overwintering as both buried vegetative tubers andseed.
Seed germinates early in the year after its production, between January and March following adormancy-breaking chilling period (Roberts 1979). In an experimental study of 121 specimens, the meannumber of mericarps produced per plant was 215.5 ± 9 with a range of 40-958 (Salisbury 1942). Seedsurvival in soil is transient or short-term persistent (ie they persist for at least one year, but forless than five) (Thompson et al. 1997).
Fermanagh occurrence
It is very common and widespread in Fermanagh, having been recorded in 326 tetrads, 61.7% of those in theVC. Pignut appears ± everywhere throughout the county, except on strongly acidic peat, regularly wetground or heavily grazed pastures.
Cotyledons and fundamental biological concepts
C. majus is one of the few dicotyledonous species that actually produces just one embryoseed leaf, the other presumably being suppressed or aborting early in its development (Metcalfe 1936;Grime et al. 1988, 2007; Thompson 1988). It is not unique in this respect: the same is true ofthe quite closely related species, Bunium bulbocastanum (Great Pignut), plus in other families,Ranunculus ficaria (Lesser Celandine) and some Corydalis spp. (Thompson 1988). Thisobservation should forcefully remind us that even the most fundamental and revered biological concepts,like this example linking us back to the work of the great John Ray (1627-1705), may be somewhat 'leaky'(Morton 1981, p. 203; Raven 1986, p. 189). There are 'natural laws' in physics, but none in biology. Ourdescription of the natural world and the categories we use to subdivide and label organisms and ideasare often a convenient over-simplification of the variation and subtleties that really exist (Proctoret al. 1996, p. 330; Forbes 2000). It is fundamentally important that as scientists we recognisethat the plants we study have not read the books, and when we read any material, no matter howmuch we respect the author, we should never assume that everything printed is infallible 'holy writ'. Asthe current author's mother (Jean Forbes) occasionally used to remind him, "pen, ink and paperrefuse nothing".
Herbivory
Due to its vernal phenology and short photosynthetic period, C. majus is very susceptible to heavygrazing pressure in the spring. Badgers commonly dig up the globular brown Pignut tubers and in someareas of B & I they are an important part of their diet. Even recently weaned badger pups have beenobserved digging for them (Neal & Cheesman 1996). In Speyside, in Scotland, the tubers represented14% of the badger's summer diet; they dig as deep as 40 cm, but no lower. Badgers dig for the tuberswhen the plants are flowering, probably because they are unable to detect them in winter (Kruuk 1989).
British and Irish occurrence
Common throughout most of Britain, but absent from the English Fens and scarce on chalk soils. It wasintroduced in Shetland in the 19th century, but its status on Orkney is uncertain. It is less common inIreland, but especially so in the area of 'the English Pale' around Dublin, where agriculture is mostintensive, and in the far west coastal areas where soils are most likely too wet and too acidic (M.Southam, in: Preston et al. 2002).
In Britain, the recent BSBI Local Change survey of 1987-2004 showed that C. majus has suffered awidespread, statistically significant decline in recent years: when the species distribution wasexamined at the tetrad level the weighted Change Factor calculated was -23. Most probably the decline isdue to grassland 'improvement' measures, involving the ploughing up, reseeding and fertiliser andherbicide spraying of old grasslands, plus a widespread move from hay- to silage-making (Braithwaiteet al. 2006).
There are no comparable data to suggest that Pignut is suffering any decline in Fermanagh, but sincesimilar agricultural grassland management changes have been applied locally in lowland areas of thecounty, it is quite likely that some decline in population size and frequency must have occurred.
European and world occurrence
C. majus belongs to the Oceanic Temperate biogeographic element and is confined to W Europe fromNorway southwards and eastwards to NW Italy (Sell & Murrell 2009). The map prepared by Hultén &Fries (1986, Map 1392) shows C. majus very much confined to western oceanic parts of Europe,extending from the Faeroes to N Morocco and east as far as Corsica, although absent from Italy. Pignatti(1997) plots the species as present in two districts of NW Italy in or near the Maritime Alps. The textassociated with Hultén & Fries map indicates they believe C. majus is introduced in theFaeroes and in Germany, and suggests "probably [introduced in] more places" (Hultén &Fries 1986).
The current author (RSF) suggests the means of introduction is almost certainly seed, transported eitheras a contaminant in commercial grass seed mixtures, or in hay, a form of fodder that has become muchrarer than previously was the case, having been replaced by silage and manufactured 'nuts'.
In parts of Scandinavia (including Denmark), C. majus was introduced either as a minor rootvegetable, or later as a grass seed contaminant. The species is often only casual in Scandinavia,although it forms large, dense populations in W Norway in open fields and pastures along the coast(Jonsell & Karlsson 2010).
The current author (RSF) is sceptical about the extent to which the tuber was used as food, since it isgenerally small in size and would be difficult to gather in sufficient quantity to make a meal for morethan one individual. Also, C. majus apparently does not lend itself to garden cultivation andhorticultural improvement despite 18th century trials to that end which found, "the Earthnut willnot thrive in tilled land" (Bryant 1783).
Names and uses
'Pignut' and 'Earthnut' are only two of 38 English common names and variants listed by Grigson (1955,1987), both of which refer to the small, but edible underground tuber of the plant. Similarly, Vickery(2019) has a count of at least 50 English common names indicating great familiarity with the species.However, the derogatory connotations of some name elements such as 'pig', 'fare' in 'Farenut' ('fare' isa young pig, from the Old English 'fearh'), 'hog', 'swine', 'scabby', 'lousy', 'hare', 'cat', 'devil'and 'jack', all suggest a degree of contempt or dislike for the plant and its use, similar to that notedfor Anthriscus sylvestris (Grigson 1955, 1987) (see that account on this website). The diggingand eating of the tubers is described by Vickery (2019) in terms of children's play and he makes nomention of any other associated folklore.
There is very little evidence of use in herbal medicine and it is not mentioned at all in modern herbalsby Grieve (1931) and Darwin (1996). It is said to have been employed as a diuretic on the Isle of Man,for cleansing the blood in Co Donegal and as a tea substitute in Co Fermanagh, although it is not knownif this was considered medicinal or not (Allen & Hatfield 2006).
Threats
None.
Native, very rare. Eurosiberian temperate, introduced in eastern N America and New Zealand.
1884; Barrington, R.M.; Lower Lough Erne.
April to August.
Growth form and preferred habitats
This wintergreen, deeply rooted, semi-rosette perennial arises and grows erect from a stout rootstock.Burnet-saxifrage prefers well-drained, dry to damp, often shallow calcareous or base-rich soilssupporting unproductive, short-turf grassland, such as occurs in old, lowland, species-rich,hay-meadows, pastures, roadsides verges and riverbanks. In Fermanagh, similar unproductive roughgrassland habitats often lie close to many of the county lakeshores, although P. saxifraga isdefinitely neither a wetland nor a woodland species, or even one tolerant of more peaty, acidicconditions. It can sometimes be found elsewhere in B & I, however, in more acid soils of lowfertility in older low-lying grassland sites and in rocky habitats, especially those on calcareous orother basic rocks. P. saxifraga can grow in full sun or half-shade and it prefers warmer soils.It can grow and survive in dry, but not severely droughted situations thanks to a deeply penetratingtaproot (Sinker et al. 1985; Grime et al. 1988, 2007; Rich et al. 1996).
P. saxifraga is low-growing, stems (15-)30-100 cm tall, and it tends to lack the ability towithstand competition in both fertile soils and moderately disturbed habitats. The established strategyof the species is described as intermediate between a Stress-tolerant Ruderal and a more general abilityC-S-R (Competitor-Stress-tolerant-Ruderal) species. Essentially, P. saxifraga is only able tocompete successfully in short turf situations, where taller, more vigorous species are restrained byless favourable growing conditions, whether that be determined by soil nutrients, moisture, depth, orother factors such as shade or disturbance level.
Burnet-saxifrage has a low colonising ability and is considered a useful indicator species of oldspecies-rich grasslands, especially old calcareous grasslands that have not been heavily disturbed,ploughed or re-sown, a greatly diminished vegetation type that remains extremely threatened (Grime etal. 1988, 2007; Braithwaite et al. 2006).
Flowering reproduction
The plant is only conspicuous between July and September when flowering and fruiting, since the veryvariable leaves are 1-2 pinnate and are held low on the stem, around 10 cm from the ground, while bycontrast the flowering stem can reach up to 1.0 m in height. The basal leaves wither early and often arenot present when the plant flowers, their role being taken over by a small number of stem leaves.
The inflorescence is a compound umbel of 10-22 rays bearing small, white or rarely pinkish flowers, theouter petals not radiating. The peduncle is longer than the rays and the majority of the 100 plusflowers per inflorescence are hermaphrodite (bisexual), although not perfect since they lack sepals.Pollination is by insects (Hymenoptera, bees), attracted by openly presented nectar (Fitter 1987).Fruits are produced quite late in the growing season in August and September, each being a schizocarp,2-3 mm, ovoid, laterally compressed, and the two mericarps each have slender ridges, paler than thereddish-brown vittae between them (Tutin 1980; Sell & Murrell 2009).
Seed (ie mericarp) of P. saxifraga can survive passage through the complex alimentary canal ofcattle, remaining viable when cast in dung, thus achieving a limited degree of local secondary dispersal(Salisbury 1964).
Being a deep-rooted perennial species, individual plants of Burnet Saxifrage are capable of surviving inunmown meadow grassland for many years as Tamm (1956) showed in C Sweden. In permanent quadrats comparedover a twelve year period, Tamm found that one individual or clone of P. saxifraga persisted andgrew substantially, although five other clones disappeared. Only one small individual was recruited anewto the plots during the long period of observation (Tamm 1956, Fig. 12).
Variation
There is considerable variation within P. saxifraga and Sell & Murrell (2009) list threesubspecies and five varieties of one of them (subsp. saxifraga). The other two subspecies are:subsp. alpestris (Spreng.) Vollm. and subsp. nigra (Mill.) Gaudin.
Irish and Fermanagh occurrence
In Ireland, P. saxifraga is very noticeably much more frequent in the drier and warmer S & Eof the island than elsewhere. While it is recorded at the coast from the very northern tip of Irelandnear Malin Head in Co Donegal (H34) and also occurs locally on the basalt scarps of Co Antrim (H39)(FNEI 3), Fermanagh appears to be close to the NW limit of this species for Irish inland sites.It strikes the current author (RSF) that the occurrence and unusual Irish distribution pattern of thisspecies is most similar to that of Primula veris (Cowslip), another plant near its natural limitsin Fermanagh.
While the Fermanagh Flora Database contains records of P. saxifraga from eight tetrads, there areonly five post-1986 records at four widely scattered sites and RHN and the current author consider itvery rare in the VC.
Another occasional habitat P. saxifraga occupies is in drier parts of old disused quarries, suchas one at Clonmackan in SE Fermanagh, near Clones in Co Monaghan (H32). It was first found here byMeikle and co-workers in 1951 associated with Daucus carota (Wild Carrot), Anagallisarvensis (Scarlet Pimpernel), Centaurium erythraea (Common Centaury) and four orchidspecies including Anacamptis pyramidalis (Pyramidal Orchid) and Ophrys apifera (BeeOrchid). RHN found it again nearby at Clonmackan bridge in 1990 and the only other recent records arefrom Belleek village (1987) and from limestone meadows south of Clonatty Bridge (1991 & 2003) where,incidentally, Primula veris (Cowslip) is also recorded.
Meikle and co-workers discovered Burnet-saxifrage in limestone sites at Knockmore and the Marble Arch(Meikle et al., 1957), but later very detailed surveys of these areas have not seen it. Possiblythe species is being overlooked and its status is worth further careful investigation.
British occurrence
P. saxifraga is widespread in Britain, except NW of the Great Glen in Scotland, the overallpattern again, to some extent, mirroring the distribution in the warmer S & E of Ireland (NewAtlas).
European and world occurrence
Burnet-saxifrage is also widespread in most of temperate Europe, although thinning northwards. It is alsoindigenous in SW Asia and is a rare introduction in eastern N America and New Zealand (Hultén &Fries 1986, Map 1394).
Uses
The Modern Herbal (Grieve 1931) appears a little confused and confusing regarding the medicinaluses of the three Burnets and especially regarding Burnet-saxifrage which ends up with two entries inthe work (pp. 146 & 720). The leaves and roots of the plant are used by herbalists, the whole herbbeing collected in July and dried.
The root is described as very hot and acrid, burning the mouth like pepper, although on drying and ageingpungency is considerably diminished. It contains a bitter resin and a blue coloured essential oil, bothbeing useful to relieve flatulent indigestion. The fresh root chewed promotes the flow of saliva and isrecommended for easing toothache and paralysis of the tongue. A decoction is said to dissolve mucus andis used as a gargle to treat hoarseness and throat infections. It is also prescribed for asthma anddropsy.
Small bunches of the leaves and shoots tied together and suspended in a cask of beer impart to it anagreeable aromatic flavour and are also thought to correct tart or spoiled wines.
Cows that graze the plant are thought to have an increased milk flow (Grieve 1931, p. 720). Other modernherbals mention its use as a wound dressing to stop bleeding and prevent infection. It is also said tobe used to treat fever, gout and rheumatism (Darwin 1996). As an astringent, Burnet-saxifrage was saidto be used for cleansing of freckles, although Allen & Hatfield (2004) voice their considerablescepticism.
Names
The genus name 'Pimpinella' is of obscure derivation, possibly from the medieval Latin 'bipinella'meaning '2-winged', referring to the simply pinnate arrangement of the leaflets (Hyam & Pankhurst1995). The name was first used by the Italian herbalist, Matthaeus Sylvaticus in the early 14th centuryin his pharmacopoeia published under the Latin title Pandectarum Medicinae or PandectaeMedicinae (Encyclopedia of Medicines) (Gilbert Carter 1964).
Grigson (1974) has it that the 15th century name of the plant was 'pympernol' or 'pympernelle' (modern'pimpernel'), derived from the French 'pimprenelle' or Old French 'piprenelle', from medieval Latin'pipinella', derived ultimately from Latin 'piper' meaning 'pepper', owing either to the taste of theleaves of P. saxifraga or the resemblance of its ripe fruits to peppercorns. The Latin specificepithet 'saxifraga' is from 'saxum', a rock, and 'frango', to break. In herbal medicine, it was supposedto be capable of breaking up a stone in the bladder (Stearn 1992).
The English common name 'Burnet-saxifrage' is derived from a resemblance of the basal or root leaves tothe pinnate leaves of Sanguisorba officinalis (Great Burnet) and Poterium sanguisorba(Salad Burnet). The flowerheads of both the latter named are a dark crimson-brown or mahogany, or'burnet', from the Old French 'burnete', or 'brunet'), becoming French 'brunette' (Grigson 1974).'Burnet' is a term previously applied to a brown cloth and it was given to the two plants on account oftheir brown flowers (Prior 1879). As Mabey (1996) points out, 'Burnet-saxifrage' is a rare example of aplant named after two other unrelated families: 'burnet' from the shape of the leaves, and 'saxifrage'from its traditional herbal use in treating kidney and bladder stones.
Threats
P. saxifraga populations have suffered a rapid and severe decline due to the widespreaddestruction of old calcareous, neutral and acidic grasslands across B & I associated withintensification of agriculture since the 1950s and the eutrophication of many previously suitablehabitats which encourages tall, ranker swards in which Burnet-saxifrage cannot compete or survive(Braithwaite et al. 2006).
Introduction, archaeophyte, a common and very widespread, invasive garden escape, locally abundant.Eurosiberian temperate, but naturalised in eastern and central N America.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This is a notorious rhizomatous perennial garden weed of great vegetative vigour. A. podagraria iseasily recognised by its glabrous, 1-2-ternate leaves that resemble the familiar Elder shrub(Sambucus nigra), and hence the English common name 'Ground-elder'. It has stems up to 100 cm,which are hollow and grooved, and the absence of both bracts and bracteoles is very distinctive, as isthe possession of its long creeping rhizomes up to 9 mm thick bearing lateral buds (Tutin 1980; Garrard& Streeter 1983; Jonsell & Karlsson 2010). The established strategy of the species is rated CR(Competitive Ruderal) by Grime et al. (1988, 2007) and there is no doubt of the accuracy of thecategorization.
Archaeological evidence indicates that Ground-elder was originally introduced and cultivated in Britainduring Roman times, around 150 AD, probably both as an edible pot-herb and for medicinal use. The firstwritten record of its presence in Britain dates from 1578 when it was prized as a useful medicinal plant(Ivens 1966). It was not held in high regard for very long, however, since Gerard wrote of it,"Herbe Gerard [one of its many names] groweth of it selfe in gardens without setting or sowing, andis so fruitfull in his increase, that where it hath once taken root, it will hardly be gotten outagaine, spoiling and getting every yeere more ground, to the annoying of better herbes." (Gerard1597, 1633, p. 1001).
A. podograria very readily escapes from cultivation – most likely as small fragments of the verybrittle rhizome transported amongst garden waste and also, accidentally, in adhering soil or mud. Underthese circ*mstances it invades and establishes itself very successfully in both native and artificial,disturbed habitats, often in shade or half-shade, near habitation and usually in lowland sites. Soilswhere the species performs best are moderately damp, nutrient-rich and well supplied with mull humus(Jonsell & Karlsson 2010).
In addition to a range of disturbed artificial habitats including roadside verges, waste- anddisturbed-ground, principally near habitation, A. podagraria also occurs widely in woodlands andalong shady stream and riverbanks, where it presents a very convincing (however misleading) impressionof behaving like a native species, on occasions becoming locally dominant (Clapham 1953; Sinker etal. 1985).
Flowering reproduction
A. podagraria flowers in May and June, the inflorescence being a compound, slightly convex umbelof 10-20 smooth rays and the flowers small, white and slightly irregular (zygomorphic). All the flowersare hermaphrodite (bisexual), but in some peripheral flowers the stamens may drop early (Jonsell &Karlsson 2010). The plant flowers freely and is pollinated by insect visitors attracted by both pollenand nectar. Fruiting takes place in July and August, but seed production is usually poor and seedlingsare rarely observed. Sometimes, however, the plant does fruit abundantly and plentiful seedlings canthen result. Nevertheless, fresh colonies rarely establish from seed and therefore they do not greatlyadd to the plant's invasive aggression (Salisbury 1962, p. 499). In any event, the seed is transient,usually surviving in soil for less than one year (Thompson et al. 1997).
Sexual reproduction is, of course, significant in the longer term, maintaining the species adaptivevariation and vigour and contributing to the species' dispersal, so that overall the combinedreproductive capacity of the species, sexual and asexual or vegetative, is very powerful –despite the plant very often growing in shady wayside sites.
Vegetative reproduction
The most characteristic feature of Ground-elder undoubtedly is its far-reaching rhizome system, thebranches of which are white or cream when young and are described as aromatic (Hyam & Pankhurst1995). From two to five of these underground shoots spread out from the base of each tuft of leafyfoliage in all directions, sending up aerial shoots at intervals from nodes that vary in length fromjust 6 mm to over 10 cm apart, each one supplied with a potential growth bud (Ivens 1966). Experimentshave shown that very small rhizome fragments, kept moist in Sphagnum moss, provided they includeda node, no less than 84% of them regenerated and developed into flourishing plants. Old rhizome segmentswere less successful at regenerating, but juvenile segments, even those less than 6 mm in length,developed leaves within a few weeks (Salisbury 1962, p. 499).
The rhizomes of A. podagraria usually remain near the surface, rarely penetrating more than 15 cmdeep, but in fertile, moist, not-too-compacted soil, they can spread up to 100 cm horizontally in asingle growing season. Even a compacted soil will not stop the spread of the species, however, as manygardeners know! In fact, Ground-elder is often referred to as 'the gardeners' number one enemy', beingimpossible to extricate from among desirable cultivated plants without resort to repeated application ofchemicals (glyphosate) that very much risks the lives of the latter (Salisbury 1962, p. 499).
Not only does A. podagraria invade and tangle with the roots of other plants, it also has a habitof growing under path edges and into the foundations of walls, where digging it out is impossible, andconstant re-invasion of cleared areas made inevitable (Ivens 1966).
If it were not for the fact that most of the leaves are basal or radical, which effectively restricts thecanopy height of the plant to around 25-30 cm, the species would shade-out more species and dominate farmore ground than it currently does (Salisbury 1942, p. 221). As it is, it can form extensive stands,mainly by strong vegetative growth of its rhizomes.
Variation
There is very little variation, although in parts of Denmark, Sweden and Finland an unnamed form existswith once pinnate leaves and oblong to almost orbicular leaflets with an obtuse apex (Jonsell &Karlsson 2010). A variegated cultivar 'Variegatum' has leaves edged and splashed with ivory (Griffiths1994). As it possesses less chlorophyll, it is not quite as vigorous, competitive and invasive as thenormal fully green form.
Fermanagh occurrence
A. podograria has been recorded in 294 Fermanagh tetrads, 55.7% of those in the VC. In coldwinters, the plant dies down and goes dormant, but in milder, more sheltered conditions it remainswintergreen. It has been recorded during every month of the year in Fermanagh, but it is certainly moreprevalent between March and early November.
British and Irish occurrence
The same story of gradual spread from cultivation applies over almost all of B & I, the only hectadsfree of the grip of this pernicious weed in the New Atlas map being wetlands, highlands andextremely acidic peatlands, chiefly located in NW Scotland and in W Ireland (Preston et al.2002).
European and world occurrence
As with other introduced aggressively colonising weeds, it is not easy to identify just where A.podagraria originated, but probably it was in the moist deciduous forests of C Europe (Clapham1953). Nowadays, it appears native in moist woodland in most of temperate Europe, south to S France andItaly and eastwards to the W Caucasus and W Siberia. However, it has spread with man much wider thanthis, even to Arctic Scandinavia as well as to temperate N America (Tutin 1980; Hultén & Fries 1986,Map 1395; Jonsell & Karlsson 2010).
Uses
One of the most widely used English common names for A. podagraria is 'Gout-weed', sometimesamended to 'Goatweed'. This name has arisen as the plant has for centuries been recommended byherbalists as a cure for the very painful ailment gout. Indeed, the botanical name 'Aegopodium' is basedon the Greek 'aix', 'aigos' meaning 'a goat' and 'pous', 'podos' meaning 'a foot', allegedly referringto the shape of the leaves (Gilbert-Carter 1964). The Latin specific epithet 'podagraria' is alsoderived from 'podagra' meaning 'gout', or 'good for gout' (Gilbert-Carter 1964).
Gout is a painful form of arthritis that can arise suddenly as an inflammation of the joints due to uricacid crystals accumulating around them. It usually affects people over 50 years of age, especially oldermen, and it often first appears as a red, swollen and acutely painful big toe. An interesting modernendorsem*nt of the herbal efficacy of the species for treating gout appeared in BSBI News82: p. 51 (Senior 1999), although the current author (RSF) suggests it might be better torub an extract of the leaves on the sore joint – rather than to drink it, as Mr Senior reports he did.
The white rhizome is described by Grieve (1931) as, "pungent and aromatic, but the flavour of theleaves is strong and disagreeable". The plant is diuretic and astringent and, "can besuccessfully employed internally for aches of the joints, gouty and sciatic pains, andexternally as a fomentation for inflamed parts." Also, "the roots and leaves boiledtogether, applied to the hip, and occasionally renewed, have a wonderful effect in some cases ofsciatica" (Grieve 1931).
Oddly, A. podagraria gets no mention in the Medicinal plants in folk tradition: an ethnobotanyof Britain and Ireland (Allen & Hatfield 2006), which suggests these authors had little orno faith in the use of the species.
In addition to its history in herbal medicine, A. podagraria has also been used as a pot herb formany centuries. The leaves were boiled and used like spinach. They are described by Mabey (1996) as,"making a stringy but tangy dish". The leaves were also eaten as a spring salad (Grieve 1931).
Names
The undesirability of a weed can often be judged by the number of names applied to it by gardeners andothers. Grigson (1955, 1987) lists 20 names and Vickery (2019) also manages 20 (not identical toGrigson's list), including 'Farmer's plague' and 'Gardener's plague', plus other names that indicate theability of the plant to spread rapidly, such as 'Jack jump about' and 'Jump about'. Grieve (1931) givesan additional nine names not mentioned by Grigson and seven absent from Vickery's list. Assessed in thisway, A. podagraria must qualify as one of the worst weeds in the world, at least as far asgardeners and farmers are concerned.
At a glance, the leaves are not unlike those of Elder (Sambucus nigra), and hence names such as'Ground Elder', 'Dog Elder', 'Dwarf Elder', 'Dutch Elder', 'Bishop's Elder', 'Wild Elder' and so on(Grigson 1955, 1987). Many of the names have 'ash' as an element, eg 'Ashweed', but this is associatednot with the tree, but rather with 'ache' = parsley. So we have mention of 'Ground Ash', 'Pot-ash','White Ash', 'Achweed', 'Wild Esh' and 'Weyl-ash' (Grieve 1931; Grigson 1955, 1987; Vickery 2019).
Threats
A persistent and vigorous ground cover herb, mainly in shady habitats both 'artificial' and more natural,A. podagraria is so common, widespread and thoroughly naturalised, that in suitable vegetation itcan look perfectly native.
Native, varying from locally common to scarce. Eurosiberian temperate.
1806; Scott, Prof R.; Co Fermanagh.
May to October
Growth form and preferred habitats
S. latifolium is a hairless, robust, bright green, hollow stemmed perennial, 8-150(-200) cm tall,growing erect from a short, horizontal rhizome, 5-10 mm thick. It occupies shallow nutrient-rich, oftenstill alkaline water, over clay or mud substrates (Tutin 1980). The hollow stem is strongly ribbed andthe aerial leaves, usually 6-8 on the stem, can grow to 30 cm. The stems smell rather strongly ofparaffin or petrol when bruised. The aerial leaves have stalks which are also hollow and which clasp themain stem. They are usually simply pinnate, often with only 5-7 pairs of finely-toothed leaflets.
S. latifolium seed germinates underwater, produces 1-2 large, typical aquatic, finely divided, 2-3pinnate, submerged leaves, ie the species is heterophyllous, producing two quite different leaf types.It also dies down in the autumn and overwinters underwater. Thus, there is absolutely no doubt thatS. latifolium is a member of the aquatic flora of B & I, rather than an emergent terrestrialspecies (Cook 1998). Some Flora writers go halfway and describe it as semi-aquatic (eg Jonsell &Karlsson 2010), but the current author (RSF) believes the species is a good example of an aquatic plant,pure and simple.
The typical habitat is very wet, species-rich, tall-herb fen, developing as a floating mat of vegetationon the margins of large lakes and slow-flowing rivers (J.O. Mountford, in: Stewart et al. 1994).In England, it grows along the edges of lowland dykes, ditches and drains in fens and alluvial levelswhere the water is shallow and calcareous or base-rich (Garrard & Streeter 1983). In lowlandFermanagh, it grows in slow streams and muddy ditches. It prefers still or slow-moving, shallow waterthat is not acidic, is nitrogen-rich, and where the soil is formed from sedge peat, or has beendeposited by rivers (ie alluvial).
Greater Water-parsnip can compete with and tolerate the shade of other tall emergent herbs inspecies-poor reed-swamp, including reeds (Phragmites) and bulrushes (Typha), but itbecomes shaded out if wet woodland (ie swampy fen-carr) with species such as Alder (Alnus spp.),Ash (Fraxinus excelsior) and Willow (Salix spp.), encroaches upon the fen or ditch habitat(J.O. Mountford, in: Stewart et al. 1994).
The only other umbellifer of similar size to S. latifolium in this type of habitat is Cicutavirosa (Cowbane), which has much narrower, more finely cut leaf segments, so the two plantscannot readily be confused, even when not in flower. Another comparison is that Cicutavirosa is extremely poisonous and is avoided by cattle, whereas Sium latifolium is readilygrazed and stands of it are regularly subjected to trampling.
Since it is intolerant of grazing and frequent cutting, S. latifolium is often most common inditches adjacent to un-reclaimed fen or arable land, provided the water is kept open by occasional useof a weed bucket or scythe (J.O. Mountford, in: Stewart et al. 1994).
Toxcity and herbivory
WARNING – parts of the S. latifolium plant are poisonous (particularly the roots) and directphysical contact with it should be avoided at all times. Having said this, there are no recent reportsof poisoning by this species (Cooper & Johnson 1998).
Cattle and other stock seem to be immune from the plant's poison, but farmers discourage dairy cows frombrowsing it, since it gives their milk an unpleasant taste. S. latifolium can survive moderatebrowsing and trampling by cattle. However, drainage and excessive mechanical cleaning of waterways cankill it off.
Flowering reproduction
S. latifolium flowers in July and August. The inflorescence is a compound umbel of c 16-40 smoothrays, 6-10 cm across, each umbellule bearing c 16 small, 4 mm diameter white flowers. The terminal umbelconsists of hermaphrodite (bisexual) flowers and the lateral umbels are almost entirely of male flowers.The outer flower petals are not radiating. Bracts number 2-6 and are often large and leaf-like. Theflowers are pollinated by a range of insects including beetles, bees and flies (Fitter 1987). In theabsence of pollinators the flowers probably self-pollinate(https://pfaf.org/user/Plant.aspx?LatinName=Sium+latifolium, accessed 18 November 2021).
The fruit is ovoid, 3-4 mm and laterally flattened. The mericarps have distinctive thick ridges and shortrecurved styles (Tutin 1980; Blamey & Grey-Wilson 1989). The ridges on the fruits contain large,air-filled cavities that allow the mericarps to float for up to ten days, greatly enhancing thedispersal ability of the species (Egholm 1951). Otherwise, the seed is transient, surviving for lessthan one year (Thompson et al. 1997). Soon after flowering the aerial part of the plant dies downand disappears.
S. latifolium can live totally submerged for several years without flowering. In some years, itmay produce abundant seed, but opportunities for seedling establishment are few in the tall fencommunities that it typically frequents. In ditches, particular individuals can survive in tall reed forover ten years, but fresh recruits to the population are only seen following major work to cut down thetall vegetation, or following the use of a weed bucket to open up the community and create opportunitiesfor re-colonisation (J.O. Mountford, in: Stewart et al. 1994).
A history of rapid decline
Previously, this large umbellifer was widespread and typical of very wet, shallow, marshy watersidevegetation in B & I, but it suffered a catastrophic decline during the last 200 years and has becomescarce over much of its range. Changes in land use, particularly urban encroachment and theintensification of agriculture, resulted in the rapid loss of ponds and wetlands, leading to habitatfragmentation and isolation, and deterioration of both habitat and water quality.
Fermanagh occurrence
This previously quite widespread marsh umbellifer of soggy lakeshores and ditches has declinedsignificantly and become scarce in both B & I so that it is now listed as a 'Priority Species ofconservation concern' in both UK and N Ireland Biodiversity Action Plans (J.O. Mountford, in: Stewartet al. 1994; An Irish Flora 1996). However, it remains extremely frequent and abundantaround Upper Lough Erne, which must now be its current stronghold in these islands. The Fermanagh FloraDatabase has records of Great Water-parsnip in a total of 66 tetrads (12.5%), only one of which does notcontain a post-1975 record.
The very common and abundant occurrence of S. latifolium around Upper Lough Erne reflects theideal nature of the habitat for this tall and distinctive perennial, which although it appearsphysically robust, must in fact be very sensitive and demanding in terms of its physiological ecology.As the tetrad map clearly indicates, around Lower Lough Erne to the NW and along the River Finn in theSE of the VC, the plant is very much more local in its occurrence.
British and Irish occurrence
Elsewhere in B & I, the downward spiral of this species presence is very marked and obvious. InBritain, Stewart et al. (1994, p. 389) plotted records from 149 hectads with pre-1970 dates whereS. latifolium was present and only 66 hectads with subsequent finds. The New Atlas mapshows a total of 297 hectads with British records, but 175 of them (almost 59%) are pre-1970 in date. InEngland, it is still found in fair numbers south and east of a line drawn between the River Humber andthe Bristol Channel, but appears confined to Lincolnshire, the East Anglian Fens, the Norfolk Broads,the coastal levels of Kent and Sussex and the Somerset Levels. It appears most common in the coastallevels and flood plains of major rivers in Britain (J.O. Mountford, in: Stewart et al. 1994).Previously it was widespread along the Thames valley (Garrard & Streeter 1983).
In Ireland as a whole, there has also been a marked decline: from the 14 vice-counties listed by Scannell& Synnott (1987), mainly strung along the River Shannon up through the midland counties fromLimerick (H8) to Fermanagh (H33), the latest information in the New Atlas indicates that S.latifolium survives in just ten VCs (Preston et al. 2002). The New Atlasrepresentation in Ireland totals 34 hectads, eight of which are pre-1970 (ie 23.5%), a rather morecomfortable figure in terms of species survival. However, no one should be complacent when the totalnumber of squares is as low as this and there is any evidence of a decline. This message isconfirmed in the seventh revised edition of An Irish Flora (Webb et al. 1996), where theauthors regard S. latifolium as 'rare', whereas previous editions of the work described it as'rather rare'.
The Irish records are basically in two areas: on Lough Erne and along the course of the River Shannon inthe RoI.
Current conservation status in B & I: Classified as Nationally Scarce and endangered inthe UK, S. latifolium is actually too common around Upper Lough Erne to be listed for specificprotection in Northern Ireland under Schedule 8 of the Wildlife and Countryside Act and, similarly, itis not protected in the Republic of Ireland. It is listed as a NI Priority Species since more than 20%of the UK population is in Northern Ireland.
The Great Water-parsnip is listed in the UK Biodiversity Action Plans (UK BAP) and is included in EnglishNature's Species Recovery Programme involving translocation studies in several areas that previouslysupported populations of the species. In England, The Environment Agency, who are the lead partners inwork to recover this species, have produced plans to improve river and floodplain management that willbenefit other plants and animals that are features of these important habitats.
Seed from the Millennium Seed Bank, managed by the Royal Botanic Gardens, Kew at Wakehurst Place, iscurrently being used to try to restore populations of S. latifolium at suitable sites within itsformer range. Conservation projects involving translocations of S. latifolium have occurredindependently in at least seven counties of England, re-introducing the species in regions where it hasbeen lost or declined; however, the success of these translocations has so far been mixed.
European and world occurrence
S. latifolium is widespread across most of Europe, although it is very rare near the Mediterraneanregion and is absent from Portugal. Nevertheless, Hultén & Fries (1986, Map 1396) show it recordedfrom the very south of both Spain and Italy, although it is absent from all the Mediterranean islands.It is common to rare and mainly southern in Fennoscandia (common in both Denmark & S Sweden), but ithas markedly declined in Finland (Jonsell & Karlsson 2010). The indigenous range of S.latifolium extends across temperate Asia as far east as W Siberia. It is recorded as anintroduction in south-eastern Australia and Chile, but is otherwise unknown in both Japan and N America(Hultén & Fries 1986).
Names
The genus name 'Sium' is an old or classical Greek name for an unknown water plant, possibly a name givento at least two plants, thought to be Sium angustifolium (= Berula erecta) and Veronicaanagallis-aquatica (Blue Water-speedwell) (Gilbert-Carter 1964). The Latin specific epithet'latifolium' means 'broad leaved'.
The English common name 'Water Parsnip' appears to have been shared with S. angustifolium (=Berula erecta, Lesser Water-parsnip), a stoloniferous species. The name is inappropriate for boththese species, since neither possesses anything remotely similar to the fleshy rootstock of the truevegetable Parsnip (Pastinaca sativa).
Threats and causes of decline: The tall-herb fen vegetation that supports S.latifolium grows in a sensitive aquatic habitat which has recently suffered major decline acrossmuch of Britain and Ireland due to development, drainage and pollution, including nutrient run-off fromagricultural land. S. latifolium is sensitive to disturbance and cannot survive regular cuttingor ditch clearance. It can only tolerate very occasional dredging or reshaping of its ditches, ormoderate levels of grazing and trampling by stock. The neglect of ditches, allowing scrub and youngwoodland to invade and become established, also kills off the plant. Even in conservation protectedsites, decline has been observed, and new plants appear only rarely (J.O. Mountford, in: Stewart etal. 1994).
Native, rare or occasional and rather local. European temperate, but also considered native in W & CAsia & N America; introduced in several parts of Africa and in Australia.
1806; Scott, Prof R.; Co Fermanagh.
June to September.
Growth form, identification and preferred habitats
An always lowland, glabrous, rhizomatous and stoloniferous aquatic perennial, B. erecta is veryvariable in vegetative form and it is rather similar in both its simply pinnate leaf and its habitat totwo other locally well represented umbellifer species, facts which could give rise to someunder-recording of it. The species with which B. erecta might be confused are Siumlatifolium (Great Water-parsnip) and the morphologically very variable Apium nodiflorum(Fool's-water-cress). B. erecta is by far the scarcest of the three in Fermanagh, theircomparative tetrad representation being – Berula erecta 17, Sium latifolium 66 andApium nodiflorum 120.
Overall, B. erecta is around 100 cm in maximum height, with leaves that are intermediate in sizebetween S. latifolium and A. nodiflorum. The leaves, with 4-9 pairs of leaflets, are adull, bluish- or yellowish-green in colour, with narrower, more irregularly toothed divisions than theother two species (Tutin 1980), although this in fact is another rather variable character in B.erecta! Submerged leaves are similar to aerial leaves, unlike those of S. latifolium. Thepresence on the leaf stalk of a discoloured 'ring-mark' or septum, some distance below the lowest pairof leaflets (which are sometimes quite rudimentary), is diagnostic in separating vegetative specimens ofB. erecta from A. nodiflorum (Wigginton & Graham 1981). The BSBI Plant Crib1998 (pp. 220-1), tabulates and illustrates several distinctive differences which, takentogether, allow ready separation of these three similar umbellifers (Rich & Jermy 1998).
B. erecta is typically found in full sun or moderate degrees of shade,
either as an emergent on the damp margins or shallow waters around, lakes, ponds and ditches, or elsesubmerged or emergent in seasonally flooded ground beside fast or slower running water in streams andrivers. It prefers fertile, nutrient-rich, calcareous or near neutral, eutrophic or sometimesmesotrophic environments, and fine to medium textured, organic mud or clay soils, habitat conditionswhich in Co Fermanagh are frequently provided by the long, dissected shoreline of Upper Lough Erne andthe banks of the River Finn, localities where it has most often been found (Haslam et al. 1975;Preston & Croft 1997).
Vegetative and flowering reproduction
B. erecta has a short basal rhizome, 2-10 mm thick, with stolons arising at the lowermost nodes atthe base of the hollow stem. Local vegetative spread is commonly achieved in the spring by growth of theshort-lived stolons or longer surviving rhizome. Small plantlets formed in this manner, together withvegetative fragments of the plant can become detached and float off to colonise fresh sites in the watersystem.
Little or nothing is known of the frequency or success of seed reproduction by the species, butterrestrial and emergent plants certainly flower better and submerged ones very often fail to do so(Preston & Croft 1997). In Scandinavia (Denmark, S Norway and S Sweden), when growing in water,B. erecta has been observed to produce large, sterile populations (Jonsell & Karlsson 2010).The English Centre for Ecology & Hydrology has issued an advice leaflet for the control of B.erecta as a nuisance aquatic weed. Mechanical cutting or strimming gives only temporary respiteand chemical control is recommended(https://www.researchgate.net/publication/234111350_CEH_Information_Sheet_18_Berula_erecta_water_parsnip,accessed 20 November 2021).
If it occurs at all, flowering of B. erecta takes place from July to September. The compound umbelcontains 7-17 rays and is subtended by 4-7 bracts that sometimes are leaf-like. Each ultimate umbel(umbellule) contains 14-22 white flowers that are not or slightly irregular (zygomorphic) and the petalsare emarginated (ie notched at their tip). The fruit is ± orbicular, 1.3-2.0 mm, somewhat compressedlaterally and a dark greyish-brown in colour. The mericarps bear small indistinct ridges and, like thefruits of S. latifolium, they are adapted to water dispersal, their cells having large air-filledspaces within them that enable flotation (Sell & Murrell 2009; Jonsell & Karlsson 2010).
Toxicity
The rhizome and creeping stolons are poisonous, but not especially so, to most stock animals (Grieve1931, p. 617; Cooper & Johnson 1998). The entry for Berula in The plant book(Mabberley 1997) has it that it proved fatal for cattle in New South Wales, Australia.
Fermanagh occurrence
B. erecta was one of the very first flowering plants recorded in Co Fermanagh by Prof Robert Scottin a list dated 1806. Although there are a total of 43 records in 17 tetrads for this species in theFermanagh Flora Database, 15 of the squares with post-1975 dates, only two sites have been found by theRHN and the current author (RSF). These finds were at Tully Castle (the only site for the species onLower Lough Erne), where RHN found it in July 1990, and on the shore of Derrymacrow Lough, July 2002,when RHN & RSF were accompanied by J.S. Faulkner and I. McNeill. Thus B. erecta is definitelyregarded as a rare or only very occasional species in Fermanagh. The tetrad map shows that apart fromthe Tully Castle site, B. erecta is almost confined to Upper Lough Erne, but in terms offrequency it is very much concentrated in the far SE of this wetland area, centred around Drummully Tdand Wattle Bridge near Crom and the entrance to the Old Ulster Canal.
Irish occurrence
B. erecta has declined considerably since the mid-18th century in NE Ireland (FNEI2; FNEI 3). In NI, it is now found mainly in the Lecale area of Co Down (H38) and localand sparingly along the Newry Canal in Cos Down (H38) and Armagh (H37). There is a similar sparsepresence around Lough Neagh and Upper Lough Erne, but it is rare and extremely thinly scattered orabsent elsewhere in NI (Flora of Lough Neagh). In the RoI, B. erecta is quite frequent inthe E & C and occasional elsewhere. It is very much more frequent and widespread in the RoI incomparison with north of the island (An Irish Flora 1996; New Atlas).
British occurrence
Despite a known decline in Britain since around 1950, presumably due to drainage and habitat destruction,B. erecta remains fairly common, widespread and stable in most of England south of a line betweenLancaster and Hull. However, it is rare in SW England, Wales and Scotland, becoming increasingly coastalin all these regions (M. Southam, in: Preston et al. 2002).
European and world occurrence
B. erecta is native and widespread across Europe from being local in the southern tip of bothNorway and Sweden, then widely present in Denmark, the distribution continues southwards to S Spain andeast to N Greece. Although less frequent in the Mediterranean basin, it has been recorded in Corsica,Sardinia and Sicily as well as throughout Italy. It is also present in Turkey, the Middle East, Egyptand Ethiopia, and it stretches onward into W & C Asia. It is also indigenous and widespread intemperate N America and has been introduced in Australia (Hultén & Fries 1986, Map 1397).
Names
The genus name 'Berula' is a plant name mentioned by Marcellus Empiricus (Gilbert-Carter 1964) that hasbeen borrowed, recycled and reapplied to this plant when it was reclassified and removed from the genusSium. Marcellus Empiricus is a shadowy figure also known as Marcellus Burdigalensis (meaning'Marcellus of Bordeaux'), who was a French, or Gaullish, herbal writer at the end of the 4th andbeginning of the 5th centuries. His only extant work is the Latin book, De medicamentis, atypical herbal pharmacology which drew on the work of many earlier medical and scientific writersincluding Pliny the elder, as well as on popular folklore remedies and magic of the time (Sharpe 1964).As such, 'Berula' is another example of a 'book name', more or less chosen at random to act as a label.The specific epithet 'erecta' is too obvious to translate, but the growth of the plant can be quitesprawling, not always erect.
The English common name 'Lesser Water-parsnip' simply informs the student that the plant is somewhatsmaller than the similar Great Water-parsnip (Sium latifolium) and neither plant has the rootqualifications to make it a parsnip.
Threats
None.
Native, very rare, possibly an error, but perhaps overlooked and under-recorded. Suboceanicsouthern-temperate, mainly restricted to W & S Europe.
10 June 1974; Hackney, P.; Spectacle Lough, Dresternan Td.
June only.
Growth form and preferred habitats
Parsley Water-dropwort is a glabrous, somewhat heterophyllous perennial, up to 70(-100) cm tall, withtuberous roots and rigid, solid stems and 5-7 pinnate or bi-pinnate stem leaves with narrow, linear,fine-divided leaflets. The smaller basal leaves have broader leaflets but are short-lived and witherbefore the plant flowers. O. lachenalii primarily grows in coastal habitats around the shores ofB & I, in marshy grasslands and reed swamps with fine-textured, fertile, often brackish soils, or inshallow brackish estuarine waters where it stands out on account of its greater height amongst theshorter salt-marsh vegetation (Tutin 1980; Garrard & Streeter 1983; Walls 1995).
At its much less frequent inland sites in B & I, O. lachenalii really is very local and tendsto occur in lime- or base-enriched examples of rough grassland in marshes, tall-herb fens andfen-marshes – another description of what is locally referred to in Fermanagh as 'water-meadows', ieseasonally flooded ground under permanent pasture.
Flowering reproduction
The plant relies entirely on seed for its reproduction and flowers throughout the summer from June toSeptember. The inflorescence is a compound umbel and there are up to ten secondary or partial umbels,not flat-topped, with five lanceolate bracts. The umbels all contain both hermaphrodite (bisexual) andmale flowers, the petals of which are white, slightly zygomorphic (irregular) and notched (emarginated).The umbel rays remain un-thickened after flowering when the fruit develops, and its pedicels are lessthan 0.5 mm thick, two features which together help distinguish the species from the even rarer O.pimpinelloides (Corky-fruited Water-dropwort), which in Ireland is a rare introductionconfined to the far SW. Also, O. lachenalii fruits are initially conical, but they becomebarrel-shaped when ripe, with short styles (2.5-3 mm long), compared to fruits 3.5 mm long in O.pimpinelloides, with styles as long as the fruit. The fruit is obovate in outline, greenishbrown and does not split at maturity. The mericarp ridges are five in number and the lateral ones havelarge cell cavities that aid flotation and enable short-distance dispersal in water (Tutin 1980; Garrard& Streeter 1983; Walls 1995; Jonsell & Karlsson 2010).
Fermanagh occurrence
There are only two records for this umbellifer in the Fermanagh Flora Database. The first one listedabove is on the Western Plateau and has a voucher in BEL. The second find, made on 27 June 1996by the EHS Habitat Survey Team, was recorded on or near the shore of Rosskit Island, Lough Melvin. Thereis no voucher for the latter, although it really requires one to be fully accepted as a valid SecondCounty Record. Paul Hackney is not entirely sure that his identification is correct, casting doubt onthe reality of this species occurring in Fermanagh.
There is a plentiful supply of suitable seasonally flooded water-meadow habitats for O. lachenaliiin lowland Fermanagh, particularly around the Upper Lough Erne basin, and RHN and the currentauthor (RSF) as joint BSBI County Recorders suggest this species should be more actively searched for infuture, since it could well be present and is perhaps simply being overlooked and under-recorded.
British and Irish occurrence
The New Atlas hectad map demonstrates that there are very few inland records of O.lachenalii anywhere in Ireland, although in Britain, by comparison, inland sites are thinlyscattered in the English Midlands and as far south as Bristol and Hampshire. It is absent from N & EScotland, but does occur on several of the Western Isles, including Rum and the Outer Hebrides(Pankhurst & Mullin 1991; Preston et al. 2002; Pearman et al. 2008). However, thenumber of suitably moist inland sites in England and Wales has been declining for some time due todrainage and land-infill operations (M. Southam, in: Preston et al. 2002).
European and world occurrence
O. lachenalii belongs to the southern temperate phytogeographical element and is almost completelyrestricted to W & S Europe. It extends from coastal Denmark and the southern tip of Sweden south toS Spain and Portugal, and eastwards to Corsica, Italy, Sardinia and Sicily and onwards as far as Polandand Yugoslavia. It also occurs in Algeria, but is very rare there (Hultén & Fries 1986, Map 1400).
Names
The Latinised specific epithet was given to honour the 18th century botanist, Werner de Lachenal(1736-1800) (Sell & Murell 2009). The English common name 'Parsley Water-dropwort' is another rathermisleading 'book name' associated with this genus.
Threats
Drainage and land use change, including coastal and inland development and the intensification ofa*griculture have reduced the availability of suitable sites for this species (M. Southam, in: Prestonet al. 2002).
Native, uncommon, yet locally frequent. European temperate.
1881-2; Barrington, R.M.; Co Fermanagh.
January to September.
Growth form and preferred habitats
A tall, robust and vigorous, tuberous, wintergreen perennial that produces lush amounts of its 3-4pinnate, parsley-scented, basal leaves when growing in wet conditions, O. crocata is the largestand the most widespread of the seven species of the genus in B & I, four of which occur inFermanagh. It often grows luxuriantly, forming extensive clonal patches, and its thick, hollow, ribbedstems can reach a height of 150 cm when inhabiting reliably wet, nutrient-enriched sites (Walls 1995).Stem leaves are 2-3 pinnate, with narrower leaflets than the basal leaves.
It is a lowland plant of wet, moderately acid to neutral, muddy soils and shallow waters, mainly in sun,but occasionally under partial shade of woodland canopy.
Although it is generally most frequently found in or on acidic water and soils and, therefore, oftenconsidered a calcifuge, O. crocata is tolerant of lime-rich waters and can also be found in arange of calcareous habitats throughout B & I, including wetlands in the karst Burren district, CoClare (H9) and along many chalk-derived rivers in S England (Webb & Scannell 1983; Walls 1995). Incoastal sites, it can also colonise stony storm beaches and boulders at the top of beaches providedthere is flushing ground water and it can also grow on dripping or flushed parts of sea cliffs (Preston& Croft 1997; M. Southam & M.J. Wigginton, in: Preston et al. 2002).
The species can survive a limited amount of drainage and the influence of agricultural improvements andfertiliser or slurry run-off, but plants become tough and weedy if the water table drops too low (Walls1995). The plant is sufficiently ruderal, weedy and stress tolerant to successfully colonise crevices inwaterside masonry or those in rocky stream-sides in coastal parts of W Britain (Preston & Croft1997). Further south, on the Isles of Scilly, O. crocata can even grow in very dry conditions onthe tops of walls (Lousley 1971). The established strategy is categorised as C/CR by Grime et al.(1988, 2007), meaning it is ecologically intermediate between a Competitor and a Competitive Ruderal.
Fermanagh occurrence
Locally, O. crocata has been recorded from January to September in a total of 79 Fermanaghtetrads, 15% of those in the VC. Hemlock Water-dropwort is unusual amongst locally widespread largeemergent plants in Fermanagh in being much more frequently recorded around the calcareous, open watershores of Lower Lough Erne and its wooded islands, than around the more acidic and more definitelyeutrophic Upper Lough. Around Lower Lough Erne shore area, it has post-1975 records in 35 tetrads,compared with the very sheltered, dissected shores of Upper Lough Erne where it has been recorded injust eight tetrads during the same 35-year period (1975-2010). The species is also frequent along thebanks and ditches linked with the Colebrooke and the Ballycassidy Rivers that flow into Lower Lough Erneand in the west of the VC with Lough Melvin and adjacent Upper and Lower Lough Macnean.
The Fermanagh populations of O. crocata are stable, or possibly even spreading somewhat withintheir well-defined area and there have been 25 additional records since 2010 (BSBI Database accessedRSF, 27 January 2022).
Toxicity
All species of the genus Oenanthe are toxic to man and livestock to some extent, as the 'dropwort'portion of their English common name implies. However, O. crocata is by far the mostdangerous and lethal of them all. The active principle, 'oenanthetoxin', is chemically verysimilar to 'cicutoxin' found in Cicuta virosa (Cowbane). A convulsant poison, the toxin isunaffected by drying or storage, making it very dangerous, even when decomposing parts of the plant areleft exposed on diggings after drain clearing. The roots tubers are the most poisonous part of theplant and ingestion of just a small portion of these is sufficient to kill a cow (Walls 1995).Cattle, horses, sheep and pigs have all been poisoned on occasions. Death is often sudden, sometimeswithout the appearance of any clinical signs. People mistakenly cooking the stems for Wild Celery(Apium graveolens), or making soup of the tubers, thinking it is Wild Parsnip (Pastinacasativa), are occasionally poisoned (Cooper & Johnson 1998).
Flowering reproduction
In line with other Oenanthe species, effective reproduction in O. crocata appearsheavily dependent on seed, dispersal being usually achieved by floatation in water flushing or drainingthe habitats it tends to occupy. Flowering occurs in June and July, the inflorescence, 5-10 cm indiameter is a compound umbel with 10-40 rays, numerous 2 mm white flowers and linear bracts. Theterminal umbel has hermaphrodite, bisexual flowers, the lateral umbels with mainly male flowers. Theouter flowers of each umbel have petals somewhat unequal, ie radiating, making the flowers moreconspicuous and attractive to insect visitors (Sell & Murrell 2009).
As with all Oenanthe species, nectar is well exposed and pollination of the protandrous flowers(anthers maturing before the stigmas) is carried out by flies and other unspecified insect visitors(Proctor & Yeo 1973; Fitter 1987). The fruit is 4-6 mm long, cylindrical or barrel-shaped, thestyles about half the length of the fruit. The pedicels are not thickened after fruiting, nor are theyconstricted at the top. The mericarps have slender ridges that possibly contain air-spaces that wouldaid dispersal by flotation, but this is not mentioned by any reference the current author (RSF) hasaccessed (Tutin 1980; Clapham et al. 1987). There does not appear to be any available informationon seed longevity, but seeds of other Oenanthe species are transient, surviving for a maximum ofone year.
After flowering and fruiting the stout, hollow flowering stems often collapse, rot and disappear (Preston& Croft 1997)
Vegetative reproduction
Finger-like, cylindrical-obovoid tubers that overwinter the species are formed from buds at the base ofthe plant at or close to the soil surface in the autumn. At the same time, the current tuber orrootstock that gave rise to the aerial shoot, withers and dies off. The overwintering tubers can readilydetach and spread the species, especially in flood waters in the spring (Walls 1995; Preston & Croft1997).
British and Irish occurrence
The New Atlas hectad map of B & I displays a strong southern and western and essentiallylowland distribution. O. crocata is uncommon in areas with a mean January daytime temperature ofless than 5ºC (Rich et al. 1996). Having said this, its distribution stretches beyond Invernessand it has colonised as far north as Orkney, where it was first recorded in 1988 (Walls 1995; NewAtlas).
The Irish distribution appears rather curious, the plant being frequent (sometimes abundant) in the N, S& E, but scarce or absent in large areas of the lowland C & W. Although farmers would likely bekeen to eradicate such a poisonous species, they rarely make the attempt, probably regarding the task asnear impossible (Preston & Croft 1997).
European and world occurrence
Hemlock Water-dropwort is a member of the Suboceanic southern temperate phytogeographical element, itsdistribution being strongly oceanic (or Atlantic), western and southern in Europe and the Mediterraneanbasin. The most northerly station on the continental mainland is at Voorne, in Holland, where it wasfirst noted in 1975. It extends south to Spain, Portugal and Morocco and eastwards in the WMediterranean to reach Italy (Preston & Croft 1997). O. crocata appears to be increasing andspreading northwards from its warmer southern strongholds, probably as a result of recent and continuingglobal warming (Walls 1995).
Names
The Latin specific epithet 'crocata' means 'citrus yellow' or 'saffron-like', referring to the sap of theplant that turns yellow on exposure to the air when it is cut (Gilbert-Carter 1964; Gledhill 1985). TheEnglish common name 'Hemlock Water-dropwort' is another invented 'book name' that is less useless thanmost such names because it alerts people to the deadly poisonous nature of this species by making aconnection to the related, extremely dangerous genuine Hemlock, Conium maculatum.
Threats
Stable, or possibly spreading within its well-defined area.
Native, very rare and perhaps declining, but also very possibly under-recorded. Oceanic temperate,occurrence limited, disjunct and declining.
1 August 1986; EHS Habitat Survey Team; shores of Inishcollan Td and Creaghanarourke Island, Upper LoughErne.
August.
Growth form and preferred habitats
A truly aquatic, wintergreen perennial with stems 30-100 cm long, floating, procumbent or ascending whenflowering, O. fluviatilis produces fibrous roots rather than forming tubers and its lower leavesare always submerged and are deeply cut into filiform lobes. Being entirely aquatic and confined tolowland waters, O. fluviatilis has a longer growing season than related terrestrial orsemi-aquatic Oenanthe species and thus does not require tuberous storage organs (Walls 1995). Asa general rule, as its names suggest, River Water-dropwort is found in constantly running water atdepths varying from shallow to 1.5 m, in small rivers and in streams that usually are calcareous andmoderately eutrophic (ie meso-eutrophic). However, this is not an absolute situation, since it canoccasionally also inhabit still or sluggish waters, eg in fenland man-made ditches and canals. In termsof bottom substrate, it prefers organic-rich or sandy ones poor in calcium and avoids only deep, veryfine silty conditions. It seems to be confined to relatively cool, clear water (Cook 1983; Preston &Croft 1997).
In comparison, the very similar, closely related O. aquatic (Fine-leaved Water-dropwort), withwhich it is easily confused, requires or prefers sluggish or still waters in lakes, ponds and ditches.In addition, O. aquatica often contends with very shallow pools that dry up in summer, forcingthat species into a terrestrial mode of growth form not seen in O. fluviatilis (Walls 1995;Preston & Croft 1997; Sell & Murrell 2009).
Fermanagh occurrence
There are only five records for this aquatic in the Fermanagh Flora Database at present, all of whichpost-date 1985 and are confined to the Upper Lough Erne area and along the River Finn. RHN and thecurrent author (RSF) believe there is a possibility that two of the records might be mis-identificationsof the closely related O. aquatica. The two species are very similar in form, behaviour and areecologically alike, being found in Fermanagh in rivers, streams and ditches near lakeshores.
The original plant list for Ross Lough, for instance, records both O. fluviatilis and O.aquatica and the EHS Habitat Survey Team clearly marked O. fluviatilis on their fieldcard with the code '3', which indicates that they knew the species was noteworthy and rare. Of these twoOenanthe species, however, O. aquatica is much more common and widespread in Fermanagh andelsewhere in B & I.
Both species are definitely aquatic members of this genus, since for most of the year their plants aresubmerged, only producing aerial or, in the case of O. fluviatilis, aerial and floating stems, inthe early summer. They become conspicuous (and fully recognisable) only for a brief period in latesummer when they flower and fruit. This is why all five Fermanagh records were determined in the monthof August when the fruits are mature and recognisable.
The record details of the other four finds of O. fluviatilis are: marsh to E of Inishroosk, 1August 1986, EHS Habitat Survey Team; S shore Ross Lough, 5 August 1986, S.A. Wolfe-Murphy & L.W.Austin; Kilturk Lough, Derrymacrow Td, 14 August 1986, P.J.T. Brain & T. Waterman; S shore of TrasnaIsland, 19 August 1986, P.J.T. Brain & T. Waterman (a rather doubtful determination).
Flowering reproduction
It is well known that some stands of O. fluviatilis only rarely produce flowers and hence they arereadily over-looked, especially since they very often associate with aquatic Ranunculus species,submerged Potamogeton species and other aquatic plants with finely dissected leaves like theirs.When plants do manage to flower, it is usually in slow flowing or sluggish waters and it takes place inJuly and August (Preston & Croft 1997). In comparison with the submerged leaves, the aerial onesthat accompany the flowering stem have broader leaflets up to 10 mm long, with blunt tips. The umbelsarise from the axils of leaves and there are no bracts. Fruits are longer than in most otherOenanthe species, 5-6.5 mm, ovoid in shape and ripen in August and September (Walls 1995).However, ripe fruits are rarely found and the frequency of successful seed production is unknown (Cook1983; Preston & Croft 1997).
Vegetative reproduction
In fast-flowing waters, O. fluviatilis stems root at their nodes and often develop large, clonalmats that rarely flower. In these circ*mstances, the species reproduces by vegetative fragmentationwithout any specialised structures being involved.
British and Irish occurrence
In Britain, O. fluviatilis is completely absent from Wales and Scotland. In England, apart from afew outliers in W Yorkshire, it is virtually restricted to the area south of a line between the Wash andthe Severn estuary. In comparison with O. aquatica, O. fluviatilis is the scarcer of thetwo species, the New Atlas survey mapping it in just 160 hectads from the 1987-99 date range inthe whole of B & I.
The New Atlas map displays quite a difference between the distribution of O. fluviatilis inEngland and in Ireland. In the latter, the species is thinly but rather widely scattered, mainly in CIreland, but it extends much further north than it does in England, reaching Mountsandel, near Colerainein Co Londonderry (H40).
O. fluviatilis appears to be slowly declining throughout these islands, probably due to graduallyincreasing eutrophication as a result of pollution and agricultural fertiliser run-off, together withexcessive disturbance from mechanised stream channel clearance operations and pleasure boatingactivities (Preston & Croft 1997).
European occurrence
Beyond our shores, O. fluviatilis is endemic to NW Europe – being confined to N France, Germanyand W Denmark, where it is nowhere common. In fact a significant proportion of the species totalpopulation resides in B & I (Cook 1983; Walls 1995).
Threats
O. fluviatilis is sensitive to pollution, including excessive eutrophication, and also tolarge-scale disturbance from over-zealous drainage and boating.
Native, uncommon yet locally frequent. Eurosiberian temperate, also disjunct in mountains of SE Asia andan introduction in New Zealand.
1806; Scott, Prof R.; Co Fermanagh.
May to November.
Growth form and preferred habitats
An erect to sprawling floating marginal aquatic plant, O. aquatica differs from other members ofthe genus Oenanthe in being a short-lived annual or biennial. Young plants initially developtubers on their fleshy roots, but these storage organs are re-absorbed and disappear whenever the plantmatures and produces a flowering stem. This tuber behaviour is also the pattern in the closely relatedperennial species, O. fluviatilis (River Water-dropwort).
In the summer months, the shallow, still or sluggish, marginal waters of the ecologically open, butsheltered shores of lowland lakes, ponds, rivers, drains or swampy fens that this normally aquaticspecies occupies, frequently dry out or have markedly reduced water levels in summer. When this happens,the previously submerged or floating plant produces a terrestrial form with feathery, finely divided,3-4 pinnate aerial leaves that differ considerably from its submerged filiform leaves (ie it isheterophyllous) (Jonsell & Karlsson 2010). Under these drier conditions, the terrestrial plant thenproduces erect or sprawling, stout, flowering stems, 60-150 cm tall (Walls 1995).
O. aquatica tends to prefer shallow ponds and ditches in damp or marshy pastures, where it growson deep, silty, clay- or mud-based, often eutrophic soils. This type of site is kept open by grazing andtrampling cattle (but see Toxicity section below) (Preston & Croft 1997).
Although it prefers fully illuminated situations, the species can tolerate moderate levels of shade fromnearby trees. However, it avoids closed vegetation in fen carr, tall fens or reed swamps.
Fermanagh occurrence
The typically tall, hollow, striate, 1 cm wide flowering stems of O. aquatica make it adistinctive feature of many of the nutrient-rich, muddy, organic soils of drainage channels that connectthe multitude of lakelets around Upper Lough Erne to one another and to the shores of the eutrophiclarge lake itself. Elsewhere in Fermanagh, O. aquatica features along the slow flowing lowerreaches of rivers and around suitable shallow ponds in pastures and water meadows, where water levelsfluctuate considerably with prevailing precipitation. Currently, the Fermanagh Flora Database containsrecords of O. aquatica from 56 tetrads (10.6%), 50 of them with post-1975 dates.
Flowering reproduction
Flowering usually occurs from June to September. The inflorescence is a series of small compound umbels,2-4 cm diameter, all leaf-opposed, plus a terminal one. Each umbel consists of 4-16 rays and pedunclesare usually shorter than the rays. Most umbels contain both hermaphrodite and male flowers, the bisexualones being in the majority (Jonsell & Karlsson 2010). The flowers are white and the outer petalsscarcely radiating. The leaflets of the aerial leaves are smaller than those of the closely related, butvery much rarer O. fluviatilis – only about 5 mm long and they have acute tips, making the aerialleaves of O. aquatica look daintier than those of O. fluviatilis. Pollination is carriedout by a range of insect visitors, including beetles, flies and bees (Fitter 1987).
In August, the mature fruits of O. aquatica are also smaller, being only 3-4.5 mm long, comparedto 5-6.5 mm in O. fluviatilis. These features, plus the more terrestrial habitat, are the mostreliable characters for separating these two Water-dropwort species (Tutin 1980; Walls 1995). The fruitmericarps have five prominent but low, greenish to yellow-brown, rather thick ridges and the pedicelsare not thickened (Sell & Murrell 2009). The mericarp ridges contain cells with large air spaces toassist flotation and water dispersal (Jonsell & Karlsson 2010).
The seeds of O. aquatica germinate soon after they are shed and 'swarms' of seedlings can be foundon ground from which water has receded. Seedlings emerging in autumn behave as winter annuals, floweringthe next summer, while those that spring germinate will complete their life-cycle the same year and thusare summer annuals. The longer-lived winter annuals accumulate more photosynthetic reserves in theirtemporary tubers and produce larger aerial plants and more seeds (Preston & Croft 1997).
Overwintering plants and temporary tubers
Both O. aquatica and O. fluviatilis are wintergreen and they have fibrous rather thantuberous roots. Since the main bulk of the plants spend most of the year submerged in water, they have amuch longer growing period than the more clearly terrestrial or semi-aquatic species of the genus. Thisexplains why they do not require or form any specialized overwintering organ, such as root tubers. Youngplants, however, do develop temporary root tubers before they reach flowering condition and thephotosynthate energy stored in them becomes completely consumed powering the sexual reproduction processof the mature plant (Tutin 1980).
Vegetative reproduction
The plant has a procumbent stem which roots at the nodes acting like stolons and producing vegetativeoffsets. The individual flower stems are annual or biennial, ie they are monocarpic. Overwinteringsurvival is achieved by the annual populations of vegetative offsets and seed (Clapham et al.1987).
Toxicity
Like all Oenanthe species, O. aquatica is considered poisonous, the active principle beingoenanthotoxin, a polyunsaturated higher alcohol, chemically similar to cicutoxin found in Cicutavirosa (Cowbane). It is a convulsant poison that is unaffected by drying and storage. There havebeen reports of cattle deaths caused by it in Poland and Sweden (Cooper & Johnson 1998).
British and Irish occurrence
Despite its hectad frequency in the New Atlas map, this is an uncommon species throughout most ofits B & I range. The British distribution shown in the New Atlas is rather odd, the plantbeing mainly southern in England plus in E Wales, but more or less absent from a broad tract down thecentre of England. Presumably, this vacant area represents the outcome of a combination of factors thatmake habitats unsuitable for the species, including higher ground and both dense urban and widespreadfarmland development. The distribution only just reaches the extreme SE of Scotland, so that temperaturelimits must clearly also apply (New Atlas). In fact, O. aquatica populations have declinedin England in recent decades and have become rather scattered (Rich & Woodruff 1996).
In NI, O. aquatica is chiefly found in Lough Neagh, the River Bann and Lough Erne. In the RoI, itis much more widespread, although again it is locally frequent in the centre and much rarer elsewhere(An Irish Flora 1996).
European and world occurrence
The species is widespread throughout middle latitudes in Europe and also into W Asia plus a few scatteredareas in the SC Asiatic mountains. On the Mediterranean islands, it is only present on Corsica andSicily. It has been accidently introduced to New Zealand, probably as a seed contaminant (Hultén &Fries 1986, Map 1401).
Threats
Drainage operations and excessive disturbance are potential, but not really actual threats in theFermanagh area.
Introduction, possibly an archaeophyte, declining and now very rare. European temperate, an introducedweed in both N Europe and N America.
1892; Praeger, R.Ll.; cultivated field by the Ballycassidy River.
Few recording dates with months, but mainly July.
Growth form and preferred habitats
A ruderal annual or biennial, glabrous herb, very variable in size and many other characters, A.cynapium has smooth, thin, dark green, finely dissected leaves that give off a distinctiveacrid, unpleasant odour when bruised, totally unlike the familiar, culinary Garden Parsley(Petroselinum crispum). The most characteristic distinguishing feature of the plant when inflower is the downwardly directed, strongly reflexed bracteoles beneath the smaller umbels of whiteflowers. They are usually three in number, but can be four or five (Salisbury 1964; Jonsell &Karlsson 2010).
A. cynapium is chiefly a weed of lowland, arable and disturbed ground, neglected, often boundarywaste areas, hedge banks and gardens. Less commonly, it is found on river-banks and in poached, heavilytrampled pastures and around gateways. It generally prefers more basic or lime-rich, sandy or loamy,cultivated or disturbed soils and sunny, warm ground situations. The established strategy of the speciesis categorised as R/CR (ie intermediate between Ruderal and Competitive-Ruderal) by Grime et al.(1988, 2007).
Flowering reproduction
Small, white, bisexual flowers are produced June to August, the ultimate umbel divisions, uniquely3-bracted; after pollination by flies and bees of various sorts, the typical sized garden form of thisweed may produce up to 3,000 green, oval, flat, heavily ribbed fruits, which split in two before theirrelease (Salisbury 1964). Germination can occur in autumn or, more commonly, in the spring. As is oftenthe case, and especially so in this family, little or nothing is known about its real powers ofdispersal, but in the past it has certainly been regularly (frequently) transported by agricultural manas a contaminant of commercial crop and pasture seed.
Many references regard it as long-persistent in the soil seed bank, surviving burial for five or moreyears (Thompson et al. 1997).
Variation
In Britain, where the species is still widespread in the lowlands, the plant is very variable andsufficiently polymorphic for four subspecies to be recognised, although they are not always separated byfield botanists. The most common form, subsp. cynapium occurs throughout the range of thespecies, while the dwarfish subsp. agrestis (Wallr.) Dorstál., which is very probably anarchaeophyte, is most frequent on arable land in S England, since it can escape the reaper thanks to itsreproductive shoots being shorter than the crop stubble (Salisbury 1964; Tutin 1980; M. Southam, in:Preston et al. 2002).
Two additional taxa, sometimes considered varieties of subsp. cynapium (Stace 2019), are subsp.gigantea (Lej.) P.D. Sell, annual, stem 100-200 cm, bracteoles 1-2 times as long as partialumbels, and subsp. cynapioides (M. Bieb.) Arcang., biennial, 100-200 cm, bracteoles 2-3(-4) timesas long as partial umbels (Sell & Murrell 2009).
Toxicity
A. cynapium is a very poisonous plant, but the polyacetylene toxins it contains (conine andcynapine) are a danger to livestock only when it is eaten fresh and in large quantity. Fortunately, theplant gives off a repulsive warning odour, sometimes described as acrid and mouse-like (the genus namederives from the Greek 'aitho', 'I burn'). The stink increases the more the tissues are bruised, readilydiscouraging humans from consuming it (Salisbury 1964, p. 289).
That said, however, the finely dissected, darkish, blue-green leaves can be mistaken for Parsley, and theroots for Radish. Symptoms of poisoning by this species include nausea, vomiting, diarrhoea and musculartremors (Cooper & Johnson 1998).
Fermanagh occurrence
There are just six records in total for A. cynapium in the Fermanagh Flora Database. There is onlyone recent record, from July 2000, which was observed by RHN in disturbed conditions on the roadside Nof Killymackan. Apart from this, Fool's Parsley had not been recorded in the VC for almost half acentury.
As listed above, the first record of A. cynapium in Fermanagh was made by Praeger, at the end ofthe 19th century, and Meikle and his co-workers added four more stations during the 1946-53 period oftheir recording activity. The details of these latter records are: Lisbellaw, 1946-53; quarry nearDonagh Crossroads, 1946-53; weed in garden of Melvin Hotel, Garrison, 1949; and in cultivated ground,Belcoo, 1952.
A. cynapium also appears to be a declining weed in NE Ireland where it was once a frequentspecies. FNEI 3 recorded very few sightings since 1920 in Cos Down (H38), Antrim (H39) andLondonderry (H40). Having said this, the absence of new data may be a matter of effort or direction inrecording; Beesley (2006) has listed seven additional weedy sites in Co Antrim (H39), all with post-1989dates.
Irish occurrence
Probably mainly for the mentioned soil, weather and arable farming associations, A. cynapium islargely a southern and eastern species in Ireland, the distribution and frequency thinning verynoticeably towards both the W and N, where damp, acidic pasture grasslands or peat bogs heavilypredominate in the landscape and arable agriculture has become of rare occurrence.
British and Irish status
Aside from the above subspecies distinction, in Britain A. cynapium is regarded as native as farnorth as Edinburgh and Glasgow. North of this, however, plus on the Isle of Man and throughout Ireland,Fool's Parsley is everywhere considered a definite or very probable introduction (Scannell & Synnott1987). The current post-glacial fossil record shows A. cynapium first appearing late in thecurrent interglacial (in Britain, the Flandrian, in Ireland, the Littletonian) at zone VI and,subsequent to this, at archaeological sites: a single record in the Bronze Age, followed by twoappearances in the Iron Age and three in the Roman period. Godwin (1975) comments, "In the BritishIsles too, it becomes less frequent in Ireland, and much less so in Scotland, a pattern seen in otherUmbelliferae with a late Flandrian expansion and response to anthropogenic influence." RHN and thecurrent author (RSF) believe this weed species most probably is an ancient agricultural introductionthroughout B & I.
European and world occurrence
A. cynapium probably originated in C Europe, but it is now widespread in temperate Eurasia and NAfrica. Man has certainly been instrumental in its spread as a weed of cultivation, both in N Europe andN America (Hultén & Fries 1986, Map 1403).
Threats
None.
Introduction, archaeophyte, occasional. Eurosiberian southern-temperate, but widely naturalised.
1864; Dickie, Dr G.; Newtownbutler.
April to October.
Growth form and preferred habitats
C. maculatum is a large, bushy, glabrous, hollow stemmed biennial that grows up to c 200 cm talland is leafy and heavily branched mainly above. It looks rather like a taller form of the much morecommon roadside umbellifer, Anthriscus sylvestris (Cow Parsley), but the bright green stem isalmost always well covered with dark purple blotches. The plant usually gives off an unpleasant, foetid,mousy odour when bruised, crushed or broken. The leaves are up to 30 cm long, bi-pinnate and the finelydivided leaflets have deeply serrated margins that give the plant a rather delicate appearance, similarto Garden Parsley (Petroselinum crispum).
The species is sometimes erroneously said to be tuberous, but it has a fleshy white or pale yellowtaproot, that is usually unbranched and rather like a small, narrow Parsnip (Pastinacasativa). The current author (RSF) has found that many Floras (and not just B & Iones), describe the species without mentioning its underground parts at all! There are no tuber(s)present (Grieve 1931; Melderis & Bangerter 1955; Sell & Murrell 2009) and their lack means thespecies is entirely dependent on seed for its reproduction.
C. maculatum is a pioneer coloniser of bare, recently disturbed, nutrient-rich, damp to moderatelydry soils. Very suitable open conditions of this nature are regularly provided by waterway dredgingoperations and in these situations C. maculatum can, for a time, form large colonies (Brewiset al. 1996). Hemlock is a classic nitrophile, ie a 'nitrogen-lover', or really a 'nitrogen andphosphate demander'. It shares this characteristic with the very much more common perennial Urticadioica (Common Nettle), with which it regularly occurs, especially on farmland. The establishedstrategy of C. maculatum is categorised as C/CR meaning it is intermediate between a Competitorand a Competitive Ruderal species (Grime et al. 1988, 2007), reflecting the tall, vigorous natureof the plant and its ability to form large colonies in suitable open, fertile habitats.
The most typical C. maculatum habitat is among other tall weeds growing in disturbed ground alonglowland roadside hedgebanks, preferably in damp, heavy, but fertile, moderately acid to neutral soil.However, it also frequents other forms of open, disturbed or waste ground, especially when these arenear water channels of any sort. Although it can occasionally be found in open areas within dampwoodland as, for example, locally at Ely Lodge woods and in Brookeborough Deerpark, Hemlock really doesnot tolerate very much shade. It really prefers sunny, warm growing conditions.
Flowering reproduction
C. maculatum flowers in June and July, the compound umbel inflorescences, terminal and axillaryare 2-5 cm across and have 10-20 rays bearing numerous white flowers 2 mm in diameter. The terminalumbel consists entirely of hermaphrodite (bisexual) flowers, while the lateral ones contain a mix ofmale and hermaphrodite flowers (Tutin 1980). The white petals are unusual in having their tips shortlyinflexed (ie bent back) (Clapham et al. 1987). As in other members of the umbellifer family,nectar and pollen are well exposed and pollination is carried out by unspecified insect visitors(Proctor & Yeo 1973; Fitter 1987), which very probably include beetles, flies and bees.
The fruit is ovoid, almost globose, about 3 mm long when ripe. It is laterally compressed and isdistinguished by having five prominent wavy or crinkled longitudinal ridges (Tutin 1980). The seed(mericarp) can persist in the soil for more than five years and germination is mainly, but notexclusively, in the spring (Roberts 1979).
Species status in Britain & Ireland
Until recently, in all published Britain and Ireland Floras and botanical literature, C. maculatumwas believed or assumed to be a native species throughout. In the 6th edition of his An IrishFlora, Webb (1977) suggested it might possibly be introduced in Ireland and in the 2002 NewAtlas the status was revised to a probable archaeophyte, ie a pre-1500 AD introduction. Thedecision to change the status of the species was based on evidence assembled and carefully analysed byPreston et al. (2004).
The evidence reviewed in deciding the status of the species in B & I included the fact that ancientbotanists of Classical Greece knew of the plant's narcotic poisonous nature and it was, therefore,cultivated in herb gardens for many centuries and used with care in minute doses in folk medicine as asedative and anti-spasmodic. Hemlock has been detected in archaeological deposits from the Late Bronzeage and the Roman period; documentary proof (albeit pre-dating the taxonomic treatment of plants thatdeveloped from the 16th century onwards) exists that Medieval herbalists grew the plant in England asearly as the tenth century, and the earliest botanical record dates from 1548 (Salisbury 1964; Prestonet al. 2004).
Toxicity
The deadly poisonous nature of Hemlock is due to the presence of a range of alkaloids including coniinethat act by attacking the central nervous system, inducing paralysis, convulsions and death fromrespiratory paralysis. All parts of the plant contain the alkaloids, the roots at all times of yearproving the least poisonous. Before flowering, the leaves contain the most alkaloids (up to 2%) but thegreatest concentrations are found in the flowers and the fruit. When dried, the plant loses most of itstoxicity (Grieve 1931; Cooper & Johnson 1998).
The unpleasant, nauseating, mousy smell of Hemlock appears to make it unattractive to grazing animals,warning them off. Poisoning mostly occurs in spring when grazing is in short supply and the young leavesare growing along hedgebanks where the hungry animals may be able to reach them. However, this is alsothe period of the year when the plant is at its least toxic. Sheep are more resistant to the toxins thancattle or horses. Poisoning has also been reported in goats, pigs, rabbits, chickens, turkeys andquails.
Fatal poisoning in humans has occurred through people mistaking the plant for Wild Carrot (Daucuscarota) or Garden Parsley (Petroselenum crispum) (Cooper & Johnson 1998). Childrenhave been poisoned by making the hollow stems into whistles and pea-shooters (Grigson 1955, 1987). Thebest advice with regard to umbellifers is, unless the identity is absolutely definite, AVOID EVENHANDLING THE PLANT, let alone playing with it, putting it to the mouth, or consuming it.
Use in herbal medicine
The use of C. maculatum in orthodox medicine ceased in the 19th century due to theuncertain actions of the preparations and the danger of lethal poisoning (Grieve 1931; Cooper &Johnson 1998). The purple blotched stem is sufficiently obvious that C. maculatumcan easily be identified by anyone interested in using it medicinally, but it needs to be handled withgreat care. Unfortunately, there are a number of other relatives with 'Hemlock' used aspart of their English common name (eg Oenathe crocata (Hemlock Water-dropwort)) and they are alsovery poisonous. Hemlock has a long history of its leaves being used as a poultice on external cancersand it was also widely used in this way to treat general sores and swellings. The hemlock poultice hasalso been used in Ireland for treating rheumatism, burns and perhaps wounds (Allen & Hatfield 2004).
Fermanagh occurrence
When recording in the 1947-53 period, Meikle and his co-workers considered this tall biennial a frequentspecies in Fermanagh. However, since very nearly half of the 40 records in the current Fermanagh FloraDatabase are from the pre-1953 period, RHN and the current author (RSF) believe C. maculatum hassuffered a local decline since then and it is now regarded as merely occasional. As the distribution mapshows, records are thinly scattered across 36 Fermanagh tetrads (6.8%) but only 22 of them containpost-1975 dates.
In Fermanagh quite of number of the sites are listed in the database as 'lakeshore'; however, in suchcirc*mstances, the plant is really associated with disturbed areas on the banks of ditches and streamsfeeding into these larger water bodies.
British and Irish occurrence
The New Atlas hectad map indicates that Hemlock is fairly common and widespread in much of lowlandBritain, becoming less frequent and more coastal towards the N & W and in Scotland. In Ireland, itis more scattered and less frequent than in S & E England and it becomes even more scattered towardsthe N & W and decidedly coastal.
European and world occurrence
C. maculatum originated in the Mediterranean region and W Asia, but has certainly been introducedfar beyond its natural range in N Europe and many other temperate places around the world. Despite, oron account of, its deadly poisonous properties and herbal medicinal use, Hemlock is still commonthroughout temperate Eurasia (except the extreme north), plus in N Africa and Macronesia. The specieshas been introduced into temperate regions of N & S America, S Africa, Ethiopia, S Australia and NewZealand (Clapham et al. 1987; Hultén & Fries 1986, Map 1406).
Names
The genus name 'Conium' is from the Greek 'komion' or 'kōneion', either meaning 'small cone' (Hyam &Pankhurst 1995), or just a label for both the plant and the poison (Stearn 1992). The Greek 'konos'means 'a cone', which does not appear to the current author (RSF) to have any connection with thisspecies. Another possibility is the Greek 'konas' meaning 'to whirl about', because the plant beingeaten causes vertigo and death (Grieve 1931). The Latin specific epithet 'maculatum' translates as'spotted' or 'speckled' (Gilbert-Carter 1964).
The English common name 'Hemlock', or 'Homlocke' as Gerard (1632) has it, is said to be derived from theAnglo-Saxon 'hem', meaning 'border' or 'shore' and 'leac', meaning 'leak' or 'plant' (Grieve 1931, p.392; Harvey 1981). Prior (1879) gives a different name origin, 'healm' meaning 'straw', and hence theterm 'haulm', and 'leac', 'plant', so called from the dry hollow stalks that remain after flowering.Britten & Holland (1886) show that numerous poisonous umbellifers are called 'Hemlock', or have itas a name element. Grigson (1974) takes the view that there is no clue to the meaning of the name, whichhe points out is found only in English.
Grigson (1955, 1987) lists a total of 30 alternative English common names for C. maculatum,several of them with 'devil', 'bad man' of 'gypsy' word elements, indicating the dangerous nature of theplant. Even Grigson's list is not exhaustive, as other writers mention additional, obscure local names(eg Grieve 1931 and Mabey 1996). Vickery (2019) lists a total of 38 English common names but,surprisingly, there does not appear to be much folk-lore associated with the species.
Threats
None.
Native, common. Eurosiberian southern-temperate.
1881; Stewart, S.A.; Co Fermanagh.
January to November.
Growth form, identification and preferred habitats
A. nodiflorum is a glabrous, polycarpic, patch-forming perennial with much-branched, slender,hollow, low-growing, trailing stems 30-90 cm long (not tall!). The basal roots of the plant are shallowand vegetative stems are mostly creeping and procumbent, while flowering stems are more ascending,reaching around 30-100 cm tall. The flowering stems also produce adventitious roots at the nodes nearthe ground. The simply pinnate leaves are sheathed at the base and divide into several pairs ofopposite, toothed, stalk-less leaflets.
The leaves of A. nodiflorum are very variable and can easily be confused with those of Berulaerecta (Lesser Water-Parsnip), the two species sometimes occurring together. The trueWater-cress, Rorippa nasturtium-aquaticum s.l. is also ecologically very similar to A.nodiflorum and all three species can occur together or in pairs. The similarity between A.nodiflorum and R. nasturtium-aquaticum, and the fact that they can occur intermingled,means Fool's Water-cress can readily be mistaken for genuine Water-cress. Although A. nodiflorumis not really poisonous, it is rather unpalatable and consequently it is called 'Fool's Water-cress'.The Plant Crib 1998 provides a useful table to distinguish Apium nodiflorum, Berulaerecta and another umbellifer, Sium latifolium (Greater Water-Parsnip) (Rich & Jermy1998, p. 220). Apium species do not have the node-like ring-mark towards the base of the petiolesof the lower leaves that B. erecta and S. latifolium both have, the ring-mark denoting apair of leaflets that have not developed. The ring-mark is not always present in B.erecta, however, as it may sometimes be replaced by a pair of small leaflets.
A. nodiflorum is a wetland and waterside species that can develop dense, sometimes pure clonalpatches and may dominate shallow, still or slow-flowing, moderately nutrient-rich (mesotrophic toeutrophic), often calcareous waters. It grows in ditches, streams, ponds and in marshes and fen swampsbeside seasonally reduced rivers and lakes, especially where there is sufficient disturbance to limitthe growth of taller herbs. Colonies can be either emergent on bare mud in more shallow situations, orlargely submerged in deeper, faster flowing waters, although the latter is seldom or rarely the case inFermanagh. Occasionally the plant spreads out from the waterside environment to invade or engulfadjacent, drier, marshy grassland vegetation (Preston & Croft 1997).
Although the taste of the plant is described as nauseous, stock animals will still browse on it andreduce its presence accordingly.
Shoots die down completely in the autumn and reappear in late spring (Grime et al. 1988, 2007).
Flowering reproduction
In July and August, plants produce more erect flowering stems up to 100 cm tall. The inflorescences arecompound umbels that look as if they are arising on short peduncles in the axils of leaves, but oncloser examination they are actually leaf-opposed. The flowers are minute, greenish-white, almostsessile (pedicels just 1-2 mm) and they are borne on 5-8 rays in short-stalked umbellules (partialumbels, subdivisions of the compound umbel) (Tutin 1980). Pollination is by insects and is sometimesdescribed as 'promiscuous' since the pollen and nectar are openly displayed and freely available to awide range of unspecialised flower visitors, including beetles, flies and bees (Fitter 1983; Knees1989). However, as is also usual in this family, the flowers are self-fertile (Knees 1989).
Fruiting takes place in September and October. The fruit is small (2-2.5 mm), rounded, the mericarpshaving five slender ridges and with a single vitta (ie resin canal) between each pair of ridges (Tutin1980). Germination occurs in spring and early summer and seedlings may become established in mud orsubmerged in water. It is believed that they rarely survive, except in very open areas, being veryvulnerable to competition from other established wetland plants (Thommen & Westlake 1981). Thus themost reliable means of reproduction and dispersal in A. nodiflorum is probably vegetative.
Vegetative reproduction
This is achieved in part by creeping growth and rooting of horizontal stems. Stem fragmentation anddispersal by flotation is also prevalent and appears a significantly more reliable means of increasecompared with the probably very rare establishment achieved by seedlings (Thommen & Westlake 1981;Grime et al. 1988, 2007).
Fermanagh occurrence
In Fermanagh, A. nodiflorum is quite common and widespread, being recorded in 120 tetrads, 22.7%of those in the VC. Having said this, it is chiefly associated with the shores of Upper and Lower LoughErne, both of which water bodies are fed by lime-rich rivers and streams and they are becomingincreasingly nutrient-enriched and eutrophic in recent years as a result of agrochemical and sewagerun-off. However, A. nodiflorum is known to be very tolerant of both eutrophication andfluctuating water levels. The incredibly dissected, sheltered, muddy shores of Upper Lough Erne inparticular, provide very many suitable sites for this species, but it is also well represented on theRivers Finn, Ballycassidy and Swanlinbar that flow into the lake and in their feeder streams andditches.
Irish occurrence
In NI, A. nodiflorum is common and widespread in Cos Fermanagh (H33), Armagh (H37), Down (H38) andAntrim (H39), but apparently less prevalent in the other two northern VCs (Cos Tyrone (H36) andLondonderry (H40)).
In the RoI, A. nodiflorum is generally widespread, but it becomes scarce or absent in the acidicinland, more upland areas of Co Donegal (H34 & H35), becoming decidedly coastal both there and inother equally wet, western counties.
British occurrence
In Britain, the species is widely distributed in lowland England and Wales, showing no decline over thelast 40 years. However, it becomes much more local and scarce northwards and especially so across theborder in Scotland, where it very quickly becomes rare and coastal (Preston et al. 2002). Thedistribution is particularly odd in the western Scottish isles, where A. nodiflorum and, indeed,A. inundatum (Lesser Marshwort) too, are absent from Mull and Skye, but not uncommon in the OuterHebrides (Pankhurst & Mullin 1991). As in Co Donegal (H34 & H35), much of this distribution maybe easily explained by the prevalence of strongly acidic, peaty soils and high ground.
European and world occurrence
A. nodiflorum is distributed throughout W, C & S Europe, although it is most frequent in thewest of the Continent and reaches its northern limit in Scotland. The species is also found in SW &C Asia and in N Africa. It is a naturalised and still actively spreading introduction in N & SAmerica (Preston & Croft 1997; Sell & Murrell 2009).
Threats
None.
Native, very rare but probably regularly over-looked.
1857; Moore, D.; Lower Lough Erne.
July.
As explained in the A. inundatum (Lesser Marshwort) species account (below), theparent species of this perennial, sterile hybrid overlap ecologically in Ireland in an almost uniquemanner. As a result, this is a widespread but nevertheless still rather thinly recorded hybrid acrossthe whole of Ireland, but as the 2002 New Atlas and 2015 Hybrid Flora hectad mapsindicate, it is extremely rarely found in Britain. The hybrid is also totally unknown elsewhere inEurope. It is unfortunate that in Ireland, the hybrid is probably often overlooked and the currentauthor (RSF) and RHN assume that we are among those guilty of under-recording it in Fermanagh.
A detailed account of this hybrid and its differentiation from the parent species is provided by O'Mahonyin Stace et al. (2015) and will not be repeated here. The hybrid is recorded from places wherethe parent species overlap ecologically, typically in shallow water in swamps and at the edges of lakes,rivers, steams, canals and ditches and on damp ground near water. Well developed hybrid plants have beenobserved forming tangled, floating, felted mats with slightly submerged stems bearing aerial leaves andflowers (Praeger 1951). The F1 hybrid is absolutely sterile and reproduces and spreadsentirely by vegetative growth and subdivision (T. O'Mahony, in: Stace et al. 2015).
Fermanagh occurrence
There are only four records of this hybrid in the Fermanagh Flora Database, all of which are confined tothe shores of Lower Lough Erne. This very large body of water has a very much more open, rockycalcareous shoreline than the dissected Upper Lough Erne, which is shallower and is littered with amultitude of islands, both small and large. Both these large Fermanagh lakes are somewhat polluted withagricultural chemicals and sewage, leading to rather strong and steadily increasing eutrophication. Thedetails of the remaining three local records are: Carrickreagh Bay, Lower Lough Erne, 1900 & 1934,R.Ll. Praeger; and Brockagh Point, Boa Island, Lower Lough Erne, 25 July 1976, Miss N. Dawson.
It is clear that only a small number of field botanists are able to distinguish this hybrid and RHN andthe current author (RSF) have no personal finds of it ourselves as yet. Despite the three records abovebeing confined to the Lower Lough, the most likely place to find the hybrid today would appear to be atthe southern end of Upper Lough Erne, where both parent species are fairly common.
Apium inundatum (L.) Reichb. f., Lesser Marshwort
Native, locally frequent. Suboceanic temperate, including in NW Africa.
1806; Scott, Prof R.; Co Fermanagh.
May to November.
Growth form
This small, glabrous, heterophyllous, perennial umbellifer really is amphibious; it can survivecompletely submerged as a true aquatic with typical flaccid, hollow, branched stems, 100 cm or longer,bearing finely divided submerged leaves, plus broader, pinnate or trifoliate floating leaves. Thecreeping basal part of the stem, which is usually submerged, sometimes roots at the nodes touching thesubstratum (Jonsell & Karlsson 2010). However, should the shallow water bathing the plantsignificantly lower or temporarily disappear in the summer months, the plants of A. inundatum canassume a small-scale or even tiny terrestrial or semi-terrestrial mode, modifying their form andproducing a crop of somewhat broader, singly-pinnate, flattened aerial leaves with up to seven sessilelobes c 5 mm long (Tutin 1980).
Identification from associated species and preferred habitats
In spring, the finely divided submerged leaves of Sium latifolium (Greater Water-parsnip),although differing in colour, are very similar in form to those of A. inundatum. In Fermanagh,both of these species are very frequent and they often coincide in sites around the sheltered,dissected, muddy lowland shores and nearby ditches of Upper Lough Erne in particular.
In Britain, A. inundatum is regarded as a species associated with moderately acid to weakly basic,nutrient-poor, moderately oligotrophic soft water conditions whereas, in comparison, Siumlatifolium always occupies decidedly enriched, moderately to strongly eutrophic situations (Hillet al. 1999; M. Southam, in: Preston et al. 2002). The two species are thus ecologicallywell separated. However, in Fermanagh, and indeed throughout much of its widespread Irish occurrence,A. inundatum tolerates much more nutrient-enriched wetland habitats than it does elsewhere in itsrather restricted western European range. This allows it to overlap ecologically not only with S.latifolium, but also with its near relative A. nodiflorum (Fool's-water-cress). BothApium species flower freely and the outcome of this unusual juxtaposition is the sterileApium hybrid, A. × moorei, which is widespread in Ireland, ofvery restricted occurrence in Britain and completely unknown in continental Europe (T.G. Tutin, in:Stace 1975, pp. 268-9; Preston & Croft 1997).
Part of the reason why these normally ecologically separate species are so very successful at co-existingin Upper Lough Erne in particular, may be the relatively high levels of calcium in the inflow waters ofthe lake. Perhaps this creates a base-rich but nutrient-poor environment, or an unbalanced supply ofnutrients as far as plant growth is concerned (Gibson et al. 1980). Without being a limnologicalexpert, the current author (RSF) suggests this potential explanation might be worth investigating.Another possibility, however, might lie in the restricted range of competitor species that exists inIreland in comparison with British waters, and even more so when compared with Continental Europe.
Fermanagh occurrence
A. inundatum has been recorded in 64 Fermanagh tetrads (12.1%), 59 of which have post-1975 dates,so there is no pressing evidence of a decline locally.
Flowering reproduction
In B & I, A. inundatum flowers and fruits freely in both shallow water and in bare, recentlyexposed terrestrial ground around lakes, ponds, rivers, streams and ditches. It is believed thatreproduction in this species is largely by seed (Preston & Croft 1997), rather than by vegetativefragmentation and re-establishment.
When flowering occurs, from June to September, umbels are reduced, very small and few flowered. Thecompound umbel is short-stalked or almost sessile, attached opposite the leaf supporting the leadinglateral shoot. There are few umbellules (1-4), each bearing 3-7 very small, white flowers, stamens withpurplish anthers, and 4-6 conspicuous, persistent bracteoles (Jonsell & Karlsson 2010). The fruits,2.5-3.0 mm long, are elliptic to oblong in outline and laterally compressed with five thick ridges oneach mericarp and a vitta in each groove (Tutin 1980; Stace 2019).
While there is some circ*mstantial evidence from disturbed sites which suggests seed may be longpersistent in the soil seed bank, the survey of the literature on this topic in NW Europe by Thompsonet al. (1997) uncovered no data.
British and Irish occurrence
The New Atlas map indicates A. inundatum is widespread and locally frequent in NI,and likewise in the RoI along the very long River Shannon, and in lakes in both Connemara and Co Cork.It is much more scattered and scarce elsewhere on the island.
The distribution remains widespread in Britain, showing a definite western bias and becoming more coastaland island-based further north and in Scotland. The New Atlas hectad map shows the species hasdeclined considerably in Britain since the 1962 BSBI Atlas. This is particularly obvious ineastern and southern England and is said to reflect drainage and eutrophication associated with changesin land use including building and other development, including agricultural intensification (M.Southam, in: Preston et al. 2002).
European and world occurrence
A. inundatum has a restricted distribution in W Europe stretching from the Iberian peninsulaeastwards to Sicily, northwards to SE Sweden and southwards to N Africa (Hultén & Fries 1986, Map1411). A similar pattern of decline mirroring the situation in Britain appears to be happening incontinental Europe, where the species is listed as 'vulnerable' in the Netherlands. It also appears tobe declining in Germany (Preston & Croft 1997).
Threats
None really, but potentially drainage and excessive dredging of ditches could affect regeneration.
Introduced, archaeophyte, a very rare garden escape.
1884; Barrington, R.M.; on the walls of Old Crom Castle.
May and July.
This very familiar glabrous, heterophyllous, tap-rooted, aromatic biennial or short-lived perennial hasbeen in cultivation as a potherb in B & I gardens since at least 995 AD (Harvey 1981). Records of itnaturalised as an established garden escape are widely scattered around these islands on cliffs, rocks,banks, walls, ruins and waste ground, especially in coastal regions (Clement & Foster 1994; M.Southam, in: Preston et al. 2002; Cat Alien Pl Ir). Many or most of the naturalisedparsley plants do not have the crisped leaves of the culinary herb cultivars, but presumably they havearisen from cultivated stock nevertheless, and have reverted to the flat-leaved wild form, the gene(s)for crisped leaves having segregated out (P.M. Smith, in: Simmonds 1976). The variation in leafmorphology appears to be dependent on a few genes only (Jonsell & Karlsson 2010).
Having remarked on the widespread B & I occurrence, the New Atlas hectad map indicates thatrecords are very thinly sprinkled across Ireland. There are 22 hectads of the most recent date class(1987-1999) on the map of Ireland and they are mainly coastal, the plant being very rarely recorded atinland sites.
Records prove that Garden Parsley has persisted at its only Fermanagh site, on the walls of Old CromCastle, since at least 1884. It is still plentiful on the castle walls today, having perhaps rathersurprisingly survived the re-pointing of the stonework in the 1990s.
In Britain, P. crispum is much more frequent than in Ireland, the New Atlas map displaying169 hectads with the 1987-1999 date class. However, English and Welsh records are greatly concentratedto the SE of a line on the map drawn between Cardiff and Hull, which represents the sector with the mostContinental climate on the island. Also noticeable is that towards the N & W of Britain, GardenParsley becomes decidedly coastal in its occurrence, a fact that suggests its growth and reproduction isinfluenced by milder temperatures in maritime regions.
Although the plant is long-persistent at its only Fermanagh site, commonly Garden Parsley is a merecasual in most of B & I. Even when long survival does arise, which appears to happen most often incoastal sites, the plant shows little tendency to spread in the wild, even though it can seed itself andit obviously 'jumps the garden wall' quite frequently, especially in the warmer areas of these islands.In cultivation, it is raised from seed, but it is always slow to germinate.
The centre of origin of P. crispum is obscure on account of its long period in cultivation, but itis probably native in the warmer parts of SE Europe, possibly in the Mediterranean basin from Sardiniato Greece (Clapham et al. 1962). An alternative suggestion is in W Asia, where it is widespreadtoday (Clement & Foster 1994). If either of these possible origins is correct, it helps to explainwhy the species is not fully adapted to life in the wild in B & I.
Threats
None.
Native, common and locally abundant. Eurasian boreo-temperate, but part of a circumpolar species complex.
1726-72; Anon (C. Threlkeld); Co Fermanagh.
April to November.
Growth form and preferred habitats
This robust, tall, glabrous, much branched perennial umbellifer has hollow stems 0.5-1.5 m high, oftenstreaked with purple. It grows from a somewhat misshapen globose, rootstock or rhizome that is developedat ground level and has short, narrow roots growing from its base. Leaves are 2- to 3-pinnate, the lobes3-9 cm linear-lanceolate, remotely and deeply serrate (Tutin 1980; Mulligan & Munro 1981; Jonsell& Karlsson 2010). The only other large aquatic, white-flowered umbellifer with which C.virosa might be confused is Sium latifolium (Greater Water-parsnip), which differs inhaving simply pinnate leaves and umbels with leafy bracts (NB both these species have small, linearbracteoles at the base of the ultimate subdivisions of the flower-head) (Tutin 1980).
The international expert on aquatic plants, Prof C.D.K. Cook (1990, 1998) regards C. virosa as atrue aquatic species on the basis of its life history, but Preston & Croft (1997) take a narrowerdefinition and disregard it in their British Isles treatment of aquatic plants.
Despite its large, robust appearance, C. virosa is very particular in its habitat requirement,being tied to wet ground sites with permanently available water, usually shallow and either still orslow-flowing. Typical habitats include lake margins, ponds, the banks of rivers, streams and ditches,marshes, wet-meadows, tall-herb fens and, occasionally, in slightly brackish water and along saltmarshes in parts of its wider range (M.F. Watson, in: Preston et al. 2002; Jonsell & Karlsson2010).
The species is extremely sensitive to drainage operations and the major and rapid decline of C.virosa noticed in other areas of B & I is undoubtedly the result of habitat loss due towidespread drainage designed to enable or expand intensive agriculture practices and other forms ofdevelopment. On the other hand, unlike many aquatic species, C. virosa is tolerant ofconsiderable tree or shrub shade in fen-carr, being capable of withstanding conditions down to almost50% of full sun.
While it prefers moderately fertile, nutrient-rich, base-rich, silty or muddy organic substrates andalkaline or near neutral waters, C. virosa has a wide pH tolerance, is indifferent to lime andcan also survive high levels of eutrophication where many submerged species have disappeared (M.E.Braithwaite, in: Stewart et al. 1994; Jonsell & Karlsson 2010). Eutrophication has developedand become very prevalent in all of the larger lakes in NI (Eutrophication in Northern Ireland'swaters: proposals for a strategy to control nutrient enrichment., Anon., undated, c. 2000,quoting 1997 data).
Fermanagh occurrence
There are post-1975 records of C. virosa from 132 tetrads in Fermanagh (25%), while by comparisonM.E. Braithwaite in Stewart et al. (1994, p. 115) map a total of just 107 tetrads with post-1970records for the whole of Britain!
C. virosa is typically found in still or slow-flowing water in the shallows around lakes, ponds,rivers and ditches. In Fermanagh, it is chiefly associated with the fen-fringed shores of the largerlakes. Although it is present throughout the whole of the Lough Erne system, Cowbane is especiallyfrequent and abundant around the sheltered, peaty, often fen-carr wooded bays of the drowned drumlinlandscape of Upper Lough Erne. In this generally shallow, hard-water lake, the intricate shoreline andvery large number of small to medium sized islands, which together cover approximately 40% of the totallake area, provides many suitable sites for this species (Gibson et al. 1980).
In deeper waters, C. virosa is said to grow on floating mats of vegetation, but while this may bethe case, personally the current author (RSF) and RHN have never observed it happening in Fermanagh(M.E. Braithwaite, in: Stewart et al. 1994).
Overwintering mechanism and vegetative reproduction
In the autumn, several globose rootstocks form from buds near the base of the current year's rootstock asit dies and rots. These freshly formed rootstock storage organs serve as the overwintering organ of thespecies and they also contain air chambers. The new rootstocks generally grow just above ground leveland during winter or spring flooding events they can readily detach and float downstream, spreading theplant vegetatively throughout the water catchment (Mulligan & Munro 1981; Jonsell & Karlsson2010).
Flowering reproduction
The mature plant produces a succession of large white-flowered inflorescences, 8-15 cm wide, in July andAugust. The numerous flowers are tiny, 2 mm diameter, the petals with an incurved point (Butcher 1961).Nectar and plentiful pollen attracts a wide variety of unspecialised insect visitors which transfer thepollen. The flowers are also self-compatible and if not crossed will self-pollinate. The schizocarpfruit consists of two large, single-seeded mericarps joined laterally and each surrounded by a spongyfruit coat with five blunt ridges containing large air spaces that keeps them buoyant, enabling theirwater dispersal (Mulligan & Munro 1981; Jonsell & Karlsson 2010). The seed is transient in thesoil seed bank, surviving for one year or less (Thompson et al. 1997).
Toxicity
All parts of the plant are poisonous, but the rootstock and lower stem are especially toxic. Consumptionof even a portion of the latter is sufficient to kill a cow or a horse. The poison is a yellow oilyjuice called 'cicutoxin' which rapidly attacks the central nervous system, causing painful convulsions,nausea and death. Not surprisingly, measures are very often taken by farmers to eradicate the plant fromaround waters where animals drink or might feed. The toxin turns reddish-brown after exposure to theair, but it survives drying very well. Thus Cowbane roots excavated by ditch cleaning operations andleft lying around on the soil surface, can still cause the death of animals long after they have beendug up. Cattle are particularly susceptible on account of their feeding habits, and hence the Englishcommon name 'Cowbane' is very apt, although a name like 'Farmer's Bane', would be even more appropriate.Cows are known to pull the highly toxic rootstocks and roots out of the soil, especially in the springwhen the ground is soft (Mulligan & Munro 1981).
The leaves and stems are poisonous to a lesser extent and they are sometimes eaten by hungry stock whengrass or other fodder is in short supply (Cooper & Johnson 1998). Fortunately, the strong, ratherbitter, unpleasant (but to some people celery-like), smell of all its parts, is usually sufficient todeter both browsing animals and people; human poisoning as a result of misidentification is rare (Knees1989). However, since 1900, a total of 83 cases of Cicuta poisoning in man have been reported inliterature for the United States of America (Starreveld & Hope 1975). There are four Cicutaspecies in N America, of which only one is less than deadly poisonous to humans and stock animals (C.bulbifera L.) (Mulligan & Munro 1981).
British and Irish occurrence
C. virosa was previously quite widespread in Britain, particularly in the eastern half of thecountry. The distribution has become very local and fragmented, being mainly confined to meres inShropshire (VC 40) and Cheshire (VC 58) and the Norfolk Broads. It occurs more rarely from S Scotlandnorthwards to Easterness (VC 96) and the Outer Hebrides (VC 110) (Sinker et al. 1985; Prestonet al. 2002). Cowbane is considered 'Nationally Scarce' in Britain, but the distribution isstable.
In Ireland, C. virosa is much more abundant, yet within NI it is still quite locally confined tothe southern portion (ie the Lough Neagh and Lough Erne basins and connecting waterbodies). Altogetherthis very poisonous perennial is represented in a total of 16 Irish VCs (Cen Cat Fl Ir 2). Thereis a solitary outlying station in Co Clare (H9), remote from the main Irish distribution in the northernmidlands.
European and world occurrence
C. virosa forms part of a circumpolar species complex that includes closely related species orsubspecies including C. maculata L., C. douglasii (DC.) Coult. & Rose and C.bulbifera L. in N America (Mulligan & Munro 1981). The complex is mapped by Hultén &Fries (1986), Map 1412. C. virosa extends through mid-latitude, temperate, continental Europe andAsia, from France to Japan, mainly north of 45°N. It becomes more scattered in the western part of itsEuropean range and is virtually absent from the whole Mediterranean region (Tutin 1980; M.E.Braithwaite, in: Stewart et al. 1994; Jonsell & Karlsson 2010).
Names
The genus name 'Cicuta' is the Latin name of Hemlock (Conium maculatum) which, in the opinion ofthe current author (RSF), is more than a little bit confusing and not at all helpful (Gilbert-Carter1964; Gledhill 1985). The connection is very probably the exceedingly poisonous and dangerous nature ofboth plants. The Latin specific epithet 'virosa' means 'slimy', 'rank', 'foetid' and 'poisonous'(Gilbert-Carter 1964).
Five English common names are listed for the plant, the most frequent being 'Cowbane' (from the 18thcentury) and 'Water Hemlock', but other interesting ones given by Britten & Holland (1886) are'Deathin', 'Brook-tongue' and 'Scoots'. 'Deathin', a name of Scottish origin, is described as being,"peculiarly obnoxious to cows". 'Scoots' is a name associated with this and any otherumbellifers frequenting wet places (Britten & Holland 1886, p. 419).
Threats
Drainage is the main threat to this erect, emergent (or rarely floating) aquatic species, since generallyit is strongly tied to permanently shallow water. Provided the root of the plant is kept permanentlydamp, the soil surface may dry out temporarily.
Cultivated introduction, archaeophyte, rare, casual and locally extinct. Considered native in N, C &E Europe and much of boreo-temperate Asia, but widely naturalised and now disjunct circumpolar.
1900; Praeger, R.Ll.; Co Fermanagh.
Growth form and preferred habitats
Carum carvi is a glabrous, erect, much-branched, aromatic, biennial, 25-60 cm tall, with 2- to3-pinnate leaves and a fusiform taproot. The umbels are small, 2-4 cm in diameter, rays 5-16, bracts andbracteoles absent or few. The flowers, produced in June and July, are white or pink and the fruit is 3-4mm in diameter and ellipsoidal. The twin mericarps smell strongly of aniseed when crushed and have longbeen used for flavouring in cookery, including bread, cakes and liqueurs (Grieve 1931; Clapham etal. 1987).
In cultivation, C. carvi prefers warm, sunny locations and well-drained soils rich in organicmatter. In warmer regions, it is treated as a winter annual, while in temperate situations it is plantedas a summer annual or biennial.
The plant was introduced from Europe to garden cultivation in B & I some time prior to 1375 AD(Harvey 1981). It was in widespread kitchen use here for flavouring purposes for centuries, but thispractice has declined to rarity in recent times (M. Southam, in: Preston et al. 2002). Caraway isstill available and in regular culinary and other use, but it is imported from commercial growers mainlyin the Netherlands, Poland and Germany.
Fermanagh occurrence
Only recorded once by Praeger as listed above. In his monumental book, Irish Topographical Botany,Praeger listed the VCs in which C. carvi did not occur and Fermanagh was not on his list.There is no other reference to this monocarpic species occurring in Fermanagh, either present or absent.
Irish occurrence
Throughout Ireland, Caraway has long been considered a rare casual introduction of disturbed ground,generally occurring in waste ground near houses and gardens where seed was accidentally introduced orescaped from garden cultivation (Cybele Hibernica 1866). During the 19th century and early 20thcentury, Caraway was a widespread casual in Ireland, recorded in 30 of the 40 Irish VCs. However, it wasalways occasional, never common, except possibly in parts of the NE of the island. In 1901, Praeger(Irish Topographical Botany) commented that it, "seems naturalised in [Cos] Armagh andAntrim".
By the 1950s, however, records of C. carvi were becoming very scarce or rare, even in those twoVCs (FNEI 3). Reynolds (Cat Alien Pl Ir) listed only three post-1986 Irish records for thespecies, all of them from coastal counties in SE Ireland. Interestingly, the New Atlas map plotsfour widely spaced post-1986 hectads for Caraway in Ireland, only one of which might overlap withReynold's listing.
The most likely origin of plants beyond cultivated plots would have been Caraway seed used in the kitchenas a culinary flavouring. Caraway bread, for example, was once very much more popular than it is today.The current author (RSF) remembers with nostalgic affection Brewster's 'seedy loaf' produced and sold bythe long-gone Londonderry bakery of that name during his 1950s boyhood.
British status and occurrence
In her Modern Herbal, Grieve (1931) described C. carvi as a naturalisedspecies in Britain and considered it an escape from cultivation. The third edition of the standardFlora of the British Isles of the time (Clapham, Tutin & Moore 1987) considered C.carvi, "perhaps native in some south-eastern counties and naturalised in wasteplaces". The species was, "accepted, with reservations, as native" by Clement &Foster (1994) in their survey of Alien Plants of the British Isles and not dealt with further bythem. Stace (1997) recognised Caraway as an introduction; the New Atlas first described it as anarchaeophyte throughout B & I (ie a pre-1,500 AD introduction), a status further argued andconfirmed by Preston et al. (2004).
C. carvi is widely distributed but rather uncommon throughout Britain, apparently being bestnaturalised in Shetland (VC 112) (New Atlas). The distribution displayed in the New Atlashectad map suggests it is most frequent in the English southern Midlands, in areas of greatestpopulation density. However, the species is much less commonly cultivated now than previously and inmany occurrences the plant is a mere casual ruderal on waysides, waste places and rubbish tips, ratherthan truly naturalised (M. Southam, in: Preston et al. 2002).
European and world occurrence
C. carvi occurs throughout most of Europe, temperate Asia and NW Africa, but it is rare in theMediterranean region (Tutin 1980). It is usually considered native in C, N & E Europe and largeparts of Asia (including Turkey, Iran, N India and Siberia), but it has long been cultivated as acondiment both inside and outside the supposed native territory. Rather surprisingly, the main areas ofcultivation are or were restricted to SE England (Essex, Kent and Suffolk), Holland, Germany, Finland,Norway, Russia and Morocco (Grieve 1931). In 2011, it was reckoned that Finland supplied around 28% ofthe world's caraway seed production, this high output reflecting the country's dry continental climateand its high latitude providing long summer days that favour the species' growth and reproduction.
Caraway also occurs as a rather common adventive introduction in the flora of many scattered parts of theworld, including N America, Brazil, C & S Africa, Japan and New Zealand. As a result of itswidespread introduction, numerous uses and subsequent naturalisation, C. carvi is nowdiscontinuously circumpolar (Hultén & Fries 1986, Map 1414).
Uses
Caraway is another member of the aromatic umbelliferous plants characterised by carminative and stimulantproperties, like Anise, Cumin, Dill and Fennel. In ancient times, caraway oil was recommended byDioscorides to be taken by pale-faced girls. At one time in the past, it was used as a carminativecordial recommended for use in dyspepsia and for symptoms around hysteria. It has some tonic propertiesand was also used as a pleasant stomachic. It is nowadays grown, however, chiefly for the use of itsfruits as a spicy flavouring agent in association with purgatives and for flatulence. Nowadays, it ismuch more used for flavouring in cookery, confectionary and liqueurs, rather than for any medicinalproperties. (Grieve 1931).
Threats
None.
Native, common, widespread and locally abundant. Eurosiberian boreo-temperate.
1881-2; Barrington, R.M.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
A rather large semi-rosette, short-lived, tap-rooted, glabrous, hapaxanth (ie a plant that is eitherannual, biennial, or requiring several or even many years' growth to reach flowering and fruitingstate). [See below for discussion/explanation of this topic.] A. sylvestris produces hollow,purplish, striate stems up to 200 cm tall. The glossy, dark green bi-pinnate or tri-pinnate lower leaveswith their inflated stalk and the distinctive regularly serrate margins of the large leaf-lobes, setwith sharp-looking cartilaginous teeth, make A. sylvestris very easily recognisable even in thevegetative state (Tutin 1980). The only other Irish umbellifer it might just possibly be confused withis Ligusticum scoticum (Scots Lovage), which is a maritime species and, therefore, does not occurin Fermanagh (Tutin 1980; Jonsell & Karlsson 2010).
Although it is a very common and widespread plant throughout B & I, the biology of A.sylvestris does not appear to have been the subject of much detailed study. The speciespossesses a stout taproot 8-20 mm thick, yet there is uncertainty about just how perennial andlong-lived individuals may be (Jonsell & Karlsson 2010). For instance, Grime et al. (1988,2007) reported that observations in winter suggest that around 50% of plants are monocarpic, dying afterflowering, the other 50% being polycarpic perennials. After a natural history study, comparing twopopulations on a coastal headland and in a marsh in Wales, Knight (1997) concluded that the plants inhis study were entirely monocarpic. Mowing or grazing prevents individuals reaching the threshold sizenecessary to initiate flowering, a feature found both in many short-lived polycarpic perennials andmonocarpic biennials, which frequently prolongs their vegetative life almost indefinitely (Forbes 1989).The latter plants (either annuals or biennials), with delayed flowering and fruiting, are now describedas 'hapaxanthic' (Jonsell & Karlsson 2010) and this appears to be the situation in A.sylvestris.
A. sylvestris prefers well-lit growing conditions, although it can tolerate partial shade inopenings or margins of woodland, in fen-carr, or on N-facing cliffs and slopes (Grime et al.1988, 2007; Hill et al. 1999).
The chief ecological demands Wild Angelica makes are for constantly moist but not submerged soil thatprovides a moderate supply of nutrients and that in reaction varies between moderately acidic tocalcareous. A. sylvestris is quite frequently recorded in the Fermanagh Flora Database in habitatlists along with species of strongly acidic Sphagnum bog, but it never grows on waterlogged mosspeat, being confined to the sloping fen margins of such sites.
The characteristically wide range of habitats in B & I which supply the modest requirements of thisspecies includes sea cliffs, although these are not represented in landlocked Fermanagh. The establishedstrategy of the species is described as C/CR, ie intermediate between Competitor and Competitive Ruderal(Grime et al. 1988, 2007).
Wild Angelica is not a gregarious, patch-forming species and it never (well, hardly ever!) forms a largeor even very conspicuous component of the vegetation cover in the plant communities in which it grows.The typical 'Domin cover value' of the species in many vegetation communities is less than four percent(ie, Domin values 1-3, whether this is made up of few, several or many individuals) (Rodwell etal. 1991 (b) & 1992).This is true even whenever there is protection from grazing or otherforms of disturbance, or when it is a 'constant species' in the vegetation community, eg in mesotrophictall-herb grassland or on montane cliff ledges (NVC MG2 and U17) (Rodwell et al.1992).
Angelica sylvestris certainly does not make the same visual impact that other equally robust,common umbellifers such as Anthriscus sylvestris (Cow Parsley) and Heracleum sphondylium(Hogweed). This is probably because those species are so abundant and extremely successful atmaintaining large populations, or very frequent individuals, in conspicuous situations along roadsideseverywhere throughout these islands. In the roadside verge habitat in particular, while it is oftenpresent, Wild Angelica is but a minor player compared with these two big hitters!
Fermanagh occurrence
A. sylvestris is the eighth most frequently recorded vascular plant species in Fermanagh, a sureindication, if one did not already know it, of the enormous amount of damp (or regularly wet butdrained) rough grassland found in all but the most extreme environments in the county. It is present in491 tetrads, almost 93% of those in the VC, making it the ninth most widespread species we have in theFermanagh Flora Database – just after Urtica dioica (Common Nettle).
Typical habitats of this species in Fermanagh include marshy or damp rough grassland, open areas ofwoods, scrub, meadows and ditches and in tall herb vegetation beside lakes, rivers and streams. It alsooccurs to a lesser extent on N-facing cliffs, in quarries and on damp roadside verges.
Flowering reproduction
Large enough plants with sufficient photosynthetic reserves flower from July to September. Speciesregeneration is entirely by seed, there being no means of horizontal vegetative spread or reproduction.The inflorescence is a large, domed, compound umbel 4-7 cm high and 8.5-17 cm wide with 15-40 sub-equalrays. The umbel rays and peduncle are densely puberulent or papillose (Tutin 1980; Jonsell &Karlsson 2010). The greenish-white or pinkish-white flowers are very numerous, each inflorescencebearing more than 1,000. They are hermaphrodite, protandrous, offer copious, unconcealed nectar andpollen, and, typical of the family, they attract a range of unspecialised insect pollinators includingbeetles, flies and bees (Grime et al. 1988, 2007). The schizocarp fruit, 4-5 mm long, is oblongto almost rectangular in outline, dorsally flattened and the two mericarps have broadly winged lateralridges (Tutin 1980). The wings are wider than the mericarps themselves, undulate and not closelyappressed to one another (Sell & Murrell 2009).
Seed dispersal
Seedlings are seldom observed, but that is quite normal in very many plant species. Since the mericarpsare very distinctly winged, dispersal in A. sylvestris is probably to some degree effected bywind. Ridley (1930) reported that the 'seeds' (ie the mericarps) float, so presumably they may also bedispersed in moving water.
Seed survival
In respect of the species longevity in the soil seed bank, again there is no consensus available: of 22estimates in the survey made of data from across NW Europe, 15 references suggested A. sylvestrisseed is transient; three believed it short-term persistent (surviving 1-5 years); one study consideredit persisted long-term (more than five years); and three estimates were indeterminate (Thompson etal. 1997). It is all too obvious that a little more systematic observation could clarify many ofthe basic life-history details which are at present missing for this significant indicator species.
British and Irish occurrence
The New Atlas hectad map shows A. sylvestris is very common and widespreadthroughout the whole of B & I.
European and world occurrence
A. sylvestris belongs to the Eurosiberian boreo-temperate phytogeographic element and is commonand widespread in most of Europe and parts of temperate W & C Asia, although having said that, itbecomes scarce or absent in drier, warmer parts of S Europe. It has also been occasionally or rarelyintroduced to easternmost N America (Hultén & Fries 1986, Map 1421; Sell & Murrell 2009).
Threats
None.
Introduction, archaeophyte, a rare garden escape, locally extinct.
1950; MCM & D; roadside bank between Tempo and Brougher Mountain.
Origin and reproduction
Introduced by medieval herbalists from the mountains of C & S Europe, this rather large, perennial(up to 100 cm) grows from a rhizome or tuberous rootstock and has distinctive ternately divided leavesand umbels with numerous papillose rays (Tutin 1980). Previously it was valued and cultivated for manycenturies, both as an edible potherb and for herbal medicinal and veterinary use. Nowadays, it hasceased to be employed in herbalism and is rarely cultivated in Britain and even less so in Ireland(Stace 2019). Masterwort flowers regularly, and while to some extent it must have escaped fromcultivation by means of seed dispersal, it could also be spread by transported rhizome or tuberfragments or through discards. It reproduces locally by vigorous rhizome or tuber growth and can producepersistent, clonal patches several metres across (Jonsell & Karlsson 2010).
Fermanagh occurrence
There is only one record for Masterwort in the Fermanagh Flora Database, when Meikle and co-workers foundit in 1950 in some quantity along the hedgerow between Tempo and Brougher Mountain, along a stretch ofabout 1300 m (Revised Typescript Flora). It has not been recorded again since then in Fermanagh.
The properties of P. ostruthium appear to have been especially highly regarded by Scottishsettlers, themselves 'planted' in Ulster in the early 17th century. It was very probably they whointroduced the plant to Fermanagh along with a number of other similarly valued pot and medicinal herbs.In addition to Masterwort, Praeger (1915) listed Sweet Cicely (Myrrhis odorata), Tansy(Tanacetum vulgare) and Elecampane (Inula helenium) as being three other NI alienspossibly first introduced by these Scottish settlers. However, the species may have been in cultivationvery much earlier than this, since seeds believed to be P. ostruthium have been excavated at anarchaeological dig at a ring-fort in Co Antrim (H39) dated much earlier, at around 850-950 AD (Allen& Hatfield 2004).
P. ostruthium was always local and uncommon in NE Ireland during the 19th and early 20thcenturies, but it declined to definite rarity in the last 80 or 90 years (FNEI 3). Masterwort isnot as thoroughly naturalised as another of these potherbs, Myrrhis odorata (see the speciesaccount on this website), and the only really recent records of P. ostruthium anywhere nearFermanagh are from along the Glenelly River and along some other riverbanks and roadsides in Co Tyrone(H36) where it is regarded as a persistent relict of cultivation (Preston & Stone 1999; NewAtlas; McNeill 2010).
British and Irish occurrence
P. ostruthium can escape from garden cultivation (or perhaps parts of its large creeping rhizomeare eventually discarded whenever the plant becomes too large and invasive). As the New Atlas mapshows, it has become locally naturalised in mesic to rather dry soils in sun or half-shade on roadsideverges and banks and hedgerows near houses, and along banks beside streams and rivers and in dampmeadows in hilly country at relatively low altitudes in northern England, Scotland and N Ireland (Knees1989; M.F. Watson, in: Preston et al. 2002). It usually becomes established in moist groundaround houses and farm buildings near where it previously has been cultivated. Many of the populationsappear to be very persistent and the species has not noticeably declined since the 1962 BSBIAtlas (M.F. Watson, in: Preston et al. 2002).
European occurrence
Native only in the C & S European mountains from N Portugal eastwards to Yugoslavia but introducednorthwards to southern parts of Scandinavia (Hultén & Fries 1986, Map 1427). The closely relatedP. caucasium (M.B.) Koch occurs in the Caucasus mountains. P. ostruthium has beenintroduced into Newfoundland.
Names and uses
The genus name 'Peucedanum' is a name of an unknown plant in the work of Theophrastus, borrowed andreused (Gilbert-Carter 1964). A quite different suggestion is that the name is from the Greek'pĕukĕdanŏn' meaning or referring to 'parsnip' (Stearn 1992). The Latin specific epithet 'ostruthium'means 'purplish', probably referring to purple stem markings or flower colour.
The English common name 'Masterwort' is a translation of a previous Latin genus name for the plant,'Imperatoria', a name that has recently been revived and reapplied to this plant and its relatives(Prior 1879; Stace 2019). Another explanation or suggestion by Grigson (1974) is that 'Masterwort' is atranslation made by Turner (1548) of the German 'Meisterwurz', meaning 'Master root', from the medievalLatin name 'Magistrantia'.
This suggests the plant was important in herbal medicine, ie a master wort of medicine. It was used as analexipharmic, or as an antidote to poison or for warding off infection. It was also regarded assudorific (ie a drug for inducing sweating) and, "a great Attenuater and Opener" (ie a stronglaxative) (Grigson 1955, 1987).
Alternative English common names include 'Fellon-grass', Fellon-wood' and 'Fellonwort', where a 'felon'or 'fellon' is a sore place swollen with pus of bacterial origin, in humans, often affecting a fingertip, but also found on the skin of sheep and cattle and hence the veterinary connection with the plant.Examples of such Masterwort use included, "the rootes and leaves stamped, doth dissolve and cureall pestilential carbuncles and blotches, and such other apostemations and swellings" (Gerard1633). Grieve (1931) lists the herbal use of the plant in treating asthma, dyspepsia and menstrualcomplaints. It was also considered effective in dropsy, cramp, falling sickness, kidney and uterinetroubles and gout (Culpeper 1653). As Grigson (1955, 1987) points out, "The demand was great, andMasterwort was regularly supplied by 18th century market gardeners."
Threats
None.
Introduction, archaeophyte, an infrequent casual escape from cultivation, probably locally extinct.Eurosiberian temperate, but very widely naturalised.
1900; West, W.; roadside or waste ground at Drumskew Td, W of Enniskillen.
Growth form and preferred habitats
The fleshy, white taproot of this large, biennial, monoecious, yellow-flowered species has been incultivation as a culinary vegetable grown from seed throughout the Mediterranean basin and temperateEurope since the Classical Greek period; even earlier than this it was used for animal fodder. Goodedible selected forms were first developed around the Middle Ages (Zohary & Hopf 2000), and as thefood value of parsnip exceeds any other vegetable except potato, and it is very easy to produce, itshould be more extensively grown than it is.
Escapes of the plant from cultivation in fields or gardens tend to revert to the 'wild condition', withtough, dry, much thinner roots than the usual cultivated rootstock (Smith 1976). The erect stem can beup to 180 cm tall and it may be either hollow or solid, its surface furrowed, terete or angled. It mayalso vary greatly in hairiness, either having sparse, short hairs, or numerous long, flexuous ones.Variation in the species is considered sufficient for three subspecies to have been recognised recentlyin Britain (see below).
Wild Parsnip typically occurs in unmanaged grassland, roadsides and other waste or rough ground nearhabitation, especially on chalk and limestone soils, or near the coast (Sell & Murrell 2009).
Fermanagh occurrence
There is just the single record detailed above in the Fermanagh Flora Database as a roadside gardenescape near Enniskillen dated over a century ago.
Variation
Since 2009, three subspecies are distinguished in Britain, if not in Ireland (Sell & Murrell 2009;Stace 2010; Parnell et al. 2012). The form escaped from cultivation is subsp. sativa, withits swollen rootstock that is not found in the other two subspecies. Subsp. sativa also hassparse, short, straight hairs on its stems and leaf upper surfaces. The form most like this is subsp.sylvestris (Mill.) Rouy & Camus which has long, soft, flexuous hairs on its stems and upperleaf surfaces grey-hairy. The third form, subsp. urens (Req. ex Godr.) Čelak., has stems terete,with short straight hairs or nearly glabrous (Sell & Murrell 2009).
Irish occurrence
In Ireland, Wild Parsnip is an infrequent casual weed of rough grass on roadsides, waste ground and inold quarries. Unfortunately, all of these are places where people frequently 'fly tip' garden rubbish.P. sativa has been recorded at least once over the years from 33 of the 40 Irish VCs (CatAlien Pl Ir). The New Atlas map of it for Ireland plots just 13 hectads with post-1986records. All but two of the sites plotted are coastal, undoubtedly reflecting this species preferencefor sandy, neutral and/or calcareous ground.
Fossil record
Evidence from pollen and fruit fossils in Britain is sparse, but nevertheless it shows P. sativahas been present right back into the early record of the Cromer Forest Bed series and in allinterglacials since then, plus in the last glacial period, the Devensian (or Weichselian). As Godwin(1975) commented, "It thus appears that P. sativa is a long-persistent native, at least insouthern England where most of the fossil records come."
British occurrence
Wild Parsnip is common on calcareous soils in lowland Britain, particularly S of a line between Newcastleand Preston, and it is even more frequent if we draw the line from the mouth of the River Humber to thatof the Severn. In addition, it does occur north of these two lines of demarcation, chiefly in coastal ordisturbed urban sites; in these northern areas, it is recognised as being an alien. Based on its fossilrecord, south and east of the Humber-Severn line P. sativa is traditionally believed to be anative species (Godwin 1975; New Atlas).
There are three subspecies in Britain: subsp. sativa is usually recognised to be an escape fromcultivation, whereas subsp. sylvestris is the form most usually met in the SE of England and in SWales and is considered native in those areas. The third subspecies, subsp. urens, is confined tothe Suffolk coast, where it just might be native, although its main distribution is in S, C & EEurope (Sell & Murrell 2009).
It is also known that in recent years, P. sativa has sometimes been accidentally introduced as aseed contaminant in 'wild flower seed mixtures' when roadside verges and banks are resown by LocalDistrict Council Roads Departments.
Seed survival is rather impressive: in a buried seed experiment, germination occurred after 17 yearsincorporation at a depth of 120 cm (Toole & Brown 1946).
European and world occurrence
Wild Parsnip is widespread throughout temperate Europe except the extreme north, plus in western Asia.However, as is the case in England and Wales, because the plant is widely naturalised, its native rangeis difficult or impossible to discern. The cultivated form, subsp. sativa, has been introduced toN & S America, S Australia and New Zealand, where it regularly escapes from gardens into roughground habitats (Hultén & Fries 1986, Map 1428).
Uses
Apart from being cooked and used at table as a nutritious vegetable, Parsnip has also been used as aquality fodder for fattening pigs; mixed with bran it has been fed to horses, and it has also been givento cattle, including milking herds (Grieve 1931). The roots (ie rootstocks) contain starch and sugar andhave been brewed with malt to make beer, wine and spirits.
In the past, herbal medicine has used P. sativa to treat cancer, consumption, asthma and similardiseases. It has also been claimed to have cleansing and opening qualities, provoking urine, easing painand stitches in the sides, expelling wind from the stomach and bowels and also useful for colic(Culpeper 1653; Grieve 1931).
Threats
None.
Native, common throughout. Eurasian boreo-temperate, but widely naturalised, including in eastern NAmerica and New Zealand.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Despite many modern Floras categorising H. sphondylium as a biennial (Tutin 1980; New Flora ofthe BI 1991), it is more accurately described as a (relatively) short-lived, semi-rosetteperennial. Frequently H. sphondylium is polycarpic, only becoming monocarpic if, or when, itexhausts itself after excessive seed production (Kilburn 1983; Sheppard 1991). Hogweed is non-aromaticand extremely variable. In Europe, it has been divided into nine subspecies and numerous varieties orforma, although considerable intergradation exists between them (R.K. Brummitt, in: Tutin et al.1968; Sell & Murrell 2009; Jonsell & Karlsson 2010). H. sphondylium stems vary greatly inheight, occasionally reaching up to 200(-300) cm. They are hollow, ridged and are often covered withbristly hairs (ie hispid). Leaves, which number 2-3 at the base and 3-5 on the stem, are simply pinnate,the leaflets lobed, with toothed or crenate margins. Leaves are usually densely hairy beneath (Tutin1980).
H. sphondylium is frequent and conspicuous in a very wide range of habitats but it is mostfrequent in more or less disturbed ground on roadsides, waste ground, scrub and woodland (except underthe densest canopies) and in poorly grazed or infrequently mown grassland and recently disturbed baresoil. Although absent from swampy or extremely acid soils (pH 4.3 or less), it does occur frequently inrough grassland on riverbanks and in water-meadows, provided the roots of the plant do not becomewaterlogged for periods long enough to induce rot. H. sphondylium tolerates a high range ofhumidity, wind exposure and salinity, but it does not tolerate more than transient waterlogging orprolonged drought, both of which destroy seedlings and young plants. Near the coast it can toleratemoderate salinity from sea spray, but not salt-water inundation. Light requirements are also low and itcan survive woodland shade down to c 5% of daylight (Sheppard 1991).
Although in fertile growing conditions it rapidly establishes a stout, branching, perennating taprootsystem that can penetrate to depths well below 100 cm, and above ground the plant is so vigorous it canreach heights of 2 or even 3 m, Hogweed does not reproduce vegetatively. Only the upper 'collar' portionof the taproot is capable of regeneration after ploughing or other uprooting disturbance (Sheppard1991). The foliage and stem die right back in autumn. The lack of vegetative reproduction means thatalthough H. sphondylium is common in disturbed, man-made or semi-natural habitats, it is notclump-forming, typically occurring as individual established flowering plants and only abundant whenpresent as seedlings or young plants (Grime et al. 1988, 2007).
H. sphondylium relies entirely on seed for its reproduction and dispersal and the species isconsidered a competitive-ruderal without a persistent seed bank (Grime et al. 1988, 2007). Thecompetitive status of the species has yet to be experimentally defined and, as it displays highmorphological variability and plasticity throughout its range, this would be a major undertaking.Observation suggests that the competitive ability of Hogweed is derived from its large taproot energystore, which gives it the ability to expand its leaves early in the growing season before associatedgrasses and other neighbouring competitors can do so.
Germination of the large seeds (flat mericarps) peaks in March, early in the growing season whenvegetation in most grassland is still short. Hogweed thus seems capable of establishing new plants evenwithin ± closed perennial communities (Grime et al. 1988, 2007; Sheppard 1991).
Occasional cutting or grazing favours Hogweed by reducing the advantage of clonal competitors such asRanunculus acris (Meadow Buttercup) and large, tufted grasses, including Dactylisglomerata (co*ck's-foot). However, without some form of major uprooting disturbance, competitivedisplacement of H. sphondylium by other perennials appears extremely unlikely (Sheppard 1987).
Individual Hogweed plants can persist for many years with estimates of average life span ranging from 12to 16 years depending on the habitat.
Fermanagh occurrence
Hogweed is very common in Fermanagh and has been recorded in 368 tetrads, 69.7% of those in the VC.Hogweed is really only entirely absent from areas of upland peat and fully aquatic habitats, where it isconfined to wetland margins, but it is probably also under-recorded to some extent in well-managedfarmland in eastern lowland parts of the county.
Flowering reproduction
The flowering period is from June to September. In individual plants, flowering can occur from about thethird year of growth onwards, but the average pre-reproductive stage is 6-7 years long. Most individualsdo not have the stored photosynthetic resources to flower in successive years, but they can survive toflower repeatedly about eight times, although it appears that seeding two or three times is more normalfor the species (Sheppard 1991). The flowering stem is determinate, with one large terminal umbel up to25 cm across, and 1-9 lateral umbels below it. Smaller tertiary umbels can branch from the peduncles oflateral umbels. Each inflorescence is a compound umbel, flat or slightly convex, composed of 10-20somewhat unequal rays, measuring 4-25 cm in diameter. Very large plants can bear up to 30 umbels ofinsect- or self-pollinated flowers, capable of producing 10-20 thousand seeds in a season. Theschizocarp fruits (each of two mericarps) are relatively large, measuring 6-10 mm long and equipped withlateral wings. Fortunately for the landowner and gardener, the vast majority of plants produce very manyfewer seeds than this estimate, the mean total being c 850 per flowering plant.
The single-seeded fruits (ie twin mericarps, each one an achene) are passively dispersed by wind in lateSeptember to early October. Being relatively large and flat, while most fruits scatter near the parentplant, a small proportion are carried up to 50 m downwind. The seed bank is transient and, afterchilling, virtually all seeds germinate the following spring (Sheppard 1991).
Toxicity and herbivory
Despite containing a wide range of aromatic oils, resins, terpenes, saponins and coumarins as toxicprotection against herbivores and microbes, H. sphondylium supports a wide spectrum of feedinginsects (Sheppard 1991). Insect herbivory can seriously restrict flower and seed production as can also,of course, grazing by large vertebrates including sheep, cattle and pigs.
The only fungal disease Hogweed commonly suffers from is a white powdery mildew, caused by Erysipheheraclei in late summer and autumn, but this too can seriously reduce flowering and seedproduction.
British and Irish occurrence
Common and widespread throughout B & I so as to be almost ubiquitous in soils of suitable pH onground up to around 1,000 m altitude (New Atlas). Subsp. sphondylium is the common form inB & I, while subsp. sibiricum (L.) Simonk has been found in Norfolk and in wild areas aroundthe Cambridge Botanic Garden.
European and world occurrence
The H. sphondylium species complex is common and widespread throughout temperate Europe andAsia, becoming less frequent towards the Mediterranean and N Africa (Hultén & Fries 1986, Map 1429).Subsp. sphondylium occurs mainly in NW Europe, extending into Scandinavia, east-central Europeand the mountains of the Mediterranean region (Sheppard 1991). Subsp. sibiricum is mainly a plantof NE and east-central Europe but also occurs in C & SE France (Sell & Murrell 2009).
Uses
Young Hogweed stem 'spears' have been eaten as green vegetables and are described as very succulent ifcooked briefly in boiling water like broccoli (Mabey 1972, 1996). However, it should be pointed outthat, like the alien Giant Hogweed (H. mantegazzianum), the plant contains furanocoumarins whichcan sensitize human skin to sunlight causing severe allergic reactions that are akin to scalding water,causing permanent scars. These toxins are not inactivated by cooking. A condition known as 'strimmerrash' is caused by skin contact of allergic field workers with sap from Anthriscus sylvestris(Cow Parsley) and Heracleum sphondylium (Cooper & Johnson 1998).
Hogweed has been traditionally used in herbal medicine as a mild expectorant and in a few proprietarymedicines for laryngitis and bronchitis (Launert 1981). In homeopathy, the plant has been used to treatkidney complaints (Sheppard 1991). The seeds have been used as the basis for an alcoholic beverage(French 1971), and foolish young boys and gypsies have even smoked the stems as a tobacco substitute(Vickery 1995).
Names
The genus name 'Heracleum' is Greek for 'Hercules' (or 'Heracles'), 'healer', which was a name given toit by Theophrastus (Gilbert-Carter 1964; Gledhill 1985). The Latin specific epithet 'sphondylium'translates as 'rounded', but it is not clear to what aspect of the species this refers (Gledhill 1985).
The English common name 'Hogweed' derives from the fact that the plant previously was collected as fodderfor pigs and rabbits (Grigson 1974), and the alternative common name, 'Cow Parsnip', also alludes to itsuse for domestic animals. Grigson (1955, 1987) lists a total of 70 English common names from around B& I, many of which refer to animals that presumably were known to graze the plant or were fed it byfarmers. For instance, Grigson (1955, 1987) reported that H. sphondylium is (or was) used to feedpigs up and down the country, people collecting the plant and, "carrying it home in bundles as afree harvest for the sty".
Threats
None.
Introduction, neophyte, an invasive, occasional garden escape.
1938; Praeger, R.Ll.; streamside at Maguiresbridge.
Throughout the year.
Fermanagh occurrence
This hugely tall biennial or perennial (ie in reality hapaxanthic – requiring 2-7 years before achievingflowering), monoecious, strong- or sharp-smelling herbaceous umbellifer species up to 5 m tall wasrecorded on the banks of the Tempo, Maguiresbridge and Ballinamallard rivers prior to 1951. While thesesites remain the centres of its Fermanagh distribution, it is now considerably more widely scattered inlowland areas and has been recorded in a total of 26 tetrads (4.9%), principally in the eastern lowlandsof the VC.
The primary colonisation sites of this invasive alien are typically in the deep, loamy or boggy soil ofrelatively undisturbed waterside habitats along the banks of rivers and streams, where it may then formlarge, dense stands (Tiley et al. 1996; Jonsell & Karlsson 2009). Fortunately, so far inFermanagh, it has not succeeded in establishing anything more than a token presence around or close toUpper and Lower Lough Erne. It has begun to appear, however, in other disturbed, ruderal, or unmanaged,damp lowland sites further away from water bodies, which means it has already started the process ofsecondary colonisation in Co Fermanagh.
Growth form, introduction and preferred habitats
Giant Hogweed is a native of the NW Caucasus mountains between Russia and Turkey, where it is a plant ofwet places at forest edges and in glades. In its native region, it often grows by stream-sides inmontane areas which have quite a damp, humid climate and an annual rainfall of between 100-200 cm. H.mantegazzianum was introduced into B & I in the 1820s for horticultural purposes, beingregarded as very dramatic and especially suitable for waterside plantation in larger gardens. Unusually,for an introduced species, it escaped into the wild almost immediately (by 1828 in Cambridgeshire) andit has been spreading in both B & I ever since (Tiley et al. 1996). In Ireland, the NationalBotanic Garden at Glasnevin was offering seed of H. mantegazzianum for exchange in 1889, whichappears to be the first mention of the plant in print in the country (Wyse Jackson 1989).
Throughout Britain and Ireland, it colonised new lowland territory extremely rapidly in the last 30 yearsof the 20th century. Surveys carried out in the late 1980s and 1990s suggested that if left unchecked itwould embark upon a secondary phase of invasion, colonising sites in the general countryside remote fromwater courses, mirroring the situation it has already accomplished in Sweden and the Czech Republic(Lundström 1984; Pyšek 1991; Tiley et al. 1996).
Secondary colonisation involves the invasive occurrence in low-lying rough grassland on roadsides,hedgerows, railway embankments, refuse tips, wasteground, cemeteries and disused farmland, in some ofwhich habitats H. mantegazzianum has already appeared in Fermanagh.
Flowering reproduction
The reproductive potential of the plant is prolific and almost as enormous as the physical scale of theplant itself! It is monocarpic in B & I (ie it flowers once and dies). The plant typically takes upto four year's growth to reach flowering capacity. The flowers are self-compatible, requiring nopollinator(s), and a vigorous individual plant may then produce up to 50,000 seeds in a season, althoughestimates of this vary from 5,000 upwards, peaking at the higher figure (Tiley et al. 1996). Theseed is dispersed by wind and water and can remain viable in the soil for up to 15 years.
Variation and hybrids
Little intra-specific variation has been detected in H. mantegazzianum and nosubspecific taxa have been described although there appears to be some confusion as to the specificlimits and a number of other apparently very similar taxa have been named (Page et al. 2006; Sell& Murrell 2009).
Hybridisation is unusual between species in the family Apiaceae, but occasional hybrids between H.mantegazzianum and H. sphondylium have been reported in Europe where both species grow inproximity. Fertility in hybrids is low (seed set less than or equal to 1%) and introgression has notbeen detected (Weimarck et al. 1979). The hybrid is multi-stemmed like H. sphondylium, butclearly larger. The hybrid is convincingly intermediate in all characters, although rather closer toH. sphondylium in plant height, stem diameter and ray number, and to H. mantegazzianum infruit length (Weimarck et al. 1979; Stace et al. 2015). The paucity of fruits and presenceof some malformed ones is an additional hybrid character.
British and Irish occurrence
H. mantegazzianum distribution is currently biased towards both the SE and the NW of both islands,but the species is spreading at an amazing rate – the change index calculated between 1962 and 2002 forthe New Atlas being one of the highest of any species at +2.09. This suggests that many of thepresent gaps in the hectad map could soon be filled unless weed control can be achieved (Preston etal. 2002).
The species' preference for the cooler, moist oceanic climate of N and W Ireland and N Scotland maypossibly be associated with the requirement for sufficient soil moisture at the seedling stage, but anecological preference for the more continental climatic conditions of SE England is more difficult toaccount for, except perhaps in terms of the prevalent warmer environment fostering its rapid growth rate(Tiley et al. 1996).
On the other hand, there may be an economic, historical and social reason for the current greaterfrequency of Giant Hogweed in SE England, since this is undoubtedly the region where the plant was firstintroduced and became a popular plant with the wealthy owners of large gardens. The impression createdby its grand scale will have appreciated quite some time before its overly aggressive and dominantnature became apparent. The danger posed by the allergic blistering sap is a further drawback which onlycame to light much later, but is now very well known and the general public has been frequently alertedto avoid the plant.
European and world occurrence
The species was similarly introduced, is actively spreading and becoming widely naturalized in 13 otherEuropean countries; it has also followed the same invasive pattern during the last 50 or 60 years inboth Canada and the United States (Tiley et al. 1996; Page et al. 2006). In Sweden, humaninterest and activity, for example collecting the impressive fruiting heads, also contributed to itsspread (Lundström 1984).
Toxicity
The sap contained in the hollow stem, leaves, branches and surface hairs contains substances calledfurocumarins, which on contact with the skin, but only in the presence of strong sunlight, can cause asevere allergic blistering reaction. The damage to skin is permanent, since its ability to filterultra-violet light is compromised, and future exposure to the sun will give rise to recurring severesunburn, a condition that will last for several, or perhaps many years. The sunburn will be accompaniedby rashes, blistering and swelling of the skin (Powell 1988). The painful blisters engendered by the sapdevelop into pigmented scars that can last for up to six years, but which more typically disappear aftera few months (Page et al. 2006).
On account of this danger the plant should never be touched without protective clothing. Having saidthis, the health threat from the plant can easily be exaggerated: for instance, while H.mantegazzianum is common and locally abundant on the banks of the River Clyde, a survey ofGlasgow hospitals and general practices close to large stands of the plant strongly suggested that thesap is not the serious medical problem it is often purported to be (Dickson 1998).
Weed control
Eradication is difficult and expensive, since early season spraying with glyphosate may not be totallyeffective for five to ten years (if ever). Manual cutting must be done below ground level otherwisevigorous growth is only encouraged. Ploughing is effective, but again it needs to be repeated since thelarger roots will regenerate the plant.
Rather amazingly, cattle, sheep, pigs and goats can browse the plants perfectly safely. Foraging by pigsthat damages the roots or heavy and prolonged grazing by sheep both eventually eradicated Giant Hogweedfrom pastures in Denmark (Tiley et al. 1996).
On account of its invasive nature and potential danger to humans, especially children, it is now illegalto plant Giant Hogweed or to transport its seeds. Dead plant material and the soil around the plant mustalso be disposed of in a recognised landfill site run by an appropriate local government authority.
Threats
Eradication requires sustained, deliberate destruction.
Native, frequent. Eurasian temperate and quite widely introduced to eastern N America.
1881; Stewart, S.A.; Co Fermanagh.
April to January.
Growth form and preferred habitats
This is an erect, semi-rosette, hairy, winter annual that rarely or occasionally behaves as a biennial.It grows on solid, striate stems 5-125 cm tall, pale green, sometimes tinted purple, clad with closelyappressed deflexed bristles. The plant has a slender tap-root (up to 8 mm thick) and produces muchdivided, 1- to 3-pinnate, fern-like, hairy leaves on slender petioles that are also covered withdeflexed appressed bristles (Tutin 1980; Grime et al. 1988, 2007; Sell & Murrell 2009;Jonsell & Karlsson 2010).
T. japonica appears on dry banks on woodland margins and clearings, in rough grass along roadsideverges and hedgerows, and on dry stony banks, rocks and cliffs in a wide variety of other mainlylimestone or base-rich, somewhat disturbed habitats. The soils it prefers are dry to moderately dry andfertile, mildly acid to neutral in reaction. However, it prefers situations where vigorous growth ofcompeting species is limited by periodic drought, instability of steep terrain, or some other negativeenvironmental factor(s). T. japonica prefers full sun but can tolerate moderate shade. Many, butnot all, of the wide range of habitats the species occupies are artificial, wayside, or 'man-made', suchas quarries and roadside banks and verges and it tends to be confined to lowland areas, below 400 m(Grime et al. 1988, 2007; New Atlas).
Upright Hedge-parsley is capable of growing to its maximum height (c 125 cm) if it finds deeper pocketsof suitable soil and some form of support to climb. However, very often it is much less tall than this,depending upon a combination of the severity of the growing conditions and particularly upon the densityand height of its competitors. Its competitors often include established, deeper rooted, perennialspecies, with which it is unusual for an annual ruderal to manage co-habitation. The fact that it cangrow in more open areas in woods or on their margins, in hedgerows and in tall verge grasslands that arenot cut or only annually mown, proves that T. japonica is able to compete and persist amongmainly perennial companions. The commonest associates in the Sheffield area of England were Dactylisglomerata (co*ck's-foot), Festuca rubra (Red Fescue) and Arrhenatherum elatius(False Oat-grass).
T. japonica is generally absent from regularly cut grassland and from well-managed pastures. Theestablished strategy of the species is classified as intermediate between C-S-R (ie a balance ofCompetitor, Stress tolerator and Ruderal) and Competitive Ruderal (Grime et al. 1988, 2007).
Fermanagh occurrence
While T. japonica is quite frequently found in Fermanagh and is widely scattered, having beenrecorded in 94 tetrads (17.8%), in most situations it is only sparsely present. Eight tetrads havepre-1976 records only. Occasionally, however, it is locally abundant along roadsides. In the absence ofChaerophyllum temulum (Rough Chervil) in Co Fermanagh, this bristly hairy, rather tall winter orspring annual is the last of the three most common wayside umbellifers in B & I to flower in the VC.
Flowering reproduction
The compound, flat umbels 2-4 cm across of pinkish- or purplish-white, distinctly radiate (zygomorphic)flowers make their appearance from July onwards. Fruiting begins in early September, following insect-or self-pollination and fertilization and the 'seed' (ie mericarp fruits) are retained on the plant asit dies off (Fitter 1987; Jonsell & Karlsson 2010). In a mild season, however, late flowering plantscan still be seen well into the autumn and the dead stems with their characteristic spine covered, ovalfruits are still perfectly identifiable in January (Tutin 1980). The fruits in this family aresingle-seeded mericarps formed in pairs, separating and hanging apart when fully mature and ready fordispersal (S.L. Jury, pers. comm., March 2005).
Compared with the seeds of most other annual species of similar habitats, the 'seeds' (ie fruitmericarps) of T. japonica are large, measuring 3-4 mm long. Relatively long bristles or spines(0.4-0.5 mm long) cover the mericarps, and they are all curved at their tip (but not hooked), whichenables them to be transported by attachment to the coats of passing animals. When there is no animalcontact, 'seed' dispersal would appear to be severely limited. Some populations of T. japonicahave fruits with the mericarp spines reduced to small tubercules; it has been suggested that thesefruits serve to maintain the local population, while the spiny ones travel further and colonise newareas (Jury 1980).
Seeds germinate mainly in the autumn and the plantlets overwinter as small leaf rosettes with a slendertaproot (Grime et al. 1988, 2007). Alternatively, some seed overwinters and germinates in thefollowing spring after a chilling requirement has been met. Naturally enough, the resultant plantsmature and fruit later in the season. There are conflicting reports of the longevity of seed in the soilbank, estimates varying from transient to long-term persistent for over five years (Grime et al.1988, 2007; Thompson et al. 1997).
British and Irish occurrence
Of the three most common roadside umbellifers in B & I, the hectad maps in the New Altasdemonstrate T. japonica is the least widespread. In comparison with the other two, it is slightlyless well represented than Anthriscus sylvestris (Cow Parsley) and more so than Heracleumsphondylium (Hogweed), especially in N & W Ireland, and much less so on the predominantlywet, acid, peaty soils of N & W Scotland (Preston et al. 2002).
European and world occurrence
T. japonica is widespread throughout temperate Europe from Ireland eastwards to W Russia andsouthwards just into N Africa. It extends in an extremely scattered, disjunct manner across to Japan, asits name suggests, where it has a secondary, and very much more minor, centre of distribution. Thespecies is a quite widely scattered alien introduction in N America, especially in eastern states of theUS (Hultén & Fries 1986, Map 1433). T. japonica is regarded as an invasive species insouthern parts of Canada and in the Mid-western, Eastern, Southern parts of the US, including Oregon.
Uses
T. japonica does not feature at all in western herbal medicine (Grieve 1931; Allen & Hatfield2004), but it has a long history going back centuries in traditional Chinese medicine. The plantcontains a range of bioactive ingredients that give it antiviral, antibacterial, anti-inflammatory andanti-oxidative properties. It is used today in the treatment of Lyme disease, influenza and a number ofother inflammatory conditions.
T. japonica may also have potential to fight several cancers through a terpene it produces calledTorilin that is extracted from its fruits. This compound has been shown to inhibit the growth of bloodvessels in tumour development from benign to malignant and thus has a toxic effect on tumours. It alsohas been found to inhibit the conversion of testosterone to androgen, which is being studied further inthe treatment of both prostate cancer and alopecia.
Names
The genus name 'Torilis' is an example of a pure label invented or dreamt up by the 18th century Frenchtaxonomist Michel Adanson (1727-1806), that in common with other names he provided (eg Apera,Cicendia, Kalanchoe and Tolpis) is meaningless and has no translation leading to oralluding to anything else (Gilbert-Carter 1964). As such, and with this knowledge, the current author(RSF) finds the label name surprisingly refreshing! The Latin specific epithet 'japonica' means 'ofJapan', but this is misleading and inaccurate, since the plant has a widespread range right across thewhole of Eurasia and Japan is merely a secondary centre of distribution, rather than a genuine point ofspecies origin.
The English common name 'Upright Hedge-parsley' is a typical book name. Grigson (1955, 1987) lists atotal of ten additional common names, several of which allude to the lace-like, heavily dissected leavesand white flower heads, eg 'Honiton Lace', 'Lace Flower', 'Lady's Lace' and 'Lady's Needlework'. Thesenames, and several other English common names, are shared with Anthriscus sylvestris (CowParsley), as Grigson says, "the companion plant of white lace along the roads, though Torilisjaponica blossoms as the true Cow Parsley fades. It is the roadside lace of high summer."
Threats
None.
Native, rare. Eurosiberian southern-temperate, but very widely naturalised and now circumpolar.
1950; MCM & D; Clonmaulin near Clontivrin.
April to November.
Growth form and preferred habitats
This subspecies is a monocarpic, biennial or short-lived perennial (perhaps very rarely, an annual), thathas a slender, whitish tap-root, less than 5 cm in diameter, becoming woody with age – quite unlike thefamiliar, closely-related Cultivated Carrot (subsp. sativus (Hoffm.) Arcang.), with its moreswollen, fleshy, orange, edible root (Dale 1974; Sell & Murrell 2009). The flowering stem of subsp.carota is solid, frequently coloured red or purple where it arises from the basal leaf rosette,and it varies in height from 10-120 cm. The basal rosette leaves are heavily dissected, 2- to 3-pinnate,fern- or parsley-like, but ± bristly hairy, and they range from 5-40 cm in length.
Wild Carrot appears on moderately disturbed dry, open or bare, sandy, gravelly or stony ground insand-pits, waste ground and quarries, or in short, open, well-lit, patchy turf. Usually this means itappears in unmanaged rough grassland on dry or well-drained, rather infertile, mainly calcareous,typically wayside situations. It avoids acid soils below pH 5 and has a bias towards infertile habitats(Grime et al. 1988, 2007). Experimental work suggests Wild Carrot grows faster, becomes largerand regenerates and survives clipping or grazing better on fine textured, somewhat damper soils(Harrison & Dale 1966).
Until it is mature and capable of initiating flowering, the Wild Carrot plant exists as a compact rosetteof basal leaves that is easily overgrown by taller, more robust dominant species. Thus, D. carotasubsp. carota usually only manages to persist in short turf oropen sites in ungrazed or under-used pastures in situations where dominant species are somewhatrestricted by other environmental factors, such as low nutrient levels or lack of moisture. Theestablished strategy of the subspecies is categorised as intermediate between stress-tolerant ruderaland the more balanced C-S-R (ie Competitor-Stress-tolerant-Ruderal) (Grime et al. 1988, 2007).
Requiring a growth period of more than 120 days frost-free, in B & I it is confined to lowlandsituations below 450 m. Wild Carrot is indifferent of photoperiod length, being adventive throughoutwarmer parts of the world and occurring from C America and Mexico to ground north of 65°N in Sweden(Dale 1974). It prefers full sun and when shaded its vigour drops. In deeper shade, its reproductivemethod may change to that of a short-lived perennial (Harrison 1965). In Europe, Wild Carrot showed apreference for medium to high levels of nitrogen, but was indifferent to phosphate levels on wellfertilised sites in hay meadows in Germany (Kϋhner 1951, cited inWilliams 1968, quoted in Dale 1974). It is also much less closely associated with calcareoussoils in Europe than is the case in Britain, so that it might even be described as indifferent to limeon the continent (Dale 1974).
Herbivory
The characteristic carrot odour is perceived when tissue of any part of the plant is crushed. Milk ofdairy cattle can become tainted when large amounts of Wild Carrot are present in pasture, but this isvery unlikely today and it has never been reported that cows graze upon it from choice. However, thedigestibility and nutritive value of Wild Carrot foliage is similar to that of legumes and, therefore,it should be tolerated in pastures among plants of low nutritive value. Cattle, horses and sheep willall graze Wild Carrot (Harrison & Dale 1966; Dale 1974). Compared with the leafy basal rosette, theflowering shoot is particularly conspicuous and therefore much more vulnerable to grazing damage ordestruction (Dale 1974).
Flowering reproduction
D. carota subsp. carota is completely dependent on seed forsurvival, increase and dispersal. The timing of flowering in a maturing plant appears to be triggered bya combination of rosette size and recent growth rate. An increase in size but a decrease in relativegrowth at the end of the summer is associated with an increased probability of flowering in thefollowing growing season for short-lived monocarpic species, including D. carota subsp. carota. Year of flowering also has a genetic component (iematernal phenotype – annual, biennial or triennial) and nutrient supply certainly also plays a part indetermining growth and the timing of the switch between vegetative and reproductive growth (ie bolting).In an experimental study, maternal phenotype influenced the year of flowering both directly andindirectly through growth. Plants die after bolting regardless of whether or not they set seed (Lacey1986).
The compound umbels are terminal and andromonoecious (ie they contain both male and bisexual flowers);the bisexual blossoms are protandrous, ie the pollen matures before the stigma does, thus favouringcross-breeding. Umbels normally consist of over 1,000 white flowers, developing from a convex dome intoa flat circle at time of fertilisation. The flat white inflorescences often have one or more centralflower(s) that are slightly enlarged and coloured black, red or purple. This observation and theirexistence is something of a puzzle to explain, and since at least the middle of the 19th century therehas been an ongoing search for their purpose, if any (Hoekstra 1997). It has been suggested the darkflorets may have adaptive functions, such as mimicking insects (including beetles), visiting theinflorescence, possibly to discourage herbivory, pollen and nectar theft, or alternatively, to attractadditional pollinators to the plant, perhaps by indicating the presence of food or opportunities formeeting and mating. Charles Darwin dismissed the dark flower spot as a vestigial remnant of someredundant function, and he just might be perfectly correct. However, the debate continues and thecurrent author (RSF) cannot yet see where it is taking us, since insect visitors attending theinflorescence are already legion, without the need for any enhancing mechanism, simple or complex, thatrequires further explanation (Dale 1974).
After fertilisation the inflorescence becomes contracted into a concave, cup- or bird's nest-like massduring the period of fruit maturation. Eventually, the nest-like mass flattens out again during seeddispersal.
The outer flowers in umbels are surrounded by a whorl of green, finely-divided bracts. A large floweringplant individual may produce up to 100 umbels during the flowering season from June to August. Axillaryleafy, flowering stalks are produced in succession until the autumn, when the plant dies (Dale 1974;Sell & Murrell 2009).
Cross-fertilisation is the norm and a very wide range of unspecialised insect visitors are attracted byabundant, freely available pollen and nectar to carry out pollination. Should crossing fail for anyreason, the flowers can self-fertilise (Dale 1974).
As with other umbellifers, the schizocarp fruit splits into twin, single-seeded mericarps (or achenes),3-4 mm long × 2 mm wide, for dispersal (Dale 1974). Man and his activities represent the major vectortransporting the bristly, spiny-looking mericarps over long distances, although wind and attachment toanimal coats are also definitely involved in local dispersal (Salisbury 1964; Lacey 1981). Lacey showedexperimentally that animal coats rougher than those of mice and rabbits can hold the spiny, barbed seedlong enough to assist dispersal. She also proved that wind in autumn and winter can carry seed longdistances across snow and frozen ground (Lacey 1981). Seed can also be ingested by browsing animals andcan pass undamaged through the digestive tract of a horse (Salisbury 1964). Seed is long-persistent insoil, surviving for more than five years (Thompson et al. 1997).
Variation
D. carota is an extremely variable, environmentally plastic and genetically polymorphic species,to the extent that eleven subspecies are described in Flora Europaea (V.H. Heywood, in Tutinet al. 1968, 2, p. 374). Most wild Daucus forms occur in SW Asia and theMediterranean region, a few in Africa, America and Australia. All wild forms investigated in Asia, AsiaMinor, Japan and the USA have the same chromosome number as their European relatives (2n=18). Neitherpolyploidy nor structural changes in chromosomes seem to have played a role in the differentiation ofthe species (Whitaker 1949; O. Banga, in: Simmonds 1976). The Iranian Plateau (Afghanistan, Iran andPakistan) is one of the areas where subsp. carota shows the greatest variation and has beensuggested as a possible source of the species origin. The roots vary in their degree of ramification,fleshiness and colour: some are white and others coloured in varying degrees by anthocyanin (O. Banga,in: Simmonds 1976; https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:841063-1website viewed 29 January 2022). Subsequent breeding and development of the modern, larger, orange,fleshy, western carotene carrot (subsp. sativus (Hoffm.) Arcang.) was carried out in theNetherlands in the 16th and 17th centuries (Heywood 1983 – abstract only viewed).
Only subsp. carota and the cultivated subsp. sativus occur in NI and there is very littlechance of them inter-breeding. There are a few stations for the coastal subsp. gummifer (Syme)Hook. f. in SE Ireland and again in W & S Britain (Preston et al. 2002).
Fermanagh occurrence
In Fermanagh, Wild Carrot has been recorded in 16 tetrads (3%), ten of them with post-1975 dates. Inlandlocked Fermanagh, only twelve stations for Wild Carrot have been recorded since 1988. The two mostrecent finds, where the plant was seen in any quantity, were beside a stream in Cross Td, on thenorthern outskirts of Enniskillen, where 25 plants were counted in September 2001, and outside thequarry gate at Coagh, 5 km N of Enniskillen, where ten seedlings were found in October 2001.
British and Irish occurrence
Wild Carrot is a common and familiar plant in coastal sites in NI, but rather rare and possibly declininginland. The distribution pattern of the plant elsewhere in Ireland is common on coasts, rarer inland,but subsp. carota becomes more widely recorded southwards.
D. carota subsp. carota is widespread in lowland Britain, althoughagain it becomes increasingly scarce and coastal north of the Humber-Severn line. The hectad map in theNew Atlas suggests a decline in Scotland, both inland and along the E coast. A smattering of redsymbols on the same map reflects the fact that nowadays D. carota subsp. carota is sometimes included in wild flower seed mixturesthat are increasingly sown in amenity and garden situations (Preston et al. 2002).
European and world occurrence
Carrot is a cool weather plant (O. Banga, in: Simmonds 1976). Wild Carrot is widespread in Europe southof the Baltic, plus in temperate Asia and North Africa (Tutin 1980) but it has been introduced to manyareas around the world and become so widely naturalised that D. carota s.l. is now adiscontinuous circumpolar southern-temperate taxon (Hultén & Fries 1986, Map 1434; M. Southam, in:Preston et al. 2002). Having said this, it is such a variable species aggregate with so manysubspecies distinguished, it is difficult or often impossible to determine whether an occurrence isindigenous or not. It is known, however, that subsp. carota has been introduced and becomenaturalised (sometimes abundantly) in N America and in New Zealand (Dale 1974; Webb et al. 1988).
Uses
In Classical times, the white-rootstock carrot we know as Daucus carota subsp. carota was well known and is mentioned by Greek and Latinwriters, although there sometimes is a measure of ambiguity regarding whether they are referring tocarrot or parsnip. However, by the time Dioscorides described the plant in the first century AD, he gavean accurate description of the carrot dealt with here (Grieve 1931).
The western, bright-orange coloured, fleshy, carotene Cultivated Carrot was first introduced to Englandby Fleming refugees in the reign of Elizabeth I (1553-1603). Previously, the Wild Carrot was cultivatedand used both as a vegetable and for medicine although the root was small and spindle-shaped, whitish,slender, tough and had an acrid, unpleasant taste.
In herbal medicine, the whole Wild Carrot plant had uses and was considered more efficacious than thecultivated form of the plant (Allen & Hatfield 2004). It was considered a diuretic, stimulant and'deobstruent'. A tea-like infusion of the whole herb was considered an active and valuable remedy in thetreatment of dropsy, chronic kidney disease and diseases and affections of the bladder. A strongdecoction was used to break up stones and gravel and was good in treating flatulence. The seeds weredescribed as carminative (ie used for flatulence) and stimulant and were useful in treating flatulence,windy colic, hiccough, dysentery and chronic cough. They were also believed to clear obstructions of thevicera and useful for jaundice and scorbutic disorders (ie the treatment of scurvy). An infusion of theroot was also used to ease constipation (Grieve 1931).
Grieve also reported the old belief that a poultice made from the roots helped relieve the pain ofcancerous ulcers, and the leaves applied with honey, cleansed running sores and ulcers (Grieve 1931).
Names
The name 'Carota' was first used by Athenaeus around 200 AD and it was Galen (second century AD) whoadded the name 'Daucus' to distinguish the carrot from the parsnip (Grieve 1931). An additional elevenEnglish common names for 'Wild Carrot' are listed by Grigson (1955, 1987). 'Bird's Nest' is a name takeninto English from the German, referring to the nest-like contracted umbel fruit. 'Bee's Nest' and'Crow's Nest' are obvious alternatives derived from the same idea. The names 'Kex' and 'Keggas' areassociated with plants with hollow stems in general and often to other umbellifer species in theApiaceae. A Scottish name, 'Curran-Petris', denotes a carrot root, 'curran' being Scots gaelic for'carrot' (Britten & Holland 1886). The name 'Eltrot' or 'Eltroot' is given to several umbellifersincluding D. carota and is said to be from the Anglo-Saxon 'eald', meaning 'old', and root, orfrom 'elt', meaning 'a young pig', and root. The name 'Rantipole' refers to the bunch of leaves thatcompose the basal rosette (Britten & Holland 1886).
Threats
None.
Native, occasional. European southern-temperate, but also widely naturalised in N America, S Australiaand New Zealand.
1882; Stewart, S.A.; Co Fermanagh.
May to December.
Growth form and preferred habitats
This glabrous, rosette-forming, tap-rooted biennial or winter-annual has 4-6 pairs of opposite, ovatestem leaves that are more than 5 mm broad, clustered pink flowers, usually in at least five cymes, andproduces one (or rarely more) erect flowering stem(s) that can vary greatly in height, from 9-30 cm tall(Ubsdell 1976a; Garrard & Streeter 1983).
C. erythraea is a mobile, small-seeded, locally frequent, grassland colonising species thattypically occupies relatively infertile, unproductive, lowland soils that are characterised by amoderate degree of disturbance. These conditions together create an open or semi-open habitat withlittle competition and a near absence of grazing since the plant contains a number of very bittersubstances that deter most browsers. As an example of this, C. erythraea has persisted in apaddock heavily grazed by two horses for 24 years on shallow-soiled land owned by the current author(RSF) and his wife.
The species is rather variable and ecologically wide-ranging and it also occurs thinly scattered onlakeshores, riverbanks, woodland glades and margins, open scrub, roadsides and quarries, where it growson suitably drained, open sandy soils of a somewhat more acidic nature. It can tolerate trampling,compacted soils and light grazing, but cannot compete successfully with vigorous grasses (Sinker etal. 1985). The established strategy of C. erythraea is thus well categorized as aStress-tolerant Ruderal species (Grime et al. 1988, 2007).
In coastal areas of B & I, C. erythraea is very frequent on sheltered, dry or well-drained,mildly acid to calcareous dune grasslands. Studies in the Sheffield area showed C. erythraeaoccurring on soils down to pH 4.8, but no lower, and the majority of sites were above pH 5.5 (Ubsdell1976a; Grime et al. 1988, 2007).
Variation
Phenotypic variation within C. erythraea with respect to the environment in B & I isconsiderable and no less than five varieties have been proposed (Ubsdell 1976a; Sell & Murrell2009). Only one of these, var. erythraea, occurs inland as well as on sheltered coasts (AnIrish Flora 1996).
Fermanagh occurrence
C. erythraea has been recorded in 67 Fermanagh tetrads, 12.7% of those in the VC. There are 16Fermanagh tetrads where the species has pre-1976 records only, which reflects the transitory nature ofsome of the more open habitats it colonises. The well-drained but not overly dry, infertile soils of aneutral to mildly acidic reaction that Common Centaury most typically frequents are common in limestonedistricts of Fermanagh.
British and Irish occurrence
In terms of frequency, C. erythraea is chiefly, yet far from exclusively, a coastal plant in thenorth of Ireland (FNEI 3). Further south in Ireland, it is more generally distributed inland,occupying relatively dry, but essentially well-drained, short, unproductive pastures with quite highlevels of bare soil. Although it has been recorded in every Irish VC, C. erythraea is still muchmore abundant at the coast (Cen Cat Fl Ir 2; An Irish Flora 1996).
In Britain, the New Atlas map shows the distribution is widespread and perfectly stable, oractually increasing in frequency in lowland areas south of a line between Sunderland and Lancaster, asshown in the 1987-2004 monitoring survey (Braithwaite et al. 2006). It becomes scarce and muchmore coastal further north beyond this line and it only very rarely occurs inland in Scotland where ithas been recorded as far north as the isle of Lewis on the W coast.
Phenology and ecological separation from a closely related species
Seed of C. erythraea germinates in the autumn, produces a taproot and overwinters as a tightrosette of leaves flattened close to the ground. Whether C. erythraea is regarded as awinter-annual or a biennial, certainly it is monocarpic (ie plants fruit only once and then die), so thedistinction between these two life forms rests purely on the length of time the individual plant needsto achieve the size and accumulate the photosynthetic resources needed to produce flowers, fruit andseed. After seeding in the late autumn the plant dies. Shed seed germinates in vegetation gaps duringthe autumn (Grime et al. 1988, 2007).
After winter, growth resumes in the spring, producing an erect stem up to 30 cm tall. Height is veryvariable, however, and in the north of Ireland the plant is frequently only around 15 cm tall. Itdevelops lower stem leaves which are ovate and more than 5 mm broad, a feature which distinguishes thisspecies from C. littorale (Seaside Centaury, or sometimes known as Dwarf-tufted Centaury), whichhas a shorter stem and narrower leaves. The latter is a closely related species which rather rarelyoccurs intermingled with C. erythraea on sheltered coasts of Britain and N Europe (Ubsdell 1979),but it is confined to a narrow portion of the Co Antrim coast in Ireland (H39). Both these species aretetraploid (2n=40) with regular meiosis (Ubsdell 1976b). While there are no effective barriers to geneexchange between the two species, ecological isolation operates through slight differences in floweringtimes and the tendency for C. erythraea to inbreed (Ubsdell 1979).
Since Fermanagh lacks a coastline, C. littorale has not appeared and is not likely to arise inthis area (Ubsdell 1979; Garrard & Streeter 1983; Webb et al. 1996; Stace 1997).
Flowering reproduction
C. erythraea flowers from June to October, small individuals having only a few flowers (usuallyless than 10), while larger plants produce a profusely branched, laxly spreading or densely crowdedcymose inflorescence, with many pale pink, star-like flowers. The anthers and stigma are exserted beyondthe floral tube, so the flowers are open to many insect types. They produce no free nectar, but areinsect-pollinated, most likely by small black thrips. They open early in the day, between 5 am and 7 amand begin to close around noon, or earlier if the sky becomes overcast. Failing insect pollination, indull weather when the flowers close up for example, self-pollination is highly effective and plants arehighly self-fertile with 77-100% seed set (Melderis & Bangerter 1955; Ubsdell 1979). Flower numberand seeds/capsule measurement means found the average individual produced around 10,800 seed (Ubsdell1979). The fruit is a two-valved capsule that splits to release numerous seeds. All seed set inexperimental measurements by Ubsdell (1979) showed a high percentage germination.
There is no specific adaptation enabling seed dispersal but presumably, as with other plentifullyproduced small seed of species of semi-open habitats, it relies upon wind, rain wash and transfer in mud(Ridley 1930). Seeds are short-term persistent in the soil seed bank, ie present for at least one year,but less than five years (Thompson et al. 1997).
In common with other members of the Gentian Family, C. erythraea roots have a mycorrhizal fungalassociation. As the small, lightweight seed incorporates minimal food reserves, the presence of thefungal partner, open habitats featuring little or no competition for scarce nutrients and for light, areall essential, required factors for successful seedling establishment (Brown & Oosterhuis 1981).
C. erythraea shares many features of its biology and ecology with the related annualBlackstonia perfoliata (Yellow-wort) and, in many regions of B & I, they frequently occurtogether. At first glance this might suggest B. perfoliata is an obvious 'absentee species' notyet recorded in Fermanagh. However, the species distribution shown in the two BSBI atlases (Perring& Walters 1962; Preston et al. 2002) clearly demonstrates that B. perfoliata has avery much more restricted B & I distribution than C. erythraea and the former does not occuranywhere across the whole six counties of NI.
Toxicity
C. erythraea contains a bitter principle, erythro-centaurin and a bitter glucocide, erytaurin,which appear effective in discouraging most browsing animals. Of all the bitter appetising wild herbs,Common Centaury is regarded the most efficacious, sharing some of the antiseptic properties of FieldGentian (Gentianella campestris) and Bogbean (Menyanthes trifoliata) (Grieve 1931,referred to as Erythraea centaurium (Pers.)).
European and world occurrence
C. erythraea s.l. is present throughout Europe from S Sweden southwards to the Mediterraneanbasin, Asia Minor, N Africa, Arabia and the Azores. It stretches eastwards to SW Asia and is introducedand naturalised in eastern and western N America, Chile, S Australia and New Zealand (Hultén & Fries1986, Map 1492; Clapham et al. 1987).
Uses
It has a herbal reputation for ailments of the liver and kidneys and has been used to treat fevers andbleeding. Being bitter in flavour, it was said to "purify the blood" and be an excellent tonic(Grieve 1931). Used as a tonic, C. erythraea is less liable to upset digestion than most othervegetable bitters and so was a preferable alternative to Yellow Gentian (Gentiana lutea) from thecontinent. It has also been used to treat colic, coughs, jaundice and as a wound healer or bloodstauncher (Allen & Hatfield 2004).
Names
The scientific genus name 'Centaurium' and the English 'Centaury' are both derived (like 'Centaurea')from the Greek 'kentaur', referring to the legend of Chiron the centaur, one of mythology's divinehealers, who was reputed to have a wide knowledge of herbs. The Latin specific epithet 'erythraea',previously was the name of the genus and is Greek for 'reddish', which really does not describe theflower colour accurately (Gilbert-Carter 1964; Grigson 1974).
Grigson (1955, 1987) list ten English common names, some of which refer to the herbal uses mentioned, forinstance 'Bloodwort', 'Feverfew' and 'Gentian', the latter since it was a cheap substitute for importedcontinental Gentiana lutea (Yellow Gentian), the extremely bitter roots of which had many herbaluses, especially as a strong tonic. Culpeper (1653) approved of Centaury's use for a whole raft ofmedicinal problems, but aware of its rather bitter taste he wrote, "It is very wholesome, but notvery toothsome.", which, in the view of the current author (RSF), expressed it rather well (Grigson1955, 1987).
In Irish and Manx folklore, C. erythraea is a 'blessed herb', bringing good luck, since legendsays it grew where Jesus Christ trod on the way to Calvary (Grigson 1955, 1987).
Threats: None.
Native, rare, NI Priority Species of conservation concern.
European boreo-temperate.
1882; Stewart, S.A.; Legland Mountain, SW of Knockmore.
July to September.
Growth form and preferred habitats
This little, erect, branching annual or biennial gentian is nothing like as conspicuous as Gentianaverna (Spring Gentian), or other members of the family found in the European mountainspossessing unforgettable brilliant blue corollas. The flowers in this instance are a rather darkbluish-purple, the tube with a long bluish fringe at the mouth, the corolla then dividing into four,occasionally five, ± spreading petal lobes. The calyx is undoubtedly the most distinguishing feature ofthis particular species, having four lobes, the two outer ones being much wider, enveloping the otherpair.
When not in flower, Field Gentian is very inconspicuous and difficult to locate and, consequently, mustoften be overlooked. G. campestris flowers from early July to late September and appears rarelyand very sparingly in short, grazed turf in dry, rocky, heathy unimproved grassland in a variety ofterrain, from lowland pastures to cliffs, screes, to mountain (or rather hilltop) pastures. It frequentsmildly acidic to neutral, infertile soils in a variety of open habitats, often over leached limestone,or elsewhere on fixed maritime sand dunes. It is generally a lowland species in B & I, but doesreach an altitude of 915 m at Cairnwell, NE Scotland (VCs 89 & 92) (R.D. Porley, in: Preston etal. 2002; Smith & Lockwood 2011; Stroh et al. 2019). It prefers constantly moist,well-drained soils and full sunlight and it is slightly salt-tolerant (Hill et al. 2004).
Locally in Fermanagh, these dry, rocky grassland habitats are confined to the limestone areas of thecounty where the shallow soils are heavily leached, nutrient-depleted and become neutral or mildlyacidic, so that a species-rich pasture of low productivity develops and persists. G. campestrisis often found on the shallowest soils around rock outcrops on cliffs, screes and limestone pavement.Typical associated species include Antennaria dioica (Mountain Everlasting), Briza media(Quaking-grass), Campanula rotundifolia (Harebell), Danthonia decumbens (Heath-grass),Galium verum (Lady's Bedstraw), Linum catharticum (Fairy Flax) and Thymuspolytrichus (Wild Thyme).
Flowering reproduction
Regeneration is entirely by seed and the species is monocarpic. Germination takes place in the spring,dormancy being overcome by cold stratification overwinter (Milberg 1994). The low-growing, erect flowersare visited by long-tongued humble-bees and butterflies, the only insects in B & I that are able toreach into the fringed corolla tube to find the nectar legitimately, although as in the genusGentiana, robber bumble-bees may bite and bore holes through the calyx and corolla tubes andsteal the food reward. The flowers also self-pollinate very readily and full seed set often results(Clapham et al. 1962; Lennartsson et al. 2000). Measurements made of reproductive capacityby Salisbury (1942) found G. campestris plants produced an average of just 5.5 capsules, with anaverage seed content of 60, giving a calculated mean output of 325 ± 23 seeds per flowering plant.Subsequent study found fruits contained between 40 and 120 ovules, supporting Salisbury's findings (K.Walker, in: Stroh et al. 2019).
Seed has proved nigh impossible to germinate experimentally, which probably indicates a long period ofseed dormancy, a feature that is also the case in G. amarella (Autumn Gentian) (Pritchard 1959).Dispersal of the small, lightweight seed from the fruit capsule is most probably achieved by wind,especially during the winter when the surrounding vegetation dies down somewhat.
Horses, cattle and undoubtedly other wild grazing animals, eat the fruiting plants and disperse gentianseed with their dung, despite the fact that all Gentian species contain bitter glycoside substances akinto tannins. As a result of these, the species had herbal medicinal uses in the past (Ridley 1930; Grieve1931).
There is no evidence for G. campestris having long-term survival of buried seed. The soil seedbank survey of NW Europe did not mention G. campestris, but of 18 reports for eight relatedGentiana and Gentianella species, only one told of seed surviving longer than 3 years (ieshort-term persistent) for Gentianella germanica (Chiltern Gentian) (Thompson et al.1997). The seed must therefore normally be transient, surviving for one year or less.
The observation that in Britain undergrazing represents a greater threat to G. campestrispopulations than overgrazing, suggests the species cannot survive or 'out-live' unfavourable growingconditions when they occur, since it has no buried seed reserves available to revive and re-establishthe population should favourable conditions return. Undergrazing or neglect of grassland quickly leadsto invasion and overgrowth by shrubs, vigorous herbs and coarse grass species (Walker et al.2017; K. Walker, in: Stroh et al. 2019).
Fermanagh occurrence
In Fermanagh, there are records of small populations of G. campestris from just nine post-1975tetrads, plus one tetrad with a 1970 date at Oweyglass upland. As the tetrad distribution map clearlydemonstrates, its Fermanagh sites fall into two distinct districts, around theMonawilkin-Knockmore-Legland area and S of Lower Lough Macnean in the limestone uplands around theCrossmurrin NR and the Gortmaconnell Rock area.
Irish occurrence
In NI, G. campestris to some extent mimics (without actually belonging to) the group ofso-called Irish 'Mountain plants', a listing similar to and based upon Watson's 'Highland Type' (Watson1859). This grouping was very loosely defined by Watson as, "plants chiefly seen about themountains" (Praeger & Megaw 1938). The definition does NOT carry the inference that 'mountainplants' are confined to upland sites, and in Ireland (especially as one travels westwards), manyof these mountain species also occur in utterly different growing conditions. For instance, some of themgrow right down to sea-level in coastal, often sandy habitats, including Dryas octopetala(Mountain Avens) and Gentiana verna (Spring Gentian) in the Burren and Connemara (H9 & H17)(Moore & More 1866; Colgan & Scully 1898; Praeger 1934; Webb & Scannell 1983).
In parallel with this so-called 'mountain group' of Irish plants, G. campestris occurs and,indeed, is much more frequent and prevalent in coastal sites in the N & W of Ireland, while alsobeing found down the E coast as far south as Wexford (H12). However, G. campestris is also foundgrowing at high elevation in the Wicklow hills (H20) (Perring & Walters 1962, 1976; Hackney etal. 1992).
G. campestris populations have been in steady decline in NI over the last 60 years or more, andNIEA has listed it as a priority species of conservation concern. In NI, the number of hectads withrecords of any date total 36, but only 13 squares have post-1986 observations (NI Vascular PlantDatabase, accessed 2010).
British occurrence
Previously widespread in Britain, G. campestris is now rare and localised in S England and SWales, but still relatively frequent in the Pennines, Scottish mountains and around the Scottish coastand islands (Smith & Lockwood 2011). However, the species has suffered a severe decline across itswhole European range since about 1930 and some lowland regions have lost the majority of theirpopulations in the last 50 years (K. Walker, in: Stroh et al. 2019). This has also been theexperience in B & I.
In England and Wales, the decline of the species is very marked, especially in SE England (Walker 2007).It is now extremely rare in southern England, except in the New Forest (VC 11), where populations arestable or possibly expanding. Cumbria (VCs 69 & 70) remains the last major English stronghold –although even here it has disappeared from over half of the hectads for which there are post-1930records (Halliday 1997; Preston et al. 2002).
Possible explanations for species decline
G. campestris and the closely related Autumn Gentian (G. amarella) both require an open,short turf habitat. The demise of much of the rabbit population following myxomatosis in the 1950sundoubtedly affected the extent of open grassland habitats suitable for both these Gentianellaspecies (Pritchard 1959). This detrimental situation has not yet been reversed by the development ofalternative grazing practices. Surveys and monitoring of surviving populations by governmentconservation bodies and subsequent listing in Red Data Book terms as 'Vulnerable' or 'Endangered'species of concern (Cheffings & Farrell 2005), together with listing of sites of special scientificinterest, should certainly have helped alleviate environmental pressures to some extent. However, activemanagement measures, including reintroduction programs and the careful addition of monitoredlow-intensity, autumn to spring grazing by sheep and rabbits, may also be required in the long term toreplace or restore extinct or near-extinct populations (Smith & Lockwood 2011).
The widespread decline of G. campestris and G. amarella populations in B & I isprobably associated with the loss of hay-meadows (including their conversion to silage) and, in lesssteep or hilly areas, agricultural pasture 'improvement' measures, involving ploughing, reseeding andthe regular application of fertiliser (often as slurry). Together with the widespread use of herbicides,farmland management practices such as these have increased grass production and threatened the survivalof these two interesting Gentianella species.
Populations of G. campestris are still being lost in Britain due to overgrazing in the uplands andthe neglect or mis-management of lowland pastures. Cessation or a marked reduction in grazing or mowingof grassland, especially in more fertile mesic sites, can quickly result in local extinction of G.campestris (Stroh et al. 2019).
European occurrence
A European endemic species, G. campestris has a boreo-temperate phytogeographic distribution, butis declining in presence throughout its range (Preston & Hill 1997; Smith & Lockwood 2011). Itis widespread across N Europe, including Iceland and extends southwards to the Alps and more rarely tothe Pyrenees, Picos de Europa and Apennines (Hultén & Fries 1986, Map 1504; Smith & Lockwood2011).
Uses
In herbal medicine, G. campestris fulfilled a similar role to Centaurium erythraea (CommonCentury) in acting as a replacement for the continental Gentiana lutea (Yellow Gentian) root. Assuch, it was used (especially in the north of England and in Scotland, where it was more frequent) as atonic and for treating gravel, digestive complaints, and as a cure for jaundice (Allen & Hatfield2004). In the Scottish Highlands, Field Gentian was used to treat a rickets-like disease in cattle thatenforced crouching. Nowadays, this is recognised as a phosphorus deficiency (Vickery 2019).
Names
The genus name, 'Gentianella', is a diminutive of 'Gentian', which is a name in Pliny of a plant calledafter Gentius, a 2nd century Illyrian king, who is reputed to have discovered the medicinal use ofGentiana lutea (Yellow Gentian) (Gilbert-Carter 1964; Hyam & Pankhurst 1995). The specificLatin epithet 'campestris' means 'of the pasture' or 'from flat land' (Gledhill 1985).
Threats
Despite conservation listing and management measures over at least a decade, G. campestrispopulations remain small and declining. Most likely this is due to changes in land management andintensification of farming involving reseeding of old pastures, overgrazing of upland pastures and theuse of fertiliser which encourages growth of stronger competitors. Atmospheric pollution and climatechange may also be significant, but little information of these factors exists yet (K. Walker, in: Strohet al. 2019).
Native, very rare. Circumpolar boreo-temperate.
1884; Barrington, R.M.; Lower Lough Erne.
August and September.
Growth form and preferred habitats
A very variable species, G. amarella is sometimes considered a summer annual, ie it germinates inthe spring and produces a flowering individual later in the year. However, if insufficientphotosynthetic resources are accumulated to permit reproduction in this manner, and in G.amarella this is usually the case, then the plant behaves as a biennial, producing anoverwintering underground bud in the first autumn after germination and flowering in the second summer.The aerial leaf rosette dies away completely in the winter, but the plant will re-grow and flower duringits second season (Pritchard 1959). G. amarella has calyx lobes all ± equal in size and a corollathat is usually 5-lobed. The plant varies greatly in size, from 3-30 cm tall, stems simple or branchedfrom the base. The flowers are slightly smaller than those of G. campestris (Field Gentian),usually bluish-purple, though occasionally pale blue, pink or white.
G. amarella is a more definite calcicole than G. campestris, being closely associated withdry or well-drained, nutrient-poor, infertile, grazed, short-turf, calcareous or basic pastures, drybanks and sandy or gravelly places, including maritime grey sand dunes and dune slacks, coastal machairand limestone quarries. Sometimes it is also found on similar well-drained basic soils in roadsidecuttings, quarries and spoil-heaps (Kelly 1984; R.D. Porley, In: Preston et al. 2002). It is amostly lowland species, but does occur in NW England at up to 750 m in Westmorland (VC 69) (Pritchard1959; Sinker et al. 1985; Sell & Murrell 2009; Parnell & Curtis 2012).
Variation
There is considerable genetic variation within G. amarella, eg the flowers can be four- orfive-parted, even on the same plant. Petal colour can range from the usual rather dull, dark purple to amuch rarer white, pink or pale blue (Clapham et al. 1987; New Flora of the BI 1997). As aresult, several local endemic subspecies have been described for B & I (Pritchard 1959).
Measurements of seven plant characters in Irish G. amarella plants made by Pritchard found theyformed a hom*ogenous group that lay outside the range of British and European material examined. TheIrish plants differed in the size of their corollas and this was correlated with other small butdefinite differences. All Irish G. amarella plants are now assigned to the endemic subsp.hibernica N.M. Pritch., having slightly longer corollas and narrower and less tapering leavesthan subsp. amarelle, to which it most closely lay (New Flora of the BI 1997; Sell &Murrell 2009).
However, Pritchard subsequently reconsidered the Irish material and decided to reassess its taxonomicrank, referring it then to "a race of G. amarella". Examination now confirms that thedifferences between Irish and British material are relatively small, and the separate subsp.hibernica is not worth retaining (Parnell & Curtis 2012). Stace (2019) continues to key outsubsp. hibernica, but does admit it is, "possibly not worth separation from subsp.amarella". There currently are four subspecies recognised in B & I, subsp.amarella, subsp. septentrionalis, subsp. anglia and subsp. occidentalis, ofwhich only the first is known to occur in Ireland (Stace 2019).
Hybrids
Rare hybrids are formed in Britain with three other Gentianella species, G. germanica(Chiltern Gentian), G. anglica and G. uliginosa, but none of these three species occurs inIreland and naturally the hybrids are also absent (Stace et al. 2015). Two of the three 'species'(G. anglica and G. uliginosa), have been reclassified as subspecies (Stace 2019).
Fermanagh occurrence
The two or three Fermanagh stations for G. amarella are the only ones known in NI and thus RHN andthe current author (RSF) believe it should be given the conservation status and protection of theWildlife (NI) Order, Schedule 8. The sites are on limestone pastures at Screenagh (August 1970, L.Farrell, not refound), at Rahallan Td, S of Belmore Mountain (where patches can vary from a few plantsto 400, 16 September 1990 to 4 October 2003, RHN), and on the SE and NE shores of Monawilkin Lough(where it occurs on up to 20 south-facing slopes, with total populations numbering up to thousands,September 1991 to September 2003, RHN).
G. amarella is rarer and more definitely lime-tolerant or calcicole than the closely related G.campestris (Field Gentian). However, both species occur in very similar shallow, leached,heavily grazed, species-rich, heathy limestone grassland situations. The two species overlapconsiderably in terms of their biology and ecology and sometimes they occur in mixed populations. Todate, they have only been found together at five subsites in Fermanagh, around Monawilkin Lough on 13September 2009. On 28 September 1991, thousands of G. amarella plants were observed on the SEslopes of Monawilkin, while in September 2009 the total number on all slopes around the lake was just 85plants (RHN & HJN).
At the Rahallan site on Belmore, G. amarella is accompanied by Neotinea maculata(Dense-flowered Orchid), another extremely rare species in NI. Other typical lime-tolerant species whichregularly associate with G. amarella include Ophioglossum vulgatum (Adder's-tongue),Arabis hirsuta (Hairy Rock-cress) and several further orchids, for example, Listera ovata(Common Twayblade), Gymnadenia conopsea (Chalk Fragrant-orchid), Plantantherabifolia (Lesser Butterfly-orchid) and Orchis mascula (Early-purple Orchid).
Flowering reproduction
G. amarella is completely dependent on seed reproduction for increase, dispersal and overwinteringsurvival. The throat of the tubular flowers opens and closes with surprising speed, depending on changesin temperature. Within 20 seconds of clouds rolling away and the sun emerging, the flower may open(Lousley 1969). They are said to be pollinated by bumble-bees, which are sufficiently strong and able toreach beyond the fringe of stiff hairs guarding the mouth of the corolla (Clapham et al. 1987).Soon after fruiting the plants die (Pritchard 1959). Measurements of reproductive capacity made bySalisbury (1942) found the seed output was between 800 and 900 per plant. The seeds are small and, incommon with other members of the family, require a mycorrhizal partner in order to germinate.
Field observations show that G. amarella populations are sporadic, varying in numbers and locationand this, together with experimental attempts to germinate seed, suggests a long period of seed dormancyexists (Pritchard 1959). An example of population fluctuation in Fermanagh was observed at Rahallen,where in 1996 Ian Rippey found 'a few', while in 2003, after a good warm summer, Robert and HannahNorthridge estimated 2,000 plants at the same site.
As in G. campestris, dispersal of the small, lightweight seed from the fruit capsule is mostprobably achieved by wind, especially during the winter when the surrounding vegetation dies downsomewhat. Horses, cattle, rabbits and undoubtedly other wild grazing animals, eat the fruiting plantsand disperse gentian seed with their dung (Ridley 1930).
The loss of rabbit populations since the 1950s is one feature that has destroyed previously suitablesites that supported Gentianella species on downs and dunes across B & I. This is largely dueto the subsequent invasion of herb-rich grass heath by species such as Arrhenatherum elatius(False Oat-grass) and Brachypodium pinnatum (Tor Grass). Tall perennial grasses such asthese rapidly oust gentians that are both short in stature and short-lived, and that need to seed everyyear in order to survive. Gentians are not ecologically flexible, and have suffered local extinctions inmany sites as a result (Pritchard 1972).
Toxicity and herbivory
The presence of somewhat toxic bitter glycoside substances in the plant tissues does not greatly detergrazing animals (Ridley 1930). Previous to their decline from myxomatosis, rabbits frequently nibbledyoung gentian shoots to ground level, which would then react by re-growing into small, very bushy,tufted plants (Lousley 1969).
British and Irish occurrence
G. amarella remains widespread across lowland Britain in suitable basic and calcareous soilconditions. It reaches its highest altitude of 750 m at Knock Fell in Westmorland (VC 69) and is muchmore coastal in distribution in Wales and Scotland than is the case in England (R.D. Porley, in: Prestonet al. 2002). In Ireland, Autumn Gentian is less frequent and more thinly scattered than in mostof England, at least. It stretches across country from Co Dublin to Cos Galway and Clare and thennorthwards from there to Sligo, W Donegal and Fermanagh in a very disjunct manner (New Atlas). The IrishCensus Catalogue lists the species as having been recorded at least once from 31 of the 40 VCs (Scannell& Synnott 1987), but, as is also the case in Britain, there have been numerous populationextinctions due to changes in land management in the last 50 or more years, so that this no longeraccurately summarises the present distribution.
European and world occurrence
The species s.l. is widely distributed in Eurasia and N America and is considered circumpolarboreo-temperate in its phytogeographic distribution. Subsp. amarella occurs across N & CEurope to E Ukraine, the Caucasus and the Yenisei region and W & C Siberia, but is scarce and verydisjunct in S Europe and, apart from SE France, completely absent in the Mediterranean region (Hultén& Fries 1986, Map 1505; Clapham et al. 1987).
Uses
Like related members of the Gentianaceae, G. amarella has been substituted in B & I at timesfor the more famous Yellow Gentian (Gentiana lutea) in herbal medicine as a tonic, to cleanse theblood and kidneys, and for indigestion and colic, jaundice, wounds, sores and rheumatism (Grieve 1931;Grigson 1955, 1987; Darwin 1996).
Names
The name 'Gentianella' is a diminutive of 'Gentian', which is a name in Pliny of a plant called afterGentius, a 2nd century Illyrian king, who is reputed to have discovered the medicinal use of Gentianalutea (Yellow Gentian) (Gilbert-Carter 1964; Hyam & Pankhurst 1995). The species name'amarella' is the feminine diminutive of the Latin 'amarus', meaning 'bitter' (Gilbert-Carter 1964).
The English common name 'Felwort' originally referred to the important imported medicinal herb,Gentiana lutea, but the name became transferred to the related native herb with similar medicinalproperties. The name is derived from the Old English or Anglo-Saxon 'feldwyrt', meaning 'field plant' or'field-wort'. There may possibly have been a mistaken connection of the Old English or Anglo-Saxon'feld', with the Latin, 'fel', meaning, 'gall', referring to the bitter taste of the plant (Prior 1879;Grigson 1955, 1987; Grigson 1974). In Shetland, G. amarella has the interesting and unusual name'Dead Man's Mittens', because the half-open flower buds are likened to livid finger-nails protrudingfrom the turf (Grigson 1955, 1987).
Threats
Rarity is always a threat to long term survival and this species requires statutory protection and activemonitoring in NI.
Introduction, archaeophyte, a rare garden escape.
30 May 1990; Northridge, R.H.; Nutfield Estate.
April to September.
This very familiar, usually blue-flowered, semi-woody, evergreen garden herb spreads vegetatively by itscreeping or arching stems rooting as the tip touches the soil. It thus 'frog-hops' along the ground,making it a very effective garden ground-cover species, especially when grown under damp, shadyconditions. It can tolerate dry soils and steep slopes, especially when it becomes established, butfoliage can become burnt if exposed to full sun. It tolerates a wide range of soils, but is unsuitablefor heavy, very wet or compacted ground. Although it regularly flowers, V. minor does not oftenform fruit (An Irish Flora 1996). Flower colour can vary from blue to purple, lavender or whiteand there are variegated leaf varieties also.
Typically, V. minor is found near houses, frequently along woodland margins or hedgerows, or onroadsides where it may have been deposited with garden waste, or as plant fragments capable ofestablishment and survival.
In Fermanagh, there are records of it from seven tetrads, thinly scattered in the E & S of the VC.Oddly enough, as the tetrad map indicates, almost all of the records are close to the county boundary!At Nutfield, near Brookeborough, the plant is very well established in woodland, which is quite a rarecirc*mstance in Ireland.
Both species of garden periwinkles, V. minor and V. major (Greater Periwinkle), wereintroduced to B & I, probably from S, W & C Europe and the Mediterranean region. V. minoris believed to have been in British gardens by the year 995 making it an archaeophyte (Harvey 1981).V. major was introduced considerably later, around 1550 (Ellis 1993). Of the two, V. minoris frequently preferred ground-cover since it is smaller, lower growing, more shade-tolerant and lessinvasive than V. major (Stace & Crawley 2015). Both Vinca species contain toxicalkaloids that can poison pets. Since it is the most widely cultivated of the two, V. minor moreoften escapes the garden and is more frequently recorded (T.D. Dines, in: Preston et al. 2002).
Both Periwinkle species are much more frequently recorded in the warmer southern half of Britain, whilein Ireland they are only occasionally listed by botanists (BSBI Atlases; FNEI 3; Floraof Co Dublin; New Atlas). Nevertheless, the Cen Cat Fl Ir 2 lists V. minoras having been found in 19 of the 40 Irish VCs, without including Fermanagh.
Threats
None.
Introduction, possibly an archaeophyte, but only a very rare casual. As a native, it appears Eurasiantemperate, but it is also very widely naturalised.
29 July 1986; Corbett, P., Austin, L.W. & Wolfe-Murphy, S.A.; waste ground on S section of Corrardshore, Upper Lough Erne.
July and August.
Growth form and preferred habitats
S. nigrum is a white-flowered, rather hairy, much branched annual, or short-lived, monocarpicperennial with fibrous roots and oval leaves. It is sometimes quite woody at the base. It is a rare,occasionally abundant, casual alien in Ireland, where it most often occurs in cultivated ground,gardens, waste ground, at ports and among suburban planted shrubs, along with which it may have beenintroduced (Cat Alien Pl Ir). It mainly occurs in southern VCs (Cen Cat Fl Ir 2). BlackNightshade is also called Garden Nightshade, since it is a notorious garden weed of fertile, nitrogen-and phosphorus-rich soil, manure heaps and on waste ground where there is organic enrichment (Garrard& Streeter 1983). It prefers moist environments and thrives in regions with low rainfall only wherethere is irrigation (Holm et al. 1977). Although often low and spreading (10-30 cm), S.nigrum can grow 60-100 cm tall. In flower, it bears small, ivory-white, star-like flowers inclusters of 3-12 (but usually numbers 7-9) (Salisbury 1964; Sell & Murrell 2009).
In terms of established strategy, S. nigrum is classed as an R/CR, ie intermediate between Ruderaland Competitive Ruderal (Grime et al. 1988, 2007).
Variation
S. nigrum is a very variable species both in terms of physical or geographical forms and itsphysiology. So much variation is involved that it is probably best to consider S. nigrum as acomplex of subspecies, varieties and forms with varying degrees of similarity, rather than as a single,clearly defined species (Cooper & Johnson 1998). In the critical Flora of B & I, Sell &Murrell (2009) describe two subspecies (subsp. nigrum and subsp. schultesii (Opiz)Wessely), plus three varieties of the former.
When taken in the broad sense as the S. nigrum species complex, the plant has an almost world-widedistribution as an arable weed. It is reported as an important weed in 61 countries and 37 crops (Holmet al. 1977). Even when considered in the more restricted Eurasian form of the species, it ismorphologically and physiologically variable in different locations and between populations, some ofwhich are in the process of developing resistance to herbicides (Cooper & Johnson 1998).
Fermanagh occurrence
This is a very rare plant in Fermanagh, there being records from a total of only five sites in separatetetrads. The details of the earliest two Fermanagh records, both from around Upper Lough Erne, arefirstly as given above and, secondly, waste ground at Killynubber Lough, 15 August 1986, L.W. Austin& S.A. Wolfe-Murphy. The recorders of the first Fermanagh record, however, questioned its occurrenceat the time. Consequently, both it and the subsequent Killynubber record require further verification.Regrettably, since there are no voucher specimens for either discovery, they fail to fit the BSBIcriteria for first and second county records and, strictly speaking, they are unacceptable. Onthe other hand, Black Nightshade is easily recognised, its stem being entirely herbaceous, up to 60 cmtall, the leaves oval, sometimes deeply toothed and the flowers are familiar and potato-like. Thus it isvery likely that the two Fermanagh identifications were perfectly correct.
The details of the remaining three sites are: in hedge at Knockninny Quarry, 29 August 2004, I. Rippey;Killymackan Lough ASSI, 24 June 2007, ENSIS Lake Survey; Galloon Td, Upper Lough Erne, 17 August 2006,ENSIS Lake Survey. Again, the database does not mention vouchers for these sites.
Fossil history, species status and occurrence in Britain
Despite its weedy behaviour and the wide range of variation mentioned above, appropriate fossils (seedfrom the Cromerian and pollen from the Flandrian interglacial periods) prove S. nigrum isundoubtedly of early native status in SE Britain, if not too likely so in the rest of Britain, or inIreland (Godwin 1975). Having said this, there are two mediaeval fossil records from Ireland, and thetwo earliest Flandrian records in England lie near the presumed northern limit of its range (Godwin1975). Later fossil records from the Roman and mediaeval periods confirm S. nigrum's associationwith cultivation and with other weedy species (Godwin 1975).
S. nigrum is widespread and common in England and parts of Wales, S & E of a line from theHumber to the Severn, but it is scarce elsewhere in Wales and in SW England and absent or purely acasual alien in both Scotland and Ireland (Garrard & Streeter 1983; T.D. Dines, in: Preston etal. 2002).
Irish occurrence
In NI, until recent years, S. nigrum had only been recorded on a total of five occasions in CosDown (H38) and Antrim (H39) during the pre-1925 period (FNEI 3). With the increased recordingeffort associated with a flora survey of urban Belfast in the 1993-5 period and the subsequentwide-ranging BSBI New Atlas 2000 survey, a total of twelve more records were discovered; threerecords each in Cos Antrim, Down and Tyrone (H36), plus one in Co Armagh (H37) and the two in Fermanagh(H33) already mentioned. Since the New Atlas was published in 2002, two additional Tyrone recordshave been listed by McNeill (2010), one of them from a maize crop. Interestingly, S.nigrum was discovered for the first time between September 1998 and 2000 in Co Cavan (H30)(which shares a border with Fermanagh), at no less than five different sites (Reilly 2001).
Elsewhere in the RoI, S. nigrum is very thinly and widely scattered, with a cluster of hectadsaround Dublin (H21) and Wexford (H12) (New Atlas). In Co Waterford (H6), S. nigrum appearsto be increasing in recent decades; although still a casual ruderal of open habitats, it also appears asan arable weed of sugar beet, maize and carrot crops (Green 2008).
Flowering reproduction
S. nigrum relies entirely on seed reproduction for increase, dispersal and survival. Seedsgerminate in the spring, from early May onwards, reaching a peak emergence in late May or June (Roberts& Lockett 1978). Plants reach flowering condition from July to September, the regular(actinomorphic), perfect (bisexual), star-like flowers, 10-14 mm in diameter have a calyx of five erect,fused, oblong sepals, not enlarging in fruit. The five ivory-white, fused petals are about twice as longas the calyx, the lobes spreading at first, then rolling back on themselves (ie revolute) as the flowersage. The long, bright yellow anthers are inserted (ie attached) on very short filaments to the corollatube, so that they poke out of the corolla. They lie close to one another, forming a cone around thelong, well exerted, solitary style and stigma.
The anthers open through pores at their tips to release the pollen that is the main flower foodattractant, since the very little nectar is produced. Insect visitors (mainly bees) cling to the cone ofanthers and by vibrating their wings rapidly, they draw pollen out through the anther pores onto theirhairy bodies. Since the style protrudes well beyond the anther cone, the stigma tends to becross-pollinated (Proctor & Yeo 1973; Hickey & King 1981).
Fruit and seed production
Fruits ripen in succession from September to November if the weather remains mild. The plant is killed,however, by the first frosts. The plant produces globular fleshy berries, green at first, then turningshiny, black when ripe, 5-13 mm diameter, with the slightly enlarged sepals attached, curving backwards.The fruits are more watery than fleshy and they do not persist for long if not eaten: they tend to burstopen and then dry out (Snow & Snow 1988). Each fruit contains 40-50 flattened seeds which can passunharmed through the digestive tract of farm stock and other animals. The average number of fruit perplant measured by Salisbury (1942) was 238, so it is very possible that an average-sized individualplant could produce around 9,500 seeds. In the same study some really huge plants were calculatedcapable of producing up to 130,000 seeds (Salisbury 1942). A maximum seed number of 178,000 per plantwas published by Holm et al. 1977).
Seed dispersal
Birds are undoubtedly one of the main vectors of S. nigrum seed. Studies using marked berriesshowed that birds took ripe berries during the second half of September and in October. Blackbirds andRobins are the species most involved, with Starlings the third possibility (Snow & Snow 1988).Observation suggests that storm rain-wash along paths and bare ground is another short-range method ofS. nigrum dispersal (Ridley 1930). Probably man has become the most significant vector of all,since in historic times, S. nigrum has been introduced with crop seed worldwide and has becomeone of the most noxious alien weeds in many countries (Ridley 1930; Holm et al. 1977).
Seed longevity
Establishment after dispersal is assisted by the relatively long survival in soil of undisturbed buriedseed. Roberts & Lockett (1978) reported survival rates akin to those of Capsellabursa-pastoris (Shepherd's-purse), Chenopodium album (Fat-hen) and Poa annua(Annual Meadow-grass); in the long-running Duvel experiment (Toole & Brown 1946), seeds placed atdepths of 20, 56 and 107 cm showed little loss of viability after 39 years.
European and world occurrence
The black-fruited, rarely green-fruited S. nigrum s.s. is believed to have originated in S & CEurope and is distributed throughout most of Eurasia including N Africa and stretches eastwards to theYenisej River and C Asia. It has been spread with agriculture and is commonly introduced in E Asia andfrom Nova Scotia to Florida and westwards in America, although mostly along the Atlantic seaboard(Hultén 1971, Map 258; Hultén & Fries 1986, Map 1622; Clapham et al. 1987).
The species s.l. (including S. americanum Mill. and other different but very closely related taxa)is almost cosmopolitan, occurring throughout Europe, Asia, N, C & S America and the tropics of boththe old and new world. In many areas of Europe, Asia and N America, both native and introduced taxawithin the S. nigrum complex are present. The distribution of the S. nigrum complex isdiscontinuous circumpolar and it has been introduced to many isolated islands around the world,including Cape Verde Islands, Réunion, Madagascar, Australia, New Zealand, Tasmania and Hawaii (Hultén1971; Hultén & Fries 1986).
Toxicity
All parts of Black Nightshade plants, but particularly the green unripe berries contain amounts of thepoisonous steroidal glycoalkaloid, Solanine. Nitrates and nitrites also occur in variable amounts andcan contribute to the toxic effects of the plant. Ripe, shiny black berries are the least poisonousparts of the plant, the poisonous principle being chiefly associated with all green parts (Grieve 1931;Cooper & Johnson 1998). Although the amount of toxin varies with growing conditions and plant age,Black Nightshade should be considered dangerously poisonous to all farm stock and to humans.Cattle will not eat the plant and sheep rarely do (Grieve 1931).
Names
The name 'Solanum' is a classical name, first given by the Roman naturalist Pliny to one of theNightshade species. It possibly may be derived from the Latin 'solamen', meaning a solace, referring toits supposed medicinal virtues (Johnson & Smith 1931). On account of its shiny black berries, S.nigrum was called by older herbalists 'Petty Morel' to distinguish it from its even moredangerously poisonous relative, 'Deadly Nightshade' (Atropa belladonna), often known as 'GreatMorel'. The dried whole plant or fresh leaves are (or were) used in herbal medicine. Black Nightshadehas narcotic properties and is a strong sudorific (ie it causes sweating).
The chief uses are in treating skin diseases, for instance, dealing with obstinate skin eruptions, burnsand ulcers using bruised fresh leaves externally to ease pain and abate inflammation. However the actionis variable and even the herbalists considered it a rather dangerous remedy, except when used in smalldoses. There are reports of the leaves being eaten like spinach and of it being a famine food, but thisshould never be countenanced (Grieve 1931).
Threats
None.
Native, locally frequent to quite common. Eurasian southern-temperate.
1860; Smith, T.O.; Co Fermanagh.
April to November.
Growth form and preferred habitats
A climbing or scrambling rhizomatous perennial liana with stems up to 6 m or more in length, but oftenonly half this, and woody at least at the base. The alternate, stalked leaves are of two types: lowerleaves are deeply 3-lobed, a large ovate middle and two small side lobes; upper leaves are simple,ovate, cordate at their base, with margins entire. In summer, the stems bear star-like flowers of abright purple hue with a projecting and strongly contrasting yellow anther cone, making Bittersweet orWoody Nightshade quite conspicuous, distinctive and readily recorded (McClintock 1965).
Typically, Bittersweet occurs in shaded, lowland, marshy ground, including tall herb fens on lakeshores,along riverbanks, stream-sides, plus on damper woodland margins and in hedgerows, thickets and wasteground. S. dulcamara can grow under a rather wide range of lowland, environmental conditions,including in relatively dry and fully exposed sites, but it thrives in damp to wet, semi-shaded, fertilehabitats. Under these circ*mstances it can clamber over and occasionally dominate other herbaceousplants in woodland (Pegtel 1985). A very distinct succulent, prostrate form of the species, var.marinum, grows on maritime shingle (Sell & Murrell 2009).
The established strategy of the species is categorized as intermediate between the generalist C-S-R (ieCompetitor/Stress-tolerator/Ruderal) and Competitor. In the herb layer, under woodland shade, it canclamber over and dominate other herbs, but in unshaded habitats it may itself be suppressed bytaller-growing herbaceous species (Grime et al. 1988, 2007).
Vegetative reproduction
The plant is long-lived, individuals persisting for c 20 years (Salisbury 1942). It is therefore capableof forming large clonal patches, since when stems touch the ground, or when they fragment and disperse,they can readily root and establish new plants. Fresh shoots can also arise directly from the roots andthe creeping rhizome (Salisbury 1964). The frequency of S. dulcamara along stream and river banksmay well reflect the ability of the plant to fragment and re-establish new colonies downstream (Grimeet al. 1988, 2007).
Flowering reproduction
Flowers are borne in a much-branched, flat-topped inflorescence generally containing around 15 flowers,although there can be up to 25 or more (Tutin et al. 1972). Technically, the inflorescence typeis described as a corymbose dichasial cyme! The star-like corolla is 15 mm in diameter, dark purple(rarely white), with five lobes bent backwards, each petal having two greenish spots at the base. S.dulcamara flowers all summer from June to September and fruits from August onwards until thefirst frost.
The flowers offer visitors plenty of protein and mineral-rich pollen but no nectar and they are adaptedto 'buzz pollination' by bumblebees. The bees hang on to the anther cone and vibrate it with theirthoracic muscles, releasing the very small pollen grains through apical pores on the anthers (Proctoret al. 1996).
The fruit cluster consists of 6-8 mm, many-seeded, oval, green berries that become bright shiny red whenmature. The fruit stalks recurve as the berries develop, finally hanging downwards. Reproductiveperformance data were calculated by Salisbury (1942, p. 79) from a small sample of eight plants, but hewas able to show that the seed output is potentially enormous: the average plant produced around 1,400berries, each typically containing 38 kidney-shaped seeds, so that the annual seed total per plant liessomewhere between 40,000 and 70,000!
Berries are produced from late July onwards and while those formed in relatively dry, exposed conditionsmay shrivel after only a month, plants in more typical damp, sheltered habitats can carry fruit in goodcondition until December, long after the leaves have fallen. A range of birds eat the berries anddisperse the seed. Chief amongst these are Blackcaps, Blackbirds, Song Thrush, Robin and Starling. Acertain amount of seed predation has been observed, the main culprits being Bullfinches (Snow & Snow1988).
Seed requires low temperature after-ripening, or a period of natural weathering involving fluctuatingtemperature. Germination occurs in the spring (Pegtel 1985). Little or no persistent seed bank exists(Grime et al. 1981; Roberts 1986). Despite the considerable range of variation found in thespecies and its wide ecological tolerance, Pegtel (1985) could not detect genetic differences in thegermination of seed from contrasting habitats and the case for delimiting ecotypes does not seemconvincing.
Fermanagh occurrence
In Fermanagh, the plant is really rather local, but very frequent where it does occur, particularlyaround both parts of Lough Erne as the tetrad map indicates. It has been recorded in 119 tetrads,representing 22.5% of those in the VC. All but seven of the tetrads contain post-1975 records. A totalof 90 tetrads are concentrated in and around Lough Erne.
British and Irish occurrence
In Britain, S. dulcamara is widespread and common in England and Wales, but is absent fromland lying above 310 m (ie around 1,000 ft.). It becomes rare and more or less coastal in N England andScotland. In Ireland, S. dulcamara is much more frequent in the N & E of the island,stretching as far south as Dublin. However, it becomes scarce to rare in the far W & SW of the RoI(Rich & Woodruff 1990; An Irish Flora 1996; New Atlas).
There did not appear to be any appreciable change in the presence of S. dulcamara from theevidence contained in the BSBI Atlas 2000 survey (T.D. Dines, in: Preston et al. 2002),but the subsequent second monitoring survey also carried out by the BSBI in Great Britain only,Change in the British Flora 1987-2004 found a definite and statistically significant decline witha 'Change Factor' measuring -11 (Braithwaite et al. 2006).
European and world occurrence
S. dulcamara belongs to the Eurasian southern-temperate phytogeographic element and is widespreadthroughout most of Europe and Asia except the extreme north with the native range extending into NAfrica and the Middle East. It has been introduced to N America (Hultén & Fries 1986, Map 1624;Grime et al. 1988, 2007).
Toxicity
All parts of the Bittersweet plant contain a mixture of steroidal glycol-alkaloids often referred tocollectively as 'solanine'. The alkaloids are degraded during the maturation of the fleshy fruit. It issuggested that this makes the berries more attractive food for birds and other animals that eat them andtransport the seeds in their gut. The ripe berries should not, however, be regarded as safe to consumeas they have been known to cause poisoning (Cooper & Johnson 1998).
In the 1960s and early 1970s, poultry and sheep are known to have died after grazing the plant andchildren who ate the attractive-looking berries were very sick, although thankfully only one of themdied (Cooper & Johnson 1998)
Uses
S. dulcamara has a history of use in herbal medicine dating back to classical times and there arefew ailments and complaints for which it has not at some time been recommended. The main uses were intreating rheumatism, skin diseases and as a purgative. Gerard (English), Boerhaave (Dutch) and Linnaeus(Swedish) all spoke highly of its medicinal virtue and S. dulcamara continued to feature in theBritish Pharmacopoeia until 1907 for skin complaints, after which it was dropped from modern pharmacy(Grieve 1931).
N.B. No part of the plant should ever be eaten, as even the ripe black berries may contain the verypoisonous alkaloid, solanine.
Names
The name 'Bittersweet' was coined by the 16th century English herbalist, William Turner, who translatedthe medieval Latin names for the plant, 'dulcamarum', 'dulcis amara', or 'amara dulcis' (dulcis, sweetand amarus, bitter), referring to the woody stem bark, which if tasted is at first sweet and then turnsvery bitter in the mouth (Britten & Holland 1886; Grigson 1974). The modern botanical species name'dulcamara' given by Linnaeus refers to the taste change more correctly. Do not attempt to test thistaste change, for fear of dangerous ill effects! The medieval herbalists also referred to S.dulcamara as 'Woody Nightshade' to distinguish it from its relative, 'Deadly Nightshade'(Atropa belladonna).
There are numerous alternative, local English common names. Grigson (1955, 1987) lists a total of 22, sixof which refer to the plant's poisonous nature. Numerous other names refer to supposed magicalproperties that garlands of the plant, on account of its red berries, offered against witches and suddenillness.
Threats
None.
Introduction, neophyte, a very rare casual escape or discard from cultivation.
21 August 1986; Wolfe-Murphy, S.A.; waste ground, E shore of Lough Head.
This extremely familiar vegetable grows occasionally on waste ground and rubbish tips from potato tubersin transported soil, discarded agricultural waste, garden rubbish or domestic refuse, but the high levelof disturbance usual in such sites means that it does not persist and, therefore, it is regarded asmerely casual. The tuberous perennial nature of the plant, however, must enable it to reproduceeffectively in more stable situations.
There are only two records in the Fermanagh Flora Database for S. tuberosum, but while they areboth from waste ground, the situations appear rather more semi-natural than for most records of thisspecies elsewhere. The details of the first record are given above, the details of the second site whereit was found are: Drumbuleen, S of Irvinestown, 16 August 1994, RHN.
The current author (RSF) and his H33 joint BSBI Recorder colleague, RHN, consider it very probable that,as is the case with other crop casuals, S. tuberosum has in the past often been simply overlookedor ignored by field botanists in Fermanagh and it is, therefore, under-recorded. It could be argued thatit is insignificant, in conservation terms at least, and so why bother with recording it!
As if to prove that this is the case, the Cen Cat Fl Ir 2 completely ignores S. tuberosum.The Cat Alien Pl Ir described S. tuberosum as a, "fairly frequent casual".However, it lists only eleven VCs with records – Fermanagh not being included. Altogether, Reynolds, theauthor of A catalogue of alien plants in Ireland, gives details of very few occurrences, and noneof the sites mentioned could be regarded as being even semi-natural.
The accounts in the two inner-city Floras of Dublin and Belfast showed the potato is oftenestablished from domestic or garden rubbish, typically being found near habitation. It was, forinstance, recorded in twelve of the 14 subdivisions of inner Dublin (Wyse Jackson et al. 1984),while in Belfast it was found in 30 of the 76 one km squares in the city survey (Beesley & Wilde1997).
The Flora of County Dublin (Doogue et al. 1998) reckoned that, "some clumps (ofpotato) appear to persist for a few years". Several editions of the Flora of the NE ofIreland described potato as a, "frequent casual escape, never permanent", an overallposition the current author and RHN would happily endorse (Praeger & Megaw 1938; Hackney etal. 1992). The New Atlas shows that on a B & I scale, records exist in 17.5% ofhectads across the two islands (Preston et al. 2002).
Threats
None.
Native, rare. Eurosiberian southern-temperate, but widely naturalised and now circumpolar.
1872; Brenan, Rev. S.A.; Co Fermanagh.
March to October.
Growth form and preferred habitats
Convolvulus has small bracteoles, borne some distance below the calyx, which readily distinguishesit from all Calystegia species. The funnel-shaped flowers of Convolvulus arvensis are muchsmaller than those of any Calystegia species we have in Ireland (only 15-40 mm in diameter) andtheir colour varies from white to deep rose-pink, or striped with both colours. Leaves are glabrous,alternate, stalked and arrow-shaped (ie hastate) (An Irish Flora 1996; New Flora of the BI1997).
Like the several Calystegia species in B & I, C. arvensis is a deeply-penetrating,slender, extensively-branching, rhizomatous, perennial of warm, light, fertile, less acidic soils (pHgreater than 4.5), which dies down completely in the autumn. Stems are numerous and are eitherprocumbent, or erect and twining on support provided by adjacent plant stems or slender inanimatestructures, usually reaching a height of around 60 cm. It never twines around anything of bulkydimensions, such as for instance, gateposts (Grieve 1931, p. 220). Twining is anti-clockwise, the stemtaking about two hours per revolution, and often strangling its support. C. arvensis is mostabundant when colonising recently disturbed sites, particularly in cultivated ground and, since arableagriculture has almost completely ceased in Fermanagh, locally this now really means plots and plants ingardens (Melderis & Bangerter 1955; Grime et al. 1988, 2007).
The typical habitat of Field Bindweed is lowland, disturbed waste or cultivated ground and it isfrequently found along less disturbed, more-or-less unmanaged linear habitats, such as hedgerows andalong lakeshores, roads and railways, a pattern which can just be detected in its scarce or rareFermanagh occurrences. It demonstrates a strong preference for fertile soils and is absent from veryinfertile ones (Grime et al. 1988, 2007).
Roots and rhizome can penetrate very deeply (down to 9 m), twining like the aerial stems around the rootsof other garden plants and making eradication of the species very difficult or impossible. Theunderground parts can spread extremely rapidly in cultivated soil, infecting almost 30 m2 ina single season (Salisbury 1964). Since it roots so deeply, C. arvensis is absent from wet groundand it readily survives drought when growing on cliffs and rock outcrops. It is sufficiently rare inBritain above c 200 m that a temperature limitation is suspected (Grime et al. 1988, 2007).
The plant can regrow from the tiniest fragment of root or rhizome left in the soil, making it, as manygardeners know, almost impossible to eradicate and very difficult to control. The established strategyof the species is categorised as Competitive-ruderal. It is absent from regularly grazed or mown groundand also from situations providing heavy shade (Grime et al. 1988, 2007).
Variation
A very variable species, no fewer than eleven forma have been described in C. arvensis,based largely on colour variants of the corolla, anthers and filaments (Sell & Murrell 2009).Corolla colour appears to be constant when plants are grown from seed (Proctor et al. 1996). InEurope, ecotypes based on leaf shape and flower colour have been recognised (Weaver & Riley 1982).
Fermanagh occurrence
There are records from a total of ten tetrads in the Fermanagh Flora Database, eight of which are ofpost-1975 date. The sites are thinly scattered in the lowlands and they range from roadside hedgerows,to waste ground and gardens.
Flowering reproduction
C. arvensis is a polycarpic perennial, the decorative, funnel-shaped flowers of which are producedin axillary cymes of 1-4 flowers from June to September. Each blossom last just a single day, thecorolla unfolding around 7 a.m. and closing about 10 p.m. or earlier in bad weather. The faintvanilla-like perfume of the flower attracts bees and a variety of flies including hoverflies whichcollect a nectar reward when transferring pollen. Nectar is secreted at the base of the ovary, but canonly be reached through five narrow passages between the broad bases of the stamens. This is referred toas a 'revolver flower', since the insect is delayed inside the funnel-shaped corolla, visiting theseparate 'chambers', to reach all of the nectar reward (Proctor et al. 1996). It is believed thatself-pollination may also take place.
Irrespective of the pollination mechanism, seed set, which takes place from August to October, isprobably restricted in B & I by low temperatures, only happening in hot summers (Thurston 1960). Thefruit capsule is small, about 6 mm in diameter, ovoid and contains between 1-4 relatively large, 3.5 mm,minutely pitted seeds. The seeds can survive burial for at least six years and may indeed be aslong-lived as those of Calystegia sepium subsp. sepium (Salisbury1964), a view supported by evidence presented in the soil seed bank survey of NW Europe (Thompson etal. 1997).
Seed dispersal
Seed of C. arvensis can float for about twelve hours and can therefore be water dispersed alongstreams or in rainwater runoff. Alternatively, they may be eaten by birds and carried in their crops(Ridley 1930). Nowadays, man probably is another major agent of dispersal however, with seed and smallfragments of root or rhizome being all that is necessary to transport the species in soil, on the rootsof other plants, or in mud on boots or on vehicles.
Species management
The plant is very difficult to control, even with modern herbicide sprays, since the hairless leaves andstems are rather waxy and difficult to wet. The plants also show ecotypic resistance to 2,4-D herbicide.While the tops may be killed by the chemical spray, portions of the underground system usually manage tosurvive. The best level of control (actual eradication is almost impossible) is achieved byspraying just prior to the plant flowering (Muzik 1970).
British and Irish occurrence
In Britain, there is fossil pollen evidence from the Flandrian zone V proving C. arvensis isnative. Later fossil records from zone VIII show the species present in Anglo-Saxon and Norman levelsamongst other weeds of arable ground (Godwin 1975). Field Bindweed, which probably originated in theMediterranean region, is very common in Britain throughout the southern lowlands, but it becomes muchless frequent and more coastal N of a line between Hartlepool and Heysham.
In Ireland, C. arvensis is very widespread and scattered and while it has been recorded in everyIrish VC (Cen Cat Fl Ir 2), there is a greater prevalence of it near coasts and it isnoticeably more frequent in the eastern half of the island (New Atlas).
European and world occurrence
Thought to have originated in the Mediterranean region, C. arvensis is widespread in temperateEurasia, including N Africa, the Azores and Macronesia, but it is absent in more northerly boreal parts.In phytogeographical terms, it is described as Eurosiberian southern-temperate. A very variable species,C. arvensis has spread widely as a weed of arable cultivation to C & S Africa, Ethiopia, SArabia, N & S America, Japan, S Australia and New Zealand, so that its distribution is nowdiscontinuously Circumpolar (Hultén & Fries 1986, Map 1538).
Names
The genus name 'Convolvulus' is from the Latin 'convolvo', to entwine or roll around and it firstappeared in Pliny (Gilbert-Carter 1964; Stearn 1992). The Latin specific epithet 'arvensis' means 'ofploughed fields', or 'of cultivated fields' ('arvum' (solum) translates as 'arable land')(Gilbert-Carter 1964; Gledhill 1985).
There are a large number of local English common names – around 30 in Grigson (1955, 1987), many sharedwith Calystegia sepium (Hedge Bindweed), for example, 'Bellbind' and 'Old Man's Nightcap' (theold man being the Devil). Due to its twining habit, the plant has acquired names such as 'Bearbind','Bedbind', 'Wheatwind', 'Withwind', 'Cornbine' or 'Cornbind' and 'Ropebind', but the current authorlikes 'Devil's Guts' and 'Hellweed' best, since they conjure up the gardeners' deep-seated hatred ofthis pernicious, extremely persistent weed.
Threats
None.
Native and/or a possible archaeophyte, common. Circumpolar temperate, very widespread in bothhemispheres.
1866-72; Stewart, S.A.; Enniskillen Town.
April to November.
Growth form and preferred habitats
Hedge Bindweed is a climbing, twining, tenacious perennial with large, heart-shaped leaves, thoroughlyloathed by gardeners for its persistent, deeply penetrating roots and white rhizome that tangles itselfamong the roots of more desirable decorative subjects in the tended plot, always proving extremelydifficult to eradicate. Herbicides such as 2,4-D offer some hope, since the species is moderatelysusceptible to its effect (Salisbury 1964).
The species, as currently understood, was united with Calystegia silvatica until the 1930s.Praeger (1934, 1936) seems to have been the first botanist in the British Isles to distinguish C.silvatica at Lough Gill, Co Sligo (H28) and the species group was further delimited and C.pulchra recognised and named in 1960 (Brummit & Heywood 1960).
Apart from the garden, typical habitats of the species range from hedges by roads, streams and rivers,waste ground, old quarries and in scrub woodland, both terrestrial and wetland fen carr. It also climbs(always twisting in an anti-clockwise direction) in reed-beds and on other tall vegetation in wetditches, marshes and fens. It is in the latter, wetter habitats mentioned, that C. sepium subsp. sepium appears to be most truly indigenous in B & I(FNEI 2).
Under favourable growing conditions, the twining motion can be very rapid, the climbing stem completing arevolution in one to two hours. If no support is sufficiently close to climb upon, the Hedge Bindweedstems often twine around one another, producing a thick, rope-like structure that explains the name'Ropewind' (Salisbury 1964). The species is a good indicator of fertile soil conditions and thus itnever occurs in very acidic peat habitats (Hill et al. 1999). Grime et al. (1988, 2007)considered it was restricted to situations where the pH was above 5.5, which is the reaction typical offertile topsoil in NI.
C. sepium subsp. sepium uses either the stout stems of tall herbs,shrubs and hedges, or artificial, man-made structures such as fences to climb and ascend to heightsaround 3 m or so above ground level. The plant can then sometimes form a dense, smothering, curtainingcanopy over supporting vegetation in the latter part of the growing season. Moist, fertile growingconditions and a moderate degree of disturbance appear necessary to sustain a competitive balancebetween Hedge Bindweed and the potentially dominant tall, perennial herbs and shrubs on which it climbs.If the habitat is left undisturbed, shade builds up and C. sepium subsp. sepium tends to decline in vigour. It is then eitherincapable of flowering, or flowering ceases altogether if it becomes subjected to dense shade, as forinstance frequently happens in undisturbed tall-herb fens and reed-beds. In autumn, the aerial shootdies back, giving competing species some months of respite. The established strategy of C. sepiumis categorised as intermediate between Competitor and Competitive-Ruderal (Grime et al. 1988,2007).
Variation
Calystegia sepium s.l. is described as a polymorphic and widespread species that forms an"unusually complicated complex of closely related taxa, taken as species, subspecies, varieties orforms by different authors, who rarely agree on their limitation and geographical range" (Hultén1974, Circumpolar plants II, p. 282). The critical Flora of B & I recognises threesubspecies of C. sepium and two forma of subspecies sepium ; forma sepium with awhite, undivided corolla and forma colorata (Lange) Dörfl., having a corolla with five pink'windows' and undivided (Sell & Murrell 2009). 'Windows' appear to be coloured sections of thecorolla, separated by white rib-like regions associated with the joins between the five fused petals.
Vegetative reproduction
In disturbed sites (and especially in gardens), the plant displays vigorous vegetative regeneration. Evenvery small fragments of rhizome or root prove capable of forming new plants and by this means thespecies can develop spreading clones that gradually build upon the growth achieved in previous years,thanks to its long-term persistence in disturbed ground and a very rapid growth rate in moist, fertile,productive sites, including recently abandoned or neglected corners of gardens (Grime et al.1988, 2007).
Flowering reproduction
Seed reproduction is often secondary in importance to vegetative increase and spread in C. sepiumsubsp. sepium populations The solitary, axillary, bisexual,flowers are borne on flexuous, pedicels 2-21 cm long. The flowers are self-incompatible and when formedin small, single- or few-cloned colonies, developed by vegetative increase, seed set is frequently poor.However, where more numerous separate colonies of differing vegetative origin and strain occur in alocality, seed production is more frequent, as would be expected (Salisbury 1964).
The glamorous, white, funnel-shaped flowers, 3-7 cm in diameter, produced from June to September are thelargest of any species in the flora of B & I. The five united petals that make up the corolla funnelhave the five stamen filament bases fused to them at their base (ie they are epipetalous, meaning 'uponthe petals'). Below the corolla base is the well-developed disc at the base of the superior ovary thatsecretes nectar. The flower is thus a 'revolver', with five narrow openings at the base of the corollathat allow insects to probe and suck up the nectar reward. On entering the corolla an insect bearingpollen from another flower will readily effect cross-pollination as it comes in contact with the 2-lobedstigma which projects beyond the surrounding stamens. Each solitary, open flower typically lasts foraround a single day (but see below), and they open in regular sequence along the stem (Stace 1965).
The large, pure white corolla might suggest the flowers are evolved to attract night-flying (iecrepusclar) pollinators, such as moths. However, the corolla trumpet opens soon after sunrise andusually closes after 12-28 hours, depending upon whether or not pollination has been achieved and alsoupon the prevailing weather conditions – dull, wet or cold days prolong the duration of flower openingto a limited extent. The blossom is scentless, or only faintly scented, although it does offer bothnectar (well concealed at the bottom of the corolla tube) and more freely obtained pollen food sources(Salisbury 1964; Stace 1965). Thus, although daytime visitors are relatively scarce, considering howconspicuous the large flowers are, they must be the important pollinators for the species (Salisbury1964). The daytime visitors are chiefly bees (including bumblebees) and hoverflies, although if theflower has not been pollinated by nightfall, it remains open to visits from moths and other night-activepollinating insects until at least the second sunrise. Some flowers may thus remain open for around 48hours. Flowers never close at night and re-open the next day. Not all insect visitors are pollinators,others may be seeking shelter, or are foragers and flower pests (Stace 1965; Proctor et al.1996).
The fruit is a dehiscent, capsule and its development continues from September into October. The twolarge enveloping bracteoles and five small sepals appear to have no biological significance whenever theflowers are open and functioning, although they undoubtedly conceal and protect the developing flowerwhen in bud, and the young fruit. This gains the species the English common name 'Hooded Bindweed' andalso the genus name (see Names section below) (Grieve 1931; Stace 1965).
Fertile fruits contain from 1-4, large, rounded, dark brown or black seeds, 4-7 mm in diameter. The factthat the seeds are large and heavy limits their ability to disperse, although streamside populations maybe transported by water (Ridley 1930). Were it not for sparse seed production, the species would be evenmore prevalent than is the case, particularly since the seed has great longevity. C. sepium seed,buried at a depth of 105 cm for 16 years, produced 70% germination. After 39 years burial, anothersample achieved 53% germination (Salisbury 1964).
Hybrids
C. sepium readily forms hybrids with two introduced relatives, C. pulchra (Hairy Bindweed)and C. silvatica (Large Bindweed) that colonise hedges and waste places, mainly in urban areasand are widely naturalised in parts of B & I (New Atlas). These hybrids can form at leastpartially fertile clones, obviously complicating plant identification where they occur (Stace 1961,2015). Fortunately these hybrids are rare or very rare in Ireland and none of them have ever beenrecorded in Fermanagh.
Fermanagh occurrence
In Fermanagh, subsp. sepium has been recorded in 191 tetrads, 36.2% of those in the VC. As thetetrad map clearly indicates, it is frequent and widely scattered throughout the more fertile lowlandareas of the county to the N and W of Lough Erne, but rather scarce or absent from most of the groundlying to the SW of the major lakes, including the western plateau uplands.
British and Irish occurrence and status
In the not so distant past, the seed of this form was an occasional contaminant in sown agriculturalmixtures, especially those that included vetches. Together with the strong association the species haswith artificial rather than with natural habitats in B & I, this raises some doubt as to its nativestatus (eg in Ireland, in Connemara and the Burren (H9 & H16) (Flora of Connemara and theBurren)).
Hedge Bindweed is common and is generally regarded as native throughout southern regions of B & I butin Scotland, north of Edinburgh and Glasgow, it becomes much more local and there it could be anaccidental introduction.
There does not appear to be any unequivocal fossil pollen evidence from pre-Neolithic deposits in B &I, so very possibly it is a long-standing accidental introduction here too (Salisbury 1964; Godwin 1975;Webb 1985).
European and world occurrence
C. sepium is also considered native and is common throughout temperate regions of Europe, W Asia,N Africa and N America (Clapham et al. 1962). C. sepium subsp.sepium occurs as a casual introduction in Madeira (where it is very rare) and the species nowoccurs world-wide with many geographic subspecies, certainly in many instances occurring purely as anintroduction (Ridley 1930; Hickman 1993). The species s.l. belongs to the discontinuous circumpolarplants, the map showing it introduced in the southern hemisphere in Africa Cape Province, Java,Australia, New Zealand and temperate S America (Hultén & Fries 1986, Map 1537; Sell & Murrell2009).
Uses
All members of the Convolvulaceae have purgative properties to a greater or lesser degree and C.sepium subsp. sepium is used in both allopathy and hom*oeopathyto treat constipation and fevers (Launert 1981). The acrid roots of C. sepium are much moreviolently purgative than those of C. soldanellla (Sea Bindweed), and despite their drasticaction, they were used by poorer people in place of the milder C. scammonia (Scammony) importedfrom the Levant. The use of C. sepium as a purgative goes right back in herbal literature toancient Roman times and Dioscorides (Allen & Hatfield 2004). Having said this, although Grieve(1931) describes the plant in her book A Modern Herbal, she makes no mention of the medicinal useof the species.
The rhizome is boiled and eaten as a vegetable in China, and young leaves are likewise eaten as salad inIndia (Mabberley 1997).
Names
The genus name 'Calystegia' is derived from two Greek words, 'kalyx', a calyx or 'cup', and 'stege', 'acovering', probably referring to the two bracts that cover the calyx of some bindweeds (Johnson &Smith 1946; Gilbert-Carter 1964). The specific epithet 'sepium' is from the Latin, 'sepes', 'a hedge',referring to one of its most common habitats (Gledhill 1985).
The C. sepium complex as a whole has a great many common names. Grigson (1987), for example, listsa total of 60, most of which refer to its climbing habit (eg 'Bindweed', 'Bearbind', 'Ropewind' and'Woodbine'), but the remainder of them stress either its beautiful flower ('Hedge Lily' and 'Old Man'sNightcap'), or else its troublesome character as a weed ('Devil's Garter', 'Devil's Vine' and'Hellweed').
Threats
None.
Introduction, history unknown, a very rare casual, but possibly an identification error.
1 October 1995; McNeill, I.; edge of laneway, Gushedy Beg Td, N of Ederny.
This subspecies differs from subsp. sepium in its deep pink, white-striped flowers and in havingshort hairs on stems, leaf-stalks and flower-stalks (An Irish Flora 1996). It is principally acoastal plant of salt-marshes and maritime sands, native of W Europe and local on W coasts of B & I(R.K. Brummitt, in: Tutin et al. 1972, Flora Europaea 3, 78-9; New Flora of theBI 1997). It also occasionally occurs elsewhere, apparently introduced and naturalised. Thistaxon has previously been recorded in the NE of Ireland at two coastal sites in the extreme N of Antrim(H39) (FNEI 3), but never inland or indeed landlocked as is the situation in county Fermanagh.There are also a number of sites for this subspecies in W Donegal (H35), along the northern shore ofDonegal Bay, which is not very distant from Fermanagh (Perring & Sell 1968, map 406/1b).
The similarity of the plant to C. pulchra (Hairy Bindweed) means that care in identification isrequired, but the latter has bracteoles below the calyx that are overlapping and strongly inflated,while subsp. roseata has bracteoles that are not inflated and that scarcely overlap (An IrishFlora 1996).
Although Ian McNeill is an excellent field botanist, without a voucher to obtain verification, we cannotbe certain that the identification of the solitary Fermanagh record listed above is correct.Accordingly, with regret, we can only regard this as a tentative First County Record.
Introduction, neophyte, a very rare garden escape.
17 September 1993; Northridge, R.H.; grounds of Portora Royal School, Enniskillen.
July to September.
This large pink-flowered garden species of bindweed has only been found in three widely spaced sites inFermanagh. The details of the other two records are: near houses, Clabby Village, 21 June 1997, I.McNeill; on waste ground between road and lake in front of house at Bleanalung Bay, NE Lower Lough Erne,4 October 2010, RHN & HJN.
C. pulchra is a hairy stemmed climbing perennial of unknown geographical origin. It may eitherhave been introduced directly from NE Asia, or perhaps it may have arisen as a hybrid during Europeangarden cultivation (Griffiths 1994). It is rather similar to C. sepium subsp. roseata, both bindweeds having either pure pink or pink andwhite striped flowers and short hairs on their stems, leaf-stalks and flower-stalks. The principaldifference between them is the form of the bracteoles, which in C. pulchra are strongly inflated,pouch-like and overlapping one another, whilst those of C. sepium subsp. roseata are about as long as the calyx, not inflated andscarcely overlap at all (An Irish Flora 1996).
C. pulchra was probably first introduced to B & I around the 1850s or 1860s, since the firstEuropean herbarium specimens known are both dated 1867 (one from Twickenham Park (VC 21) and one fromSweden) (Brummitt & Heywood 1960). C. pulchra very frequently escapes from gardens and it hasbecome naturalised locally. Nowadays, it is widely scattered throughout the whole of these islands andvery probably it is still spreading (Clement & Foster 1994; G.M. Kay, in: Preston et al.2002). It is also naturalised in Holland, France, Germany, Poland, Austria, Czechoslavia, Denmark andSweden, and possibly elsewhere in Europe (Perring & Sell 1968). In Britain it appears to be mostfrequent in SE and NW England, and in the Glasgow and Edinburgh regions of Scotland, usually occurringclose to habitation (Garrard & Streeter 1983; G.M. Kay, in: Preston et al. 2002).
The Cen Cat Fl Ir 2 indicates C. pulchra has been recorded from 25 of the 40 Irish VCs, andCat Alien Pl Ir listed seven additional VCs, which brings the total to 32. Inspection of theNew Atlas hectad map suggests that C. pulchra is more frequent and widespread in NI thanis the case in the RoI.
Threats
The species does not set much or any seed, but it is actively spreading, probably as rhizome fragmentstransported with soil. It could easily become a nuisance over time, having the ability to overgrownative vegetation.
Introduction, neophyte, very rare, but probably under-recorded.
1951; MCM & D; hedgerow, Tattenabuddagh Bridge, Colebrooke River.
June and July.
Growth form and preferred habitats
C. silvatica is a rather glamorous, very large white-flowered bindweed, originating from theMediterranean region and SW Asia. It differs from both subspecies of its close relative C. sepiumin that the two large bracteoles at the base of the flower overlap one another and completely concealthe calyx. It was not until the 1930s that the two species were separated in the field (Praeger 1934;Lousley 1948; Stace 1961), but it is now recognised that C. silvatica is a garden introduction toB & I from S Europe, which probably arrived in Britain sometime in the early 19th century, admiredas a cultivated decorative flowering plant.
An illustration of C. silvatica produced in 1777 suggests it was in Great Britain by that earlydate (Salisbury 1964, p. 292). However, another more or less contemporary commentator, John ClaudiusLoudon (1783-1843), the founder and 'conductor' of The Gardener's Magazine and The Magazine ofNatural History, suggested C. silvatica was first introduced to British gardens as lateas 1815 (Loudon 1830; Walters & Webb 1956). Some authorities feel it was not reliablyrecorded in Britain until 1867, the date of a BM herbarium specimen (Ellis 1993). However, the1867 specimen in the BM from Twickenham Park (VC 21), regarded by Lousley (1948) as C.silvatica, was later re-examined by Brummitt & Heywood (1960) and pronounced by them,"a poor specimen, ... almost certainly C. pulchra". Herbarium specimenscollected from the wild in Britain have now been reported dating from 1863 (Stace & Crawley 2015).
Whatever date Large Bindweed was first introduced to B & I as a decorative garden subject, itsubsequently escaped 'over the wall' into the wild, and was already growing widespread in lowlandBritain and the more populated parts of Ireland when Lousley was writing of it in 1948. One wonders justhow long it took gardeners to realise that they had a pernicious weed on their hands, and exactly whendid they begin discarding it on waste ground, old quarries and the like?
C. silvatica is a plant of drier habitat conditions than C. sepium (Hedge Bindweed)avoiding the marsh and fen situations so characteristic of the latter (Garrard & Streeter 1983). Itoccurs most commonly near habitation in gardens, on fences, in hedgerows and on waste ground includingin old quarries (An Irish Flora 1996). It can occasionally also colonise semi-natural habitatsfurther from habitation, presumably spread by seed, although vegetative fragments of root and rhizomecan also travel in mud, eg attached to vehicle tyres (G.M. Kay, in: Preston et al. 2002). LikeC. sepium, with which it shares many ecological characteristics, the established strategy ofC. silvatica is categorised as intermediate between Competitor and Competitive Ruderal (C/CR)(Grime et al. 1988, 2007).
Variation
There is sufficient variation in C. silvatica to merit the recognition of two subspecies (subsp.silvatica and subsp. disjuncta Brummitt) based on differences in the bracteoles, and onevariety, var. quinquepartita N. Terrace, which has a corolla divided into several definite lobes.The variety is rare, having only been recorded in Britain on a few occasions and it might be referableto either of the two subspecies (Sell & Murrell 2009).
Strong vegetative and limited flowering reproduction
In common with C. sepium subsp. sepium, the even larger and moreglamorous pure white flowers of C. silvatica are completely self-incompatible, a condition whichlimits the species' ability to set seed, especially in small populations. To an unknown extent, plantsof C. silvatica must sometimes (often) be derived from fragments of the fleshy, almost succulent,brittle rhizomes and roots transported with soil, with the result that local stands may represent singlestrain clones (Salisbury 1962). Whenever a local colony of any dimension consists of just one or a fewgenetic strains, bee-transported compatible pollen may prove to be too distant, making it unavailablefor fertilisation of the flowers.
Thus in both of these common species, the native C. sepium and the alien C. silvatica,vigorously spreading vegetative populations frequently fail to produce fruit and set no seed (Stace& Crawley 2015). Of course, some larger colonies will contain more than one strain and they canfreely set fruit and develop abundant seed (Baker 1957a), but fortunately the reduction created byhom*omorphic incompatibility limits the dispersal and colonising abilities of the species to aconsiderable extent.
It is remarkable that three of the alien species most feared and hated by gardeners (C. silvatica,Veronica filiformis (Slender Speedwell) and Fallopia japonica (Japanese Knotweed) shouldbe mainly or entirely vegetatively propagated due to their inability to set seed (Stace & Crawley2015).
Any flowering that takes place is from July to September and pollination of the self-incompatible flowersis similar to C. sepium (Garrard & Streeter 1983) (see the C. sepium subsp. sepium account on this website). The fruit capsule of C.silvatica develops from September onwards into October. The capsule is 10-15 mm in size andsub-globose in shape, splitting when ripe to release the seeds, which are larger (4-5 mm in diameter),smoother and blacker than those of C. sepium subsp. sepium. Likethe latter, they are hard-coated and presumably have similar powers of extended dormant longevity(Salisbury 1964; Sell & Murrell 2009). Having said that, there is a solitary observation of thespecies in the soil seed bank survey of NW Europe which suggests C. silvatica seeds aretransient, surviving in the soil for less than one year (Thompson et al. 1997). However onerecord in that survey is insufficient to base a definite case on, since four of a total of nine recordsfor C. sepium suggest it also has transient seeds, whereas other data findings in the same surveyindicate buried seed survival in soil for up to 40 years (Thompson et al 1997).
Fermanagh occurrence
While elsewhere in B & I this is a rather common and widespread naturalised garden escape, there areonly six widely spread records of it across Fermanagh spanning a 59-year period.
Apart from the first record listed above, the details of the other five Fermanagh finds are: roadsidehedge at Gorteen, below Slieve Rushen, 18 July 1991, RHN; disused quarry on the outskirts of Edernyvillage, 3 October 2002, I. McNeill; Cloncoohy, 3 km SW of Teemore, 11 September 2010, RHN & HJN; Sof Clonelty, 3 km SE of Newtownbutler, 15 September 2010, RHN & HJN; Necarne near Irvinestown, 18September, RHN.
Since so few colonies of C. silvatica have been recorded in rural Fermanagh, seed production andits successful dispersal appears very much less likely here than in eastern NI. In urban Belfast, forinstance, C. silvatica has become sufficiently common for Beesley & Wilde (1997) to record itin 40% of the 1-km squares in the inner city, some colonies demonstrating high levels of vegetativevigour and dominance, although not as common as C. sepium anywhere across the city, andespecially so on waste ground.
Irish occurrence
The pattern of occurrence or recording of this clambering rhizomatous perennial in Ireland is very patchyand uneven. The Cen Cat Fl Ir 2 lists it as having been recorded in 30 of the 40 Irish VCs, towhich Cat Alien Pl Ir adds two more; neither of these sources includes Fermanagh. After theannouncement here of its rare presence in Fermanagh, the situation remains that C. silvatica hasnot yet been reported in seven other Irish VCs. The New Atlas hectad map shows C.silvatica present in most maritime and a proportion of inland squares up along the Irish Seacoast from E Cork (H5) to Co Londonderry (H40). However, it is much more thinly scattered up theAtlantic west coast and inland, stretching northwards from Mid-Cork (H4) to E Donegal (H34).
Although it may sometimes be mistakenly recorded as either C. sepium or C. pulchra (HairyBindweed), the existing pattern of Irish records suggests that C. silvatica might still be slowlyspreading on the island and may well be under-recorded.
British occurrence
In most parts of lowland, populated B & I the species is now well established and naturalised inwoods and hedgerows. This is especially so in lowland England and Wales as far north as Cumberland (VC70) and Cheviot (VC 68), although it remains very much more scarce and patchy in lowland Scotland exceptup the E coast, and in the major conurbations of Glasgow and Edinburgh. The distribution appears to bestable, rather than increasing or decreasing (Rich & Woodruff 1990; Clement & Foster 1994;Preston et al. 2002). There is a separation in distribution between the two subspecies, subsp.disjuncta being the more common form throughout most of Britain, whereas subsp. silvaticaappears the more frequent of the two on the Celtic fringe of Cornwall, Wales, parts of Scotland and allof Ireland (G.M. Kay, in: Preston et al. 2002; Sell & Murrell 2009).
European and world occurrence
Subsp. disjuncta is a native of the W Mediterranean region while subsp.silvatica is from the E Mediterranean. As C. sepium subsp.silvatica (Kit.) Maire, the range of C. silvatica is described as S Europe including theMediterranean basin, N Africa and SW Asia including the Caucasus (Hultén & Fries 1986, Map1537; Clapham et al. 1987).
Threats
Since only six colonies of C. silvatica have been recorded in rural Fermanagh, seed production andsuccessful species dispersal appears very much less likely here than in urban parts of eastern NI.
Threats
None.
Native, common and widespread. Circumpolar boreo-temperate.
1882; Stewart, S.A.; Co Fermanagh.
April to November.
Growth form and preferred habitats
An emergent aquatic or semi-aquatic glabrous perennial with a creeping rhizome, Menyanthestrifoliata used to be considered a member of the Gentian Family, but it has now been transferredto the Menyanthaceae, a small family of five genera and 40 species, all of which are aquatic(Preston & Croft 1997). Whatever its relationships or taxonomy, M. trifoliata has to be oneof the most strikingly beautiful wild flowers in the flora of B & I.
It is a significant member of numerous marsh, fen and bog plant communities on the shallow margins oflakes, pools and ditches, where, depending on depth, it occasionally forms either single-species standsaround the edge of open water in lakes and tarns, or floating mats in deeper water further offshore. Inpoor- to medium-nutrient fen-meadow conditions, or in slow-flowing drains of similar nutrient status, itis more usually associated with a wide range of other common emergent shoreline species such as Carexrostrata (Bottle Sedge), Equisetum fluviatile (Water Horsetail), Galium palustre(Marsh-bedstraw), and Potentilla palustris (Marsh Cinquefoil).
M. trifoliata also commonly occurs in shallow, acidic, nutrient-starved pools on raised or blanketbogs, or in cut-over versions of these, where it is frequently accompanied by species such asEriophorum angustifolium (Common Cottongrass) and Potamogeton polygonifolius (BogPondweed). Few plants can compete with the vigorous growth of Bogbean on deep, oxygen-deficient(anaerobic), organic mud or peat. It can also be found on the wettest areas of moorland flushes, whichbeing fed with ground water are of slightly richer nutrient status and where also root aeration is verymuch better than in the surrounding ground (Hewett 1964; Preston & Croft 1997).
Fermanagh occurrence
Bogbean is a very common, widespread and locally abundant species in Fermanagh as the tetrad mapemphasises, having been recorded in 242 tetrads, 45.8% of those in the VC. It is particularly common,indeed omnipresent, around the shallows of Upper Lough Erne and other inter-drumlin lakes. It is alsofrequent in pools and flushes on raised and blanket bogs, except at the highest levels.
Flowering reproduction
The deciduous, fleshy, trifoliate, bean-like leaves are overtopped by a thick, short raceme of up to 29flowers on a leafless stalk or peduncle from May to July (Hewett 1964). The corolla tube, broadlycampanulate, c 15 mm diameter, has five deep recurved lobes, the petals being rose pink on their outersurface, pure white on the inner, 'much fribriate', which is the technical botanical shorthanddescribing the incredibly beautiful, deeply-cut, lace-like fringing, which makes the flower soremarkable it really becomes unforgettable (Clapham et al. 1962; Hewett 1964). The early Britishherbalist, John Gerard described the flower very poetically in language of Shakespeare's time as,"... towards the top of the stalks standeth a bush of feather like flowers of a white colour,dashed over slightly with a wash of light carnation" (Gerard 1597).
It is difficult to imagine why the Bogbean petals are so deeply fringed, or indeed why any flowershould be so decorated (eg Dianthus superbus (Fringed Pink) or Gentiana ciliata (FringedGentian)). One possibility is the fringe may deter small crawling insects, too small to effectpollination (Melderis & Bangerter 1955). Or does it perhaps play some part in a water-pollinationmechanism (ie hydrophily), as in other aquatics such as Najas spp., Callitriche hamulata(Intermediate Water-starwort) and Ceratophyllum spp. (Faegri & van der Pijl 1971)? The fringemight somehow enable rain-pollination, where the flower fills with water to a certain level and pollengrains float on the surface until they eventually reach the stigma (Hagerup 1950).
These latter suggestions are ruled out by the fact that M. trifoliata flowers areself-incompatible (Hewett 1964) and, furthermore, they exist in two forms, 'pin' and 'thrum', ie theyare dimorphic heterostylous, like Primrose (Primula vulgaris) and Cowslip (P. veris),having styles and filaments of two different lengths, a very positive out-breeding mechanism (NicLughadha & Parnell 1989; Proctor et al. 1996; Richards 1997a).
The flowers secrete nectar at the base of the globular ovary and attract a variety of small insects thatact as pollinators (Garrard & Streeter 1983). In an Irish study, on average, 17 ovules were producedper flower and, on average, six seeds developed, although seed production was found to be very uneven.Thirty-five percent of sampled capsules contained no ripe seed whatsoever (Nic Lughadha & Parnell1989).
Seed dispersal
While fruit set may well be rare and the number of seed produced few (Hewett 1964; Nic Lughadha &Parnell 1989), seed dispersal, even as an occasional event, is of vital importance to the long-termsurvival of the species. Water-borne dispersal is bound to be significant and, as expected, Bogbean seedis hard coated and extremely buoyant, the seed-coat integument containing, "aeriferous tissue withintercellular spaces" (Ravn 1894), so the seeds can often float for up to two months and remainviable, or rarely longer than this (Sculthorpe 1967). According to Hewett (1964), however, many seedsare still buoyant and remain viable when kept in water for up to 15 months. The current author (RSF)finds the latter claim rather incredible, and would like to see the detailed data on which it is basedbefore fully accepting it.
Aerial seed travel after capsule dehiscence is a very short distance. Nic Lughadha & Parnell (1989)measured distances in the field that ranged from 4 to 49 cm, with a mean of only 16.3 cm.
Since M. trifoliata is regularly recorded in isolated lakes and tarns, the plant must be capableof some additional form of jump-dispersal. Birds, particularly Mallard ducks, have been implicated inobservations reported by Ridley (1930) and, as some of these birds migrate, seeds in their crop might betransported, especially those with a resistant testa like that of Bogbean.
Experimental studies on seed dispersal by fish involving M. trifoliata are also described byRidley (1930). The fish mentioned include Roach and Perch. Ridley even moots the remote possibility offish with seed in their gut being taken and transported by birds, chiefly ducks, including Mallard. Thismight seem a tad ridiculous as a mechanism, but there is plenty of evidence from island biogeographythat proves long-distance or jump-dispersal does occur in Nature (Carlquist 1974) and generally it is aprocess involving the highly improbable to overcome the otherwise impossible!
Vegetative reproduction
Probably like other common and successful rhizomatous species, eg Calystegia sepium subsp. sepium (Hedge Bindweed) and Tussilago farfara(Colt's-foot), Bogbean seed production is of secondary importance to vegetative reproduction,except in terms of long-distance transport and long-term species evolutionary survival. Vigorousrhizome spread is a highly successful means of transport for a species living in a less-than-idealnutrient environment. Indeed, vegetative reproduction tends to become of paramount importance in everyenvironment where survival is severely tested, such as arctic, alpine and desert conditions. Bogs andacidic waters fed by runoff from siliceous, nutrient-poor rocks, also provide such a testing environmentand clearly M. trifoliata must be well-adapted to cope with the problem of competition for scarceresources.
Fossil record
The British fossil record of M. trifoliata is remarkable for its extensiveness and completeness(Godwin 1975). The seeds are regarded as the most easily recognised Quaternary plant fossils, but pollenand rhizome fragments are also found. It occurs in both full-glacial and early post-glacial deposits(Hewett 1964).
Bogbean appears on a list of 34 ancient aquatic plants native in the British Isles, all represented inlate-glacial sediments, some of them possibly periglacial survivors (ie species considered to have beenpresent in the near vicinity of major ice sheets during the glacial phase(s)) (Sculthorpe 1967).Confirmation of the periglacial presence of M. trifoliata in Ireland comes from a study of a siteat Derryvree, near Maguiresbridge, Co Fermanagh, where a fossil seed of the species was recovered in aMiddle Midlandian full glacial deposit radio-carbon dated to 30,500 BP. This deposit contained a floraand fauna of open tundra, and a periglacial climate prevailed when it was laid down (Colhoun etal. 1972).
Of the ancient aquatic species listed by Sculthorpe (1967), only two do not currently appear inFermanagh, Scheuchzeria palustris (Rannoch-rush) (extinct at its previous solitary site in CoOffaly (H18)) and Subularia aquatica (Awlwort), an extremely rare plant in Ireland with post-1970records from just four hectad squares (Preston & Croft 1997).
British and Irish occurrence
M. trifoliata is a common, locally abundant plant in B & I, most prevalent in the N & W ofboth major islands, although it has suffered centuries of depredation in the S & E due to drainageand development, so that in those parts of the country it has become scarce or absent (NewAtlas).
Possessing such a beautiful flower, it is not surprising that M. trifoliata is regularlycultivated in water gardens. In turn, this can lead to plant material either escaping, or the excessbeing irresponsibly dumped in the wild. Burton (1983) believed that some occurrences of Bogbean in theLondon area originated in this manner and the same could very easily occur elsewhere. In Middlesex (VC21), where Bogbean was previously local, it has become rare, and Kent (1975) believed that on numerousoccasions it had been deliberately planted in semi-wild situations.
European and world occurrence
M. trifoliata belongs to the circumpolar boreo-temperate phytogeographical element. The main areaof distribution lies in the northern boreal region, between 40°N and the Arctic Circle. The southernlimit of the range lies between C Portugal and Spain, S France, Corsica, N Italy, Greece, the Caucasusinto temperate Siberia and on to Japan. It has also been recorded in N Morocco. In N America, thesouthern limit of the distribution runs from California in the west to Maryland in the east. It grows Nof the Arctic Circle in W Greenland, Iceland, Norway, Siberia and Alaska (Hewett 1964; Hultén &Fries 1986, Map 1510).
Uses
Apart from its decorative use in water gardens, Bogbean contains a small quantity of volatile oil and abitter glucoside called 'Menyanthin' that makes it very unpalatable. It is possibly this feature thathas made it one of the most prized herbs available to folk use and herbal medicine (Allen & Hatfield2006). M. trifoliata has been used in herbal folk medicine as an astringent tonic and for avariety of ailments including digestive problems, rheumatism, constipation, arthritis, scurvy, asthma,persistent coughs, pulmonary tuberculosis, fevers, jaundice and kidney trouble. A decoction of the roothas also been used to relieve pain of stomach ulcers and the leaves were used as a poultice for boilsand skin troubles (Grieve 1931; Vickery 1995; Allen & Hatfield 2006).
In other uses, the leaves, being bitter, have been used in the past in N England as a substitute for hopsin beer-making and hence the local name 'Bog Hop' (Grigson 1955, 1987). The rhizome has also been driedand powdered for use as a substitute for bread flour by Esquimaux, and also in N Eurasia (Mabberley1997).
Names
The genus name, 'Menyanthes' originated with the Greek herbalist, Theophrastus, who appears to have givenit to the related aquatic, Fringed Water-lily (currently Nymphoides peltata). The word elementsof the name are, 'mene' in Greek meaning 'moon' or 'month', and 'anthos', 'a flower' (Johnson &Smith 1946; Gledhill 1985). The suggestion, applied to M. trifoliata, is that the floweringperiod lasts one month, but in reality, the flowering phase continues for three months of the summer.
The English common name 'Bogbean' is a late 18th century modification of 'Buckbean', the 16th centuryname translated by Henry Lyte (1578) from the Flemish 'Bocksboonen' in the herbal of Rembert Dodoens(1554). Lyte took this to mean 'Goat's beans', the leaves resembling young leaves of the Broad Bean(Grigson 1974). 'Buckbean' was eventually transformed to 'Bogbean', presumably to better fit the typicalhabitat and become more comprehensible.
Threats
None.
Introduction, neophyte, a very rare garden escape or discard. Eurasian temperate, but widely naturalisedincluding in N America.
1988-90; NI Lakes Survey; Cargin Lough, Upper Lough Erne.
Native range and English occurrence
The native range of this Water-lily-like species stretches from SE England and the Baltic southwards andeastward through Asia to Japan (Preston & Croft 1997). It was recorded from the Thames in 1570 andEast Anglia in 1660 and is considered indigenous only in these two local areas, although it is widelyrecorded in S & C England and thinly scattered over a wider area of Britain as far N as Inverness(Preston & Croft 1997).
In England, where the species is a very much more widespread introduction beyond its supposed nativeoccurrence, N. peltata is often found as a broad band around the edges of lakes, slow-flowingstretches of rivers and associated backwaters, plus in fenland lodes and disused canals (C.D. Preston,in: Stewart et al. 1994). Although they appear quite different in form, in their biology N.peltata and M. trifoliata (Bogbean) share many features, especially with respect to theimportance of rhizomatous vegetative growth and their reproduction by fragmentation. The flowers alsoshare dimorphic heterostyly, ie they both produce 'pin' and 'thrum' flowers. The seeds of both speciesfloat for quite prolonged periods, weeks and possibly months. They may also be dispersed, at leastexternally in mud, by birds (Preston & Croft 1997). Fringed Water-lily flowers are also veryattractive in appearance: individually they last a single day, but clonal colonies flower over a longperiod.
Irish occurrence
With small, yellow, dimorphic heterostylous flowers and 12 × 10 cm floating leaves, N. peltata isknown as an introduction in eastern NI, having been planted in the Lagan Canal by J. Campbell, one ofthe earliest curators of the Belfast Botanic Garden in the early 19th century (FNEI 1).Early introductions like the Lagan Canal one in various parts of Ireland, including Lough Neagh, appearinitially to have naturalised, spread and thrived for a number of decades, but they suddenly declinedand disappeared sometime later in the same century (Flora of Lough Neagh; Cat Alien PlIr).
Fringed Water-lily has been recorded a few times from a total of six Irish VCs in the post-1985 period,namely in NI from Cos Fermanagh (H33), Armagh (H37), Down (H38) and Antrim (H39), and in the RoI fromMid- and East-Cork (H4, H5) (Cen Cat Fl Ir 2; FNEI 3; Cat Alien Pl Ir). All Irishrecords have been known or suspected introductions, most probably originating from accidental gardenescapes or discarded surplus material. Fortunately the species did not persist long at any of the siteswhere it occurred.
Fermanagh occurrence
N. peltata was unknown in Fermanagh until 1988-90, when it was discovered by the NI Lakes Surveyat both Cargin Lough, Upper Lough Erne and in the much more isolated Watson's Lough, 6 km NE ofEnniskillen. The behaviour of this rhizomatous perennial at the two known lake sites and at any furthersites that may emerge should be carefully monitored to see whether, in the currently warming climate,growing conditions have changed and the plant might soon become capable of persisting and spreading.
In contrast with the only other member of this family in the B & I flora, Menyanthestrifoliata (Bogbean), N. peltata is a plant of open, relatively deep waters (ie 50-200cm), that are calcareous, nutrient-rich (or enriched) and eutrophic. This habitat description suggeststhat both Upper and Lower Lough Erne could well provide a very suitable environment for this aquaticintroduction, a rather alarming notion since it might successfully compete with native floating speciesand begin to oust them.
In northerly sites in Britain, plants flower only sparingly, a situation that may also pertain in NI.Happily, to date, there have been no further sightings of N. peltata anywhere in Fermanagh, butno one can be complacent – the threat that this plant poses is a real one.
Threats
Not a problem at present, but could possibly increase dramatically and become a weed, endangering nativeaquatic plants.
Introduction, neophyte, a very rare casual garden escape. Eurosiberian boreo-temperate, but also widelynaturalised including in both W Europe and N America.
1947; MCM & D; Carrickreagh Bay, Lower Lough Erne.
Only recorded once in Fermanagh, in a much-frequented rocky limestone lakeshore situation on Lower LoughErne, in Ireland this decorative perennial is always a garden neophyte and, at this location, it wasprobably derived from dumped garden waste. Jacob's-ladder has not been seen again at Carrickreagh, whichis a frequently visited site, so we can assume that it did not persist. Hackney et al. (FNEI3) made a similar statement about this species' occurrence in NE Ireland.
Pollen grains of P. caeruleum have very characteristic surface sculpturing that makes them readilyidentifiable to species level. Fossil pollen has been identified in deposits in England from all zonesof the last Ice Age (Weichselian) and from the early Flandian Post-glacial, proving the species isnative (Pigott 1958; Godwin 1975). Thus, at least in a number of sites in N England, Jacob's-ladder isone of the special, rare and very beautiful native plants of constantly moist, flushed soils of thePennine Carboniferous limestones, on cool, N-facing, steep but stabilised, upland calcareous screeslopes at the base of cliffs (Wiggington 1999). Jacob's-ladder also occurs in two sites further north inNorthumberland, on andesite clay soil cliffs along river banks, where it is also considered native.Altogether there are a total of around 20 native P. caeruleum stands in England and it is a RedData Book species of conservation concern (Swan 1993; Wigginton 1999).
The prime requirements of P. caeruleum are for a moist, fertile soil and freedom from competition.It can tolerate a certain degree of shade and grows in tall herb vegetation with associated species thattypically include Arrhenatherum elatius (False Oat-grass), Dryopteris filix-mas(Male-fern), Epilobium montanum (Broad-leaved Willowherb), Festuca rubra (Red Fescue),Filipendula ulmaria (Meadowsweet), Heracleum sphondylium (Hogweed) and Urticadioica (Common Nettle) plus a thick carpet of moisture-holding mosses and liverworts.Jacob's-ladder plants are intolerant of prolonged water loss, so must have a regular seepage ofgroundwater, or a frequent rainfall supply (Pigott 1958). Seedlings are particularly vulnerable towilting and a light, open canopy of Fraxinus excelsior (Ash) helps to maintain a humidenvironment. Grazing is needed to keep the Ash in check, although this means flowering performance isseriously suppressed. A careful balanced management regime involving light, occasional grazing isessential to maintain the scattered, though sometimes considerable-sized native P. caeruleumpopulations (I. Taylor, in: Wiggington 1999).
However, apart from these English native sites, Jacob's-ladder is also a deservedly popular,clump-forming decorative perennial, widely grown by discerning gardeners across B & I. It hasfern-like, elegantly pinnate, radical leaves with 17-27 leaflets that give it its English common nameand in May and June it bears terminal and axillary cymes of clear blue, or rarely white, small, open,campanulate (bell-like) flowers with strongly contrasting orange stamens. It performs best in almostfull shade and has stems 20-60 cm tall.
In the garden setting, it is described as short-lived, but it seeds itself prolifically (Hansen &Stahl 1993). The native plant, on the other hand, is considered a long-lived, polycarpic perennial,reproducing by seed. It flowers a month or so later than the garden form, in June and July, butexceptionally can continue into September. It is pollinated by bumble-bees, although it is alsoself-fertile. Seed is released from the dry capsules from early autumn onwards, and germination ismainly in spring. Mature plants die down in winter, but the dead flower stems remain erect, eventuallyshedding the seeds (I. Taylor, in: Wigginton 1999).
As demonstrated by the New Atlas hectad map, garden forms of Jacob's-ladder very commonly escapefrom cultivation and can become naturalised in Britain from E Cornwall to Shetland (VCs 2-112). Thishappens very much more rarely in Ireland: the Cat Alien Pl Ir lists P. caeruleum'soccasional, non-persistent 19th century occurrences from a total of just six Irish VCs, mainly in thefar south of the island. There have also been 20th century discoveries from Cavan (H30), where it isplanted but spreading, and from the three NE VCs (Cos Down (H38), Antrim (H39) and Londonderry (H40)).Thus, the solitary Fermanagh record listed above is not an isolated occurrence, but it ispreviously unpublished.
The escaped or discarded garden material is morphologically different from the native species and the twoare easily distinguished (Pigott 1958). Garden escapes usually have pale, purplish-blue, lavenderflowers, yellow pollen and narrower leaflets (Sell & Murrell 2009).
European and world occurrence
P. caeruleum is a mainly boreal species that forms part of a species complex along with some ±critical taxa (Hultén & Fries 1986, Map 1530). It occurs in N, C & E Europe and adjacent Asia,especially in the mountains. Garden cultivation occurs across Europe with the inevitable escapes intowild and semi-natural habitats and, often, native and escaped specimens are mixed together as inEngland. Thus it is almost impossible to ascertain the true native distribution area.
The native distribution of P. caeruleum itself occurs throughout the uplands of C & N Europe,with strongholds in the Alps, Finland, Norway and Russia, extending eastwards to Lake Baikal. The rangeof habitats occupied by the species on mainland Europe is much wider than in England, including alpinemeadows in Switzerland, birch woodland in Scandinavia, Picea abies (Norway Spruce) forest inSlovakia and even lowland fens in N Germany (I. Taylor, in: Wiggington 1999).
P. caeruleum has also been introduced quite widely in eastern N America and the whole speciescomplex has a circumpolar distribution, although with a large gap in E Asia and N America. As acomparison of two maps shows, the closely related P. acutifolium Willd. almost fills that gap(Hultén & Fries 1986, Maps 1530, 1531).
Threats
None.
Introduction, neophyte, a very rare casual, either a garden escape or from wild bird seed mixture.
23 June 2003; Northridge, R.H.; south corner of Holme Bay, Lower Lough Erne.
This attractive pale-blue-flowered N American member of the Waterleaf family has been found only twice inFermanagh, in a heap of discarded garden gravel on the lakeshore, as detailed above, and secondly (againfound by RHN, 24 August 2004) on a roadside bank near the Spence farm at Moyglass, 5 km NW ofEnniskillen. The plant, which is an annual, has distinctive dissected, Tansy-like leaves. It is a rarecasual alien introduced in seed mixtures either for feeding wild birds or pheasants, or for growingso-called 'wild flower meadows'. It may also be deliberately grown as an ornamental garden annual, or asa 'green manure' in allotments, or in arable fields as a nectar source to attract bees and hoverfliesthat prey on aphid pests. P. tanacetifolia may also appear as a seed contaminant in re-seededlawns and other grasslands (Clement & Foster 1994; New Atlas; Cat Alien Pl Ir). Theplant is found typically in cultivated or disturbed ground, often where waste is discarded. It isgenerally regarded as a casual and most populations are not expected to persist.
P. tanacetifolia was first cultivated in Britain in 1832, but was not found there in the wilduntil 1885. In Ireland, the first discovery was in a field near Downpatrick, Co Down (H38) in 1962 (H.J.Killick, in: Preston et al. 2002; Cat Alien Pl Ir). The recent history of its occurrencesuggests it has increased quite considerably in Britain. There may also be a much less obvious increasein Ireland. The Cat Alien Pl Ir details finds in a total of six Irish VCs prior to our 2003Fermanagh record, but only one of these is featured in the New Atlas hectad map.
Threats
Although this is a casual, it appears it is being more frequently introduced in recent years, eitherdeliberately sown or as a seed contaminant. It would be wise to research its reproductive capacity now,before it becomes a more significant weed.
Native or possibly a garden escape, very rare and now locally extinct. Eurosiberian temperate, but widelynaturalised.
1939; Praeger, R.Ll.; roadside near Crom Castle Estate.
Growth form and preferred habitats
L. officinale is a shortly rhizomatous perennial with erect stems, 30-100 cm tall, much branchedabove and leafy, bearing alternate, sessile, lanceolate leaves with blades 2-8 × 1-2 cm. The leafsurfaces have distinct lateral nerves and are pubescent and rough to the touch (Launert 1981; Sell &Murrell 2009). It flowers during June and July, producing dense, leafy, terminal and axillary cymoseinflorescences of bisexual flowers. The inflorescences elongate after flowering as the fruits develop.
The yellowish- or greenish-white corolla is not much longer than the calyx and is 3-4 cm in diameter. Theflower parts are in fives, the corolla funnel-shaped, its throat containing hairy longitudinal folds orscales. The five stamens are attached to the middle of the corolla tube; the filaments and anthers areyellow. The ovary is superior, of two carpels, each containing two ovules. There is a solitary shortstyle, 1-2 mm and a bifid stigma. Pollination is mainly by bees and flies (Fitter 1987). The fruitconsists of four, pale pearly grey, shining, ovoid nutlets or achenes – ie single seeded dry fruits(Hickey & King 1981; Clapham et al. 1987; Sell & Murrell 2009). There is no explanationas to how the nutlets are released and dispersed in Ridley (1930) and the current author (RSF) is at aloss to come up with any dispersal mechanism, other than possibly birds feeding on them and some of thehard fruits passing through the gizzard and alimentary canal intact (van der Pijl 1972, p. 27).
Common Gromwell is a plant of dry, grassy, lowland vegetation in semi-shade on woodland margins, inscrub, hedges and on more open ground on sandy banks, roadsides and quarries. It almost always occurs oncalcareous or base-rich, sometimes rocky, soils (Garrard & Streeter 1983). It can tolerate moderatelevels of shade but prefers full sun conditions (Fitter 1987). The established strategy of L.officinale is categorised as intermediate between Stress-tolerant Competitor and a generalistCompetitive, Stress-tolerant, Ruderal species combining features of all three types of ecologicalapproaches (SC/CSR) (Grime et al. 1988, 2007).
Fossil record and species status
The solitary B & I fossil of this species was found in Norwich in a post-medieval context, datedbetween 1500 and 1700 AD (Tomlinson & Hall, website article, last updated Aug 1996: http://intarch.ac.uk/journal/issue1/tomlinson/toc.html)(website accessed 18 July 2022). Similar suggestions of doubtful native status apply in Scotland and, inthe absence of any pre-Neolithic fossil material of such a hard-seeded species (Godwin 1975), in theview of the current author (RSF), the question remains wide open in both these countries, or even indeedthroughout the whole of B & I.
Fermanagh occurrence
In Fermanagh, L. officinale has only ever been recorded twice: both discoveries were made in theSE corner of the county, in 1939 and 1949. As it has not been seen for over 50 years it is assumed to belocally extinct. There is a voucher in BEL for Praeger's first record listed above and thedetails of the second station are: at the Old Ulster Canal near Wattle Bridge, 1949, MCM & D.
Irish occurrence
The Cen Cat Fl Ir 2 indicates that previously it has been recorded (at least once), in 34 of the40 VCs on the island, but as is often the case, this is not the whole story. In reality, Common Gromwellis very rare, local and widely scattered in Ireland and populations of the species have been decliningsince before 1930 (BSBI Atlas 2; New Atlas).
The New Atlas map for Ireland shows L. officinale sparsely scattered, mainly around theBelfast area of Co Antrim (H39) and in eastern Tyrone (H36), with a few isolated hexads in Co Armagh(H37) and Fermanagh (H33). The date classes of these records, however, show that it is even less presentthan a first glance might suggest. Across NI only one record in Tyrone has the most recent 'post-86'date class; three records date from 1970-86 and eight other hexads are of pre-1970 vintage.
L. officinale appears to have declined everywhere in Ireland and it most certainly has in NI.The Flora of Northern Ireland web site describes it as a rare plant of quarries and rocky places,ie strictly open habitats where it can avoid competition to some degree. Nowadays, it exists only as apioneer colonist of disturbed ground, or as a ruderal species. Mackay (1825, 1836) pointed in much thesame direction, "Dry, waste and uncultivated places, and among rubbish.", although at the timehe added, "frequent".
Being confined to dry, disturbed ground makes it very likely that Common Gromwell is at its 'last stand',on the downhill road towards local extinction. The species appears insufficiently competitive tomaintain even this role in the long term without repeated fresh introductions to the wild from medicinalherb garden stock, and the plant is no longer commonly cultivated, if at all.
British occurrence
In Britain, it is also widely distributed but a decidedly local species of lowland areas lying Sof a line from the Tyne to the Severn. It is scarce and coastal in Wales and the SW of England, while inScotland it is again very rare and thinly scattered (Perring & Walters 1976; Preston et al.2002). There has been a marked, prolonged decline in the species across B & I, especially inScotland and N England where major losses occurred pre-1930 (D. Welch, in: Preston et al. 2002).
European and world occurrence
L. officinale appears to be native in large parts of temperate Europe and W & C Asiastretching eastwards to Siberia and the Himalaya, although some occurrences are certainly adventive herbgarden escapes (Hultén & Fries 1986, Map 1539). It is absent from Crete, the eastern Mediterraneanisles and N Africa, but has been introduced in C & S Africa, Ethiopia and also in N America,including Barbados. Although it has been recorded well towards the north in coastal Norway, it is absentfrom Orkney, Shetland, Iceland and the Faeroes (Hultén & Fries 1986).
Uses
The plants contain silicic acids and calcium compounds that are the active ingredients in its medicinaluses. The Latin specific epithet of this plant informs us that L. officinale had uses in herbalmedicine and it appears it was Dioscorides himself who recommended it as a cure for internal stones,referring to the fruits as 'stone seed'. The hard-coated seed provided the clue to this medicinal use,following the well-known 'Doctrine of Signatures'.
The mature nutlets are the useful part of the plant and, when ground to a powder, they are both diureticand labor inducing (Launert 1981). If seed is not available, the whole plant can be substituted, but itis not as effective (Culpeper 1653). Report of the medicinal treatment of kidney stones appears inIreland (Co Londonderry) by 1834, but its use for this purpose must have been practised very muchearlier. A record from Meath – a region where L. officinale was at one time locally abundant –referred to it as 'Grumble seed'. It is reported that it was collected along the banks of the RiverBoyne and the fruits boiled to treat kidney trouble, presumably again treating kidney and bladderstones. Mixed with a little water, it was also taken for arthritis and febrile conditions (Launert 1981;Allen & Hatfield 2004). "The powdered seed mixed in women's breast milk, is very effectual toprocure a very speedy delivery to such women as have sore pains in their travail, and cannot bedelivered." (Culpeper 1653).
Common Gromwell appears in a list of medieval garden plants of NW Europe, making its first writtenappearance in 1265 AD (Harvey 1981). Thus there are good grounds for suggesting that L.officinale might have been introduced and cultivated by herbalists throughout B & I, and itmight possibly have been used medicinally for many centuries.
Names
The genus name 'Lithospermum' is from Greek 'lithos', 'stone' and 'sperma', 'a seed', given byDioscorides. The Latin specific epithet 'officinale' meant the plant was kept at the druggist's 'shop'(officina), indicating that it was used medicinally (Gilbert-Carter 1964). The English common name'Gromwell', in more modern books, was also spelt 'Gremil' and 'Gromall' by Lyte (1578). Furtheralternatives were 'Grommell', 'Gromaly', 'Gromyl', 'Gromyll' and 'Greymile' or 'Grey Myle' (Turner1548). Turner says that Lithospermum was called 'Herbaries Milium solis', 'Millet of the sun', or'Granum solis' on account of its hard, glistening or shining seeds.
Grigson (1955, 1987) gives the Old French name derived from this as 'Gremil', and in modern French as'Grémil'. The first syllable 'gré-' suggested 'grey', and the second syllable '-mil' is from the Latin'milium', 'millet', referring to the hard seed. Other English common names include 'Lichwale','Lithewale' or 'Lychwale' and 'Lychworte' (from the first edition of the Grete Herball (Anonymous1526)), 'Little Wale', or 'Littlewale'. Some of these are probably misprints or mis-copying in oldherbals! 'Pearl-plant' and 'Stonyhard' both refer to the hard, white nutlets of the species (Britten& Holland 1886).
Threats
None.
Introduction, neophyte, a very rare casual garden escape or discard. European temperate, absent as anative from much of W Europe but frequently cultivated and naturalised.
24 December 1996; Northridge, R.H. & Northridge, Mrs H.J.; wet woods at the base of low but steepportion of cliffs of Pollaphuca (or Poulaphouca).
Growth form
Lungwort is an low, erect, tufted, shortly creeping, bristly, herbaceous perennial up to 30 cm tall withlarge, oval basal leaves, heavily marked with large, conspicuous white blotches and spots (supposed bythe 'Doctrine of Signs' to resemble diseased lung tissue) and hence its scientific and English commonnames. Basal leaves 10-20 × 4-10 cm, elliptic to broadly ovate, margins entire but undulate and narrowedor cordate at the base with a winged petiole. The leaves and stems are densely covered with short, hard,stiff hairs, making them feel very rough to the touch. The inflorescence is a short, congested cyme ofbisexual, heterostylous flowers, 13-18 mm long, 10 mm across, pink in bud, turning bluish when open(Huxley 1967).
Variation
There is a considerable range of variation in the genus Pulmonaria, and in P. officinalis,for instance, the development of the white blotches on the leaves is variable, even within a singlepopulation. Other variable matters are the rosette leaves (summer leaves) that develop during flowering,the effect of dimorphic heterostyly on flower size and stamen insertion, and the interior of the corollatube, which may be glabrous or hairy below the tufts of hairs at the corolla tube mouth (H. Merxmϋler& W. Sauer, in: Tutin et al. 1972, Flora Europaea 3).
Some taxonomic treatments consider P. officinalis contains two subspecies (subsp. obscura(Dumort.) Murb. and subsp. maculosa (Hayne) Gams.), but they are also regarded by others asseparate species, respectively P. obscura and P. officinalis. The key point between thetwo is the degree of white spotting on the leaves, P. obscura being unspotted or with faint greenspots (Sell & Murrell 2009). The sterility of the hybrids between these two is considered sufficientto recognise the two as species (H. Merxmϋler & W. Sauer, in: Tutin et al. 1972).
Origin, introduction and preferred habitats
Lungwort is a popular garden plant introduced from C Europe to gardens in B & I sometime around thelate 16th century or earlier. Despite its botanical and English common names, it was more grown fordecoration than medical or culinary use. In English folklore, Allen & Hatfield (2004, p. 207) couldfind only rare mention of its medicinal use in Hampshire and Norfolk, and no reference to it at all inIreland. The species was first recorded in the wild in Britain in 1793 and it now regularly crops up asan established, naturalised escape or discard in damp, ± open, shaded or semi-shaded conditions,preferring but not confined to rich, deep, humus soils, generally over limestone (Blamey &Grey-Wilson 1989). Habitats include woodland, scrub, roadside hedges and banks, and on rough ground,always in lowland situations (FNEI 3; Clement & Foster 1994; D. Welch, in: Preston etal. 2002).
Fermanagh and Irish occurrence
In Ireland, it is only ever a casual garden escape or discard and it does not persist for long. The tworecords in the Fermanagh Flora Database are most remarkable for the fact that they were recorded in aquite remote site in 1996 around Christmas time! The second record was made four days after the firstdiscovery listed above, further west along the same range of cliffs. The details of the second recordare: wet woods at the base of cliffs, W of waterfalls at GR H0577, Cliffs of Magho, 28 December 1996,RHN.
Taken together, the Cen Cat Fl Ir 2 and the Cat Alien Pl Ir list at least one record ofP. officinalis from five other Irish VCs. These include ground around Co Dublin (H21), but aremainly in the north of the island (ie in Cos Tyrone (H36), Down (H38), Antrim (H39) and Londonderry(H40) (FNEI 3).
British occurrence
The New Atlas hectad map showed P. officinalis much more widely recorded in Britain thanwas the case in the 1962 BSBI Atlas. It is now particularly frequent in S England, where thereare very many active recorders.
The increased coverage displayed in the New Atlas hectad map undoubtedly reflects both increasedabundance of the plant, as well as more intensive recording of introduced alien species than previouslywas the case (D. Welch, in: Preston et al. 2002).
European and world occurrence
P. officinalis, rather than P. obscura, is the form that is found across most of B & I.It is distributed in mainland Europe in a tightly restricted manner from the Netherlands and S Sweden toN Italy and Bulgaria (Hultén & Fries 1986, Map 1542). The map just mentioned has an incorrectrepresentation of the B & I occurrence of the species, showing it confined to SE England. The Hultén& Fries (1986) map for P. officinalis subsp. obscura (Map1543) shows it widespread in Europe from the Alps eastwards into W Asia and north into S Scandinavia.This latter subspecies or species is also shown as an introduction further east into C Asia, and SEaround the Black Sea area.
Names
The genus name 'Pulmonaria' is from the Latin 'pulmo' or 'pulmōnes', 'pertaining to the lungs',the idea being the blotched leaves looked like lung tissues and that this was a 'signature', suggestingthe plant was useful for treating pulmonary disorders. The Latin specific epithet 'officinalis' meant'kept at the druggist's 'shop'' (officina), ie used medicinally (Gilbert-Carter 1964).
There are a large number of English common names associated with P. officinalis, 27 beinglisted by Vickery (2019) and 28 by Brittain & Holland (1886). Many names refer to the white blotcheson the leaves, including 'Lady Mary's tears', 'Lady's Milk-sile' (sile meaning soil), 'Lady'sPincushion' (the white spots resembling pins'-heads on a cushion), 'Mother Mary's Milk', and 'VirginMary's milk-drops', 'Virgin Mary's tears', all referring to legends of Mary feeding milk or weeping overher son. Another feature often referred to in common names is the bicoloured flowers, with names such as'Adam and Eve', 'Joseph and Mary' or 'Joseph and Maries', 'Joseph's coat of many colours', 'Soldier andhis wife', 'Soldiers and Sailors'. The association with the Virgin Mary was on account of the blue andpink flowers, these two colours being the colours of Mary's clothes in medieval paintings (Vickery2019).
Threats
None.
Possibly native at some sites, but also an introduction being an occasional, well-established gardenescape. European temperate, but also in C Asia and widely naturalised elsewhere, including in eastern NAmerica and New Zealand.
1900; Praeger, R.Ll.; Co Fermanagh.
April to October.
Growth form and preferred habitats
This vigorous perennial, has a branched rootstock that is thick, fleshy and tuber-like. From this ariseserect, branched stems 10-150 cm tall, bearing large, ovate-lanceolate, entire leaves, 15-25 × 1-8 cm,bristly hairy (ie setose) and very rough to the touch. Stem leaves are smaller, sessile, the leaf basestrongly decurrent, forming wings on the stem that extend down more than one internode. It flowers inlate April or early May onwards, the inflorescence being a terminal, bract-less, scorpioid (ie curled)cyme of pendulous, bisexual flowers. The bell-shaped, campanulate corolla is variable in colour, usuallypale- or creamy-yellow, but it can also rarely be purplish or pink, or occasionally striped dark andlight purple, rarely white making identification from the more tubular, but very variable hybrid, S.× uplandicum (Russian Comfrey) somewhat more difficult (F.H.Perring, in: Rich & Jermy 1998; Sell & Murrell 2009; Stace 2019). The flowers are insectpollinated and the ripe fruit nutlets are smooth and shining (Garrard & Streeter 1983; Parnell &Curtis 2012).
There are just two estimations of seed longevity in the survey of soil seed banks of NW Europe and theyboth indicate the seed is transient, surviving one year or less (Thompson et al. 1997). The rootsare very brittle and the plant can regenerate very quickly, even from a very small portion of root(Grieve 1931).
Across B & I, S. officinale appears to prefer damp or wet waterside habitats with fertile,neutral or lime-rich soils. These include fens, marshes, river-, stream- and canal-banks, as well asbeside wet ditches and on damp roadside verges (Garrard & Streeter 1983; D. Welch, in: Preston etal. 2002). The species is vigorously competitive and it can quickly develop a sizeable clonalclump of vegetation in sun or half-shaded, sheltered situations. It is intolerant of grazing (Sinkeret al. 1985; Ellis 1993).
Occasional planted in an ornamental garden setting, although much more often the hybrid S. × uplandicum is the preferred planted subject, S. officinale isnot fussy regarding soil type, provided it is neither too dry nor too greatly disturbed. Having saidthis, once introduced to a garden, it is very difficult to subsequently extirpate it on account of itsvigorous vegetative reproductive ability.
Nowadays, Comfrey is frequently grown as a useful manure yielding plant. The plant tissues are rich inminerals including potassium, phosphate and calcium. Cut stems are steeped in water for around fourweeks and the resultant liquid used as a plant fertiliser, especially beneficial for the growth of softfruit and potatoes. Comfrey is also useful as an accelerator for compost making(https://mantis.uk.com/nettle-comfrey-perfect-organic-treatment-spring/#:~:text=Comfrey%20manure%20accelerates%20cell%20division%2C%20favours%20flowering%20and,plants%20such%20as%20red%20berries%2C%20tomatoes%20or%20potatoes,accessed 16 February 2022).
The established strategy of the species is categorised as C/CSR, ie intermediate between a Competitor anda more general non-specialist Competitor-Stress-tolerant-Ruderal (Grime et al. 1988, 2007).
Variation
Two subspecies are recognised in Flora Europaea 3, subsp. officinale with the leavestypically decurrent down the stem, and subsp. uliginosum (A. Kerner) Nyman, with leaves, even theuppermost, not or shortly and narrowly decurrent. Subsp. uliginosum is confined to east centralEurope whereas subsp. officinale is more widespread (B. Pawlowski, in: Tutin et al. 1972).On the other hand, Sell & Murrell (2009) feature two varieties, described as var. officinale(= S. officinale var. purpureum Pers.), with flowers purplish or pink, and var.ochroleucum DC. with flowers white or creamy-yellow, sometimes tinged purplish.
Stace (2019) also recognises two subspecies, subsp. officinale, often >1m tall, corolla >16mm, variable in colour – cream, purplish or sometimes striped dark and light purple, rarely white; andthe much rarer, very restricted, subsp. bohemicum (F.W. Schmidt) Čelak., stems shorter, usually<1m, corolla <16 mm, pale cream.
The Royal Horticultural Society's Index of garden plants (Griffiths 1994) describes the flowercolours as white, pink or purple-violet and lists cultivars of subsp. officinale as cultivar'Bohemicum' to 30 cm, flowers off-white, occasionally tinged purple, and cultivar 'Variegatum' withleaves edged white and cream, flowers cream, but red in bud. The Index also mentions subsp.uliginosum, describing it as, "stem and leaves sparingly setose, denselyverrucose-hispid".
Fermanagh occurrence
S. officinale has been recorded in 36 Fermanagh tetrads, 6.8% of those in the VC. It is occasionaland widely scattered throughout the county, but it is principally associated with out-of-the-waystream-sides and waste places where garden refuse is discarded. The tetrad map shows that it isparticularly frequent along the Colebrooke River and its tributary streams. It is also found in roughgrass in damp meadows and along roadside verges.
S. officinale is just possibly native along some streams and rivers in the VC, but it morefrequently and more widely occurs as an established alien, largely due to the deplorable, yet verycommon practice of unauthorised, illegal dumping of garden waste in semi-concealed sites. All too often,this dumping occurs over bridges, in ditches or on bog margins in rural districts. Discarded tubers maythen spread the species downstream along these various waterways. Trade and transport of top soil mayprovide a secondary means of dispersal for tubers of S. officinale, and the species can persistfor many years and develop large clonal patches in a variety of damp to wet, rough ground habitats.
British & Irish occurrence
In field recording, S. officinale is often confused with the more commonly planted S. × uplandicum, so that the data available in both BSBI Atlases(Perring & Walters 1962; Preston et al. 2002) need to be viewed with the possibility of thisconfusion and that S. officinale has been over-recorded borne in mind. Having said this, theNew Atlas hectad map shows S. officinale as being locally frequent and widespread acrossdamp to wet habitats in lowland parts of Britain from Cornwall to Inverness. Fossil evidence is scant ornon-existent at the species level (Godwin 1975) and the native area of S. officinale in B & Iis unknown. The species may well be an alien garden introduction across N & W Britain and inIreland, although the map depicts it as native everywhere (D. Welch, in: Preston et al. 2002).
When considering status, it is important to bear in mind that Comfrey has a long association withfolklore and medicinal herbal use (see under Russian Comfrey below).
European and world occurrence
The species is considered to have originated in SE Europe and W Asia and since it was primarily used as aherbal medicinal plant cultivated in gardens, it has been spread widely by man into N Europe (includingIceland and N Scandinavia), eastern N America and New Zealand (Hultén & Fries 1986, Map 1546).
Names and uses
The genus name 'Symphytum' is from the Greek 'symphy-' meaning 'growing together' (Gledhill 1985), areference to the wound-healing medicinal properties of the herb known to Dioscorides and herbalistssince Classical times. The Latin specific epithet 'officinale', refers to the herbalist's 'office' orpharmacy.
There are numerous English common names for the herb, 13 in Britten & Holland (1886), 12 in Grieve(1931), 10 in Grigson (1955, 1987) and 14 in Vickery (2019) and they do not all repeat oneanother and overlap! Most of the names refer to the knitting together of wounds including broken bones,eg 'Knitbone' and 'Boneset', the leaves being bandaged over the break (Vickery 2019). The name 'Comfrey'is a corruption of 'con firma' in allusion to the uniting or knitting of bones (Grieve 1931). The plantwas also used in various ways to ease the pain and heal sprains, bruising, arthritis, skin ulcers,boils, cuts and grazes (Vickery 2019). The chief constituent of Comfrey is mucilage which it contains inabundance. It is also mildly astringent and an expectorant. It was used as a demulcent in treating lungtroubles and for quinsy and whooping cough (Grieve 1931).
The name 'Abraham, Isaac and Jacob' given by Grigson (1955, 1987) refers to the variation in colour ofthe flowers, but other names he lists, such as 'Church Bells', 'Coffee Flowers', 'Snake' and 'GooseberryPie' appear more obscure. The latter name was also given to Epilobium hirsutum (GreatWillowherb), apparently a reference to the smell of the leaves (Britten & Holland 1886).
In the past, the plant was widely recommended as a green food for most animals and was said to be bothpreventive and curative of foot and mouth disease in cattle. If grown as a crop for harvest, it couldproduce sufficient growth for two large cuts per year. It was found that horses, cattle and pigs wouldeat Comfrey, but they never took kindly to it as forage. Pigs took best to it as fodder in the greenstate, but even they took some time to get used to it. The feeding value of Comfrey proved little betterthan grass and the species is not adapted for growth on dry or poor land, so its use as a fodder cropwas brief (Grieve 1931).
Threats
None.
Introduction, neophyte, garden escape, occasional.
1939; Praeger, R.Ll.; Newtownbutler.
April to November.
Rather surprisingly, in view of the (at least) eight toxic alkaloids it contains, this erect,much-branched perennial up to 1.5 m tall was introduced and cultivated in gardens as a decorative plantin the early 19th century and certainly by 1827 (Cooper & Johnson 1998; Stace et al. 2015).In addition, hybrid material was introduced from Sweden by the agronomist, Henry Doubleday, andenthusiastically promoted and marketed as the forage crop 'Russian Comfrey'. As food, the bristly hairyplant proved unattractive to most farm animals, with the possible exception of pigs (Grieve 1931). Itwas first recorded in the wild at Marlborough, N Wiltshire, in 1861 (Wade 1958). These plants, togetherwith further introductions, gave rise to many naturalised populations across B & I.
S. × uplandicum is a hybrid between the possibly native S.officinale (Common Comfrey) and the definitely introduced, sky-blue flowered, S. asperum(Rough Comfrey) which originated in NE Turkey, the Russian Caucasus and Iran. The hybrid is still foundnaturalised around farms and remains cultivated in many gardens, it is just a little more frequent andwidespread than Common Comfrey in Fermanagh. Being closely related, Common- and Russian Comfrey bothoccur in very similar ecological situations, ie in damp, rough ground where garden material isdeposited, along stream-sides, roadsides, waste ground, dumped soil and on woodland margins. The hybridcan form dense, often rather uniform stands on damp, fertile soils in these types of habitats (Staceet al. 2015).
In Fermanagh, S. × uplandicum has been recorded in 40 tetrads (7.6% ofthose in the VC. As the tetrad distribution map shows, it is widely scattered, but is chiefly found inthe lowlands E of Lough Erne. Although S. × uplandicum readilyestablishes and persists, it does not appear to spread to any great extent. This hybrid has beenrecorded in 35 of the 40 Irish VCs (Cat Alien Pl Ir) and probably it is still being introduced tonew areas and is also spreading itself to some limited extent.
The hybrid can be distinguished from S. officinale by the stem leaves being less decurrent than inthe species, ie the wings on the petiole and stem of the hybrid do not extend below the next leafdown the stem. The flower colour of the hybrid is also more variable, either pinkish-blue, violet orpurplish or starting rose, turning bluish, or whitish, ± tinted pale rose and blue (Sell & Murrell2009). Despite the corolla colours of the parent species, surprisingly the hybrid never has the blue orreddish-purple that they display, nor the cream colour of S. officinale. Moreover, the flowers ofS. × uplandicum often change colour as they mature, turning from morereddish to more bluish (Stace et al. 2015).
There is no conclusive evidence for the in situ production of S. ×uplandicum F1 hybrids in the wild in B & I. However, S. × uplandicum does backcross with S. officinale and produce arange of intermediate plants that complicate identification to some degree (Stace et al. 2015).
The hybrid distribution in B & I, as shown in the New Atlas hectad maps, is now quiteindependent of its parents and is considerably more extensive than the occurrence of S.officinale, while S. asperum is very rare (D. Welch, in: Preston et al. 2002).Unfortunately, identification confusion between S. officinale and S. × uplandicum has led to an unknown degree of over-recording of theformer and this needs to be borne in mind when comparing maps (D. Welch, in: Preston et al. 2002;Stace et al. 2015).
Like other Comfreys, the plant has long been valued by herbalists for treatment of bruises, sprains andarthritis, and as a general wound master-healer. More recently, Russian Comfrey has been favoured byorganic gardeners who grow it, and then cut and steep the foliage for up to a month to produce a liquidmanure, or else dig the foliage into the soil, or add it to developing compost heaps in order toaccelerate the decomposition process (Vickery 1995; Mabey 1996).
Threats
None.
Introduction, neophyte, a very rare garden escape or discard, presumed locally extinct.
1951; MCM & D; riverbank near Pollboy Bridge, Colebrooke River.
Like the hybrid Russian Comfrey, the single Fermanagh occurrence of this rhizomatous, hispid perennialspecies 20-50 cm tall, found by Meikle and friends is derived from material of garden origin dumped in arural riverbank situation. It flowers in June and July, the corolla is a pale creamy yellow and thecalyx is more deeply divided than that of the other two much more frequent, taller Comfreys in the VC,the lobes being free for at least three-quarters of the way towards the base (An Irish Flora1996). It is also distinguished from S. officinale by its lower stature,scarcely-branching habit and the middle stem-leaves, not the lowest, being the largest (Garrard &Streeter 1983).
There is just the one record in Fermanagh as listed above. S. tuberosum has not been seen in theVC for more than 70 years and is, therefore, presumed to be locally extinct. The species has a similarstatus elsewhere in Ireland, where it has been recorded as an occasional garden escape or relict ofcultivation in open, disturbed habitats such as roadsides, hedge-banks, woodland margins or clearings,plus on waste ground and in quarries along with obviously dumped material. The Cat Alien Pl Irlists S. tuberosum as having occurred at least once in thirteen Irish VCs, not countingFermanagh. The FNEI 3 lists S. tuberosum as having naturalised and persisted at two sitesin Co Down (H38): for at least 33 years at Conlig, and 81 years in the Clandeboye estate woods.
The survival and local increase of this perfectly hardy species anywhere in B & I is almost certainlychiefly the result of vegetative growth of its creeping rhizome, rather than seed production, sincethere does not appear to be evidence of jump dispersal. There are, however, rare reports of large,dominant patches probably developing over several decades in parts of S England, for instance in S Hants(VC 11) (Brewis et al. 1996).
The New Atlas map for Ireland plots a total of 21 hectads with the most recent date class(1987-1999) having at least one record. The hectads are rather scattered but are somewhat moreconcentrated the NE of the country and, in a general manner, in the northern half of the island.
Again, as the New Atlas indicates, the occurrence of S. tuberosum in Britain is very muchmore frequent and widespread than in Ireland. This is especially so in lowland Scotland from Glasgow andEdinburgh north to Aberdeen and Inverness. This particular swathe of Scottish records is plotted as ifthe species is native, whereas, elsewhere in Scotland, S. tuberosum is considered alien,appearing thinly scattered and becoming more coastal in the W and the far NE of the country. In itssupposed native range, Tuberous Comfrey occurs in damp woods, riverbanks and ditches. In England andWales, south of a line between Morecambe and Alnwick, S. tuberosum is thinly and widelyscattered, and, again, is considered an alien garden escape or discard. It the regions where it isconsidered alien, S. tuberosum grows mainly in disturbed, open, artificial habitats such asroadside verges and waste places (D. Welch, in: Preston et al. 2002).
Beyond B & I, S. tuberosum belongs to the European temperate phytogeographical elementand occurs in W, C & S Europe and NW Anatolia (Sell & Murrell 2009)
Threats
None.
Introduced, neophyte, a very rare, but sometimes persistent, naturalised garden escape.
21 May 1999; Northridge, R.H.; base of wall, Old Crom Castle.
April and May.
Origin, introduction and possible use
This leafy, 'almost-wintergreen', bristly hairy perennial which bears rather small bright-blue,white-eyed, flowers like a 'Forget-me-not' (Myosotis spp.) on short side branches, half-hiddenamongst the upper leaves during April to June, is a somewhat surprising, if not to say unworthy, gardensubject (Proctor & Yeo 1973; Stace & Crawley 2015). A native of SW Europe, ranging from CPortugal to SW France (Tutin et al. 1972), P. sempervirens was first introduced tocultivation in B & I at least by the late 16th century (Harvey 1981, p. 166). Possibly – or evenprobably – it was valued more for dyeing purposes than as a decorative garden ornament (Grigson 1955,1987) and, indeed, many of the herbals and older gardening books have nothing to say about its virtuesand they give us no clue whatsoever as to why it was cultivated. Neither is there much evidence of itsuse for dyeing, except perhaps as a food colorant or artists' water colour paint (Gerard 1633).
It is often difficult to decipher old herbal writers and identify which of our modern species they arediscussing. The current author (RSF) believes Gerard and Johnson in the second edition of Gerard'sHerbal (1633, pp. 796-7) refer to this perennial species as 'Borago sempervirens', andthey linked it to the herbal medicinal virtues of Borago officinalis (Borage). Other local Florawriters carrying out this exercise appear to have settled on different members of the Boraginaceae, egconnecting P. sempervirens with Gerard's 'Alkannet' or 'Wild Bugloss' and Anchusa species,although the four kinds of alkannet described in Gerard's herbal do not match this particular species atall.
However, it is still possible to find references drawing attention to the deeply penetrating and brittletaproot of P. sempervirens as a source of red dye (Phillips & Rix 1991a, 1, p. 73) andsuggesting Green Alkanet might have been a mediaeval substitute for Alkanna tinctoria (Alkanet),which itself is a poor replacement for the small tree Lawsonia inermis, from N Africa, the sourceof Henna dye (Grigson 1955, 1987).
Dispersal, colonising ability and preferred habitats
An early indication of the colonising ability of P. sempervirens was shown by the fact that by1724 it had already 'jumped the wall' and been recorded in the wild (D. Welch, in: Preston et al.2002). The species has no obvious means of dispersal and is not mentioned in Ridley's magnificent 1930survey The dispersal of plants throughout the World. Despite this lack, it appearsperfectly capable of travelling 'beyond the garden wall' to colonise open, more or less disturbed, fullylit or preferably lightly shaded ground, on mesic, moist to damp, near-neutral, moderate to richlyfertile soil (Hill et al. 1999).
Typical habitats are generally located near habitation and they include waysides, the base of walls,waste places, hedgerows and the margins of woods, scrub, rivers and streams. Some years ago, GreenAlkanet invaded RSF's Belfast garden, arriving unannounced in a moderately shaded bank under matureBirch and Rowan trees. It is hardy down to around -10°C (Phillips & Rix 1991a).
Although the species carries the Latin specific epithet 'sempervirens' meaning 'evergreen', itdoes in fact die down completely and disappear underground in the depths of winter, re-emerging after afew months in early spring.
Flowering reproduction
Flowering occurs from March or April onwards into autumn, with crop after crop of blooms on the sameplant (Hutchinson 1945, 1972). The flowers are small, bright blue with a white eye, and arranged inone-sided cymose clusters in the axils of the upper leaves. They attract bees and other insects aspollinators which work the circular 'rotate' flowers with their nectar-concealing scales in the shortcorolla tube rather like a revolver, having five separate nectar chambers, each of which has to beseparately approached from the correct angle (Proctor & Yeo 1973). The fruit of each flower consistsof four separate, shortly stalked, achenes (single-seeded dry fruits) or nutlets, which are covered withfine net-like markings (Melderis & Bangerter 1955; Clapham et al. 1962). Flowering and seedproduction is prolific and the species is highly competitive, so that colonisation of suitable openhabitats near gardens or existing colonies is often possible. The established strategy of the plant iscategorised as C/CSR, ie intermediate between outright Competitor and a more balanced mixture ofCompetitor-Stress-tolerant-Ruderal (Grime et al. 1988, 2007).
British and Irish occurrence
Nowadays, P. sempervirens is a widespread naturalised garden escape in lowland Britain, yet whilethe species is particularly widely occurring in the more densely populated areas of England and Wales,it really is frequent only in SW England and is much more local or even rare elsewhere. Until the 1960s,British Floras and flower books continued to regard the species as possibly native in SW England(Melderis & Bangerter 1955; Clapham et al. 1962). Further north, and in Scotland, the speciesdistribution becomes much more scattered, scarce and coastal (D. Welch, in: Preston et al. 2002).
The Irish distribution of P. sempervirens is very sparse and scattered compared with theoccurrence in Britain, but it again reflects the areas of greatest human population density in the E& S of the island, around the larger cities of Belfast, Dublin and Cork (New Atlas; CatAlien Pl Ir). Apart from its very rare occurrence in Fermanagh described below, further east inNI, P. sempervirens is a very much more common garden escape, fully naturalised in semi-shadedwaysides, woodland and scrub margin habitats, but still chiefly occurring close to habitation.
However, since P. sempervirens is a perennial that seeds itself readily and prolifically, andbeing deep-rooted it is persistent once established, the species has the potential to become invasive,sometimes forming monoculture stands and becoming locally dominant (Stace & Crawley 2015). Despiteits prolific level of seed production, the features that may be restricting its colonising ability arethe limited means of seed dispersal, together with the lack of any obvious means of vegetative increaseand secondary spread.
In Britain, accounts in a few local Floras indicate P. sempervirens is no longer confined todisturbed habitats near habitation but, instead, appears to have become more actively invasive since the1960s, colonising a wide variety of lightly shaded sites in woodland, scrub and along rivers (Swan 1993;D. Welsh, in: Preston et al. 2002; Crawley 2005). The change in the British flora1987-2004 survey found a marked increase in the species presence over the relevant period, arelative change of ±10%. This level of rapid increase is rather surprising, considering the length oftime the species has been present in the wild in the country (Braithwaite et al. 2006).
An alternative explanation for the current distribution is that Green Alkanet quickly becomes a difficultweed to manage in garden settings and then is dug out and discarded with other fly-tipped gardenoutcasts around village margins, in addition to self-sown individuals on roadside banks and untendedwaste ground. In London, P. sempervirens is now the ninth most frequent alien found growing onwalls (Stace & Crawley 2015).
Fermanagh occurrence
In Fermanagh, an extensive colony of Green Alkanet was first discovered in 1999 near Old Crom Castleunder trees above the southern estate boundary and on disturbed ground below this old castle wall. Ithas persisted there for at least eleven years. One theory as to its occurrence at this site assumesthat, like its relative Cynoglossum officinale (Hound's-tongue), P. sempervirens mightpossess prolonged seed longevity and therefore persist many years dormant in the soil (Roberts &Boddrell 1984). However, there does not seem to be any published evidence of seed longevity in thisspecies (eg there is no mention in the survey of species in NW Europe (Thompson et al. 1997)).
Alternatively, the plant might have survived overlooked in what is neglected ground for a long timebefore coming to someone's attention. Repair work had just been completed on the wall when the plant wasfirst noticed and it is very possible that the soil seedbank may have been disturbed during this work, resulting in germination and the reappearance of theplant. It is also feasible that the workers might have inadvertently introduced fresh seed on theirtools or attached to their clothing.
The second, and so far only other appearance of the species in the VC, at Durraghstown, SW of Ballyreagh,SE of Enniskillen in April 2000, was also discovered by RHN. The dates of the two records suggests thatP. sempervirens is just beginning to 'jump the wall', spread and establish itself in Fermanagh,in the unknown dispersal manner it employs elsewhere.
European and world occurrence
While its native range on the continent is rather restricted, stretching only from C Portugal to W &SW France (Grigson 1955, 1987; Phillips & Rix 1991a), P. sempervirens is widely cultivatedand is naturalised in NW Europe, N Italy and N America (Tutin et al. 1972).
Names
The genus name 'Pentaglottis' is derived from two Greek words, 'pente' and 'glotta', meaning 'fivetongues', a reference to the number of scales in the throat (or tube) of the corolla. The Latin specificepithet 'sempervirens' translates as 'evergreen' (Stearn 1992) although, as mentioned above, this isinaccurate. The English common name 'Alkanet' is the diminutive of the Spanish 'alcanna', which in turnis derived from the Arabic 'al-henna', the Henna tree referred to above. The 'Little Alkanna' or'Alkanet' was the name applied to the dye plant Alkanna tinctoria, the red roots of which werelong imported from the continent for many dying purposes, including colouring the red liquid in cheapthermometers! Green Alkanet, probably was a substitute for the former (Grigson 1955, 1987). Analternative English common name from Somerset is 'Pheasant's Eye', which Britten & Holland (1886)also dismissively list as, "a book name for Adonis autumnalis". The reference to thebright red eye of the bird is presumably linked to the red dyeing properties of the plant.
Threats
Currently much too rare to be considered a threat in Fermanagh, but P. sempervirens is potentiallyinvasive whenever it becomes thoroughly established at an unknown threshold number of sites. Existingpopulations therefore need to be kept under review.
Native, frequent or locally common. Eurosiberian temperate, but very widely naturalised, including inboth N & S America.
1900; Praeger, R.Ll.; Co Fermanagh.
May to November.
Growth form and preferred habitats
This very variable, rhizomatous and with creeping runners, erect but low-growing, shade-tolerant,wintergreen perennial, 15-70 cm tall is frequent to common, especially in the lowland area around bothparts of Lough Erne, plants often standing in water at the margins of larger lakes and ditches, or onwet mud in marshy, lime- or base-rich water meadows, fens and fen-carr.
M. scorpioides is best distinguished from two close relatives that often grow by the water's edge,M. laxa (Tufted Forget-me-not) and M. secunda (Creeping Forget-me-not), by its usuallysomewhat larger flowers, (3-)8(-13) mm in diameter and much longer style protruding beyond the calyxafter the petals drop (Webb et al. 1996; A.J. Silverside & T.C.G. Rich, in: Rich & Jermy1998; Sell & Murrell 2009). However, it has been pointed out that in older cymes the flowers becomesmaller, resulting in overlap between the B & I Water Forget-me-not species with respect to thischaracter. Additional useful distinguishing features of M. scorpioides are the ebracteate(bract-less) cymes and a calyx divided to only 1/3 of its length (Welch 1967).
Water Forget-me-not is regarded as a pioneer coloniser of still or slow-moving, shallow, moderately torichly productive water, occasionally forming floating clonal mats or submerged clumps in somewhatdeeper water. However, this species is also sufficiently vigorous to compete and retain its placeamongst taller growing wetland terrestrial vegetation that shades it, including sedges, reeds, Menthaaquatica (Water Mint), Ranunculus ficaria (Lesser Celandine) and other common associates(Preston & Croft 1997).
M. scorpioides is mainly absent from heavily disturbed habitats and unproductive vegetation and is± restricted to soils of pH>5.0 (Grime et al. 1988, 2007). The established strategy of thespecies is categorised as Competitive-Ruderal (Grime et al. 1988, 2007).
Variation
M. scorpioides is very variable, especially in C Europe and belongs to a species complex of fournamed species – the others being M. rehsteineri Wartm. (a very low-growing, mat forming species),M. nemorosa Besser (a biennial form, usually without stolons) and M. lamottiana (Br.-Bl.Ex Chassagne) Grau (a mountain species from SC France to NW Spain) (J. Gras & H. Merxmüller, in:Tutin et al. 1972).
In B & I, two varieties are named by Sell & Murrell (2009): var. scorpioides with thelower part of the stem glabrous or with spreading hairs; upper part of stem with appressed hairs;flowers 5-8 mm in diameter; and var. strigulosa (Rchb.) Schinz & Keller. with appressed hairson stem throughout; flowers 4-6 mm in diameter. The latter is a more upland plant than var.scorpioides. None of these taxa have been distinguished in Co Fermanagh.
In terms of cytology, two chromosome counts exist for M. scorpioides, diploid 2n=22 and hexaploid2n=66 (Merxmüller & Grau 1963).
Fermanagh occurrence
M. scorpioides is frequent and locally abundant and has been recorded in 198 Fermanagh tetrads,37.5% of those in the VC. It is particularly common, sometimes stand-forming, around the shores of UpperLough Erne and, as the tetrad map indicates, is also widely scattered in suitable damp to wet, fertileground elsewhere in the county.
A high degree of tolerance of the disturbance created by both winter flooding and a fluctuating watertable may well be the key to this species success in the extensive, fertile, lime-rich Fermanagh watermeadows that surround much of Lough Erne, since these conditions tend to restrict the dominance of mosttaller associated wetland species.
Flowering reproduction
As is frequently the case with aquatic species, submerged plants may not succeed in flowering, butemergent and terrestrial plants do so freely. Flowering begins in May and continues into September. Seedis set from August to October (Grime et al. 1988, 2007). M. scorpioides is insectpollinated and mainly out-breeding, but the degree of self-incompatibility in its flowers variesgreatly, some hermaphrodite blossom proving totally self-sterile, while other plants may be up to 95%self-compatible and regularly able to set seed. The progeny of selfed plants may, however, suffer anddisplay inbreeding depression (Varopoulos 1979).
Buried seed survival is variously estimated: of 17 studies surveyed in NW Europe, eleven reckoned M.scorpioides was transient (survived less than 12 months), two considered it long-term persistent(more than 5 years) and four could not decide either way (Thompson et al. 1997).
A study by Praeger (1913) of seed flotation found the small nutlet fruits of M. scorpioides arebuoyant only briefly, for a day or less; they can also be transported when flooding occurs and spates ofrapid flowing water scours fruiting plants (Ridley 1930, p. 218). The presence of Water Forget-me-not inand around isolated ponds is also strongly indicative of nutlet transport by ducks and other waterbirds, most likely in mud adhering to feathers and feet (Ridley 1930, p. 547).
Vegetative reproduction
Both the underground rhizomes and the surface or subterranean stolons of M. scorpioides are shortin length (Sell & Murrell 2009), but nevertheless they are sufficiently efficient to enable thespecies to compete successfully with other waterside associates and to regularly develop clonal purestands. In addition, vegetative reproduction frequently operates through stems breaking off under theforces delivered by rapidly flowing spates of water, or as a result of animal trampling, and thentransported downstream to establish new colonies (Ridley 1930, p. 182, referred to, in this case, asM. palustris). Fragmentation probably provides the most important and effective means of increaseand downstream dispersal in M. scorpioides, surpassing the significance of occasionalseeding recruitment in gaps in established populations, at least when considered in the short term(Grime et al. 1988, 2007).
British and Irish occurrence
As a native species, M. scorpioides is widely distributed across most of B & I, although inOrkney and Shetland it is recognised as an alien introduction (VCs 111, 112). The New Atlashectad map shows it chiefly absent from high ground, from less fertile and more acidic conditions, andespecially from bogland on both islands (D. Welch, in: Preston et al. 2002).
The distribution and frequency of M. scorpioides has remained stable for many decades, althoughthe Change in the British Flora 1987-2004 sampling survey noted a definite decline (survey notapplicable to Ireland), with a calculated Change Factor over the period of -27. (Braithwaite etal. 2006). Apart from loss of suitable habitat due to drainage and development, the major driverof change in wetland sites across Britain has been an increase in nitrogen levels associated witheutrophication. However, M. scorpioides is tolerant of eutrophic waters and is moderately mobile,so it is thought likely it will remain common (Grime et al. 1988, 2007).
European and world occurrence
M. scorpioides belongs to the Eurosiberian temperate phytogeographical element and is widelydistributed in most of Eurasia, but it is absent from Africa. The species s.l. has also been introducedinto N & S America and New Zealand (Hultén & Fries 1986, Map 1561).
Names
The genus name 'Myosotis' is derived from two Greek words 'mus', 'a mouse' and 'otes', 'an ear', ie'mouse ear', the hairy leaves resembling a mouse's ear. The name was given by the ancient Greek,Dioscorides, to an entirely different plant, not our familiar 'Forget-me-not' (Johnson & Smith1946). The Latin specific epithet 'scorpioides' means 'curved like a scorpion's tail' a reference to thecoiled cymose inflorescence (Gledhill 1985).
Threats
None.
Native, frequent. Oceanic temperate.
1900; Praeger, R.Ll.; bogs SW of Belleek.
May to October.
Growth form, identification and preferred habitats
This is a widespread stoloniferous, mat-forming annual or biennial plant that grows up to 60 cm tall. Itpossesses a short, scarcely creeping rhizome and decumbent or prostrate stems arise from leaf axils atthe base of the plant, the non-flowering ones rooting at the nodes and forming stolons. The densespreading hairs at the base of the stem distinguish M. secunda from both of the other wetlandMyosotis species that occur in Fermanagh (M. scorpioides (Water Forget-me-not) and M.laxa (Tufted Forget-me-not)). M. secunda also differs from M. scorpioides inhaving leafy inflorescences and longer flower stalks (3-5 times as long as the calyx), and from M.laxa it differs in having larger, emarginated (ie notched) petals (Welch 1967; Garrard &Streeter 1983; Clapham et al. 1997). M. secunda flowers from May to August, the corolla4-8 mm in diameter, pale blue with a yellow 'eye' around the throat (or entrance) of the flower.Fruiting pedicels up to 10 mm, becoming recurved; nutlets four, blackish-brown, acute, with a rim (Press& Short 1994; Clapham et al. 1997).
M. secunda is a species of wet to moist, moderately acidic to neutral, loamy or peaty soils withsome degree of flushing occurring by springs, streams, drains, pools, on heaths, bogs and marshymoorland pastures (Sinker et al. 1985). It usually occurs on poorer, more acid, colder, moreexposed upland, calcium-deficient soils and peats (Trueman et al. 1995). While it is chiefly anupland species in B & I, M. secunda can descend to sea-level in more western parts of bothislands (Welch 1967).
In moorland situations, M. secunda is tolerant of grazing and trampling, but it is a poorcompetitor against more vigorous tillering plants including grasses and sedges. The established strategyof M. secunda is categorised as a CR (Competitive Ruderal) (Grime et al. 1988, 2007).Although it has the advantage of stoloniferous vegetative spread, like most other colonising species,for establishment to be successful it really requires the open, well-lit, somewhat disturbed, bareground, habitat conditions that grazing animals help provide.
Fermanagh occurrence
M. secunda has been recorded in a total of 107 Fermanagh tetrads, representing 20.3% of those inthe VC. It is particularly frequent in wet swampy ground around upland lakes, ponds, streams and influshed areas of moors and bogs in Fermanagh, as is also the case elsewhere in B & I. However, inFermanagh, Creeping Forget-me-not is by no means confined to higher ground. As the tetrad map indicates,it is widespread and is regularly, although less commonly and much less abundantly found in lowlandsites, eg along riverbanks, stream-sides, peaty ditches and marshy pastures throughout the county.
Despite the above, in Fermanagh, M. secunda is by far the least frequently recorded of the threewetland Forget-me-not species and it has the most restricted distribution in the VC, being only locallycommon. Having said that, M. secunda is the most likely of the three to be found in uplandwetland habitats. Like M. laxa it is tolerant of infertile, acidic wetlands and it is sometimesdescribed as a calcifuge species (Flora of Lough Neagh; Trueman et al. 1995). In terms oflocal records, however, in Fermanagh it has only a third the frequency of M. laxa and it is lessthan half as widely distributed as it is across the VC. Both these wetland Myosotis species areless than half as frequent in the VC survey as the entirely lowland, more nutrient-demanding M.scorpioides, but M. laxa certainly is the most widespread of the three species.
British and Irish occurrence
Common in many parts of B & I, especially upland areas in the N & W, but rare or absent in mostof C & E England, and likewise in lowland C Ireland (D. Welch, in: Preston et al. 2002; Sell& Murrell 2009).
European and world occurrence
M. secunda belongs to the oceanic temperate phytogeographic element and is confined to W Europe,the Azores and Madeira (Clapham et al. 1997; Sell & Murrell 2009).
Names
The Latin specific epithet 'secunda' means 'one sided', probably referring to the coiled cymoseinflorescence having all the flowers facing the same way (Gledhill 1985).
Threats
None.
Native, common and widespread. Circumpolar boreo-temperate, but rather disjunct. The form in B & I issubsp. caespitosa (Schultz) Hyl. ex Nordh.
1881; Stewart, S.A.; Carrick Lough, Dresternan Td.
May to November.
Growth form and preferred habitats
Among the members of the Myosotis scorpioides agg., the B & I wetland waterForget-me-nots, M. laxa is distinct in being a summer annual or biennial, with the hairson the plant always adpressed – ie no spreading hairs, especially on the calyx – and its flowers havinga short style which does not protrude from the calyx when the corolla drops off or is removed (Welch1967; Parnell & Curtis 2012). Stems of M. laxa are pale green, leafy, either simple orbranched from near the base, grow to a height of between 20 and 40 cm and produce hermaphrodite(bisexual) flowers in a terminal cyme from May to August. The corolla is 4-5 mm in diameter, brightsky-blue with a yellow rim at the throat (Sell & Murrell 2009).
Although often occurring together, M. laxa and its common perennial relative, M.scorpioides (Water Forget-me-not) differ fundamentally in their life strategies and dominantmode of reproduction and spread. M. laxa is entirely dependent on seed production for increase,spread and overwintering survival. Both these species are common and widespread in damp to wet, fertile,open, disturbed marshy ground with a permanently high water-table around or along the margins of a widevariety of usually lowland habitats, including wet hollows in fields. Of the two, M. laxa is themore likely to occur in strongly acidic and hilly or upland sites. In lowland situations, it oftenoccurs in trampled, cattle-poached and manured mud silt, clay or peat, or by brackish ditches (Welch1967; Sinker et al. 1985).
The wider, more local distribution of M. laxa undoubtedly reflects its greater range of soiltolerances compared with M. scorpioides, the latter being confined to wetter, more muddy, muchless acidic to alkaline, lowland terrain, usually with a soil reaction above pH 5.0 (Grime et al.1988, 2007). M. laxa is tolerant of light grazing and half-shade, but not competition from tallvegetation. As a ruderal annual, it is sometimes a pioneer colonist of bare mud and associates withspecies such as Alopecurus geniculatus (Marsh Foxtail), Galium palustre (CommonMarsh-bedstraw), Glyceria fluitans (Floating Sweet-grass), Ranunculus flammula (LesserSpearwort), R. sceleratus (Celery-leaved Buttercup) and Veronica beccabunga (Brooklime)(Sinker et al. 1985).
Variation
There are two geographically distinct subspecies within M. laxa. Subsp. laxa is confined toN America, while the B & I and Old World form is subsp. caespitosa (Schultz) Hyl. Ex Nordh.(= M. caespitosa Schultz) (Sell & Murrell 2009).
Fermanagh occurrence
As noted in the account on this website of the ecologically rather similar but definitely perennialpatch-forming species M. scorpioides, this annual or biennial species is the more widespread ofthe three Fermanagh wetland Forget-me-nots, having records in 244 tetrads, representing 46.2% of thosein the VC. The comparable figures for M. scorpioides are 198 tetrads, representing 37.5% ofFermanagh's area, and for M. secunda (Creeping Forget-me-not) 107 tetrads and 20.3% of the area.Identification errors have been made in the past, where small-flowered forms of M. scorpioideshave been listed as M. laxa, over-recording which needs to be borne in mind when comparisons aredrawn.
M. laxa is, however, very comfortably the most widely distributed Myosotis species inFermanagh, but locally it still has less than half the record frequency of the perennial M.scorpioides. This is probably an artefact produced by the huge degree of over-sampling carriedout around Upper Lough Erne by EHS Habitat Survey Teams in 1986-7.
A hybrid
"The species of Myosotis series Palustris, the wetland Forget-me-nots need tobe identified with care, and few observers have learnt to recognise this hybrid." (T. O'Mahony, in:Stace et al. 2015). The hybrid between the two wetland Forget-me-not species, M. laxa subsp. caespitosa and M. scorpioides (M. × suzae Domin), has been found scattered amongst the parents in partsof England and Wales and probably it has been overlooked elsewhere on many occasions, including inFermanagh. The only Irish records are from Co Cork and areunpublished findings by T. O'Mahony (Stace et al. 2015). The hybrid is intermediate between theparent species and has been also been recorded in continental Europe (Sell & Murrell 2009). Patchforming plants that resemble M. scorpioides, but have oddly long racemes that continue to flowerwell into the autumn, should be examined to see if they belong to this partially fertile intermediatehybrid (New Flora of the BI 1997; BSBI Plant Crib 1998; Stace et al. 2015).
British and Irish occurrence
M. laxa subsp. caespitosa is widely distributed across thewhole of B & I, being represented in the New Atlas in a total of 3,175 hectads (167 more thanM. scorpioides), but generally it is not as abundant as the latter. Apart from local losses ofsuitable habitat in SE England, the distribution appears stable, showing little change since 1962 (D.Welch, in: Preston et al. 2002).
European and world occurrence
M. laxa subsp. caespitosa is found in most of Europe and in Asiaextends east to N Siberia, the Himalaya and Japan. It is also present in N Africa and has beenintroduced to New Zealand. The other subspecies, subsp. laxa, occurs in N America. Thus, takentogether, the overall species belongs to the circumpolar boreo-temperate element (Hultén & Fries1986, Map 1560; Sell & Murrell 2009).
Names
The Latin specific epithet 'laxa' means 'open' or 'loose', probably referring to the coiled cymoseinflorescence. The Latin subspecific name 'caespitosa' is derived from two words 'caespes' and 'itis'which translate as 'turf sod', meaning the plant is tufted or forms tussocks, which in this annualspecies is not really the case (Gilbert-Carter 1964).
Threats
None.
Introduction, neophyte, a rare or very rare garden escape. Eurasian temperate, but very widelynaturalised including in N America, Africa and New Zealand.
1946-53; MCM & D; shore of Upper Lough Erne at Crom Castle Estate.
May to July.
Growth form and status
This erect, 15-45 cm tall, pubescent biennial or short-lived perennial is polymorphic and very variable,occurring as a local native species in parts of Britain (see below), but entirely absent as a native inIreland, and likewise rare or absent from most of Wales, Scotland and SW England. Elsewhere in B &I, another form or variety of the species is present as a rather rare garden escape. In Ireland, M.sylvatica is always considered a rare, neophyte, garden escape. The garden form or variety iscommon as an escape in Britain, although it is often a problem distinguishing native from cultivatedforms.
Identification
Small forms of M. sylvatica and large forms of M. arvensis (Field Forget-me-not) are easilyconfused: the larger corolla of M. sylvatica is a good diagnostic character (4-)6-10(-11) mm asopposed to 2.5-5 mm in M. arvensis. In deeply shaded habitat conditions, M. sylvatica canalso be much smaller and more spindly in form than normal plants.
The diameter of the flower is generally used to distinguish M. arvensis from M. sylvatica.The critical measurement is 5 mm across: if larger than this the species is M. sylvatica, ifsmaller then M. arvensis. If the flower being examined is exactly 5 mm in diameter, then one hasto try to distinguish these two species by whether the petal lobes are flat (M. sylvatica) orconcave (M. arvensis) (Garrard & Streeter 1983; A.J. Silverside & T.C.G. Rich, in: Rich& Jermy 1998).
There are many garden varieties offering a range of flower colour, seasonality and plant size. Griffiths(1994) names eleven cultivars, which represent a mere sample of those in the horticultural trade.
Preferred habitats
Typically, M. sylvatica grows on damp, fertile, moderately acid to neutral soils in woodland shadeor in rocky grassland, both habitat situations sharing the ecological feature of sheltering herbs ofrelatively low competitive ability from the aggressive potential of taller, more vigorous species. Theestablished strategy of M. sylvatica is categorized as R/CSR, ie intermediate between Ruderal anda more balanced Competitor-Stress-tolerant-Ruderal ecological approach (Grime et al. 1988, 2007).It can also be found in more open areas of woods, in clearings and on the margins of scrub on chalkgrassland slopes, and along wooded calcareous streams. Having said this, it is not always clear whetherpopulations are native or of garden origin since the two varieties of plant are often indistinguishable.Populations where there are a number of white or pink flowers are probably of garden origin (Brewiset al. 1996).
Flowering reproduction
As one would expect of a species with numerous named varieties of garden merit, M. sylvaticaflowers abundantly, mainly in May and June, but occasionally until September. The basal or lowest leavesare scarcely stalked, usually wither at anthesis (ie flowering time) and disappear (Sell & Murrell2009). The corolla is bright blue with a pale eye, 6-8(-11) mm in diameter, the petal lobes flat, notconcave. The cultivated form has large, even brighter blue, or purple, or often pink, or rarely whiteflowers with a pale yellow eye up to 11 mm in diameter, while the native form has flowers up to 8 mm indiameter, but often much less than this (Sell & Murrell 2009).
The flowers contain nectar and are pollinated by a range of insects, or else they self-pollinate. Eitherway, they generally set large quantities of 'seed' (ie nutlets or achenes) (Clapham et al. 1987).
As with other terrestrial members of the genus (and family), there does not appear to be any obviousspecialised dispersal mechanism for the small 'seeds' (other than wind), yet the plant somehow doesmanage to regularly 'jump the garden wall', so there remains something of a mystery to be solved!
The soil seed bank survey of NW Europe makes no mention of M. sylvatica, and the current author(RSF) has not found any information suggesting seed longevity in the literature available to him. It isprobably transient, surviving less than one year.
Fermanagh occurrence
Although this freely seeding, biennial or perennial garden escape appeared thoroughly naturalised in CromCastle Estate when Meikle and co-workers first discovered it in the 1947-53 period, as the tetrad mapindicates, it has only been recorded in Fermanagh seven times at six well scattered sites since 1975. Ithas not been refound at the original Crom site in the far SE of the county.
Details of the other Fermanagh records are: lakeshore near yacht club marina, Goblusk Bay, Lower LoughErne, 1991 & 1993, I. McNeill; Necarne Estate near Irvinestown, 3 July 1995, I. McNeill; shore atDerrymullan Old Church (ruins), Upper Lough Erne, 20 May 1996, RHN; dump at Blacklands, Clabby Road,1987-99, I. & D. McNeill; roadside at Letterbreen, 21 May 2004, RHN; near Ardress House, 2.5 km SEof Kesh, 2 May 2008, I. McNeill.
Irish occurrence
In Ireland, M. sylvatica is chiefly recorded in northern VCs. Apart from Fermanagh,until recently, isolated finds of M. sylvatica garden escapes have occurred in just five otherIrish VCs (E Cork (H5), Laois (H14), Co Dublin (H21), E Donegal (H34) and Tyrone (H36)). Habitatscomprised a variety of semi-wild, mainly shaded situations and the sites included woodland, river-banks,a shrubbery, hedges and a gravel car park (Cat Alien Pl Ir).
British occurrence
The native range of M. sylvatica has become obscured by the plant mixing with the introducedhorticultural form. Always a rather local species, it is considered that the native area included mostof N England south of the Scottish Border to a line between the Wash to the Severn, plus parts of SEEngland, including Hampshire (Brewis et al. 1996). Within this area, M. sylvatica is aplant of damp woodlands, often on heavy clay or calcareous soils, and it can sometimes be abundantlypresent (Garrard & Streeter 1983; Brewis et al. 1996; Preston et al. 2002).
European and world occurrence
M. sylvatica is present in most of Europe and there are other subspecies or closely allied speciesin the Middle East and W Asia. It is therefore regarded as belonging to the Eurasian temperate element. Cultivated forms have been introducedin many areas of the world, including southern parts of Africa, further east in C Asia, Japan, NewZealand and N America (Hultén & Fries 1986, Map 1557; Clapham et al. 1987; Sell & Murrell2009).
Threats
None.
Introduction, archaeophyte, occasional or local. Eurosiberian boreo-temperate, but very widelynaturalised, including in N America and New Zealand.
1881; Stewart, S.A.; Co Fermanagh.
April to November.
Growth form and preferred habitats
Usually a slender, erect winter-annual, 15-30(-60) cm tall, rough with spreading hairs, that germinatesin the autumn and overwinters as a small leafy rosette, M. arvensis can also sometimes behave asa summer annual or as a biennial. It may very occasionally be confused with smaller specimens of the(usually) perennial, native or garden introduction, M. sylvatica (Wood Forget-me-not),particularly on waste-ground, on dumps and near gardens, from which the latter occasionally escapes.Apart from the generally smaller, more concave corolla of M. arvensis, the most satisfactorydistinguishing character in the field is the appressed (closed) calyx of freshly picked (ie undried,unpressed specimens) of M. arvensis, which conceals the seeds, while that of M. sylvaticais open (erecto-patent) in both fresh and dried specimens, exposing the seeds to view (A.J. Silverside& T.C.G. Rich, in: Rich & Jermy 1998).
M. arvensis is a weedy ruderal annual of open, moderately fertile, fairly dry conditions in a widerange of soils of pH<5.0, in situations where it is protected from heavy grazing and strong levels ofcompetition. It appears locally as scattered individuals on disturbed or otherwise open, sunny, warm,dry or well-drained, sandy, gravelly or stony soils throughout B & I (Grime et al. 1988,2007). Other habitats include woodland margins, open grassland, hedges, scrub, walls, quarries and wasteground (D. Welch, in: Preston et al. 2002).
Previously it was a familiar agricultural weed of both arable fields and grasslands, the nutlets being aregular contaminant of commercial pasture clover and grass seed mixtures (Salisbury 1964). As a resultof progress in agricultural technology and practices, including the massive move from hay towardspasture and silage meadows, better seed screening and the use of agrochemicals, this field weed hasgreatly declined in frequency and contracted slightly in its distribution. It remains mainly as a weedof agricultural land, particularly found in England in cereal crops (Grime et al. 1988, 2007).
Variation
M. arvensis is rather variable and two subspecies and two varieties of one of themexist. Subsp. arvensis has a calyx not more than 5 mm in fruit, and a small corolla up to 3 mmacross. Two varieties of this subspecies are recognised, var. arvensis is a small form, usuallywith a solitary stem up to 15 cm tall, and is the crop weed form, definitely introduced; var.dumetorum is larger, up to 30 cm tall, often with numerous long branches from the base, equallingthe central stem. It appears in more natural, often sandy places and just might be a native plant.
The second subspecies is subsp. umbrata (Mert, & W.D.J. Koch) O. Schwarz which has a largercalyx, up to 7 mm in fruit and corolla 3-5 mm across. The distribution of this form in B & I is notknown, but it appears to be widespread and is said to be confined to W Europe (Sell & Murrell 2009).
Flowering reproduction
As a ruderal annual, M. arvensis reproduces exclusively by seed, flowering in springtimefrom April to July and setting seed from as early as May onwards. M. arvensis fulfils all theproperties of a ruderal or 'agrestal' species – the latter being ruderal weeds that enter agriculturalland; they possess small size, sparse branching, rapid growth and a short life-cycle, plus early andprolific seed reproduction, very often involving self-fertilisation. Ruderal species are associated withor confined to habitats characterised by low-stress, often nitrogen-enriched and greatly altered by highlevels of human disturbance. The term 'ruderal' is derived from the Latin 'ruderis' meaning 'rubble'(Baker 1965; Grime et al. 1988, 2007).
The inflorescence is a terminal bract-less coiled (scorpioid) cyme of bisexual flowers. The calyx isshorter than the pedicel, covered with hooked spreading hairs, its teeth longer than its tube. Thecorolla is small, c 3-5 mm in diameter, pink in bud, then pale sky-blue with an orange-yellow eye at thethroat when fully open, 5-lobed, the lobes concave rather than flat (Webb 1977; Sell & Murrell2009).
M. arvensis flowers freely and sets seed abundantly. Salisbury (1964, p. 170) estimated a largeplant would produce between 1500 and 3000 nutlets, which are sometimes released as a unit, in fourscontained within the calyx. Normally it behaves as a winter annual, seed shed in summer showing delayedgermination until the autumn. Seed appears to be highly mobile, successful dispersal being both internalin the guts of cattle and horses, and external by attachment to humans and the furry coats of passinganimals thanks to the roughly hairy calyx that often contains the nutlets and acts like a burr (Ridley1930; Salisbury 1964). Seeds are also persistent in the soil seed bank for five or more years, which isparticularly significant on arable land, but also important in occasionally disturbed wayside habitats(Grime et al. 1988, 2007; Thompson et al. 1997).
Fermanagh occurrence
In Fermanagh around 2012, habitats and frequency range from rarely in gravelly screes, limestone pavementand pastures, to more frequently in cultivated or disturbed ground, including potato fields, gardens,car parks, quarries, waste or derelict ground (eg old disused railway stations), roadsides and nearhabitation. It is now regarded as occasional in Fermanagh, the plant having been recorded in 80 tetrads(15.2% of the VC total), although only 68 of them have post-1975 records. Despite this, the tetraddistribution map demonstrates that Field Forget-me-not remains scattered widely across the VC, andreflecting past agricultural land use, it remains slightly more frequent in the more fertileagricultural lowlands of the Erne basin. It is believed that the species distribution has beenmaintained by a combination of seed longevity and its flexible ruderal life-history.
British and Irish occurrence
Despite its weedy behaviour, M. arvensis has traditionally and over very many years been regardeda native species in all published Floras in these isles including Stace (1997). The publication of theBSBI New Atlas in 2002 and a subsequent revising paper in 2004 have now declared FieldForget-me-not to be an ancient introduction or archaeophyte (Preston et al. 2002, 2004).
Field Forget-me-not is a very widespread species across Britain except on the high ground and acidpeatlands of NW Scotland. In Ireland, although in the past it has been recorded in every VC (Scannell& Synnott 1987), it is more frequent in the eastern half of the island, becoming more thinlyscattered or absent in the west (New Atlas). The species is seldom if ever abundant nowadays, butany decline has been in numbers, rather than in distribution (Braithwaite et al. 2006).
European and world occurrence
Like other species widely spread beyond its native range by agriculture, the native range of M.arvensis is now impossible to accurately discern. It now ranges across all of Europe, W & CAsia and N Africa, but probably originated somewhere in S & SE of Europe. It spread as an impuritywith commercial crop and grass seed over large areas of W, C & N Europe (including W Greenland &Iceland) and E Asia (Japan), and was also introduced to N America and New Zealand (Hultén & Fries1986, Map 1552).
Names
At the end of the 16th century, Gerard (1597) knew no English names for the three species ofMyosotis for which he gave the first English records, M. palustris, M. arvensis andM. hispida. Since the cyme inflorescences of these three species are coiled, they were placedwith and confused with the leguminous plant Scorpiurus sulcata from S Europe, and all four werenamed 'Scorpion Grass'. The legume was identified with the skorpioeides in the De MateriaMedia of Dioscorides, who likened 'the fruit', the coiled pod, to a scorpion's tail. Thus'Scorpion Grass' became linked to the various kinds of Myosotis in B & I as an English bookname for centuries (Grigson 1955, 1987).
Three other English common names are listed by Brittain & Holland 1886), 'Bird's-eye' (or'Bird's-eyes'), 'Forget-me-not' and 'Blue Mouse-ear'. 'Bird's-eye' is one of the most generally usedplant names, usually applied to plants with small, bright, often blue flowers, such as Anagallisarvensis (Scarlet or Blue Pimpernel), Sagina procumbens (Procumbent Pearlwort),Veronica chamaedrys (Germander Speedwell), etc., plus numerous Myosotis species(Prior 1879; Brittain & Holland 1886).
'Forget-me-not' likewise is applied to several species other than Myosotis species, includingAjuga chamaepitys (Ground-pine) and Veronica chamaedrys. It is claimed that'Forget-me-not' was applied to V. chamaedrys on account of the fused petals and stamens fallingoff and blowing away (Prior 1879, p. 84). Prior provides a very detailed account of the variousapplications and derivations of the 'Forget-me-not' name. He suggests the reason the name was given toA. chamaepitys was on account of the nauseous taste of the plant when used in herbal medicine, soit was unlikely to be forgotten! The name was exclusively applied to this plant by many herbalists fromthe mid 15th century onwards (Prior 1879, p. 83).
'Mouse-ear' is applied to just about any species with conspicuous long hairs on smallish leaves,including Cerastium triviale (= C. fontanum subsp.vulgare), Common Mouse-ear, or Mouse-ear Chickweed; Hieracium pilosella (= Pilosellaofficinarum), Mouse-ear-hawkweed; Myosotis arvensis (Blue Mouse-ear); and Sisymbriumthaliana (= Arabidopsis thaliana), Codded Mouse-ear (a book name), or Thale Cress(Brittain & Holland 1886).
Threats
None.
Native, frequent. European temperate, but favoured by agriculture and widely introduced, including in SAfrica, India, N America and New Zealand.
1881; Stewart, S.A.; Co Fermanagh.
April to October.
Growth form, variation and preferred habitats
This small, hairy annual has two ecotypes or subspecies that occur in entirely different habitats, onedry and the other wet. The more common form (subsp. discolor) is rather similar in form andecology to M. arvensis (Field Forget-me-not), behaving as a typical winter annual and is found ina wide range of open, disturbed and bare ground where the soil is dry, warm, sandy or stony. It behavesas a small, weedy, short-lived, spring ephemeral, up to 30 cm tall, flowering and fruiting early in thegrowing season to avoid summer drought. The established strategy of M. discolour is categorisedas SR, ie intermediate between a Stress-tolerator and a Ruderal species (Grime et al. 1988,2007).
In the past, like Field Forget-me-not, M. discolor subsp. discolorwas a cornfield weed, but modern farming technology has put a stop to this behaviour. Subsp.discolor can be recognised by at least the uppermost pair of stem-leaves being oppositerather than alternate and the corolla is up to 4 mm, yellow at first before turning pink or blue. Insubsp. dubia (Arrond.) Blaise, on the other hand, none of the stem-leaves are opposite and thecorolla is smaller (not more than 2 mm) and is whitish or cream at first (Sell & Murrell 2009).
Subsp. dubia occupies bare patches in wetter, cooler meadow soils, rushy pastures and base-richflushes, where it grows much taller (up to 30 cm or more) and perhaps behaves as a spring or summerannual (Flora of Connemara and the Burren; Grime et al. 1988, 2007; An Irish Flora1996; D. Welch, in: Preston et al. 2002; Chater 2010).
Species recognition
The distinctive features of M. discolor are the very small flowers (2-4 mm across), which areyellow to cream in bud, turning from pink to blue as they open and age, thus meriting its Latinscientific epithet and English common name. The fact that the corolla tube is longer than the calyxhelps distinguish M. discolor from M. ramosissima (Early Forget-me-not), where the corollatube is shorter than the calyx and it never has yellow flowers (Garrard & Streeter 1983; NewFlora of the BI 1991; An Irish Flora 1996).
Flowering reproduction
M. discolor flowers from May to September, the inflorescence being a lax, terminal, bract-lessscorpioid cyme, not much longer than the leafy part of the stem. The flowers are bisexual and thefruiting pedicels are shorter than the calyx. The calyx is bell-shaped, up to 4.5 mm long. The flowersare pollinated by bees and flies and can also self-pollinate (Fitter 1987). As in M. arvensis,the calyx of M. discolor is covered with hooked hairs and does not open or release the four (orfewer) nutlets at maturity, but rather it detaches as a burr-like unit to be dispersed by attaching topassing animals including man (Salisbury 1964). The seed bank survey of NW Europe does not include anymention of M. discolor, but since it has a very similar ecology to M. arvensis which doesdisplay prolonged buried seed survival (five years or more), there is some possibility it might besimilar for this species (Thompson et al. 1997).
Fermanagh occurrence
In Fermanagh, M. discolor is frequent and widespread and has been recorded in 107 tetrads (20.3%of the VC total), 103 of them with post-1975 records. It is most frequently found in disturbed orotherwise open, warm, sunny, dry, sandy or stony soils in a very wide variety of habitats, but it isalso quite frequently found in damp to wet lakeshore and riverbank meadows and pastures and along damproadside verges and woodland tracks. As the tetrad distribution map shows, it is more frequentlyrecorded to the south of Lough Erne, perhaps a response to the greater prevalence here of dry, base- andnutrient-medium to rich soils on which it performs well, although it is very clearly not confined tothese.
Irish occurrence
In Ireland, M. discolor is very widespread across the whole island but, curiously, is morefrequently and more consistently recorded in NI than is the case in the RoI. The species is more widelyscattered in the RoI, with a more easterly occurrence of records, which might possibly reflect moreacidic soils in the west of the country, or perhaps merely result from an absence or scarcity ofresident plant recorders in springtime when the plant is most conspicuous (An Irish Flora 1996;New Atlas).
British occurrence
M. discolor is well distributed across Britain from the Channel Isles to Orkney (VC 111) andShetland (VC 112) in the north. It is generally lowland, but does reach an altitude of 845 m inWestmorland (VC 69). In common with the situation in Ireland, M. discolor is rather scarce andlocal in parts of N England and Wales (Garrard & Streeter 1983). The BSBI New Atlas datasetindicates a widespread decline in populations has occurred in S & E England since around 1950,undoubtedly associated with better management of weeds in agricultural fields since that time (D. Welch,in: Preston et al. 2002).
European and world occurrence
M. discolor belongs to the European temperate phytogeographic element although it shows a definitesub-oceanic tendency in its distribution. It is present in most of Europe from Iceland southwards andeast to Latvia and C Yugoslavia. It is also found in N Africa, on the Azores and on Madeira, in thelatter of which the population is distinguished as subsp. canariensis (Pitard) Grau. Subsp.discolor occurs throughout the range of the species, but subsp. dubia is confined to WEurope (Sell & Murrell 2009). Since the species is often associated with agriculture, it has beenintroduced for instance into S Africa, India, N America and New Zealand (Hultén & Fries 1986, Map1554).
Threats
None.
Introduction, archaeophyte, relict of cultivation, probably extinct. Eurasian southern-temperate, butwidely naturalised in both hemispheres and now discontinuous circumpolar.
1950; MCM & D; garden waste ground, farm at Farnaght, SE of Tamlaght.
Growth form and preferred habitats
Verbena officinalis is a square-stemmed taprooted perennial with pale green stems, 30-100 cm long,that can be either erect, procumbent or prostrate and that produces leafy branches that are lax andspreading. The opposite leaves are ovate in outline, pinnatisect or pinnatifid, the lobes broad butacute at apex, bluntly toothed (serrate) with prominent nerves. On account of its square stems andtwo-lipped, pinkish flowers, superficially Vervain looks like a member of the Lamiaceae (= Labiatae),but the inflorescence is a simple, leafless spike and the ovary of the flower is not deeply four-lobed(Garrard & Streeter 1983; Sell & Murrell 2009).
Vervain has been grown around human settlements since the Neolithic and certainly in antiquity wascultivated as a useful medicinal herb and also as a magical charm, both protective against witches anddemons and able to conjure up devilry of its own (Grigson 1955, 1987; Mabey 1996; R.M. Burton, in:Preston et al. 2002).
In uncultivated settings, V. officinalis is a plant of open or bare ground in rough grassland,hedge banks, scrub, roadsides, sunny pastures and waste ground on well-drained, often calcareous soils(R.M. Burton, in: Preston et al. 2002). The established strategy of V. officinalis iscategorised as CSR by Grime et al. (1988, 2007), meaning that it combines features of all threeprimary strategies, ie Competitor, Stress-tolerator and Ruderal species.
.
Flowering reproduction
V. officinalis flowers from July to September. The inflorescences each bear many small, pinkish topale-lilac, two-lipped flowers that offer both nectar and pollen and are either pollinated byhoverflies, small bees and butterflies, or they self-pollinate (Grieve 1931; Garrard & Streeter1983). The flowers have no perfume and the plant is slightly bitter and astringent in taste.
Fermanagh occurrence
There is only one old record for this perennial in the Fermanagh Flora Database. It has not been seen inthe VC for over 70 years and the current VC Recorders (RSF & RHN) regard it as locally extinct. Thesolitary record was listed by Meikle et al. (1975) in their Revised Typescript Flora as aweed of former cultivation occurring in neglected ground in Carrothers' garden. They added the commentthat, "the species is accredited with malign powers in Fermanagh as elsewhere". Vervain isalso listed as a rare casual introduced species in FNEI 3.
Irish occurrence
Elsewhere in Ireland, this perennial is rare to occasional, most often recorded in the southern half ofthe island and distributed in a very scattered manner. It typically occurs in small numbers in openhabitats including roadsides, limestone quarries, sand pits and on walls and ruins (Cat Alien PlIr).
Previously, in the 19th and early 20th centuries, V. officinalis was very much more commonly grownin Ireland than is currently the case, earlier published accounts of the Irish flora describing it as,"plentiful in some parts of the S & W of Ireland and frequent in many midland counties, mostlyon limestone" (Cybele Hibernica 1866), and "thoroughly established and often abundant,especially about villages" (Irish Topographical Botany 1901).
In more recent times, the Cen Cat Fl Ir 2 lists records from a total of 28 VCs, not includingFermanagh (H33). The New Atlas map for Ireland displays 38 hectads of the most recent date class(1987-99) in about 18 Irish VCs (uncertainty arises from hectads plotted on or near VC boundaries).
British occurrence
V. officinalis is a locally common plant of S England, widespread northwards to The Wash and withscattered localities as far north as Yorkshire. It is also quite frequent in S Wales and SW England, butonly thinly scattered elsewhere and almost entirely absent from N England and all of Scotland (NewAtlas).
European and world occurrence
V. officinalis is widely distributed in both temperate and tropical regions and is described ashaving a Eurasian Southern-temperate distribution. It is thought to have originated in the Mediterraneanregion from where it spread as a naturalised weed over large areas of Europe, Asia and N America, aswell as in many places scattered across the southern hemisphere including in C & S Africa, SAustralia, New Zealand and South Sea Islands. Nowadays it has established a discontinuous circumpolardistribution (Hultén & Fries 1986, Map 1567).
Uses
Vervain had very many, varied uses in herbal medicine, religious and magic rituals, and in folklorepractices as a charm from ancient times onward until the 19th century at least, as described in detailby Grieve (1931), Grigson (1955, 1987) and Vickery (2019). In herbal medicine it was recommended foraround 30 complaints, being astringent, diaphoretic and antispasmodic. It was used to treat intermittentfevers, ulcers, pleurisy and to be a good galactagogue (ie a stimulant of milk secretion) and febrifuge(ie an agent that reduces fever). Vervain poultices were used for headache, ear-neuralgia andrheumatism. Taken internally, it was used for purging and for easing bowel pain and externally it wasapplied for treating piles. It was also used in hom*oeopathy (Grieve 1931).
Names
One derivation of the name 'Vervain' is from the Celtic 'ferfaen', from 'fer', meaning 'to drive away'and 'faen', 'a stone', since the plant was used in herbal medicine to treat bladder problems includingstones in the urinary tract (Grieve 1931, p. 831). An alternative derivation is from 'Herba veneris', onaccount of the supposed aphrodisiac qualities attributed to the plant by the Ancients. Priests used itfor sacrifices and hence another attribution, 'Herba sacra', or 'Sacra herba', 'Holy herb', used insacrifice and purgation (Grieve 1931; Grigson 1955, 1987). The name 'Verbena' was the classical Romanname for 'altar-plants' in general and for this species in particular.
There are a eight alternative local English common names listed by Grigson (1955, 1987) and Vickery(2019), examples of which include 'Berbine', 'Juno's tears', 'Pigeon-grass', 'Simpler's joy' and 'HolyHerb'.
Threats
None.
Native or very possibly introduced, probably extinct. European temperate, also native in C Asia,introduced in Fennoscandia and locally in N America.
2 August 1970; Farrell, Ms L.; NE shore of Monawilkin Lough.
August.
While the validity of the solitary Fermanagh record of this distinctive wintergreen perennial specieswhich flowers from June to September is made more doubtful by the list of associated species rangingfrom those indicative of soils varying from base-rich fens to dry limestone grassland, to bogs and acidpeaty ground, experience proves that such odd things occasionally do happen! The Monawilkin site whereS. officinalis was recorded is famous for the wealth of species that it supports and it is now adesignated NR. The most likely habitat of damp, peaty soil formed over limestone, supporting roughpermanent calcareous pasture by a lakeshore, may here, as it does in the Burren, Co Clare (H9), permitotherwise unlikely associations. Unfortunately, the species has never been refound, despite an enormousamount of time spent surveying the NR prior to its designation and subsequently.
The most typical habitats frequented by S. officinalis are listed as hedge banks, grazed andungrazed infertile grassland, open woods, woodland rides and margins, heaths and occasionally incliff-top grassland. It is confined to vegetation that is both undisturbed and unproductive and that istypically species-rich since potential dominants have their vigour restricted (Grime et al. 1988,2007; K. Walker, in: Preston et al. 2002). Betony typically occurs as scattered individuals, itsseed dispersal is poor and it does not form a persistent soil seed bank (Roberts 1986; Grime etal. 1988, 2007).
Lynn Farrell, who discovered the plant, is now a very well-known field botanist of wide experience and ahighly respected conservation officer. At the time, when she was under contract to the NI DOE for alimestone grassland field survey, she was just beginning her career as a field botanist.
Although S. officinalis is currently regarded as a native species in Ireland, having beenpreviously recorded from 15 of the 40 VCs, its presence throughout the island has been in decline formany years (Cen Cat Fl Ir 2). In 1996, it was regarded as extinct in NI and therefore was deletedfrom the Wildlife Order (NI) Schedule 8 list of Protected Species by the Environment and HeritageService.
Betony was one of the great 'all-heal' plants of medieval herbalists, being used to treat a multitude ofailments including coughs, stomach upsets and kidney, bladder and spleen complaints. In fact, Betony isa fraud, having no special virtue of any kind (Grigson 1955, 1987; Mabey 1996). The plant is probablylong-lived, it has vegetative reproductive capability and its garden cultivation and propagation forherbal medicine purposes until recent years may well have assisted its local survival (Grime etal. 1988, 2007).
The New Atlas map displays the only Fermanagh record and two other old NI records, including onefrom Tyrone (H36) dating from 1909 (McNeill 2010) as introductions (pink symbol) and, althoughMonawilkin Lough is remote from habitation, the current author (RSF) is inclined to accept thisdesignation of what must surely have been a garden escape.
Threats
None.
Native, common and widespread. Eurosiberian temperate, introduced in eastern N America and New Zealand.
1881; Stewart, S.A.; Co Fermanagh.
February to December.
Growth form and preferred habitats
Hedge Woundwort is a very common rhizomatous and stoloniferous perennial with thick creeping roots anderect, softly hairy, stout, solid, much-branched, reddish square stems that grow up to 100 cm tall.S. sylvatica is readily distinguished from S. palustris by its long-stalked, ovate,heart-shaped, very coarsely toothed leaves and by a strong, malodorous smell when its tissues arebruised (Grieve 1931; Hutchinson 1972; Garrard & Streeter 1983).
S. sylvatica tends to occupy light- to medium-shaded sites on a variety of mainly fertile, damp todry soils. The plant prefers moist, decidedly fertile, mildly acid to basic calcareous soils in lightlyto moderately shaded, sheltered ground, or in open, disturbed sites where these or other factorsrestrict the vigour of taller competitors, allowing S. sylvatica to survive and form clonalpatches (Grime et al. 1988, 2007). It is particularly frequent in rough grassland on lower groundaround the margins of woods and scrub, lakeshores, riverbanks, hedgerows, lightly shaded roadside banksand verges, and in moderately disturbed waste ground including neglected areas in gardens (Garrard &Streeter 1983). It is intolerant of mowing or grazing (Sinker et al. 1985).
Flowering reproduction
Hedge Woundwort flowers during July and August, the inflorescence consisting of a long, interruptedterminal spike of 3-6-flowered whorls (or verticillasters). The reddish-purple or crimson, two-lippedflowers have a corolla 13-18 mm long, with white lines marking the three lobes of the lower lip, themiddle lobe is slightly notched, while the upper lip is shallowly hooded. The flowers are bisexual andprotandrous (ie male parts mature first), nectar being secreted by a disc at the base of the superiorovary which is characteristically deeply 4-lobed. The solitary style is inserted between the lobes ofthe ovary and is attached at the base to the receptacle (ie it is gynobasic).
Pollen and copious nectar (the corolla tube sometimes contains nectar for a quarter of its length)attract bees and long-tongued flies that carry out pollination, but if this fails to occur, the flowerswill self-pollinate (Melderis & Bangerter 1955). As with all members of the family, each lobe of theovary becomes in fruit a separate nutlet and, as there is one erect ovule in each, the nutlets are'achenes' (ie single-seeded dry fruits) (Hutchinson 1948; Hickey & King 1981; Grime et al.1988, 2007; Sell & Murrrell 2009). The nutlets are smooth, rounded at the top and are most likelydispersed by wind. The species lacks any mechanism of long-range dispersal (Ridley 1930; Grime etal. 1988, 2007). The seed (nutlets) are long-persistent, some surviving at least five years inthe soil seed bank (Thompson et al. 1997).
Vegetative reproduction
After flowering, aerial parts die down completely in winter, but vegetative reproduction and spread bymeans of the buried rhizome and surface stolons continues whenever temperatures are favourable forgrowth. This appears more immediately significant than seed production in terms of the species' abilityto increase, compete and hold onto its ground. The established strategy of S. sylvatica iscategorised as C/CR, ie intermediate between Competitor and Competitive-Ruderal (Grime et al.1988, 2007). S. sylvatica is described as a rapid coloniser of secondary woodland (Peterken 1981)and establishment from mud-transported rhizome fragments and similarly dispersed long-persistent seedare undoubtedly both important in enabling longer-distance transport by human activities and successfulcolonisation of new territory, although the species has no specially evolved mechanism to achieve thisoutcome.
Fermanagh occurrence
As the tetrad map indicates, S. sylvatica is widespread in Fermanagh and has been recorded in 240Fermanagh tetrads, 45.5% of those in the VC. It also more rarely occurs on cliffs (eg the N-facing cliffat Knockmore and those in Bolusty More Td, Cliffs of Magho) and in artificial, man-made habitatsincluding quarries, sand-pits and churchyards (eg Boho Church and Tubbrid).
A hybrid
The hybrid of S. sylvatica with the even more common S. palustris (Marsh Woundwort) (S.× ambigua Sm.) is intermediate in appearance and diagnosticcharacters between its two parents and is completely sterile (BSBI Plant Crib 1998). The hybridcan be found in the absence of one or both of its parents and it is described as being oddly locallydistributed (BSBI Plant Crib 1998). The percentage length of the petiole in comparison with thewhole leaf and petiole combined is a good diagnostic character: in S. palustris the figure is2-9%; in S. sylvatica 30-44%; and in the hybrid 9-16%.
S. × ambigua is described as occasional to frequent in NE Ireland(FNEI 3), but it has not been found, or so far has been overlooked, in Fermanagh. Harron (1986)found there were two forms of the hybrid around Lough Neagh, a more common one closer to S.palustris in character and a rarer one closer to S. sylvatica (Stace et al. 2015).
British and Irish occurrence
S. sylvatica is almost ubiquitous in lowland situations (other than arable and aquatic habitats)throughout both B & I. It is rare, scarce or absent on the more exposed acid peaty soils of theScottish Highlands and the W of Ireland (New Atlas).
European and world occurrence
The species occurs in most of W & S Europe, but is quite rare in the Mediterranean basin (present onCorsica, Sicily and the Peloponnese) and very rare on Madeira (Press & Short 1994). It extendsnorthwards along the coast of Norway to around 68°N, its most northerly point (Hultén & Fries 1986,Map 1599). Also locally present in the mountains of W & C Asia, ie the Caucasus, Kashmir and theAltai. S. sylvatica has been introduced in eastern N America and New Zealand. It belongs to theEurosiberian temperate element (Sell & Murrell 2009).
Names
Although S. sylvatica carries the name 'Woundwort' or 'all-heal', the herbal medicine virtues thatthis implies are much better displayed by S. palustris (Marsh Woundwort) and also by the hybridwith this species (S. × ambigua). The distribution of S. × ambigua extends much further north and west of that of the moretemperate parent, S. sylvatica, which suggests the hybrid may have been brought into gardencultivation on account of its ease of propagation and sterility, while retaining the valued medicinalproperties of a wound-healing, blood stanching herb. The rootstock and stout roots of the hybrid maypossibly also have been used as food in some circ*mstances (Allen & Hatfield 2004).
Threats
None.
Native, common and widespread. Circumpolar boreo-temperate.
1881; Stewart, S.A.; Co Fermanagh.
April to November.
Growth form and preferred habitats
S. palustris is a rhizomatous perennial that grows to around 100 cm tall and is similar inappearance and biology to S. sylvatica (Hedge Woundwort), but its tissues are less offensivesmelling and the stems are hollow rather than solid. Marsh Woundwort also has more lanceolate, almoststalk-less leaves with crenate margins compared with S. sylvatica and it has paler pink flowers.The two species differ considerably in their preferred habitats, S. sylvatica being a plant ofmuch more open, drier soils.
S. palustris is a very common species of damp to thoroughly wet habitats throughout lowland B& I. It typically occurs in hollows in marshy or swampy grasslands and in fens by lakeshores, alongthe banks of rivers, streams, ditches and in damp to moist, rough grassland along roadside verges. It isespecially associated with ground that intermittently floods or is poorly drained, factors that limitthe vigour of potentially dominant associated species (K. Walker, in: Preston et al. 2002).
S. palustris competes very well and persists in marshy tall herb communities, but is intolerant offrequent grazing and mowing. Although it can tolerate moderate shade, Marsh Woundwort prefers sunny,sheltered, moderately fertile, mildly acid to calcareous soils. It occasionally also occurs as a weed ofa*gricultural land (Garrard & Streeter 1983; Sinker et al. 1985). The established strategy ofS. palustris is categorised as C/CR, meaning it is intermediate between a Competitor and aCompetitive Ruderal (Grime et al. 1988, 2007).
Flowering reproduction
Marsh Woundwort flowers from July to September, the inflorescence a terminal interrupted spike withwhorls of 4- to 10-flowers each, the lower distant, the upper crowded, and bracteoles minute or absent.The 'whorls' are actually greatly condensed cymes that botanically are referred to as verticillastersand they arise in the axils of opposite leaves (Holmes 1979). Not everyone appreciates this finedistinguishing point, but it is the case!
The corolla, 12-15 mm long, has the tube cream and the lobes rose-purple or mauve-pink in colour (Sell& Murrell 2009). The tube of the corolla is twice as long as the calyx. Nectar is secreted at thebase of the superior ovary and stored in the smooth lowest part of the corolla tube. The upper lip ofthe corolla not only shelters the anthers and stigma in bad weather, it also protects the nectar fromdilution by rain. The circlet of hairs that is inside the corolla tube just above the base also protectsthe nectar and keeps out unbidden guests, such as flies (Hutchinson 1972). The flowers are protandrousand are pollinated mainly by long-tongued bees. The nutlets (or achenes) are ellipsoid and, as usual inthis family (Lamiaceae), are produced in fours. They are ovoid, finely striate and brown in colour(Butcher 1961). In the damp to wet habitats of the species, the nutlets can often float and disperse,sometimes while retained within the dry, withered calyx. The 'seed' or nutlet is either transient orlong-term persistent, both being represented in the soil seed bank survey of NW Europe (Thompson etal. 1997).
Vegetative reproduction
S. palustris has a long creeping rhizome that also produces numerous, white, fleshy subterraneanstolons that creep in all directions and develop small tubers at their apex in the autumn. Sometimes thetubers can be elongated, ranging from 8-12 cm long (Grieve 1931; Butcher 1961; Clapham et al.1987; Webb et al. 1988; Sell & Murrell 2009). The rhizome and tubers are the overwintering(perennating) organs of the species, the aerial parts of which die down in the late autumn (November)and disappear after fruiting. The tubers and rhizome sprout new aerial stems when the soil warms up inthe spring. The production of numerous sub-surface rhizomes and tubers by S. palustris helps tomaintain shoot population density in an existing population, with little or no need for seedlingrecruitment. Stolons are said to be formed less frequently than in S. sylvatica (Taylor &Rowland 2011).
Fermanagh occurrence
S. palustris has been recorded in 223 Fermanagh tetrads, 42.2% of those in the VC. It isespecially frequent in winter-flooded water meadows around the shores of Lough Erne, although it shouldbe acknowledged that this area of the VC was intensively surveyed by the NIEA NI Lake Survey. WhileS. palustris is slightly less widespread in Fermanagh than S. sylvatica, it is almost 50%the more frequently recorded of the two in the Fermanagh Flora Database.
British and Irish occurrence
S. palustris is well distributed throughout B & I, although absent from parts of NE Englandand N Scotland and also scarce or absent in the Scottish Highlands. From the New Atlas hectadmap, there appears to be little evidence of any change in the overall distribution of S.palustris since the earlier 1962 BSBI Altas (Perring & Walters 1962; K. Walker, in:Preston et al. 2002).
European and world occurrence
Like S. sylvatica, S. palustris is widely distributed across most of Europe, becoming rarerin the Mediterranean region. It extends into W & C Asia and related forms (sometimes regarded assubspecies) stretch to Japan. S. palustris has been introduced in N America, where again thereare related species or subspecies of S. palustris s.l. (Hultén & Fries 1986, Map 1600).
Uses
S. palustris was regarded by many herbalists in the past as the most effective 'wound-herb' of thegenus. Gerard (1597, 1633) describes it being used to poultice a deep scythe cut, but it was in use longbefore his time. Modern herbalists use it for its antiseptic and antispasmodic properties to relievegout, cramp, joint pain and vertigo (Grieve 1931).
Names
The genus name 'Stachys' is from the Greek 'stachus' for 'a spike', obviously referring to theinflorescence shape (Johnson & Smith 1972). The Latin specific epithet 'palustris' is 'of boggy ormarshy ground' (Gledhill 1985).
English common names include 'Clown's Woundwort', 'Clown's All-heal', two names from Gerard's Herbal onaccount of "a clownish answer" which he received from a "very poore man" who had cuthis leg to the bone and healed it with this plant (Gerard 1597). Additionally there is 'co*ck-head','Dead-nettle', 'Hound's-tongue', 'Swine's Maskert' (or 'Swine's Mosscort'), 'Swine Arnut' (possibly fromthe Dutch 'aardnut', meaning 'earth-nut'), 'Rough Weed' (a name given by Tate (1863) in FloraBelfastiensis), 'Runch' (possibly referring to the edible white roots that were sometimes fed topigs) and 'Sheep's Brisken' ("a name from Co Donegal, almost certainly a corruption of the Gaelic'briosclan' or 'brisgean', a name applied to an edible root sometimes used by poorer people to makebread" (Britten & Holland 1886)).
Threats
None.
Introduction, archaeophyte, either a very rare casual or a possible mis-identification. Suboceanicsouthern-temperate, but widely naturalised with agriculture in both hemispheres.
June 1990; Montgomery, J. & Foster, S.; Hanging Rock NR.
As detailed above, this annual woundwort has been recorded just once in Fermanagh by two members of theEHS Habitat Survey Team at Hanging Rock NR, a fairly well wooded limestone block scree. Without avoucher, this first VC record really cannot be wholeheartedly accepted. However, as a formerly frequentweed of arable cultivation that has existing records from Cos Down (H38), Antrim (H39) and Londonderry(H40), it is not an impossibility that this might be a correct identification (FNEI 3).Nevertheless, S. arvensis is rare and very local in northern parts of Ireland.
In most of its B & I range, this small annual species frequents moderately fertile, mildly acid,non-calcareous light, sandy soils in arable fields, although in W Ireland it previously did occur onlimestone soils in the 19th century (Flora of Connemara and the Burren). The New Atlashectad map shows that Field Woundwort remains quite well established, although still only occasional, inthe cereal growing SE of Ireland. It also appears locally and sporadically in parts of the far south,but it is very rare and sparsely scattered elsewhere on the island (New Atlas).
The current author (RSF) feels that the DOE recorders either made a slip of the tongue or of the pen, orperhaps they mistook the plant for a very poorly grown specimen of S. sylvatica (HedgeWoundwort), a species which previously has been recorded from this NR site.
S. arvensis was included on the list of 41 species previously assumed to be native in B & I,which Webb (1985) considered more probably introduced by man. The editors of the New Atlas havenow concurred and the species is now accepted as an archaeophyte, probably introduced to Britain inRoman times.
Across B & I, there have been many population losses of this annual, beginning pre-1950 andaccelerating since then (New Atlas). The most recent BSBI Local Change re-survey in 2003-4 of the1987 Monitoring Scheme hectads and selected tetrads in Britain, found an unexpected resurgence of S.arvensis, especially in waste ground near southern coasts (Braithwaite et al. 2006).
Threats
None.
Introduced, naturalised garden escape, rare.
21 April 1996; Northridge, R.H.; naturalised on a wooded riverbank, Derryany Bridge, NE of Cargin Lough.
April and May
There are records of this recently arrived hairy, perennial garden escape or discard from 22 tetrads inFermanagh (4.2%), typically thinly scattered near habitation. It has erect flowering stems that grow upto 60 cm tall and has broadly ovate, crenate, opposite leaves with permanent, conspicuous silvery-whiteblotches on all leaves. The flowers are yellow, about 2 cm across, two-lipped, and borne in whorls ofmore than ten per node at four nodes in leaf axils towards the top of the stem (Sell & Murrell2009).
L. galeobdolon subsp. argentatum is thought to have been firstintroduced to B & I gardens sometime in the 1960s, its rapid growth and long arching stolons quicklyrecommending it as a useful ground cover plant for difficult habitat conditions such as dry shade.Experience soon led to a reassessment and the plant is now recognised as a rampant, aggressive andextremely invasive, creeping stoloniferous garden subject, of unknown, but probable horticulturalorigin. Thanks to its stolons and successful seed production, subsp. argentatum readily escapedgarden confines and first appeared in the wild in Britain as recently as 1974 (Rutherford & Stirling1987; Clement & Foster 1994; Stace & Crawley 2015). It made its first appearance in Ireland bythe River Nier S of Clonmel, Co Waterford (H6) in 1987 (Cat Alien Pl Ir).
L. galeobdolon subsp. argentatum is a close relative of the YellowArchangel, L. galeobdolon subsp. montanum (Pers.) Ehrend. &Polatschek which is a tetraploid native plant in Ireland, confined to the SE counties that lie to thesouth of Dublin. Subsp. argentatum is also a tetraploid and the most obvious difference it showsfrom variegated forms of the native plant, with which it is very easily confused, is the conspicuouswhitish leaf blotches carried throughout the year, which turn brownish-maroon along the midrib andlateral veins in the wintertime (Rutherford & Stirling 1987; Ellis 1993). The floral parts of subsp.argentatum are also slightly larger than those of subsp. montanum and it appears likelythat it really represents a mutation of the latter, rather than being a distinct new subspecies. Theinvariable isozyme characters of subsp. argentatum suggest it may even represent a single clone(Stace & Crawley 2015).
An introduced plant of subsp. argentatum in the wild, shaded in woodland or otherwise, can coveras much as 100 m2 in two years (The Times newspaper, 22 May 2001). Being such anaggressive ground-cover plant, many gardeners on obtaining first-hand experience of subsp.argentatum may quickly decide to eradicate or severely thin it in their gardens. It thereforetends to be found where garden rubbish is dumped and all too often this means on quiet wayside vergesoff the beaten track and in shrubberies. The widespread explosion of recent occurrences of naturalisedpatches of subsp. argentatum all across these islands must nearly all derive from gardenthinnings thoughtlessly dumped where they can survive, establish and spread (K. Walker, in: Prestonet al. 2002).
Threats
An extremely invasive plant, it could become a very serious threat to natural vegetation, as has alreadyhappened in S Devon (VC 3). It should be carefully monitored, as it may need to be controlled.
Introduction, archaeophyte, very rare. Eurasian boreo-temperate, but widely naturalised, including in NAmerica.
1953; MCM & D; roadside waste ground, Rosslea.
April to October.
Growth form and preferred habitats
An erect or sometimes sprawling, perennial, wintergreen weed, up to 60 cm tall, with a creeping rhizome(or occasionally surface-running and therefore stoloniferous), L. album is most commonly foundforming loose, clonal patches on shaded, lowland roadsides, laneways, hedgerows, waste ground and onsoil heaps where the competitive vigour of taller, more potentially dominant species (such as Urticadioica, Common Nettle) is curtailed by occasional, moderate disturbance, usually involvingmowing and/or hedge trimming. It prefers fertile, well-drained, lightly-shaded sites with moderatelyacidic to basic soils no lower than pH 4.5 and most frequently pH >7.0. The established strategy ofL. album is categorised as CR, Competitive-Ruderal (Grime et al. 1988, 2007).
Flowering reproduction
L. album has a very long flowering period that begins in early summer and runs through toDecember. The main flush is in May and June when the large, white (or really, white faintly tinged withgreen), two-lipped flowers in distant whorls, each consisting of 3-10, bisexual blossoms, make thespecies most conspicuous and very recognisable. Later in the growing season, the species generallybecomes overtopped and obscured by taller-growing accompanying species. The individual flowers have agreen, tubular-campanulate calyx, 9-20 mm long and a corolla 20-25 mm, cylindrical at the base for ashort distance, but then suddenly enlarged as if swollen. The upper lip of the corolla is arched andthere is an oblique ring of hairs in the corolla tube that prevents unwanted guests obtaining the nectar(Sell & Murrell 2009).
The flowers are pollinated by long-tongued insects, mainly bumble-bees, and the four brownish fruitnutlets (or achenes) produced by each flower are trigonous and have a rough surface. When the flowerfades, the green calyx persists and protects the tiny developing fruit nutlets. The calyx becomessomewhat stiffened as it ages and when the nutlets are ripe and ready for release, any pressure exertedupon the calyx forces it back. When the pressure is removed, the nutlets are shot out with some force(Grieve 1931).
An average-sized plant can produce around 2,400 nutlets, each with an attached elaiosome oil-body thatattracts ants that help to locally transport and disperse them. As in the case of Viola species,the ant activity is thought to help the plant species avoid the worst effects of seed predation(Salisbury 1964). The survey of soil seed banks in NW Europe contains nine estimates for L.album, three of which regard its survival as long-term persistent (at least five years)(Thompson et al. 1997).
Species status and occurrence in Britain and Ireland
Until the publication of the BSBI New Altas in 2002 recognised this species as an archaeophyte,White Dead-nettle was considered native in 82% of the VCs in Britain, where the distribution thinsmarkedly towards the north and west of the island (Grime et al. 1988, 2007; Preston et al.2002). While this species is regarded by most botanical authorities as an introduction in Ireland(Cen Cat Fl Ir 2), two recent Floras covering the NE of Ireland have listed White Dead-nettle asnative without discussion (FNEI 3; Urban Flora of Belfast).
In her 2002 book, Cat Alien Pl Ir, Reynolds described L. album as being, "locallyfrequent, mainly in the E and NE; established near ruins and houses, in hedgerows, on roadsides andwaste ground". This summarises the situation well, except to add that in Fermanagh it usuallyoccurs near habitation and it has been found growing on recently transported soil. Reynolds lists 28 ofthe 40 Irish VCs where it has been found (often only rarely). The exceptions are mainly arranged alongthe W coast, although it has been recorded several times in N Kerry (H2) and in Clare (H9).
In the Flora of the Burren and Connemara, Webb & Scannell (1983) listed L. album aspresent in some abundance around Ballyvaughan harbour in 1967, and they remarked, "This species iswell naturalised in the counties N & NW of Dublin, but it is rare elsewhere in Ireland." Thisprognosis is supported by the authors of the Flora of County Dublin (Doogue et al. 1998),who found L. album, "locally abundant in some parts of the north of the county", andwho also commented that, "it can be very persistent as a garden weed". There are suggestionsthat the plant was previously used for food (especially in Sweden) and in herbal medicine (althoughthere are few records of this in B & I and its use never seems to have been popular), but it mighthave been cultivated for these reasons (see below for uses) (Grigson 1955, 1987; Allen & Hatfield2004).
Fermanagh occurrence
There are only five Fermanagh records (the VC is not included in Cat Alien Pl Ir), and the currentauthor (RSF) and RHN both regard L. album as a rare species in the VC. In addition to the firstrecord listed above, the others, all discovered by RHN, are: Topped Mountain (also known as MullyknockMt), 5 km ENE of Enniskillen, July 1987; Lisnaskea, 24 May 1990; Killypaddy, 1 km W of Lisnaskea, 27April 1991; and in loose transported earth at Tempo Road, Enniskillen, October 1991.
European and world occurrence
L. album is now widely distributed in the northern hemisphere. Its geographical origin isuncertain, but Hultén & Fries (1986) speculate that this weedy species may have originated in themountains of E Europe and W Asia. The map provided by Hultén & Fries (1986, Map 1589) shows thespecies widespread in boreo-temperate Europe and Asia from Iceland to S Italy and Turkey, although itbecomes rare towards the Mediterranean basin, and is absent from all the islands including the Azores,Canary Isles and Madeira. It has been introduced to N America. A separate form, subsp. barbatum(Sieb. & Zucc.) Mennema is widespread in E Asia (Hultén & Fries 1986).
Uses
The active medicinal ingredients of L. album are listed as mucilage, tannin, tyramine,methylamine, choline and traces of fatty oils, saponin and flavone-glycosides (Launert 1981). Theeffects are astringent and diuretic. White Dead-nettle has been put to herbal medicinal use in acurious diversity of ailments across B & I in past years. It was used to treat skin complaints inNorfolk and NW Ireland, where it was mixed with mutton suet to create an ointment for eczema. It otherregions it was recommended for treating arthritis and sciatica, or as a 'Woundwort' for treatingbleeding and deep flesh cuts. It was also used for the treatment of sore feet and even for toothache(Allen & Hatfield 2004).
In hom*oeopathy, an essence is used to treat kidney and bladder complaints. In the home, a similar aquaticherbal extract is used internally for catarrhs and, externally, for bathing burns and septic cuts(Launert 1981).
Young shoots and leaves, collected in March and April, can be used in salads or mixed with spinach orother leaf vegetables. The young leaves can also be added to soups and sauces (Launert 1981).
Names
The genus name 'Lamium' is from the Greek 'laminos' meaning 'throat' or 'gullet', a reference to thecorolla tube (Johnson & Smith 1972; Gledhill 1985).
The Latin specific epithet 'album', 'white', refers to the corolla colour. There are a total of 22English common names listed by Grigson (1955, 1987). Eight of these contain the word 'nettle' since theleaves are strongly reminiscent of the Common Nettle (Urtica dioica) with which the species oftencompetes. However, L. album has no stinging hairs and therefore is distinguished with names suchas 'Blind-nettle', 'Dea-nettle' (or 'Deaf-nettle'), or even 'Dumb-nettle'!
Other names refer to the copious nectar of the flower, such as 'Honey-flower', 'Honey-bee','Honey-suckle', 'Suck-bottle', 'Sucky Sue'. One of the most interesting names is'Adam-and-Eve-in-the-bower', and to appreciate this, it is necessary to turn the flower upside-down andlook into the white upper lip of the corolla. There one sees Adam and Eve, the black and gold stamens,lying side-by-side like two human figures against a white sheet background (Grigson 1955, 1987). Vickery(2019) gives a similar number of English common names (although not identical to Grigson's) and theirderivations.
Threats
None.
Introduction, archaeophyte, common. European temperate, but widely naturalised, including in N America.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
A common and familiar hairy, short-lived, ruderal winter- or summer-annual weed with stems branching fromthe base and growing up to 45 cm tall, but often closer to 10 cm in height. The opposite, petiolateleaves are frequently purple-tinged and this is especially the case with the inflorescence bracts.
L. purpureum is characteristic of a wide range of open, cultivated and disturbed soils, mostcommonly found in lowland areas on roadsides, waste ground, in gravel, on field margins, in gardenflowerbeds and in bare soil around specimen trees and shrubs. L. purpureum is recorded throughoutthe year. It is most prevalent close to habitation and grows vigorously, colonises and competes best insunny, warm sites on disturbed, relatively fertile, damp to dry, moderately acid to calcareous, loam andclay soils (Sinker et al. 1985).
Fermanagh occurrence
In Fermanagh, as the tetrad distribution map shows, L. purpureum is common and widespread in thelowlands, having been recorded in every month and from 130 tetrads, 24.6% of those in the VC.
L. purpureum can be distinguished from L. hybridum (Cut-leaved Dead-nettle), the only otherred deadnettle in Fermanagh, by the leaves being more bluntly and regularly (ie evenly) toothed than thelatter and by a more definite ring of hairs present on the inner surface of the corolla tube near itsbase.
Flowering reproduction
Although categorised as a short-lived, ruderal annual, Red Dead-nettle has a long flowering season,stretching from March to October, although the main flush occurs from April to June. In arable fieldsgrowing winter barley, L. purpureum behaves as a winter-annual, germinating in the autumn,overwintering as a very small, frost-tolerant leaf rosette, flowering early in spring and setting seedbefore the crop develops a dense canopy (Salisbury 1962, p. 349; Grime et al. 1988, 2007).
The terminal inflorescence on each erect stem is rather dense, but is often few flowered, containing onlyaround 4-12 oppositely arranged flowers. The corolla is 10-15 mm long, pinkish-purple, the tube beinglonger than the 5-7 mm tubular-campanulate calyx. Although the bisexual, hom*ogamous, zygomorphic,2-lipped flowers are obviously designed for cross-pollination and they attract bees by offering nectarand pollen food, they are probably often self-pollinated (Clapham et al. 1987). Despite its oftensmall size, individual plants frequently produce abundant nutlets, with plants quoted as producing inexcess of 1,000 (Roberts & Boddrell 1983a). The seed can persist buried in soil for up to five yearsor more (Thompson et al. 1997).
As in L. album (White Dead-nettle), the seed has an attached nutritive oil-body or elaiosome thatattracts ants that help to disperse it locally. This local dispersal is a significant means of avoidingseed predation, but it does not explain the observed widespread colonising ability of the species, whichremains something of a mystery. Presumably the small seeds are dispersed by wind to some extent,although the plant is rather low-growing for this to be effective over more than a metre or so. The onlyother conceivable mechanisms might involve accidental transport in soil and mud attached to man andother animals (Grime et al. 1988, 2007) or, perhaps, as a commercial crop seed impurity in pasttimes. The fact that L. purpureum has been reported as a widespread introduction in New Zealand,spreading rapidly only in the post-1940 period, does point strongly to the latter means of long-rangeseed dispersal (Webb et al. 1988, p. 773).
Vegetative reproduction
Very unusually for an annual species, in the cooler months of autumn and in milder weather during thewinter, L. purpureum can develop a limited degree of vegetative spread prior to reaching theflowering stage. In these cool, mild weeks, the immature plant leaf rosette may occasionally produceprostrate shoots which can root at their nodes and develop a clump or patch of the plant (Salisbury1964). It is also said that non-flowering shoot tips that become detached, eg after soil is rotavated,can re-root, re-establish the plant and eventually flower and set seed (Hodgson, unpublished, quoted inGrime et al. 1988, 2007).
Species status and British and Irish occurrence
Previously, most botanists in B & I assumed or regarded L. purpureum as native, but Webb(1985) listed it with 40 other species he suggested were doubtfully native and in need of reassessment.Interestingly, in this connection, it remains 'native' in the posthumously published 7th revision ofWebb's An Irish Flora (1996). Godwin (1975) noted that while the fossil finds of L.purpureum from the current interglacial in Britain (the Flandrian) conform to the present habitof the plant, favouring cultivated ground and waste places, finds from two earlier interglacial periods(Hoxnian and Ipswichian) showed it had a capacity for existence in a countryside far less affected, ifat all, by the activities of man. The editors of the New Atlas now recognise L. purpureumas an ancient introduction throughout B & I (K. Walker, in: Preston et al. 2002).
In B & I, L. purpureum is common and widespread throughout suitable disturbed lowlandhabitats, becoming scarce or absent only on very acid soils, wetlands and in woodland shade (Grime etal. 1988, 2007). The New Atlas hectad map data suggest there has been a decline of L.purpureum in Scotland, possibly due to the abandonment of marginal arable land (K. Walker, in:Preston et al. 2002).
European and world occurrence
L. purpureum belongs to the European temperate phytogeographic element and is widespread andcommon throughout most of Europe, although it becomes confined to the mountains towards the south and inthe Mediterranean basin. It is also absent in all of the Mediterranean islands E of Capri, includingSicily. Hultén & Fries (1986, Map 1590) reckon L. purpureum probably originated in the eastof its European range, but it has spread widely as an agricultural weed, including to N America,Greenland and Iceland and probably also to N Africa (Böcher et al. 1968; Löve 1983). It hasdefinitely been introduced to New Zealand and from 1940 onward has become much more widespread there(Webb et al. 1988).
Medicinal uses
Red Dead-nettle has been used in herbal medicine to a very limited extent, in much the same manner asL. album (White Dead-nettle) and Galeobdalon luteum (= L. galeobdalon) (YellowArchangel) for healing sores and ulcers. Neither of these species gets more than a passing mention byGrieve (1931) in her authoritative book, A modernherbal.
However, Allen & Hatfield (2004) report that an infusion of L. purpureum in a quart of winehas been drunk in Essex as a treatment for piles (bound to work wonders!). Elsewhere, in East Anglia, itfeatured as a cure for certain unnamed diseases of poultry and, in Co Meath, in Ireland, a decoction ofthe roots has been recommended to take out the rash in measles. In Kerry, an infusion has been drunk forheadaches. Since similar uses are reported for Betony (Stachys officinalis), it may be that L.purpureum was serving as a mere stand-in for the preferred species (Allen & Hatfield 2004).
Names
The genus name 'Lamium' is from the Greek 'laminos' meaning 'throat' or 'gullet', a reference to thecorolla tube (Johnson & Smith 1972; Gledhill 1985). The Latin specific epithet 'purpureum' wasapplied to various shades of red rather than purple (Gilbert-Carter 1964) or maybe it better referred to'reddish-purple' (Gledhill 1985).
Vickery (2019) lists a total of 17 English common names and Grigson (1955, 1987) weights in with 16, someof which do not overlap with Vickery's. Many of the names are also applied to L. album,especially those with the word elements 'Dead' and/or 'Nettle', such as 'Deaf-nettle', 'Dumb-nettle' and'Dunch-nettle'. It gets the name 'Bad-man's posey', 'Bad-man's posies' or 'Blackman's posies' in the Nof England, the 'bad-' or 'black-' man referring to the Devil. Like other species of Dead-nettle, L.purpureum was locally referred to as 'Red archangel' in Lancashire, the 'Archangel' appellationpresumably because it had the angelic quality of not stinging when touched (Grigson 1955, 1987).
Threats
None.
Introduction, archaeophyte, very rare. European temperate, but widely naturalised.
17 August 1995; Northridge, R.H. & Northridge, Mrs H.J.; potato field 300 m W of Quarry,Aghakillymaud Td.
August and September.
Growth form, identification and preferred habitats
This annual ruderal weed of cultivated ground and disturbed waste places can behave as either a winter ora summer annual. L. hybridum grows up to around 25 cm tall and, like L. purpureum withwhich it is most often confused, it flowers from March to October and is pollinated by bees.
L. hybridum appears to perform best as a pioneer colonist when exploiting, sunny, warm, dry,heavily fertilised, moderately acid soils, such as those used for broad-leaved crops, includingpotatoes. It colonises the edges of fertile, arable fields, roadsides and wayside tracks (Sinker etal. 1985; K. Walker, in: Preston et al. 2002).
As the English common name suggests, Cut-leaved Dead-nettle can be distinguished from the more common andwidespread L. purpureum by its more deeply and irregularly toothed leaves (many teeth beinglonger than 2 mm). The plant is more slender and less pubescent than L. purpureum, the leavesoften smaller, the upper ones truncate at the base (not cordate) and ± decurrent down the petiole. Inaddition, the ring of hairs on the inner surface of the corolla tube near its base, that is such awell-marked characteristic of L. purpureum, is only weakly reproduced, if present at all, inL. hybridum (Garrard & Streeter 1983; Clapham et al. 1987; Webb et al. 1996).
Fermanagh occurrence
L. hybridum has been discovered five times in Fermanagh in recent years. The record details of theother finds are: disturbed ground by roadside halfway between Newtownbutler and Galloon Bridge, 1September 2001, RHN & RSF; Lisgoole, Upper Lough Erne, 30 May 2004, RHN; Enniskillen town, 18 August2008, RHN; and Gublusk Bay, Lower Lough Erne, 19 August 2008, RHN.
The occurrence of this annual weedy species in a potato field with ten other common ruderals isinteresting in itself and suggests the possibility that it may be overlooked elsewhere. This isparticularly the case if it occurs intermingled and is confused with the very similar L.purpureum (Red Dead-nettle), which it is reputed to do (An Irish Flora 1996; K. Walker,in: Preston et al. 2002).
The first Fermanagh record listed here means that L. hybridum has now been found in 33 ofthe 40 Irish VCs (Cen Cat Fl Ir 2).
British and Irish status and occurrence
L. hybridum is a rather local weed species, widely distributed, but perhaps only casually in itsoccurrence and rather scarce and local in most of B & I. The hectad map in the New Atlasshows it recorded right across the whole range of latitude from the Channel Isles to Shetland andfrequently appearing in coastal squares, especially in the north. It also appears to be most frequentlyrecorded in the E of England, south of the River Humber. In Ireland, it is more frequently recorded inNI than in the RoI, where it is most noticeably coastal in its occurrence (Preston et al. 2002).
Previously, like other members of the genus Lamium, Cut-leaved Dead-nettle was believed to benative in Britain (Clapham et al. 1987; Stace 1997) and possibly introduced in Ireland (Webb1977; Webb et al. 1996). Along with all the other members of the genus it is now recognised asbeing an introduced archaeophyte in B & I (Preston et al. 2004; Stace 2010). The NewAtlas hectad map indicates that it has greatly increased in recent years, presumably due to thewidespread generous use of chemical fertilisers and slurry spraying on fields. Confusion with L.purpureum means L. hybridum is still very probably under-recorded and the formerover-recorded (K. Walker, in: Preston et al. 2002).
European and world occurrence
The geographical origin of this weedy dead-nettle is unknown, but it is distributed over large parts ofEurope and adjacent parts of Africa (ie Morocco and Algeria (where it is rare)). The main areas oflesser presence (or absence) are in E & SE Europe (Hultén & Fries 1986, Map 1591; Clapham etal. 1987).
Threats
None.
Introduction, archaeophyte, very rare. Eurosiberian boreo-temperate, but widely naturalised, including inW & N Europe and N America.
1947; MCM & D; peaty potato patch near Derrygonnelly.
August and September.
Growth form and preferred habitats
Previously, this coarsely hairy winter or summer annual with golden-green, ovate-elliptic, nettle-likeleaves broadly wedge-shaped at the base was a more frequent weed, especially of potato and turnipfields, but also in other broad-leaved arable crops, particularly those growing on improved, fertile,manured, sandy or peaty soils. It survives best and can be locally common on winter-wet clay and peatsoils. It can also occur very rarely on roadsides and on disturbed soil tips (Crawley 2005).
G. speciosa is one of only four cold-winter tolerant archaeophytes in the flora of B & I(Stace & Crawley 2015), the species normally germinating in late April and behaving as a summerannual. G. speciosa is a rather late spring developer, slow to germinate and not reaching theflowering stage until the height of summer.
G. speciosa is similar in appearance to the closely related native annual, G. tetrahit(Common Hemp-nettle), but it is rather larger, with more acutely-pointed leaves and much larger flowerswith a corolla 27-35 mm. The hollow, much-branched stems of G. speciosa are rough and bristlyhairy, and the nodes are swollen. The stems additionally carry yellow-tipped glandular hairs. The longyellow corolla, usually with a purple mark on the lower lip and the upper lip strongly hooded andcovered with hispid hairs, are very distinctive features (Melderis & Bangerter 1955; Blamey &Grey-Wilson 1989).
The decline in arable farming using traditional methods of rotation and manuring and the advent of modernmethods of weed control together have contributed to the decline, almost to extinction, of thisdistinctive and attractively flowered annual weed. However, as it also used to occur on other types ofmarginal and disturbed ground, these changes in farming and weed control measures do not completelyaccount for the species near total decline and disappearance since the 1960s (BSBI Atlas 2;New Atlas).
Species status
In previous years, G. speciosa was regarded by Irish botanical authorities as a definite orpossible introduction (An Irish Flora 1977; Cen Cat Fl Ir 2), whereas in recent yearsBritish Flora writers continued to regard the plant as native (Clapham et al. 1987; New Floraof the BI 1997). The New Atlas editors reassessed the status and now recognise thisagricultural weed as being an ancient introduction or archaeophyte in both B & I (Preston etal. 2002, 2004).
Variation
G. speciosa is a diploid plant with 2n=16 chromosomes and is one of the parent species of thenatural tetraploid hybrid, G. tetrahit, the other parent being G. pubescens (O'Donovan& Sharma 1987). In fertile, damp, arable situations, plants can grow up to 100 cm in height,overtopping the potato or other root-crops with which it frequently co-habits (Salisbury 1964, p. 317).There is great morphological plasticity, however, with respect to local growing conditions, and in thecurrent author's (RSF's) albeit limited experience in Donegal, the plants encountered were small,reaching only around 15-30 cm or so in height and they clearly were not capable of strongly competingwith potato plants.
The corolla colour can also vary: usually it is yellow with a purple blotch on the lower lip, but it canbe a pale yellow with deeper yellow protuberances and the purple patch on the lower lip, or it can beyellow with the lower lip variegated white, or the whole lower lip can be purplish in colour (Sell &Murrell 2009; Stace 2019).
Flowering reproduction
Flowering takes place from July to September, the flowers being produced in dense whorls in the axils ofthe upper leaves and bracts. The conspicuous bi-coloured corolla and the nectar it contains attractlarge bumble-bees with a tongue at least 10 mm long to the plant (Faegri & van der Pijl 1971, p.226). If insect-pollination fails to occur, the flowers are self-fertile and may self-pollinate,although seed yield is greatly depressed by selfing (Muntzing 1930; Garrard & Streeter 1983). Awell-grown plant may bear up to 50 or more flowers and seed production can be prolific.
Each fruit of G. speciosa contains four nutlets or achenes (single-seeded dry fruits). The nutletsmeasure 3 × 2 mm, are ovoid, trigonous, papillate, tuberculate and dark brown in colour (Butcher 1961).The seeds are originally dispersed by wind and ground water flow. In agricultural settings, they arealso scattered by farm machinery operations on the crop. In some situations, eg in grain crops such asoats, the seed may be transported over long distances as a contaminant of crop seed (O'Donovan &Sharma 1987).
There are only three estimates of G. speciosa seed (nutlet) persistence in the recent survey of NWEuropean soil seed banks. Two studies regarded it as transient (ie persisting for less than a year),while the third source could not assign a seed-base type to the species (Thompson et al. 1997).G. speciosa seed does, however, demonstrate strong dormancy (O'Donovan & Sharma 1987) and islong-persistent. There is considerable variation in germination, which generally occurs around lateApril in open conditions in cultivated or recently disturbed fertile soils. Only a proportion of theburied seed is released from dormancy each winter and manages to germinate in spring, the remaindersurviving buried to appear in later seasons (Karlsson et al. 2006).
Fermanagh occurrence
While previously this was a quite frequent weed, G. speciosa is now exceedingly rare and has notbeen seen in Fermanagh since 1988. There are a total of six records in the Fermanagh Flora Database,four from the 1947-53 period, when Meikle and co-workers were recording, and two from the 1980s. Thedetails of the other five records are as follows: field near the river below Manyburns Bridge, 1951;fields N of Corraslough Point, Cam Td, Upper Lough Erne, 1951, and seen here again on 2 September 1984by RHN; waste ground near Lisnaskea, 1947-53; and quarry beside Keenaghan Lough, 3 km E of Belleek, 17September 1988, RHN.
British and Irish occurrence
The decline of G. speciosa to occasional, sporadic or casual status appears to have occurredthroughout its range in B & I, as shown by many recent county Floras (eg Trueman et al.,1995; Brewis et al., 1996; Crawley 2005).
While the Cen Cat Fl Ir 2 indicates that Large-flowered Hemp-nettle has previously been recordedat least once from a total of 27 Irish VCs (Scannell & Synnott 1987), the species has always beenmuch more prevalent in the north of the island. The 1962 BSBI Atlas showed that G.speciosa had much greater presence in Cos Donegal (H34, H35) and Londonderry (H40) thanelsewhere in Ireland when the survey was carried out in the late 1950s and early 1960s (Perring &Walters 1962, 1976). The New Atlas hectad map confirms this northern pattern has been maintainedup to 2000 (Preston et al. 2002). The new map certainly indicates that Co Londonderry (H40) isnow the Irish stronghold of this rather attractively-flowered Hemp-nettle.
In Britain, G. speciosa was previously common and widespread throughout, although always regardedas casual in SW England. The New Atlas map shows that the distribution is better recorded than inthe 1962 Atlas, but also that the species presence has shrunk rapidly to rarity during theintervening four decades.
The distribution pattern of the surviving G. speciosa populations in B & I is a very curiousone, especially when compared with those of related annuals and other weeds of arable cultivation onacidic soils, eg Chrysanthemum segetum (Corn Marigold), Fumaria bastardii (TallRamping-fumitory), F. muralis (Common Ramping-fumitory) and Galeopsis bifida (BifidHemp-nettle), and G. speciosa has a definite northern and lowland distribution, itsremaining southern strongholds being in N Wales, Lancashire and the peaty fenland south of The Wash. Itis also prevalent in lowland Scotland, from Dumfries northwards and in the NE from Perthshire toAberdeenshire (New Atlas).
The most recent BSBI sampling re-survey (unfortunately confined to Britain) indicates that G.speciosa remains a frequent weed of arable crops towards the upper limits of cultivation, oftenon damp, peaty soils, but that elsewhere it is rare or scarce and declining rapidly. Compared with otherarable weeds, it is late in flowering and setting seed and it only really succeeds among root crops andin some game cover and exotic crops such as Phacelia tanacetifolia (Braithwaite et al.2006).
European and world occurrence
G. speciosa is widespread in Europe N of the Alps and other southern mountain ranges and itextends into temperate W Asia. The species must be cold-tolerant, since it penetrates well beyond theArctic Circle in N Scandinavia. The map published by Hultén & Fries (1986) shows it also present inS Iceland, although it is not mentioned in the Flora of Iceland (Löve 1983). It is mentioned inthe earlier The Flora of Iceland and the Faeroes as having been accidently introduced and onlyfound around the capital Reykjavik (Ostenfeld & Gröntved 1934). It has been introduced to N America,but the published map shows it very rare and scattered there (Hultén & Fries 1986, Map 1585).
Toxicity
In common with other members of the family, G. speciosa contains irritant oils which can provepoisonous – dangerously so to young stock such as pigs and horses, although there are no recent reportsof poisoning except in Russia (Cooper & Johnson 1998).
Names
The genus name 'Galeopsis' is an ancient Greek name derived from 'gaľe', 'weasel' and 'ŏpis','appearance', thus translating as 'weasel-resembling' (Stearn 1972). The Latin specific epithet'speciosa' translates as 'showy' or 'handsome' (Gilbert-Carter 1964).
Threats
Modern farming methods of cultivation and herbicide weed control have reduced this annual weed speciesalmost to extinction.
Native, occasional. European boreo-temperate.
1860; Smith, T.O.; Ardunshin.
June to December.
Growth form and preferred habitats
This bristly hairy, annual ruderal weed produces simple or branched stems, 30-75 cm tall, that areusually swollen below the nodes. The opposite, nettle-like, heavily toothed lanceolate to ovate leavesare pubescent on both surfaces, but the bristly hairs are more abundant on the upper side. The bristlyhairs that cover the plant are sufficiently stiff to penetrate the skin when handled, but they carry nosting. G. tetrahit is a tetraploid with a chromosome count of 2n=32 (O'Donovan & Sharma1987).
G. tetrahit is occasional in a wide variety of semi-natural habitats ranging from damp semi-shadeof woodland clearings, hedges, riverbanks and lakeshores to the very different situation of open,somewhat disturbed ground on agricultural field margins and damp waste places. It often grows onmoisture-retaining peaty soils, although it can grow in almost any damp, moderately fertile, mildly acidsoil with a pH between 4.5 and 7.0, such as would be suitable for arable agriculture. Its occurrence inBritain is centred on heavily disturbed, productive vegetation (O'Donovan & Sharma 1987; Grime etal. 1988, 2007).
While this summer annual favours open habitats, it produces only one tall-growing population each seasonand thus is not able to compete and survive to reproduce in more heavily disturbed ground that isregularly grazed, mown, or trampled. The phenology involves germination stimulated after low wintertemperatures in early April, followed by a first large flush of growth in early spring and later smallerflushes at regular intervals throughout the growing season (O'Donovan & Sharma 1987). Also, despiteindividuals reaching a metre or so in height, G. tetratit does not form large, conspicuouspatches or stands. The established strategy of G. tetrahit is categorised as C/R,Competitive-Ruderal, ie intermediate between a competitor and a purely ruderal species (Grime etal. 1988, 2007).
Polyploid origin
The distinctly odd variety of habitats occupied by Common Hemp-nettle may be a reflection of thepolyploid origin of this species, an amphidiploid formed by a complicated cross between two diploids,G. pubescens and G. speciosa, to produce a new tetraploid species (Muntzing 1930). Thesame two parent species at some stage also crossed to give rise to the rather similar G. bifida.The two hybrid products are inter-sterile with their parents (though not with one another) and, beingmorphologically separable, they are both given specific rank.
Since many field botanists struggle to distinguish these two species, it is not surprising that there areonly a very few, scattered records of the hybrid between them since it was first discovered in Britainin 1955. The hybrid is markedly, but not completely sterile and usually occurs as single plants withoutany back-crossing (Stace et al. 2015). Through the isolation created by occupying fundamentallydifferent open and semi-closed vegetation habitats, populations of G. tetrahit may well be in theprocess of establishing genetic barriers to gene flow, which will eventually lead to divergence andfurther sympatric speciation (Stace 1975).
Flowering reproduction
Flowering takes place from July to September, the stalkless, bisexual flowers being borne in denseclusters (contracted cymes called verticillasters), in the axils of the upper leaves. The calyx is 12-13mm long and elongates further when in fruit. It is tubular and has five unequal lobes, each onelinear-lanceolate and with a spine-like apex. The calyx is also densely covered with stiff eglandularhairs and minute glands. The corolla, which is 15-23 mm long, is rather variable in colour, ranging frompurple to mauve-pink, white or variegated (Sell & Murrell 2009). The flowers are self-fertile andalthough they produce nectar and pollen they are mainly autogamous (ie they self-fertilise). Autogamy inG. tetrahit is more pronounced than in related species (O'Donovan & Sharma 1987).
Seed output is modest for such a large plant. Each flower produces the four nutlets typical of the familyand, therefore, an average plant might produce a total of around 2,000 nutlets, although this is veryvariable with environment and will be significantly reduced at lower temperatures (Salisbury 1942;O'Donovan & Sharma 1987). Nutlets are single seeded dry fruits (ie achenes) and are often simplyreferred to as seeds. The level of G. tetrahit seed production quoted is less than a tenth of theannual seed output of Senecio sylvaticus (Heath Groundsel), for instance, while Epilobiumangustifolium (Rosebay Willowherb) produces vastly more – around 76,000 seeds/annum (Salisbury1942, pp. 188, 191).
Seed are dispersed by wind and by flowing water, being able to float for up to two days (Grime etal. 1988, 2007). They can also survive ingestion by stock animals and birds (Ridley 1930). Incultivated fields, seed is scattered by farm machinery during seeding, tillage and harvesting operationsand may also be transported over long distances as a contaminant of crop seed and livestock feed(O'Donovan & Sharma 1987).
Seeds of G. tetrahit display strong dormancy, germination being only possible after coldstratification. Germination is favoured by lower temperatures, the optimal fixed temperature being 13°C,although a fluctuating temperature regime alternating between 5° and 15° was even better (O'Donovan& Sharma 1987). Estimates of buried seed survival in soil vary greatly from transient (less than oneyear) to long-term persistent (at least five years) (Thompson et al. 1997). At least some seed isalmost certainly long-persistent (Roberts 1986)
Fermanagh occurrence
In Fermanagh, where arable fields are now very rare, the G. tetrahit agg. is thinly and ratherwidely scattered in the lowlands, but it has been recorded in 69 tetrads (13.1% of those in the VC), 57of them with post-1975 records.
G. tetrahit s.s. has only been discriminated from the species aggregate on six occasions inseparate tetrads and thus it requires much further study in the VC. The details of the records are: NEshore of Cornaleck Td, E of track, Upper Lough Erne, 23 July 1986, EHS Habitat Survey Team; ditch atCurragh and Drumguiff Tds shore, 6 August 1986, EHS Habitat Survey Team; waste ground, Killycarnan, nearLough Nacallagh, Upper Lough Erne, 4 September 1988, RHN; E of Rossmore Point, Castel Archdale, LowerLough Erne, 26 July 1995, RHN & HJN; sand pit at Pubble Bridge, Tempo River, 20 August 1999, RSF& RHN; and halfway between Newtownbutler and Galloon Bridge, Upper Lough Erne, 1 September 2001, RHN& RSF.
British and Irish occurrence
The New Atlas map shows that while G. tetrahit agg. is rather widely recorded in Ireland,at the same time it is much more consistently and frequently recorded in the north in comparison withelsewhere on the island, as is also the case with G. speciosa (Large-flowered Hemp-nettle). Thecurrent author (RSF) cannot suggest any obvious reason why this should be the case, since in Britainthis taxon is common and widespread throughout lowland regions with acidic soils.
The recent BSBI sampling re-survey effort in Britain (Change in the British Flora 1987-2004) hasfound that G. tetrahit agg. is markedly declining there (a calculated Change Factor of -23),although again no explanatory mechanistic factor is identified by the editors of the published report(Braithwaite et al. 2006).
European and world occurrence
Nowadays, G. tetrahit is widespread across most of Europe, although absent or rare in the southand on many of the Mediterranean islands (Sell & Murrell 2009). It is not clear where it originated,but Hultén & Fries (1986, Map 1586) show it more definitely present in Denmark and Scandinavia,especially around the Baltic Sea, as if this might be significant. They also question whether G.tetrahit extends eastwards to Siberia, although this is claimed by Sell & Murrell (2009).
Hultén & Fries encountered a difficulty in their mapping as they remark on the problems that exist indistinguishing G. tetrahit from G. bifida, and they map the latter as the much morewidespread of the two closely related species (Hultén & Fries (1986, Map 1587).
O'Donovan & Sharma (1987) appear clear that in their view G. tetrahit is native in Europe andtemperate regions of Asia, and that it has been found as a naturalised introduction mainly in thenorthern continental United States, Alaska and Canada, where it occurs to the northern limits ofa*griculture.
Names
The genus name 'Galeopsis' is an ancient Greek name derived from 'gaľe', 'weasel' and 'ŏpis','appearance', thus translating as 'weasel-resembling' (Stearn 1972). The Latin specific epithet'tetrahit' may be associated with 'teter', meaning 'foetid', a reference to the goat-like smell of theroot (Gilbert-Carter 1964). However, it may also refer to the tetraploid nature of the species, meaning'four times', ie four sets of chromosomes (Gledhill 1985).
G. tetrahit has a total of 17 English common names in Britten & Holland (1886), mostlyassociated with 'Dead-nettle' (ie not stinging nettle), such as 'Bee-nettle', 'Blind Nettle','Day-nettle', 'Deye-nettle', 'Dog-nettle'. A couple of names actually suggest the plant does sting, eg,'Stinging Nettle' and 'Sting Nettle'. 'Hemp-nettle' is a name first given by Gerard (1597), who alsocalled the plant 'Bastard Hemp' and 'Wild Hemp'. Other interesting names that have no or poorexplanation include 'Glidewort' and 'Hoyrope', another Gerard (1597) name more often applied toEupatorium cannabinum (Hemp-agrimony).
Threats
None.
Native, rare. Eurasian boreo-temperate.
20 August 1975; Hackney, P.; bog 1 km SW of Bannagh Bridge.
July to November.
Growth form and preferred habitats
This up to 100 cm tall, bristly hairy, annual weed occurs as a native ruderal species in damp, moderatelyacid, lowland disturbed ground in arable fields, on roadside verges, along hedge banks, waste placesand, less often, on woodland margins and clearings and on bog and ditch margins alongside its much morecommon close relative G. tetrahit (Common Hemp-nettle), with which it ecologically overlaps andundoubtedly competes (J.R. Press, in: Rich & Jermy 1998; Stace et al. 2015). G. bifidais generally regarded as somewhat more ruderal than G. tetrahit, since it is more strictly anarable weed (K. Walker, in: Preston et al. 2002). Both these Hemp-nettle species are tetraploid(2n=32) (see below), self-compatible and polymorphic (J.R. Press, in: Rich & Jermy 1998).
The current author (RSF) as not been able to discover any published information regarding G.bifida competitive ability, seed production levels and buried seed survival in soil, but thelikelihood is that it mirrors the ability of G. tetrahit and, therefore, seeding is probablyprolific and nutlets long-persistent, a proportion surviving for at least five years.
Identification
G. bifida is morphologically distinguished from G. tetrahit by the corolla being almostentirely purple in colour, the darker markings on it extending nearly to the tip, the tip clearlynotched and with middle lobe of the lower lip having its margins turned down at the sides (ie revolute)(Müntzing 1930; New Flora of the BI 1991; An Irish Flora 1996). As such it should beeasily recognised, but, since it was included within G. tetrahit s.l. in the first BSBIAtlas in (Perring & Walters 1962, 1976), more local field work is essential before anydefinitive statements about the true distribution of this taxon can be made in B & I, including theextent of under-recording and whether or not its presence is changing. G. bifida is almostcertainly under-recorded in the New Atlas survey (K. Walker, in: Preston et al. 2002;Stace et al. 2015).
Polyploid origin and a hybrid form
The distinctly odd variety of habitats occupied by G. bifida and G. tetrahit s.s.may be a reflection of their polyploid origin. They are both amphidiploids formed by acomplicated cross between the two diploid species, G. pubescens (a continental species that doesnot occur in B & I) and G. speciosa (Large-flowered Hemp-nettle) to produce two newtetraploid species (Müntzing 1930).
Although they are inter-sterile with their parents, G. bifida and G. tetratit s.s. arecapable of interbreeding with one another where they geographically and ecologically overlap, a factorlikely to complicate their distinction from one another. Having said this, the hybrid, G. × ludwigii, is very rare, is almost entirely pollen-sterile and hasvery few records in Britain and none in Ireland, despite having been first described and discovered inEngland as long ago as 1955 (Stace et al. 2015). Stace (2019) remarks on the hybrid that it isvery scattered in Britain north to central Scotland, "but is probably frequent".
Fermanagh occurrence
With only ten records so far in the Fermanagh Flora Database scattered across ten tetrads, the currentauthor (RSF) and RHN believe that this must be an under-recorded weedy species and acknowledge that notenough data exist to demonstrate the full picture of its local occurrence. However, the present evidencein Fermanagh supports the generally held notion that in B & I G. bifida appears to be more orless frequently found in semi-natural vegetation and that it is much more definitely ruderal in itspropensity and less agricultural in behaviour than G. tetrahit.
The Fermanagh record details of G. bifida are: bog 1 km SW of Bannagh Bridge, 20 August 1975, P.Hackney, BEL, Accession Number H5236; Gortcar Railway Bridge, 1 km SW of Clonelly, 25 July 1976,Miss N. Dawson; Clabby Village, 7 July 1988, RHN; roadside and blanket bog at Killee, 24 September 1988,RHN; disturbed ground, S of Knockninny, November 1989, RHN; Tullykelter Td, NE of Carran Lough, 14November 1993, RHN; waste ground, Killyvilly, 2 km NE of Enniskillen, 8 November 1989, RHN; EnniskillenTown, 9 October 1998, RHN; sand Pit at Pubble Bridge, Tempo River, 20 August 1999, RSF & RHN; andMullanaskea, 2 km E of Killyvilly, 30 November 2002, RHN.
British and Irish occurrence
G. bifida often grows alongside and among G. tetrahit and the two are probably oftenmis-identified for one another, or when they cannot be securely identified, combined into G.tetrahit s.l., or agg. As mentioned above, the true distribution of both these species requiresmore accurate discrimination in recording, but it is very likely that G. bifida is under-recordedat present (K. Walker, in: Preston et al. 2002). Having said this, the New Atlas hectadmap shows G. bifida widely scattered across all latitudes in Britain and its associated isles. Itis scarce or absent from limestone and chalk areas of S England and is almost totally absent from NWEngland. In Scotland, it is widespread as far north as the urban conurbations of Glasgow and Edinburgh,becoming much more occasional, scattered and increasingly coastal northwards, although it is present onthe N & W Isles.
In Ireland, G. bifida is shown in the New Atlas as being most widespread in the north ofthe island and in the far SE, but only very thinly and widely scattered across most of the RoI. Stace(2019) describes G. bifida as occurring in similar places to G. tetratit and,"probably at least as common and widespread [as it]".
European and world occurrence
G. bifida appears to have originated in Eurasia and is regarded as belonging to the Eurasianboreo-temperate phytogeographical element (Preston & Hill 1997). According to Hultén & Fries(1986, Map 1587), G. bifida now has a considerably wider world occurrence than the closelyrelated G. tetratit. The map provided by Hultén & Fries (1986) indicates G. bifida isnative across N & C Europe and stretches continuously well into C Asia as such, although it becomesscarce towards the south on both continents. In the S of Europe and in the Mediterranean region, it iscompletely absent on the Iberian Peninsula and has a limited presence in the very N of Italy, althoughit penetrates Greece into the Peloponnese and Crete, and is also shown as being present in Corsica andSardinia. It is absent otherwise from all the remaining Mediterranean isles.
G. bifida has also been widely introduced worldwide, probably as a contaminant of crop seed, andhas become circumpolar. Hultén & Fries 1986, Map 1587) shows G. bifida present in Iceland,but the published Flora refutes this, listing only G. tetrahit (Löve 1983). The Flora of theCretan area also fails to include any member of the genus (Turland et al. 1993).
Threats
None.
Native, frequent but rather local. Eurosiberian boreo-temperate, also native in N America.
1882; Barrington, R.M.; wood near game-keeper's lodge, Crom Castle Estate.
May to September.
Growth form and preferred habitats
This wetland perennial possesses a slender, creeping rhizome and produces erect to decumbent 4-sided,leafy stems, 15-50 cm long or tall, that are either glabrous or have scattered eglandular hairs andminute glands. The many-branched stems bear shortly-stalked, opposite leaves that are either cordate(heart-shaped) or rounded at the base and have remotely and shallowly toothed (crenate) margins. Flowersdevelop from July to September in pairs, solitary in the paired leaf axils and appear on one side of thestem only (Clapham et al. 1987).
S. galericulata is frequently found amongst wet, tussocky grass in hollows, or over-topping plantsin lowland marshy soils that are subject to regular fluctuations of the water-table. This includes inlakeside tall herb vegetation of water-meadows, marshes and fens. The soils it most commonly frequentstend to be fertile mud or clay, moderately acid to neutral or calcareous in reaction. Although it isfound most frequently in sunny situations, it is often lightly shaded and it sometimes grows inhalf-shade at the base of willows and other shrubs in fen-carr scrub woodland, behaviour that presumablyaffords it some concealment from grazing and trampling by sheep and cattle.
When growing in shade and competing, Scullcap can become a decidedly straggly plant of considerable size(Crawley 2005). It really can only tolerate light grazing pressure (Garrard & Streeter 1983; Sinkeret al. 1985). The established strategy of S. galericulata is categorised as CR/CSR,meaning it is intermediate between a Competitive Ruderal and a more general balance of all three basicstrategies – Competitive, Stress-tolerant and Ruderal (Grime et al. 1988, 2007).
In coastal areas of B & I, S. galericulata proves tolerant of salt and brackish conditions andcan therefore be found in moist areas of dune slacks or on the sides of coastal drains. It can alsooccur on boulder beaches in some parts of Scotland (K. Walker, in: Preston et al. 2002).
Fermanagh occurrence
Skullcap has been frequently recorded in 82 Fermanagh tetrads, 15.5% of those in the VC. Seventy-sixtetrads have post-1975 records. While S. galericulata is commonly found around Upper Lough Ernein particular, as shown in the accompanying tetrad map, Skullcap is much more local in its distributionwhen one leaves the Lough Erne basin. It appears chiefly on the wet to damp, intermittently floodedshores of the larger lakes in Fermanagh and along the banks of the River Finn in the SE of the VC. S.galericulata occurs in rather more acidic peaty mud in many sites around Upper Lough Erne and atSpectacle Lough, for example, higher up on the Western Plateau.
Flowering reproduction
Flowers develop from June to September and are arranged in pairs, although actually each blossom issolitary, but is carried in the axil of two opposite leaves, so they appear one sided on the stem. Theplant is made less conspicuous by having only a few of the 10-20 mm dull mauve- or violet-blue flowersout in blossom at any one time. The calyx is bell-shaped below and 2-lipped above. Behind the upper lipof the calyx there is an unusual, broad, flat, scale-like pouch attached. After the flower, which isscentless but produces nectar and pollen, has been visited and pollinated by various long-tonguedinsects, the corolla drops off and the curious pouch on the back of the calyx then falls forward,closing the top of the tube and protecting the developing fruit. As far as the current author (RSF) isaware, this is a unique feature apparently unnecessary in other members of the Lamiaceae.
The corolla is much larger than the calyx and projects well beyond it. While displaying a shade of blueoutside, the corolla tube is white inside, the upper lip concealing the stamens and the lower lipbeautifully mottled with deep-blue and white nectar guides (Melderis & Bangerter 1955). The fournutlets (single-seeded fruits or achenes) are contained within the dry calyx until they are mature.
When ripe, the wind shaking the stem causes the calyx lobes to separate, permitting the nutlets to bethrown out. They are small, yellow and densely covered with thick processes. Special buoyancy tissuecontaining air spaces is present in the walls of the nutlets that enables the lightweight propagules tofloat for up to twelve months and disperse, should they end up in the water around which the parentplant grows (Ridley 1930; Sculthorpe 1967).
The duration of seed survival in soil is probably transient, although there were only two estimatesquoted in the soil seed survey of NW Europe and one of them was non-commital (Thompson et al.1997).
Vegetative reproduction
Since S. galericulata produces slender rhizomes, once established, given sufficiently favourablegrowing conditions for it to successfully compete, the plant can locally increase and develop clumps orsprawling patches of the species. The relative importance of seed versus vegetative reproduction is notclear, however, and requires further study.
British and Irish occurrence
Well distributed throughout most of lowland Britain, but scarcer in NE England and N & E Scotland.Drainage and development have led to losses of suitable habitat in some parts of England.
In Ireland, S. galericulata has always been scarce and occasional, mainly found inland onlakeshores and along rivers, largely in the Midlands and the north of the island (An Irish Flora1996; New Atlas).
European and world occurrence
S. galericulata belongs to the Eurosiberian boreo-temperate phytogeographical element andoccurs widely and frequently across almost all of Europe and N, W & C Asia. It becomes scarce orabsent S of the Pyrenees and the Alps in W Europe and is absent from the more eastern Mediterraneanisles. It just manages to reach Algeria in N Africa and is common and widespread across N America,although it is recognised as a separate variety, S. galericulata var. pubescensBenth. Closely related species occur in E Asia, S. yezoensis Kudo and S. ikonnikovii Juz.,which taken together help make S. galericulata into a circumpolar distribution pattern (Hultén& Fries 1986, Map 1579; Sell & Murrell 2009).
Uses
Grigson (1955, 1987) refers to S. galericulata as, "another of the bitter labiates" thatwas used as a 'nervine', ie used to calm the nerves, and as a 'febrifuge', ie an agent used to lowerbody temperature and give fever relief. The American species S. lateriflora L. (VirginianSkullcap) is more used in herbal medicine than S. galericulata, being regarded as a certain curefor hydrophobia. The plants contain a volatile oil, 'Scutellarin', and a bitter glucoside that yields'Scutellarein' on hydrolysis.
Skullcap was said to have strong tonic, nervine and antispasmodic action and it was considered slightlyastringent. It was used to treat convulsions, hydrophobia, rickets, nervous headaches and severehiccoughs. It was used to sooth nervous twitches, nervous excitement and for inducing sleep whennecessary, without any unpleasant symptoms following (Grieve 1931). It is not recommended nowadays sinceoverdosing can induce ill effects.
Chemists are currently researching the species for medical uses including the creation of drugs fortreating cancer.
Names
The genus name 'Scutellaria' is derived from the Latin 'scutella', a diminutive meaning 'a small salver'or ' small dish' or 'saucer', referring to the minute pouch attached to the calyx (Gilbert-Carter 1964;Stearn 1992). The Latin specific epithet 'galericulata' is an adjective formed from 'galericulum', adiminutive of 'galerum', the leather helmet worn by Roman soldiers. It thus means 'helmet-shaped' or'small capped', like a 'skull-cap', referring again to the calyx that resembles this form of head-dress– and hence the English common name (Johnson & Smith 1946; Gilbert-Carter 1964; Gledhill 1985;Stearn 1992).
Other similarly derived English common names include 'Helmet Flower', 'Hoodwort' and 'HoodedWillow-herb', the latter given as a translation of its old name 'Lysimachia galericulata' by (Gerard1597) (Britten & Holland 1886; Grieve 1931).
Threats
None.
Native, scarce or occasional and very local. Suboceanic southern-temperate and an introduced adventive inFennoscandia and N America.
1900; Praeger, R.Ll.; Co Fermanagh.
March to December.
Growth form and preferred habitats
This pubescent, perennial herb produces erect, simple or occasionally branched, acutely 4-angled stemsfrom a much-branched, creeping underground rhizome. Stems are woody at their base and vary from 10-40 cmand may reach 50 cm when flowering. The ovate, decussate, petiolate, opposite leaves are 2.5-7 cm, withcrenate margins and rugose, wrinkled (bullate), sage-like surfaces, dark green above, paler beneath.Aerial shoots die back each autumn, but recognisable dead stems (flowering or otherwise) persistattached to the rhizome for up to two years, enabling the plant to be identified at any season.Flowering takes place June to September, the branching flowering stems taller than the barren vegetativebranches. The flowers, 9-10 mm long, are borne in pairs on long, slender, terminal, leafless spikes. Thebracts beneath the flowers are very different from the leaves and they are shorter than the flowers. Thenumerous spikes on side branches make up a loose panicle with the flowers all arranged to one side(Hutchinson 1968; Hutchinson 1972; Sell & Murrell 2009).
Throughout B & I, Wood Sage is a common plant of the less shaded parts of lowland, deciduouswoodlands, wood margins, rough grasslands, heaths and fixed sand-dunes. In suitable sites it formsextensive diffuse patches by means of its spreading rhizome. T. scorodonia prefers dry,well-drained, mineral, acid or calcareous soils and grows in sites up to an altitude of around 600 m. Itis found on a wide variety of generally infertile substrates varying in soil reaction from pH 3.6-8.2.There is evidence of ecotypic differentiation between populations exploiting acidic (pH 4.0-5.0) andcalcareous soils (pH >7.0), and between plants in shaded and unshaded habitats (Hutchinson 1968).While there are local exceptions, generally the species is rather sparse or absent in areas where soilsare derived from chalk and harder oolitic limestones, or where raw humus or acid organic peat dominates.Areas where this applies are chiefly in SE England, the Irish Midlands and parts of the far west ofIreland (New Atlas).
In Fermanagh, it has not been recorded from walls, hedgerows, quarries, or from other lightly ormoderately disturbed, dry or well-drained habitats with apparently suitable soils, other than on a fewsteep roadside banks, although elsewhere in B & I it is very often found in these kinds ofsituations, eg in lightly grazed limestone grassland and in openings in young coniferous plantations(Hutchinson 1968; Grime et al. 1988, 2007).
The established strategy of T. scorodonia is categorised as CS/CSR, intermediate between astress-tolerant competitor and a general C-S-R, a species with a combination of all three ecologicalstrategies, Competitor, Stress-tolerator and Ruderal (Grime et al. 1988, 2007).
Fermanagh occurrence
Rather surprisingly, considering the variety of soils and extent of limestone terrain in Fermanagh, WoodSage is scarce in the VC and extremely local, being absent from apparently suitable well-drained, acidmineral to mildly calcareous soils on sloping ground in many parts of the county. The Fermanagh FloraDatabase contains records from only 19 tetrads (3.6% of those in the VC), just twelve of which havepost-1975 records. As the tetrad map indicates it is almost exclusively confined to the area SW of LoughErne.
Where T. scorodonia does occur, it is associated with the typical species of more open areas ofdeciduous scrub, woodland margins, limestone pavement or rocky scree as, for example, on or below cliffsat Knockninny, Benaughlin or above Doagh Lough, and on the shallow limestone heath and pavement atLegacurragh above Florencecourt.
In sheltered sites in Fermanagh, T. scorodonia is semi-evergreen and has been confidentlyidentified from March through to December.
Flowering reproduction
Flowers in the genus Teucrium differ from all other B & I Labiates in lacking an upper lip tothe corolla. T. scorodonia flowers from June or July through to September; the calyx, 4.5-6.5 mm,is campanulate (bell-shaped), the tube of it gibbous (ie swollen, inflated or pouched) at the base, theupper end 5-lobed, with the upper lobe much broader than the other four, which are small and incurved,making it overall look 2-lobed. The corolla, 10-11 mm, is yellow or yellow-green, softly hairy on theoutside, one-lipped, the lip with five lobes and the tip deflexed (ie bent back). The central lobe ofthe corolla is large and sub-rotund, the lateral lobes small and short (Melderis & Bangerter 1955;Sell & Murrell 2009).
The flowers are protandrous and produce nectar and pollen that attracts bees and other insects. Seedproduction varies widely, insect cross-pollination being the norm although the species is self-fertile.Experimental selfing gave a marked reduction in seed set, close to 40%. Hutchinson (1968) reckonedannual nutlet production per plant could range from just 400 in deeply shaded conditions where insectvisits would be rare, to around 48,000 in favourable conditions. The four nutlets are smooth, dark brownand are mechanically shaken out of the enclosing calyx by wind, most travelling a metre or less,although on calcareous soils ants have been observed carrying them (Hutchinson 1968).
Seed longevity in soil has been surveyed across NW Europe, with a total of eight results communicated:three found it transient (one year or less), two short-term persistent (1-5 years), one long-termpersistent (>5 years) and two unassigned (Thompson et al. 1997).
The absence of Wood Sage from many apparently suitable sites in Fermanagh and elsewhere may result frompoor levels of seed set, or this plus a general lack of dispersal ability. Additionally, the speciesperhaps displays an over-reliance on vegetative reproduction, which could well play a role in limitingseed production and possibilities of long-distance dispersal. A case for more detailed study ofdispersal ability could obviously be made.
Irish occurrence
T. scorodonia is widespread in Ireland, but largely absent from the Central Plain where bogland iscommon and, also, from the boglands of the western seaboard (New Atlas). Despite a reputedlywidespread calcifuge tendency in B & I, T. scorodonia in W Ireland at least, and especiallyin the Burren, Co Clare (H9), appears indifferent to the base content of the soil. Rather the plant inIreland appears instead to demand good drainage and it is therefore absent from both heavy clays and wetpeat (Webb & Scannell 1983). As it even more obviously avoids wetlands, the observed scarcity of thespecies in Fermanagh may reflect the extreme oceanic climatic conditions and the consequent widespreadoccurrence of water bodies and wet, gleyed or peaty soils throughout the whole year.
On the other hand T. scorodonia does have an Atlantic distribution and it is said to grow bestcolonising and eventually helping to stabilize steep rocky screes, where its well-developed, deeplypenetrating and spreading rhizome allows it to survive and compete on inherently unstable ground. Inthese situations, the loose layers of rock talus provide a mulching effect, maintaining continuouslymoist soil conditions, even on warmer, south-facing slopes (Grime et al. 1988, 2007).
British occurrence
T. scorodonia is well distributed throughout Britain except for parts of the E Midlands and EastAnglia.
Despite the above general habitat features and growing conditions, Hutchinson (1968) quotes examples ofT. scorodonia growing and flowering on the edges of Scottish lochs where its roots are submergedfor several months each year. In complete hydrological contrast, the species also appears on someexceedingly dry, steep slopes on the English South Downs where there is little or no substrate otherthan pure chalk! These very weird exceptions to the general rule suggest the species encompasses widegenetic variation. However, other evidence indicates that there is only limited variation within thisspecies. Alternatively the exceptional range of habitats mentioned indicates that a certain level ofgenetic divergence has occurred and edaphic and shade-tolerant ecotypes exist in at least some parts ofB & I (Hutchinson 1966, 1967).
European and world occurrence
T. scorodonia is described as belonging to the suboceanic southern-temperate phytogeographicalelement and is endemic to W Europe, including the island of Madeira. The species is widespread in W& S Europe and has spread northwards to S Norway (where it is an alien adventive) and eastwards to WPoland and NW Yugoslavia, where it may or not be native. It is also a rare introduction in N America(Hultén & Fries 1986, Map 1577; Press & Short 1994; Sell & Murrell 2009).
Uses
Grigson (1955, 1987) says 'Wood-' or 'Wild-Sage' was not greatly valued by early botanists although itwas known as a diuretic, a wound herb and a herb against broken veins, ulcers, French pox and scurvy.The leaves were picked and used to brew tea to treat rheumatism and, since like other members of thefamily, it contains a bitter tonic substance, it was used in brewing ales and beers before Hops(Humulus lupulus) were introduced. The plant also contains a volatile oil and tannins (Grieve1931).
The medicinal action is described by Grieve as diuretic, astringent tonic and emmenagogue (ie a substancethat stimulates or increases menstrual flow). The same source indicates it was much used in domesticherbal practice for treating skin affections and diseases of the blood, also in fevers, colds,inflammations, rheumatism, gout and as an emmenagogue. It was also regarded as a first rate appetizerand tonic and useful for cleansing old sores. A snuff was also made from powdered leaves to cure nasalpolyps (Grieve 1931). Clearly Wood Sage had its supporters for many years, even if its use has declinedto near zero nowadays.
Names
The genus name 'Teucrium' is derived from a name given to an unknown plant by Dioscorides, called afterthe Greek hero, Teucer, a Trojan prince or first king of Troy, said to have first used one of thespecies in medicine (Johnson & Smith 1946; Gilbert-Carter 1964; Stearn 1992). The Latinspecific epithet 'scorodonia' is derived from the Greek 'skorodon' meaning 'garlic', a reference tothe plant's odour when bruised.
Five local English common names are listed by Grigson (1955, 1987), 'Gipsy' Baccy', 'Gipsy's Sage','Mountain Sage', 'Wild Sage' and 'Rock Mint'. 'Wood Germander' and 'Garlic Sage' are two additionalnames offered by Melderis & Bangerter (1955). 'Germander' is derived from the Latin 'chamaedrys' acorruption from the Greek 'chamai' meaning 'on the ground' and 'drus', 'oak', as if 'Speedwell' or anyother 'chamaedrys' were a 'dwarf oak', from some resemblance of the leaves (Grigson 1955, 1987). Grievegives 'Large-leaved Germander' and 'Hind Heal' as two more names. A further pair of names in Britten& Holland (1886) are 'Ambroise', or 'Ambrose', a name connected with its use in brewing.'Bread-and-Cheese' and 'Hart's-ease' (eaten by sick deer) are supplied by Vickery (2019). Grigsonsuggests the fact that T. scorodonia, "waves no brilliant colour", meant it tended tobe rather 'neglected' when it comes to local names, but the current author (RSF) things otherwise.
Threats
None.
Native, very common. European temperate, adventive in Fennoscandia and introduced in N America.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This is a low-growing, rosette-forming perennial with a short, thick rhizome that sends out slender,creeping, rooting and ascending leafy suckers or stolons. In this manner, A. reptans regularlyforms, semi-evergreen spreading patches in unimproved pasture and meadow grasslands. The aerial stemshave two lines of hairs running down opposite sides, shifting to the other two sides at each node.Leaves opposite, simple; radical leaves long-stalked, stem ones sessile and gradually shortening intocoloured bracts (Hutchinson 1945, 1972). Leaf under-surfaces darkened or reddened with anthocyaninpigment.
A. reptans occupies a wide range of habitats and is really absent only from the most acidic peatysoils, truly aquatic habitats and the highest or most disturbed ground. The most typical habitatsinvolve damp to moist, well-drained, moderately fertile, mildly acid to calcareous soils in partial- todeep shade. It commonly appears in open or marginal areas in woods, recent coppice or felled areas, butit persists well whenever shade develops. It also occupies scrub, hedges, along lakeshores and ditches,and in damp pastures, meadows and unimproved grass or more open roadside verges.
The established strategy of A. reptans is categorised as R/CSR, intermediate between Ruderal and amore balanced mix of all three ecological strategies, Competitor, Stress-tolerator and Ruderal (Grimeet al. 1988, 2007).
Fermanagh occurrence
Bugle has been recorded in 342 Fermanagh tetrads (64.8% of the VC total) and is very widely scatteredacross the whole county – too common to map!
Vegetative reproduction
The species has a short, thick rhizome but it spreads very effectively by means of long, leafy,prostrate, aerial stolons or runners that root at intervals, enabling the plant to form clonal patchesindependent of seed production. It is unclear whether vegetative reproduction is the dominant form ofplant maintenance or not but, of course, seed production remains significant when it comes to longerdistance species transport and long-term adaptation to changes in environment.
Flowering reproduction
The plant gains considerably in height when it flowers in mid-spring, reaching up to 25 cm as opposed tothe more usual 10 or 12 cm vegetative leaf canopy. The flowers, which appear from April to July, varygreatly in colour, although they are typically found in shades of blue or bluish-purple, only rarelyappearing pink or white in tightly packed whorls of usually six flowers among coloured leafy bracts, theupper ones shorter than the flowers. The calyx is bell-shaped, divided almost to the mid-point into fivenearly equal segments. The tubular corolla, five times as long as the calyx, has a very shorterect, 2-lobed upper lip and a much longer, more conspicuous, spreading, 3-lobed lowerone. The four parallel stamens project well beyond the tube and upper lip of the corolla (Melderis &Bangerter 1955; Clapham et al. 1987).
The nectar-rich flowers are hom*ogamous and have little or no scent. Nevertheless, they are usuallypollinated by bees, or they can self-pollinate. Either way, each fertilised flower can produce up tofour, ellipsoid to ovoid, brown nutlets, 3.0 × 1.8 mm, each one decorated with a coarse reticulatesurface pattern and bearing a conspicuous lateral scar (Hutchinson 1945, 1972; Butcher 1961). Manyovules do not mature to this stage (Grieve 1931). Ripe nutlets possess an attached nutritive elaiosomeoil-body, common to many, but not all, members of the Lamiaceae family. These attract ants that helpdisperse them locally, thus minimising seed predation. The nutlets are also dispersed by attachment inmud to animals, including man and his vehicles (Ridley 1930, pp. 521, 533).
British and Irish occurrence
A. reptans is common and widespread, almost ubiquitous throughout B & I, the distribution evenat the hectad scale clearly reflecting the species intolerance only of permanently wet or veryinfertile, acidic, peaty conditions which are mainly met in northern Scotland and parts of C Ireland(New Atlas).
European and world occurrence
A. reptans belongs to the European temperate phytogeographical element and is mainlyrestricted to Europe where it is widespread northwards to around 61°N, although adventive in NFennoscandia. It is also found in W Asia and in a few places in N Africa. It has also been introduced toN America and New Zealand, where it remains sporadic and nowhere extensively naturalised (Hultén &Fries 1986, Map 1576; Webb et al. 1988).
Uses
Gerard (1597) regarded Bugle, along with the related Selfheal (Prunella vulgaris), as beingamongst the very best vulneraries or wound herbs available.
The plant is bitter, astringent and aromatic. In herbal medicine, an infusion of the plant is consideredvery useful for arresting haemorrhages and is employed for coughs and for those spitting blood ininsipient consumption. It is also used for treating some biliary disorders. In action, it is somewhatsimilar to Digitalis purpurea (Foxglove), lowering the pulse and slowing its frequency. It hasbeen described as, "one of the mildest and best narcotics in the world" (Grieve 1931).
Names
The origin of the genus name 'Ajuga' is problematic, obscure and there are numerous suggestions to beconsidered. It is a name used by a physician referred to as 'Scribonius Largus' as a synonym (orcorruption) of the Latin 'abiga', said to be a plant mentioned by Pliny that produced abortion ('abigo'means 'procure abortion'), and the herb was reputed to have been used for this purpose (Gilbert-Carter1964). Another suggestion is that the name was derived as a corruption of the Greek 'abija', in turnderived from the Latin word 'abigo', meaning 'to drive away', because the plant was thought to driveaway various diseases. It was, however, chiefly valued as a wound herb on account of its undoubtedastringency (Grieve 1931). It was an ingredient, along with Sanicle and Scabious, of an ointment forulcers, wounds and bruises (Grigson 1955, 1987).
Alternatively, it is suggested 'ajuga' is a combination of the Greek, 'a' meaning 'no' and 'zuron', 'ayoke', supposedly a reference to the fact that the calyx lobes are equal, rather than bi-lipped (Johnson& Smith 1946; Chicheley Plowden 1972; Hyam & Pankhurst 1995). This seems extremely fanciful andcontrived to the current author (RSF), but all things are possible when it comes to the derivation,allusions or suggested linkages with plant names, which are, after all, simply labels.
Grieve (1931, p. 140) recognises that the names of this plant, both scientific and popular, are not easyto account for. The same sort of difficulty arises when the origin and/or meaning or allusion of thecommon name 'Bugle' is sought after. It has been suggested that 'Bugle' is derived from 'bugulus', 'athin glass pipe used in embroidery', the long thin tube of the corolla being thought to resemble this'bead bugle' (Grime 1931). Grigson (1955, 1987) agrees that 'Bugle' is an obscure name, although he saysit has counterparts in German, French, Italian and Spanish, from the apothecaries' 'bugula'. He goes onto clarify that it has nothing to do with the bugle as in a brass musical instrument that is blown,though it may be the source of the 'bugle' meaning the long, tube-shaped, glass bead, black or blue incolour, that is [or was] sewn on to clothes, a possible allusion to the, "glistening dark blue orviolet of its [ie Bugle's] own leaves" (Grigson 1955, 1987).
Prior (1879, p. 31) gives 'Bugle' as being from the medieval or Low Latin 'bugula', a diminutive of'abuga', one of the various spellings of a word given by Pliny as corresponding to the Greek'Хαμαιϖιτυς', variously written 'abiga', 'ajuga', 'iva', etc. (Grigson 1974). 'Bugle' was also referredto as 'bugula', 'bugil', 'bugile', 'bugyl', 'bugyll' 'bugill' and 'begel' in medieval England, soconfusion often raged and it is not surprising that A. reptans was sometimes confused with'Bugloss' or 'Buglossa' (Anchusa arvensis) (Hunt 1989).
A list of an additional 15 very varied English common names are provided by Grigson (1955, 1987),emphasising the fact that A. reptans was both well known to herbalists and popular in folkmedicine.
Threats
None.
Introduction, garden escape, very rare. Eurosiberian temperate, but very widely naturalised in bothhemispheres.
1872; Smith, T.O.; unspecified site on the banks of the Colebrooke River.
Growth form and British occurrence
Cat-mint is a perennial herb with thick, fleshy roots and branched leafy stems with a strongly aromatic,mint-like scent. This aroma provides a strong fascination for cats, which love to roll themselves in itand hence the English common name. In gardens, on occasions, cats may spend so much time on top of theplant, they will severely damage or even kill it (Grieve 1931).
N. cataria produces pale green stems that grow 40-100 cm tall, erect, hollow and 4-sided, withdense, white, eglandular hairs that cover the plant with a mealy down and minute glands that contain thescent. It flowers from July to September and is pollinated mainly by bees, or selfed (Garrard &Streeter 1983; Sell & Murrell 2009).
As recently as 1997, Stace in the 2nd edition of his New Flora of the British Isles was claimingthat this familiar, but rather local species was, "probably native" in England and Wales,although making no mention of its status or presence in Scotland, and recognising it as, "rarelynaturalised or casual in Ireland". It has now been declared an archaeophyte in Britain, having beenintroduced to Britain in the Middle Ages (Grigson 1974; Preston et al. 2004). Populations inBritain in open grassland, hedge banks, waysides, roadsides and rough ground, preferably on calcareousor gravelly soils, have been in decline since at least around the 1960s (K. Walker, in: Preston etal. 2002).
The New Atlas hectad map shows N. cataria is a very rare and scattered species north of aline between Lancaster and Hull. The loss of habitat it has suffered in the last 60 years in S Englandand Wales is principally associated with the intensification of agriculture, including the removal ofhedges (K. Walker, in: Preston et al. 2002).
Fermanagh and N Ireland occurrence
N. cataria is an extremely rare escape from cultivation in Fermanagh and likewise elsewhere in NI.The above listed very old record is the only one ever in the Fermanagh Flora Database. Hackney etal. (1992) also report the species similarly rare in the FNEI 3 and they note that it issomewhat more prevalent near the coast in the NE of the island, although unusually persistent on thesand-dunes on the southern shore of Lough Foyle around Magilligan – Benone and Downhill, Co Londonderry(H40).
Irish occurrence
In Ireland, in comparison with Britain, this species is merely casual on dry waste ground and it veryseldom proves long-persistent. There are no post-1970 records for N. cataria anywhere in Irelandand the New Atlas hectad map plots just six pale pink spots representing pre-1970 records, fourof which are in the province of Ulster! All but the solitary Fermanagh record listed above lie on ornear the coast.
This situation for N. cataria in Ireland did not always apply. In the mid-19th century, thespecies, although still rare, had many stations scattered across the island. Cybele Hibernica(1866) listed 17 locations from eight of its twelve 'divisions' of the island. The plant cropped up onwaysides, waste places, banks and near ruins. Many of these sites had already disappeared by the end ofthe century, however, and the plant had by that time become recognised as a definite alien, gardenescape (Colgan & Scully 1898). Oddly enough there were very few reports of Cat-mint escapes fromIrish gardens during the 20th century, perhaps because of some change in the choice of popular cultivarsbeing grown, which may have proved more casual than persistent in the wild (Webb 1980; Cat Alien PlIr).
European and world occurrence
N. cataria is believed to have originated in S Europe and W & C Asia (Grigson 1974; Hultén& Fries 1986) and it has been very widely introduced on account of its many uses as a medicinal herbto such places as the rest of Europe, S Africa, N America, S Australia, Tasmania and New Zealand. Thedistribution has consequently become discontinuous circumpolar (Hultén & Fries 1986, Map 1602).
Uses
In herbal medicine, N. cataria is regarded as a carminative (ie relieves flatulence), tonic,diaphoretic (ie induces perspiration), refrigerant (ie causes cooling) and slightly emmenagogue (ieincreases or stimulates menstrual flow), especially antispasmodic and mildly stimulating (Grieve 1931).It is considered very useful in combating colds as it produced free perspiration without causing heatingof the system. It is also useful in treating fevers for the same reason. Catnip tea is used to treatcolicky pains and to relieve pain and flatulence. It is said to be good for inducing sleep and helpfulfor treating restlessness and nervousness. One of its chief uses is in treating childrens' ailments, andGrieve (1931) lists many other uses to which Cat-mint has been put, too many to relate here. There isjust a possibility that confusion might occur as Cat-mint and Calamint (Clinopodium ascendens)might occasionally be mixed up in terms of their herbal use, since for instance they are bothrecommended for treating colds and may be drunk as herbal teas (Allen & Hatfield 2004).
In France, Cat-mint leaves and stems were regularly used to season dishes and it was specially cultivatedin kitchen gardens for this purpose. It is also used as a herbal tea. N. cataria was a populardecorative garden subject until it became largely replaced by more vigorous, more distinctive relatives,including the sterile hybrid N. × faassenii Bergmans ex Stearn. Thishybrid arose in cultivation in 1784 and quickly became popular with gardeners. It was not recorded inthe wild in Britain until 1928. The hybrid possesses a strong rhizome and it occasionally escapes fromcultivation or is discarded whenever it becomes too invasive. It is widely but thinly scattered acrossBritain on waysides, waste ground and rubbish tips, especially in coastal areas, and it is probablystill increasing (K. Walker, in: Preston et al. 2002).
Names
The genus name 'Nepeta' is a name given to an unknown aromatic, mint-like plant by the Roman writer,historian and scientist, Pliny (Grigson 1974). 'Nepeta' or 'Nepete' is also the name of a city inEtruria (Gilbert-Carter 1964). The name might also originate from 'Nepi' in Italy (Gledhill 1985). TheLatin specific epithet 'cataria' is from 'catarius', 'of cats' (Gledhill 1985).
The English common name 'Catmint' (or 'Catnep' or 'Catnip') is 13th century, recognising that cats show astrong liking for the plant. The plant was known by the medieval Latin names 'herba felina', 'herbacati' or 'herba catti', and 'herba cattaria' (Grigson 1955, 1987; Grigson 1974). Alternative namesinclude 'Dogmint', 'Nep', 'Nep-in-a-hedge' (Grigson 1955, 1987). 'Nep' or 'Neppe' is merely acontraction of the Latin 'Nepete' (Prior 1879).
Threats
None.
Native, locally frequent. Eurasian boreo-temperate, but naturalised in N America and now circumpolarboreo-temperate: also introduced in SE Asia and New Zealand.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This familiar wayside species is a creeping, stoloniferous, patch- or carpet-forming, wintergreenperennial with distinctive, hairy, opposite, long-stalked, kidney-shaped leaves. It is typically foundin a range of lightly shaded habitats in woods, scrub, hedgerows, stream-sides and roadside banks,usually on slightly disturbed, fertile soils where competition from potential dominant species isrestricted by sub-optimal growing conditions. G. hederacea has a limited ability to tolerateheavy shade from tall herb vegetation or woodland canopy and is characteristic of more open, lightly- orpatchily-shaded marginal conditions, where it itself can become the dominant species, forming extensiveclonal carpets (Clapham et al. 1987).
Ground-ivy is most frequent and grows best on damp, heavy soils in the pH range 5.5-7.5, but it can occuron the much more acidic, peaty soils down to around pH 4.0. It is principally a lowland species, but canoccur up to about 450 m (Garrard & Streeter 1983; Grime et al. 1988, 2007). While it is neverfrequent in pastures, G. hederacea contains bitter-tasting toxins and is avoided by stock andother herbivores. It is also toxic to some species of rodents, but is a food plant of many invertebrates(Cooper & Johnson 1997; Hutchings & Price 1999).
The established strategy of G. hederacea is categorised as CR/C-S-R, ie intermediate betweenCompetitive Ruderal and the more general balance of all three strategies,Competitor-Stress-tolerator-Ruderal (Grime et al. 1988, 2007).
Variation
Three varieties are distinguished and named in B & I, although their distribution and ecologicaldifferences remain unknown (Sell & Murrell 2009). The common form is var. hederacea which hasleaves up to 3.0 × 3.5 cm with few to numerous hairs and a corolla 12-15 mm long. The other two are var.minor Gilib., which has the smallest leaves of the three, up to 1.2 × 1.2 cm, often densely hairyand a corolla 8-10(-15) mm long; and var. grandiflora H. Mart., with large leaves, 4.0 × 4.5 cmwith sparse hairs and a corolla 15-20 mm long (Sell & Murrell 2009).
Fermanagh occurrence
G. hederacea has been recorded in 128 tetrads (24.2% of the total) widely scattered throughout theVC. However, having said that, it is much more local than one might predict. It definitely seems toprefer the lowland conditions in the Lough Erne basin and, very surprisingly, there is just one isolatedrecord from the Western Plateau uplands (in the Monawilkin limestone district, where it was found by RHNin 1990). On the other hand, the altitude in Fermanagh, lying mostly below 250 m, is insufficient toprovide a limiting factor. Excessive soil acidity is a more likely explanation for the distinctly localFermanagh distribution of the species, which prefers damp, heavy, fertile and calcareous soils(Hutchings & Price 1999).
Flowering reproduction
G. hederacea flowers early in the year, from March to May. The bi-lipped flowers are borne onerect stems 10-30 cm tall, in false whorls (verticillasters) of two to five blossoms, in the axils ofthe upper leaves, which are generally tinged purple. The violet flower petals are purple-spotted on thelower lip. The flowers are protandrous, contain nectar, and are pollinated by bumble-bees, honey-beesand beetles. The main period of seed set is in June, although this can continue into July (Hutchings& Price 1999). In flowering early, the species avoids the level of shade provided by full summerleaf canopy, thus achieving light conditions that permit its more successful flowering and better seedset.
Ground Ivy, like Wild Thyme (Thymus polytrichus), is gynodioecious, having both perfect bisexual(ie hermaphrodite) flowers and smaller female flowers, either on the same, or on separate bi- anduni-sexual plants (Price 1991). In a rather odd observation, although bisexual flowers areself-compatible, they were still found to require an insect visit for pollination to occur (Hutchings& Price 1999). Despite the gynodioecy adaptation designed to maximise sexual reproduction, reportssuggest that seed set is sometimes low, although many other populations do manage to set largequantities of viable seed. Seed set is especially poor in isolated, large, male-sterile clones, which isnot surprising (Hutchings & Price 1999).
When fruit is ripe, the calyx bends down and the seeds (a maximum of four nutlets per flower) arepassively dispersed. They may be further dispersed by ants (Bouman & Meeuse 1992).
On the basis of studies of the weed flora of an arable field, Chancellor (1985) suggested seed issometimes short-lived and, therefore, effective regeneration and spread may, in such circ*mstances, beprimarily vegetative. However, other observations, such as the observed rapid colonisation by G.hederacea of secondary or coppice woodland, suggest it may have a persistent soil seed bank thatmight survive for 40 or more years (Peterken 1981; Petrov & Palkina 1983; Thompson et al.1997).
Vegetative reproduction
In situations where G. hederacea manages to avoid deep shade and aggressive competition, it canrapidly develop a single species carpet of growth by means of its numerous far-creeping stolons thatregularly root at nodes at intervals of around 8-15 cm. Stolons can reach up to 2 m in length, but theydie off in the autumn, leaving the rooted ramets to overwinter as small, 8-leaved rosettes (Hutchings& Price 1999). Seed production and efficient dispersal appears in some situations to be relativelyrare, so effective regeneration in these cases must often primarily depend upon vegetative spread. Thustransport of shoot fragments may be more significant than seed in achieving long-distance colonisationin such cases (Chancellor 1985; Grime et al. 1988, 2007).
British and Irish occurrence
The New Atlas hectad map indicates that Ground-ivy is widespread throughout Ireland except in VCsalong the W coast. An odd area of absence includes the inland portion of Co Leitrim (H29). In Britain,the distribution of G. hederacea thins markedly towards the Scottish Highlands and N Scotland andit is absent from the Hebrides, Orkney and Shetland (Garrard & Streeter 1983; Clapham et al.1987; New Atlas).
European and world occurrence
G. hederacea s.l. is widespread as a native almost throughout Europe, W & N Asia to Japan. InN America, it is a naturalised alien introduction and it has also been introduced to SE Asia and NewZealand. The present distribution is discontinuous circumpolar and it is categorised as belonging to thecircumpolar boreo-temperate element (Hultén & Fries 1986, Map 1603; Clapham et al. 1987; Sell& Murrell 2009).
Uses
Before the introduction of Hops (Humulus lupulus) in the 16th century, Ground-ivy was the chiefsource of bitter for flavouring and clearing beer for the early Saxons. This is reflected in the Englishcommon names 'Alehoof' and 'Tun-hoof' (Grieve 1931).
G. hederacea has been used in herbal medicine as a diuretic, astringent, tonic and gentlestimulant. It has been regarded as one of the most popular herbal remedies for coughs and nervousheadaches, and was also useful in treating kidney diseases, indigestion and consumption. A brew of 'GillTea' made from Ground-ivy with added honey was a cooling beverage much used for treating stubborn, longstanding coughs and for clearing the blood. Gill Tea was also recommended for weakness of the digestiveorgans, being stimulating and tonic, and for treating all kidney complaints. A snuff made from the driedleaves was used for treating headaches (Grieve 1931).
Several horticultural varieties with variegated leaves occur that are popular for decorative troughs andhanging baskets.
Names
The genus name 'Glechoma' is from the Greek 'glechon', a name in Dioscorides for a kind of mint,'Mentha pulegium' (Gilbert-Carter 1964; Hyam & Pankhurst 1995). The Latin specific epithet'hederacea' means 'resembling Ivy' or 'resembling Hedera', referring to its leaves (Gledhill1985; Stearn 1992).
There are no less than 37 English local common names listed by Grigson (1955, 1987), many of which referto the spreading stoloniferous nature of the plant, eg 'Gill-creep-by-the-ground' 'Blue-runner' and'Robin-run-in-the-hedge'.
'Ground-ivy' is a rather poor name, alluding to the running nature of the plant, although its leaves inno way resemble those of Hedera helix (Common Ivy). However, it is a very old name, translatingthe 'chamaikissos' of the Ancient Greeks and Romans (Grigson 1955, 1987). There are 29 English commonnames listed by Vickery (2019), some additional to those of Grigson, again emphasising just how common,widespread and familiar the species is to the wider population.
Threats
None.
Native, common, widespread and locally abundant. Circumpolar wide-temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
P. vulgaris is a very common, widespread, almost ubiquitous, shallow-rooted, stoloniferous, flatrosette-forming, wintergreen, patch-forming perennial. It has a branching, prostrate stem from whicharise both leafy and flowering shoots, generally in the range 8-25 cm tall, but rarely stems reach 60 cm(Garrard & Streeter 1983; Sell & Murrell 2009).
Prunella vulgaris occurs in a wide range of habitats and is especially frequent in short turf,damp meadows, pastures, hedgerows, open areas by tracks or on the margins of woods or scrub, roughgrassy roadsides, around rock outcrops and on waste ground up to altitudes of around 750 m (Garrard& Streeter 1983; Grime et al. 1988, 2007). The plant is largely restricted to moist,moderately fertile, less acidic soils of pH above 5.0 and it is easily shaded out by taller growingherbs in less disturbed vegetation.
P. vulgaris is a particularly common, aggressive patch-forming weed in regularly mown, closelygrazed, or trampled permanent grassland, such as pastures, lawns, parkland and roadside verges(Salisbury 1964; Grime et al. 1988, 2007). The nutlets (ie achenes, single-seeded dry fruits) ofSelfheal were a common contaminant of commercial agriculture Clover seed mixtures in past years(Salisbury 1964), which helps to explain the species almost ubiquitous distribution in B & I andfurther afield.
P. vulgaris is also extremely tolerant of heavy grazing pressure and moderate levels of trampling,allowing it to become common and often abundant in short or open turf of older pastures and, for thesame reasons, it is also a common lawn weed. Individual leaves and stems are short-lived and, likeTrifolium repens (White Clover), P. vulgaris is rather mobile in pastures, expanding intogaps in the turf on a ± temporary basis, until ousted by more robust, competitive species. Leggy plantsare also quite frequent in woodland conditions, although here again, they are probably notlong-persistent.
Selfheal is especially associated with poorly drained, clay or loam soils, varying from moderately acidto calcareous in nature, that support short turf vegetation (Grime et al. 1988, 2007). Oncalcareous soils where P. vulgaris is most abundant, the flowers are often at their very bluestin colour (Salisbury 1964). Somewhat unusually for this flower family, the foliage is not aromatic andthe flowers are not scented (Melderis & Bangerter 1955). The established strategy of P.vulgaris is categorised as C-S-R, meaning it displays a balance of all three principalecological strategies, Competitor, Stress-tolerator and Ruderal (Grime et al. 1988, 2007).
Fermanagh occurrence
P. vulgaris ranks as the 27th most frequently recorded vascular plant species in the FermanaghFlora Database, immediately following Trifolium repens (White Clover). It is also very widespreadthroughout the county, having been recorded in 460 Fermanagh tetrads, 87.1% of the total. This ranks it26th in terms of tetrad frequency, again just behind T. repens in this respect!
Flowering reproduction
P. vulgaris flowers from June to September, the inflorescence being a short, dense, terminal,cylindrical or oblong spike of less than 50 violet-blue flowers arranged in six to eight whorls. Each ofthe tightly packed whorls contains about six bi-lipped flowers arranged between two quite largebroadly-oval bracts which are green with purple margins and fringed with hairs (Salisbury 1964). As withThymus polytrichus (Wild Thyme) and Glechoma hederacea (Ground-ivy), the flowers of P.vulgaris are of two types, small and female, or bisexual and larger on the same or separateplants (ie the plants are gynodioecious). However, unlike the other two species, the flowers of P.vulgaris are self-sterile (Warwick & Briggs 1979). The flowers in each whorl do not all opensimultaneously, so that a slightly ragged looking flower-head results (Grieve 1931). Nectar is producedinside the corolla at the base of the stigma and is protected from small, crawling insects by a thickhedge of hairs placed just above it. Pollination is carried out mainly by long-tongued bees (Garrard& Streeter 1983).
An average-sized plant produces around 850 individual nutlets or schizachenes (ie single-seeded dryfruits) or 'seeds' and, with a mean germination rate of around 70%, this gives a calculated meanreproductive capacity of 600 seedlings/plant/year (Salisbury 1942, p. 162). Nutlets can pass through thealimentary canal of birds unharmed and produce seedlings on their excreta, providing an effective meansof long-distance dispersal and establishment (Salisbury 1964, p. 104).
Vegetative reproduction
Established plants send out creeping stems and stolons that root and create daughter ramets. These becomeindependent plants by the connection with the parent plant decaying and breaking down, usually withinabout a year or so. In this way, P. vulgaris can very easily reproduce vegetatively, spreadingclonally to form ever larger, loose local patches over several years. In trampled, short-turfgrasslands, plant fragments may also become detached, transported and re-established, helping to spreadthe species locally to some extent and adding to the reproductive achievements of seed dispersal. Therelative importance of seedlings and vegetative fragments in the colonisation of disturbed groundrequires further study (Grime et al. 1988, 2007).
British and Irish occurrence
P. vulgaris is very frequent throughout the length and breadth of both B & I as shown in theNew Atlas hectad map. It is generally regarded as one of the most common and widespread speciesin the flora of B & I and, in common with the high percentage of Fermanagh tetrads in which thespecies is recorded, it is considered very likely that a similar degree of presence occurs across itswhole B & I range (K. Walker, in: Preston et al. 2002).
European and world occurrence
This polymorphic and widespread species is thought to have originated in Europe and probably adjacentparts of Africa and Asia. In the southern half of Europe, the closely related P. laciniata(Cut-leaved Selfheal) occurs. From its native European range it has spread with agricultural man to N, C& S America, SE Asia, Australia and New Zealand. The native populations of E Asia and N America aredistinguished as subsp. lanceolata (Bart.) Hult. and, in India, subsp. hispida (Benth.)Hult. occurs. P. vulgaris s.l. belongs to the circumpolar plants and the species is categorisedas belonging to the circumpolar wide-temperate phytogeographical element (Hultén & Fries 1986, Map1607; Preston & Hill 1997).
Uses
The ancients and early herbalists believed P. vulgaris to be a universal panacea for healing allsorts of ailments, but it was especially valued as one of the very best wound herbs. Grieve (1931)describes its uses as astringent, styptic (ie checks or staunches bleeding) and tonic. An infusion ofthe herb was useful as a gargle for treating sore throats or an ulcerated mouth. The infusion could alsobe used as an injection to treat internal bleeding, including for piles. The herb was also applied tofresh external wounds as an unguent (ie an ointment) or as a plaster, to stop bleeding and inducehealing. Selfheal was sometimes used in conjunction with other wound herbs such as Sanicle (Saniculaeuropaea) and Bugle (Ajuga reptans) to dress external wounds such as cuts and bruises(Grieve 1931). P. vulgaris was also used to treat colds, respiratory complaints and hearttrouble, to mention just a few of many ailments for which it was applied, especially in Ireland (Allen& Hatfield 2004).
Names
The genus name 'Prunella' may be a corruption of 'Brunella', from the German 'Bräune', meaning 'quincy'(a collection of bacterial pus between the tonsils and the wall of the throat – a rare but seriouscomplication of tonsillitis), which it was supposed to cure (Grigson 1955, 1987; Melderis &Bangerter 1955). 'Brunella' is the older spelling of the name, which was first given by Otto Brunfels(1489-1534), one of the early German herbal writers and 'Fathers of botany' (Gilbert-Carter 1964). TheLatin specific epithet 'vulgaris' simply means 'common', which in this case the plant certainly is(Gledhill 1985).
Seventeen English common names for the plant are listed by Grigson 1955, 1987), several of which like'Heart's Ease', 'Herb Bennet' and 'Wood Sage' are more frequently applied to species other thanPrunella vulgaris. Vickery (2019) goes a few better, listing 21 local names. One of these, 'Heartof the Earth' is in widespread use and, according to Britten & Holland (1886), the reason was,"it chiefly occurs on thin, poor soils, where the farmers give it the credit of eating away all thesubstance of the soil". The name 'Brownwort' refers back to the origin of the generic name,'Brunella'. Names referring to carpenters such as 'Carpenter-grass', 'Carpenter's herb' and 'Proudcarpenter' connect with the use of the plant to staunch flesh cuts (Vickery 2019).
Threats
None.
Introduced, garden escape, locally extinct. Native of S Europe and SW Asia, but native range obscured byintroductions further north.
August 1881; Barrington, R.M.; Trasna Island, Upper Lough Erne.
Growth form, origin and preferred habitats
This shortly rhizomatous perennial herb, 20-150 cm tall, has long straggly branches and stems, 4-angled,glabrous (hairless) below, finely hairy towards the top. Plants produce stolons that can spread rapidlyand the leaves smell strongly of lemon when crushed. Leaves are of two kinds, the larger only on themain stem, 6-10 × 4-7 cm, long-stalked, ovate, rounded to slightly cordate at the base, their marginscrenate or crenate-serrate. Smaller leaves, cuneate (wedge-shaped) at the base, are produced on all theother stems, especially on the inflorescence. The bisexual flowers are borne from June to August ormid-September in dense, shortly-stalked axillary whorls of four to twelve, accompanied by large, leafybracts and paired bracteoles. The persistent calyx is campanulate, 2-lipped and 13-veined. The corollais 8-15 mm, pale yellow, becoming white or pinkish as it ages, the tube curved and dilated above themiddle, 2-lipped, the upper lip erect or deflexed, the lower one 3-lobed (Hutchinson 1972).
M. officinalis was introduced to garden cultivation in B & I from S & C Europe for itsculinary herb qualities by around 995 AD (Harvey 1961). There are golden-leaved forms which often arethe preferred decorative garden varieties (eg cv. 'All Gold' and cv. 'Aurea'). The dried leaves retaintheir lemon scent and are frequently used for pot-pourri. The young shoots are used fresh for flavouringfruit salads and iced drinks (Sell & Murrell 2009).
A patch-forming archaeophyte, it was first recorded 'beyond the garden wall' in the wild in Britain by1763. The plant spreads quickly in fertile soil in garden settings and thus can become excessivelyinvasive, leading to it being cut back and excess material discarded. The discards are very probably theprincipal source of the plant appearing beyond the garden wall, where it has the ability to becomenaturalised, set seed and become self-sown (Sell & Murrell 2009).
Fermanagh occurrence
Barrington (1884) wrote of this species, "Occurs near the garden on the Island of Trasna withTanacetum vulgare and Chrysanthemum parthenium. All three probably planted." As faras the current author (RSF) knows, it has not been seen in Fermanagh for over a century and, therefore,it is considered extinct.
Irish occurrence
The New Atlas shows the species in Ireland very thinly scattered SE of a line between Dublin andLimerick, with two outlying hectads further north in Cos Monaghan (H32) and Fermanagh (H33). Having saidthat, the Cat Alien Pl Ir states that the plant has been recorded at least once from 20 of the 40Irish VCs.
British occurrence
M. officinalis is much more frequently found in the wild in S England than elsewhere in Britainand it is recognised as a long-lived established garden escape or relic of cultivation in old orabandoned gardens. However, it does have scattered outlying stations further north in England, and inScotland it stretches as far north as Dunbartonshire (VC 99) (New Atlas). An apparent increase inabundance or more widespread occurrence of this species in recent BSBI national or local county surveysis perhaps more likely due to more active, systematic recording rather than actual M. officinalisspread, since there is only occasional evidence of it spreading by seed, self-sowing, or even of itpatch-forming in the wild (Crawley 2005).
European and world occurrence
Melissa officinalis is a native of C & S Europe, W Asia and N Africa and is widely introducedelsewhere (Hutchinson 1972; Clapham et al. 1987; Sell & Murrell 2009).
Uses
Even if it is of no outstanding physical beauty, M. officinalis is worth growing in the decorativegarden for the delightful lemon fragrance of its leaves (Grigson 1955, 1987). Balm, or 'Lemon Balm' wasused as a cordial herb, said to, "remove melancholy and cheers the heart" (Grigson 1955,1987). A tea was brewed that was regarded as a tonic, a uterine and a sedative, although as Grigsonsays, "it is by no means as nice as the scent of the leaves would lead you to expect". Earlybotanists equated Balm with the 'melissophyllum', or 'bee leaf' of Dioscorides ("so called, becausebees delight in this herb") and the 'apiastrum' of Pliny was planted or rubbed on hives since itwas believed to ensure honeybees kept to their hives (Gerard 1597).
A brew of Balm was thought to keep insects away. It was also used to bathe bitches in season, to helpkeep male dogs away. An infusion was used to reduce itching in the later stages of Chickenpox. As a'green leaf tea' it was used to induce drowsiness in the middle of a sleepless night. The tea was alsotaken for stomach problems and colic (Vickery 2019). The herb was greatly esteemed by Paracelsus, whobelieved it could completely revive a man and was of great use to treat any disordered state of thenervous system. "An essence of Balm taken every morning was reputed to renew youth, strengthen thebrain, relieve languishing nature and prevent baldness." (Grieve 1931). Balm also makes a usefulflavoursome addition to omelettes. It is now perfectly clear to the current author (RSF), why the plantwas so widely cultivated.
Names
The genus name 'Melissa' refers to the Greek, 'melissa', meaning 'a honeybee', but is also believed (bysome) to be the name of a Cretan nymph or Princess who first discovered how to keep bees for their honey(Gilbert-Carter 1964; Stearn 1992). The Latin specific epithet 'officinalis' refers to the 'apothecary'soffice' or shop where herbal medicines were collected, manufactured, stored and dispensed.
English common names include 'Lemon Balm', 'Sweet Balm', 'Honey Balm' and 'Bee Balm', all easilyappreciated. The name 'Balm' is an abbreviation and comes through the French from the Latin 'balsamam','balsam', the chief of sweet-smelling oils (Grieve 1931; Grigson 1955, 1987). Variations such as 'bame','baume' and 'bawne' are recorded in several English locations (Vickery 2019).
Threats
None.
Probably introduced, garden escape, rare. Euroasian southern-temperate, introduced in N & S America.
1903; Praeger, R.Ll.; Co Fermanagh.
April, May & November.
Growth form and preferred habitats
This rhizomatous, deeply rooted, variably hairy and rarely glabrous, tufted, herbaceous perennial hasbeen used as an aromatic, culinary, kitchen garden plant and medicinal herb throughout B & I forcenturies. The long, deep root run and very numerous, long root hairs allows the species to tolerateprolonged periods of drought. Beyond the garden wall, stems can grow up to 90 cm, although veryfrequently sub-optimal growing conditions limit it to much less than this, closer to 30 cm in height.The rhizome is short and sends out runners or stolons. Despite the latter ability, the species showsonly a limited ability to spread laterally in most habitats. Stems are annual, slender, flushed purple,branched above, often with numerous short, sterile axillary branches, giving the plant a definite tuftedappearance. Stems are woody at the base and along their length they are densely clothed withlong-stalked, opposite leaves (Melderis & Bangerter 1955; Sell & Murrell 2009).
A colonist of bare or sparsely occupied, dry, infertile, calcareous soils, the range of local habitats inthe Fermanagh area includes dry hedge banks, bridges, old walls with lime-mortar and rough grass onroadsides. It generally grows where there is minimal competition and little or no pressure from mowing,grazing or trampling. The distribution of the species is centred on vegetation where competition islimited by a combination of mineral nutrient deficiencies and occasional disturbance. The establishedstrategy of the species is categorized as intermediate between C-S-R (ie a balance of all threestrategies, Competitor, Stress-tolerator and Ruderal) and Stress-tolerant competitor (SC) (Garrard &Streeter 1983; Grime et al. 1988, 2007).
Variation
In Europe, the culinary herb referred to as 'Oregano' or 'Marjoram' can be one of several relatedspecies, or a hybrid or mixture of any of these species. The species include O. vulgare, O.majorana L. (Sweet or Knotted Marjoram, a widely cultivated species), O. onites L. (PotMarjoram, a dwarf shrubby species from the Mediterranean area, also widely cultivated), O.syriacum L. and the hybrid between O. majorana and O. vulgare (Sell & Murrell2009).
O. vulgare itself is extremely variable in the colour and indumentum of its bracts and calyx, theshape and length of the inflorescence 'spicules' (see below) and corolla colour. Four subspecies arerecognised in B & I: subsp. vulgare, which is the native form in Britain and is used as botha medicinal and culinary plant; subsp. hirtum (Link) Ietsw. (= O. heracleoticum L.)(Winter Marjoram); subsp. virens (Hoffmanns. & Link) Ietsw.; and subsp. viride(Boiss.) Hayek. All four subspecies are grown in gardens in B & I and they occasionally escape(Grieve 1931; R. Fernandes & V.H. Heywood, in: Tutin et al. 1972; Sell & Murrell 2009).
Flowering reproduction
O. vulgare flowers from July to September. The inflorescence looks like crowded, terminal and/oraxillary heads, each consisting of whorls of 2-several flowers grouped into short 'spicules' or spikes.These are then tightly arranged into a branched, ± flat-topped corymb, or a slightly looser panicle(branched raceme) (Webb et al. 1988). The number of flowers in a terminal or axillaryinflorescence can approach several hundred (Grime et al. 1988, 2007).
Individual flowers are either large and bisexual (the majority) or smaller and female. The bracts exceedthe calyx and are purple, ovate and imbricate (ie they overlap one another like roof-tiles). Thereddish-purple, bell-shaped calyx, 3 mm, has five short equal lobes, spotted with yellow. The 4-8 mmcorolla is pinkish to rose-purple, 2-lipped, the tube being longer than the calyx and hairy or glabrousoutside (Melderis & Bangerter 1955; Sell & Murrell 2009). The flowers are highly aromatic andcan attract clouds of insects, some of which carry out pollination. The flowers are protandrous andself-sterile. Four nutlets are produced per fruit, each 1 mm long, ovoid and smooth (Sell & Murrell2009).
Germination takes place in spring in vegetation gaps, but some 'seed', ie nutlets or achenes(single-seeded dry fruits), can survive more than five years burial in soil (Thompson et al.1997). Plants from disturbed vegetation sites display a higher incidence of male-sterility and itappears that some populations have experienced high levels of selection pressure for out-breeding. Thishas led to the formation of the many genetic variants observed in the species. However, in B & I,O. vulgare has a modest number of subspecies (four) and occupies a rather narrow ecological rangeof habitats (Grime et al. 1988, 2007; Sell & Murrell 2009).
Fermanagh occurrence
As a glance at the Fermanagh tetrad distribution map makes clear, this familiar culinary perennial isless prevalent here now than it was previously in the 1940s and 1950s. In Fermanagh, it has beenrecorded rarely in a total of 13 widely scattered tetrads, only six of which have post-1975 records.
The range of local habitats of O. vulgare in the VC includes dry hedge banks, bridges, old wallsand rough grass on roadsides. In Fermanagh, O. vulgare has always been most frequently foundgrowing in the lime-rich mortar of old walls. While there certainly is less of it about than before, atthe same time it can occasionally become well established and long-persistent on these old walls,surviving in place for up to 50 years or more as is the case, for example, on Scarford Bridge over theColebrooke River.
The details of the six post-1975 records are: old garden wall, Inisherk Island, Crom Estate, Upper LoughErne, May 1989, RHN, still there May 2003; Scarford Bridge, Colebrooke River, 2 November 1989, RHN and12 April 1996, RHN & RSF; Aghalane Bridge, Woodford River, 2 October 1998, RHN & RSF;naturalised on N wall of garden, Belleisle ASSI, Upper Lough Erne, 26 December 1999, RHN; Long Island,Lower Lough Erne, August 2001, RHN.
Fossil occurrence and status
Nutlet macrofossils identified as O. vulgare have been rarely recorded in Britain from the Hoxnianand Ipswichian interglacial periods, as well as from the late Wechselian glacial. The species has alsobeen rarely recorded in the late stage of the current Flandrian interglacial, at Bowness Common and atEhenside Tarn (Godwin 1975). Whether this really is sufficient to recommend the species as a native isanother matter, although this is the status it is generally accorded in Britain. In Ireland, on theother hand, there does not appear to be any supporting fossil evidence and since there is a long historyof culinary and medicinal use and garden cultivation of the plant, the current author (RSF) can see nocredible scientific reason why O. vulgare should be regarded as native.
Irish occurrence
Regarded in NI at least as an escaped garden herb, O. vulgare has a quite widespread occurrencefurther south in the RoI, where it is still considered native by many botanists. While it is widelydistributed in the RoI, its distribution remains rather patchy and occasional and it is onlyconsistently found around Dublin and across parts of the Irish Midlands (Flora of Co Dublin;New Atlas).
British occurrence
O. vulgare is common in England and Wales, particularly in areas of limestone geology, but itbecomes very much more local further north and is apparently absent in NW Scotland except as a very raregarden escape (New Atlas).
European and world occurrence
O. vulgare is a polymorphic species that belongs to the Eurasian southern-temperatephytogeographical element. It is widespread across a large area of Eurasia including the Balearic Isles,other islands in the W Mediterranean, plus the Canary Isles. Subsp. viride occurs across a verywide area from Corsica to E China (Sell & Murrell 2009). The species has also been introduced intoboth N & S America, but only to a rather limited extent (Hultén & Fries 1986, Map 1610). It hasalso been introduced to New Zealand (Allen & Hatfield 2004).
Uses
As indicated above under the section on variation, several Origanum species have been in culinaryuse for seasoning for thousands of years, but only O. vulgare has a history in herbal medicine,which goes back to ancient Greece and Rome. The rather tender Sweet Marjoram (O. majorana) of thegarden was distinguished in the 16th century from the 'Common Organ', or 'Organy', also referred to as'Wild Marjoram' and 'Bastard Marjoram', which is O. vulgare (Grigson 1955, 1987). Gerard (1597)described O. vulgare as, "exceedingly well knowne to all".
Wild Marjoram was another general 'cure-all', being made into Marjoram tea and taken, for instance, totreat indigestion, earache, toothache, cough, dropsy and bladder troubles (Grieve 1931; Grigson 1955,1987). In ancient Greece, O. vulgare was regarded as a remedy for narcotic poisons and used totreat convulsions and dropsy. Marjoram contains about 2% volatile oil that can be extracted bydistillation. The properties of the oil are stimulant, carminative (relieving flatulence), diaphoretic(inducing perspiration) and mildly tonic. It was also used as an emmenagogue, ie for stimulating orincreasing menstrual flow (Grieve 1931). Externally, the dried leaves and tops could be applied as a hotpoultice for treating painful swellings and rheumatism, as well as for colic. An infusion made from thefresh plant was also recommended to relieve nervous headache by virtue of the camphoraceous principlecontained in the oil (Grieve 1931).
Names
The genus name 'Origanum' is the classical Greek name of an aromatic herb in Theophrastus and other earlyherbals (Gilbert-Carter 1964; Gledhill 1985), but it is not necessarily the same plant that we knowtoday. Another quite different theory of origin is that the name derives from the Green 'oreos' meaning'mountain' and 'ganos', 'joy', 'beauty' or 'brightness', a reference to the usual habitat of the plantand its attractive, perfumed nature (Johnson & Smith 1946; Hyam & Pankhurst 1995). The Latinspecific epithet 'vulgare' means 'common'.
Origanum vulgare was identified with the 'agrioriganos', the wild 'origanos' of Dioscorides. Thename 'Marjoram' properly belongs to the 'Sweet-' or 'Pot-Marjoram' (Origanum majorana =O. hortensis Moench.), which is taken to be the 'sampsuchon' or 'amaracon' of Dioscorides. From'amaracon' or its Latin equivalent, came the mediaeval Latin 'majorana' or 'maiorana', Old French'maioraine', or 'maiorane' and Middle English 'majoram' (Grigson 1955, 1974, 1987). Additional Englishcommon names include 'Joy of the mountain', 'Organ' and 'Organy' (Grigson 1955, 1987).
Threats
None.
Native, locally frequent. European boreo-temperate.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This prostrate, low-growing, evergreen, mat- and occasionally carpet-forming, perennial subshrub isdeep-rooting and produces creeping barren stems or runners and very short (up to 7 cm), erect,four-angled flowering ones, hairy on two opposite sides and almost hairless on the other two sides (seethe helpful illustration in Stace 1997, Fig. 570). The small, dark green, leathery, elliptic leaves arecovered in minute oil glands that give the plant its delightful, characteristic aromatic and culinaryproperties.
T. polytrichus is a plant of open, shallow, free-draining, calcareous or base-rich substrates inshort, heathy pastures, dry banks, steep rocky places on mountain sides, screes and ledges. At sealevel, it frequently occurs in calcareous maritime pastures, including on ant hills and on sea cliff topgrass (Grime et al. 1988, 2007).
In many situations, T. polytrichus is most characteristic of soils that are droughted in summer,since its deep root system can obtain water when most other plants cannot. While it is a strictcalcicole in S & E Britain, in the N & W of B & I, T. polytrichus can grow in a widerrange of habitats, including in some acidic grasslands. This may reflect the cooler, wetter climate ofthe more northern, acidic habitats, where root penetration is more shallow (Pigott 1955). Frequency andabundance of Wild Thyme are always higher on warm, south-facing grassland slopes. Under typical T.polytrichus growing conditions, the infertility and physical severity of the environmentstrongly limits competition from other species and thus the established strategy of the species iscategorised as being a Stress-tolerator. Despite its regeneration ability encompassing both seed andvegetative reproduction, T. polytrichus is a poor coloniser and is most characteristic ofsemi-natural, infertile, relatively undisturbed habitats (Grime et al. 1988, 2007).
Taxonomy and nomenclature
The genus Thymus has been in a wonderful state of taxonomic confusion and nomenclature flux formany years. As an instance of this, the Flora of Northern Ireland website lists ten previousnames of this particular taxon. Nowadays, however, T. polytrichus (= T. praecox subsp. arcticus (Durand) Jalas = T. praecox auct. non Opiz= T. drucei Ronniger = T. serpyllum auct. non L.) is regarded as the only nativeWild Thyme species in Ireland (Stace 1997; Parnell & Curtis 2012). Incidentally, the 4thedition of Stace's New Flora of the British Isles (2019) opts to take the name back to T.drucei, merely emphasising the fluidity of the nomenclature.
Thus the old Fermanagh record of T. serpyllum auct. non L., made by Stewart (1882), must inreality belong to this taxon, to which the current author (RSF) & RHN have now transferred it,making it the first county record.
Variation
T. polytrichus is so morphologically variable, in their Flora of B & I, Sell& Murrell (2009) recognise four varieties based on leaf size, inflorescence shape and size and thenumber of eglandular hairs on the leaves. They are named as follows: var. britannicus (Ronniger)P.D. Sell, var. neglectus (Ronniger) P.D. Sell, var. zetlandicus (Ronniger & Druce)P.D. Sell and var. drucei (Ronniger) P.D. Sell. In terms of distribution in B & I, var.britannicus is the most widespread form of the four. Var. neglectus seems to occur mainlyin coastal areas. Var. zetlandicus occurs only in Shetland and var. drucei seems to occurin mountainous, northern areas (Sell & Murrell 2009). In Flora Europaea 3, p. 181,this species is regarded as one of five subspecies of T. praecox Opiz (J. Jalas, in: Tutin etal. 1972), this again indicating the extent of both morphological variation and taxonomicdifficulty found within the genus.
Flowering reproduction
Plants flower from May or June to August. The species is gynodioecious (a term, incidentally, firstcoined by Charles Darwin), having separate hermaphrodite (ie perfect, bisexual) and smaller femaleflowers on different plants (Briggs & Walters 1997). This form of breeding system is an adaptationfavouring cross-pollination, which in T. polytrichus is mainly managed by bees. However, thebreeding system also allows the possibility of the hermaphrodite flowers selfing after insectpollination (Pigott 1955). It should be noted, however, that some gynodioecious species areself-incompatible, eg Plantago lanceolata (Ribwort Plantain) (Briggs & Walters 1997, p.150).
In T. polytrichus, the inflorescence is usually a tight head of few-flowered whorls; rarely it canbe somewhat elongated. The calyx is clearly 2-lipped, the upper lip with three short teeth, the lowerwith two longer teeth, while the rose-purple corolla is nearly regular, scarcely 2-lipped. Nutlets, asusual, are four in number and are ovoid and smooth (Sell & Murrell 2009; Parnell & Curtis 2012).Nutlets (achenes or seeds) are shed from July onwards, or may be retained in the calyx overwinter. Theycan also sometimes form a short-term persistent soil seed bank, surviving between one and five yearsburial, although numerous reports suggest they are more transient than this, lasting less than one year(Thompson et al. 1997).
Vegetative reproduction
T. polytrichus does also reproduce asexually by gradually forming extensive evergreen mats orcarpets of growth by long, horizontally spreading stems that root adventitiously at intervals. Theselong-lived, slow-growing clonal mats will eventually break up with age, forming individual,self-supporting plants. Plants growing in higher altitude, more exposed habitats may rely on vegetativereproduction more than those at lower levels, since seed production may be limited or restricted towarmer, sunnier years in mountain sites.
The ability to produce runners and grow long prostrate stems over shallow soils that provide littlerooting depth is important in the colonisation of ant hills which are a frequent habitat feature in SEngland (King 1977). The runners are also significant in allowing colonisation of sand dune habitats,where the stems can survive at least 50 mm of burial by shifting sand and assist in the stabilisation ofdune blow-outs (Pigott 1955; Grime et al. 1988, 2007).
Fermanagh occurrence
T. polytrichus is locally frequent and abundant on dry, shallow, infertile, lime-rich soils in thewestern half of Fermanagh, but nowadays it is completely absent elsewhere in the VC. Wild Thyme has beenrecorded in 47 Fermanagh tetrads, 8.9% of those in Fermanagh, but only 39 of them have post-1975records. The isolated older record on the tetrad map lying E of Lower Lough Erne was made by Meikle andhis co-workers at Crockanaver, S of Ederny in 1951, but the species has never again been seen in or nearthis vicinity. Elsewhere, the plant characteristically grows on dry banks, closely cropped fine-leavedpastures and in rocky places around the limestone areas of the county.
Irish occurrence
T. polytrichus has a very uneven distribution in Ireland, being very much more common at thecoast. At the same time, while it is locally abundant in some inland VCs (BSBI Atlas 2), it israre or extremely rare in other Irish counties. It has, for instance, only one or two stations in eachof the following VCs: Carlow (H13), Westmeath (H23), Longford (H24), Cavan (H30), Monaghan (H32) andArmagh (H37) (Irish Topographical Botany; Booth 1979; Cen Cat Fl Ir 2;Reilly 2001). In Co Tyrone (H36), T. polytrichus has only ever been recorded on fouroccasions, all of them in sites that suggest probable garden origin (McNeill 2010).
British occurrence
Wild Thyme remains widespread and locally abundant over most of Britain apart from the most denselypopulated and developed regions. There is little evidence in the New Atlas hectad map of anysignificant population decline since the 1962 BSBI Atlas, except in the southern part of itsrange where general habitat pressures are greatest (K. Walker, in: Preston et al. 2002).Counter-acting these losses, to a certain extent, T. polytrichus displays a limited capacity tocolonise quarries and other freshly available artificial habitats.
European and world occurrence
T. polytrichus is distributed across S, W & C Europe and is a member of the Europeanboreo-temperate phytogeographical element (Sell & Murrell 2009). It is mapped as T. praecoxOpiz subsp. arcticus (E. Dur.) Jalas (= T. drucei Ronniger) by Hultén & Fries (1986),as part of their Map 1612, where it is shown as occurring throughout B & I, Iceland and W & SWFrance to the Pyrenees.
Uses
Grieve (1931, p. 808-15) gives a lengthy but very interesting account of the medicinal uses and folkloreof both Wild Thyme (as T. serpyllum L.) and Garden Thyme (as T. vulgaris L.), the latterconsidered an 'improved' cultivated form of Wild Thyme. Thyme apparently was not in general use as aculinary herb in ancient Greece and Rome, although it was known that bees working it produced highquality fragrant honey. It was employed by the Romans to give an aromatic flavour to cheese (Grieve1931). Nowadays it is valued in the kitchen for flavouring stuffing, sauces, pickles, stews and soups,to name but a few uses.
The chief constituents of thyme are the phenols thymol and carvacrol. Cymene and pinene are also presentin Oil of Thyme distilled from the leaves, plus small quantities of menthone, borneol and linalol.
The medicinal use is as an antiseptic, antispasmodic, tonic and carminative (ie for relievingflatulence). The pounded herb, mixed with syrup and given fresh, has been used as a safe cure forwhooping cough. A similar mixture is given for catarrh and sore throat. Thyme tea was given for gastricdisorders and for colic. It also promoted perspiration at the start of a cold and it was used to treatfever and febrile conditions generally (Grieve 1931). It older herbal medicine, Thyme was used to treatshortness of breath. An ointment of it was used to reduce hot swellings and to treat warts, helpsciatica, gout and dullness of sight (Culpeper 1653). Gerard (1597) recommended it for sciatica, painsin the head, leprosy and the falling sickness. Thyme was also an ingredient of herb tobacco. Its essencewas used in cosmetics and for embalming corpses. The dried flowers were also used like Lavender topreserve linen from insects.
Vickery (1995) augments these aspects slightly, most interestingly with respect to a correspondent'srecommendation of it for retaining hair colour into old age – advice tragically obtained too late forthe current author!
Names
The genus name, 'Thymus' is a Latinised form of the Greek, 'Thumon', the name given by Theophrastus andother early herbalist writers to a Mediterranean species, most probably the dwarf woody bush, Thymuscapitatus (L.) Hoffmans. & Link (= Coridothymus capitatus (L.) Rchb. f.), which wasperhaps used in offerings, being derived either from the Greek verb, 'thuein' meaning 'that which isincluded in a sacrifice' (Gilbert-Carter 1964; Grigson 1974) or, alternatively, the name may be derivedfrom the Greek, 'thuo', meaning 'perfume' (Chicheley Plowden 1972).
The specific epithets mentioned are derived as follows; 'polytrichus' means 'many hairs'; 'praecox' isderived from the Latin 'praecoquo, meaning 'to ripen' and translates as 'early flowering'; and'serpyllum' is derived from the old Greek name 'kerpyllos', meaning 'to creep' and probably originallyapplied to T. sibthorpii Benth. or another creeping wild thyme of the Mediterranean(Gilbert-Carter 1964; Johnson & Smith 1946).
Common English names for this species apart from 'Wild Thyme', include 'Thyme (Bank, Creeping orRunning)', 'Hill Thyme', 'Pell-a-mountain', 'Puliall-mountain', or 'Penny Mountain' (corruptions ofSerpyllum montanum, a previous botanical name for the species), 'Mother of Thyme', 'Mother Thyme'(both referring to use as a uterine herbal treatment), 'Shepherd's Thyme', 'Horse Thyme' or'Brotherwort' (Prior 1879; Britten & Holland 1886; Grigson 1955, 1987).
Threats
None.
Native, frequent to common, but decidedly local. Eurosiberian temperate, naturalised in N America,Australia and New Zealand.
1881-2; Barrington, R.M.; Crom Castle Estate.
April to December.
Growth form and preferred habitats
This erect, odourless, 60-90 cm tall perennial has a slender, creeping rhizome that gives off runners.Stems are erect, unbranched or sparsely branched, acutely 4-angled, glabrous or very slightly hairy.Very occasionally, large, multi-branched plants can develop under optimal growing conditions. Theopposite leaves, 3-10 × 1-4 cm, are shortly stalked, narrowly lanceolate to ovate in outline, withmargins coarsely toothed to pinnately lobed near the base of the plant. The leaf shape and deeplytoothed margin give the plant a quite nettle-like appearance (Urtica dioica) at first glance.
L. europaeus is a lowland plant found in a wide range of wetland sites including the banks ofrivers, canals and ditches, pond and lake margins, tall-herb marshes, fens, swamps, wet fields and open,boggy places in fen-carr woods on moderately fertile, neutral to calcareous, muddy, mineral or peatyorganic soils. It does best in half-shade or sunny, sheltered conditions, but is sensitive to tramplingand grazing. Gypsywort tolerates temporary flooding and is an early colonist of nutrient-rich, exposedmud and shallow, standing or slow flowing water in newly created or recently disturbed wetlands (Sinkeret al. 1985).While it frequently fulfils this colonist role, Gypsywort is also found persistingin established vegetation and then competing with or tolerating pressure from sedges and other muchtaller and more vigorous wetland species, eg in reedswamp, fen or shaded fen carr.
L. europaeus is so frequent and characteristic a species of wet ground or shallow water habitatsthat one could argue it should be regarded as a member of our aquatic flora, rather than a purelyemergent terrestrial species (Cook 1998).
It also occurs in coastal situations at the top of beaches and in the wetter, slack areas of sand dunes(K. Walker, in: Preston et al. 2002). It is said to be quite salt-tolerant and can grow well intidal brackish marshes (Chater 2010).
The established strategy of L. europaeus is categorised as C/CR, ie intermediate between acompetitor and a competitive ruderal species (Grime et al. 1988, 2007).
Fermanagh occurrence
Very decidedly a species of nutrient-rich, lowland marshy wet ground and, as the tetrad map shows, L.europaeus is almost omnipresent around both Upper and Lower Lough Erne, but it is onlyoccasional in wet ground on other lakeshores and riverbanks in the VC. Gypsywort has been recorded in112 tetrads, 21.2% of those in the county, but it is very unevenly distributed.
Flowering reproduction
L. europaeus produces dense whorls of tiny, mint-like flowers in the axils of the upper leavesfrom June to September. Flowers are both larger hermaphrodite (bisexual) and smaller female on the sameplant (ie the plant is gynomonoecious). The number of flowers in a whorl can vary from 4-50 or more; theindividual flowers are small and sessile. The cup-shaped calyx has a series of spiny projections givingthe flower clusters a bristly appearance. The tubular corolla is white, with a faintly bluish tint andpurple spots, and is scarcely longer than the calyx lobes. The anterior petal is dotted with a rednectar guide and the corolla throat is densely hairy, protecting the nectar from rain and small crawlinginsects. There are just two functioning stamens, rather than four and they are exerted beyond the petals(Hutchinson 1972). Pollination is carried out by various small insects, including those with a shorttongue (proboscis), since the corolla tube is short and wide (Hutchinson 1972; Garrard & Streeter1983). The four fruit nutlets, 1.8-2.0 mm, are supported on a large nectar-secreting disk (Melderis& Bangerter 1955). They are 4-sided, truncate at their apex and have thickened corky margins thatgive them buoyancy in water, enabling water dispersal.
Seed dispersal
After release from the plant, the nutlets can stay afloat for 12-15 months and are readily dispersed,especially along rivers and streams, where the species is frequently found growing, even in crevices inwaterside brick walls. Birds, including pigeons also feed on the seed and L. europaeus seed hasbeen recovered from the crop of one that was shot. This does not prove that the nutlets can survivepassage through the bird and subsequently grow, but it holds out that possibility (Ridley 1930;Hutchinson 1972; Sell & Murrell 2009). L. europaeus growing around isolated ponds that werefenced off from stock animals does also suggest bird-transported seed as the most likely source (Ridley1930, p. 547).
The survey of soil seed banks in NW Europe contained three estimates of L. europaeus seedlongevity, two of which were transient (less than one year), while the third was 'present', butunassigned to any of the three types; transient, short-term persistent or long-term persistent (Thompsonet al. 1997).
Vegetative reproduction
The species has a slender creeping underground rhizome that gives off runners, so it must be capable ofsome degree of lateral vegetative spread and possibly clonal stand formation. However, the currentauthor (RSF) has not been able to detect any literature mentioning this possibility.
Irish occurrence
Apart from Lough Erne, the other major stronghold of this species in Northern Ireland is undoubtedlyLough Neagh, where Harron (Flora of Lough Neagh) described it as, "very frequent and locallyabundant". Elsewhere in Ireland, it is scarce, thinly and widely scattered, and decidedly local(An Irish Flora 1977; New Atlas).
British occurrence
A common wetland species, it has more of a southern and western distribution in Britain in both Englandand Wales, but becoming increasingly scarce, scattered and coastal northeast of a line between Lancasterand Hull and in the Channel Isles. Overall, it appears stable in both B & I (New Atlas).While the species may have lost populations in some sites due to drainage
and clearance of water courses, it has probably compensated for this by colonising ponds in quarry pitsand in other newly created artificial wetlands.
European and world occurrence
In Europe, L. europaeus is widespread and is found in Fennoscandia to 64°N. It also occurs in NWAfrica and N & C Asia. The species belongs to the Eurosiberian temperate phytogeographical elementand is also a naturalised introduction in eastern N America, together with Australia and New Zealand(Hultén & Fries 1986, Map 1613; Sell & Murrell 2009).
Uses
The plant produces a useful black dye which is fast and permanent for linen, wool and silk (Grieve 1931).Ever since the notion was first propounded by Lyte (1578) in his English translation of the Dutch herbalby Dodens (1554), it was commonly believed that gypsies used the dye produced by L. europaeus togive their faces a darker, more swarthy appearance. However this has been dismissed for many years asmere racial and class prejudice (Grieve 1931; Grigson 1955, 1987).
The fresh or dried flowering herb is described as astringent, hypoglycaemic, mildly narcotic and mildlysedative (Grieve 1931). Extracts inhibit iodine conversion in the thyroid gland and are used in thetreatment of hyperthyroidism and related disorders (Launert 1981). It is also reputed to slow andstrengthen heart contractions (Brown 1995). The plant has been used in the treatment of coughs, bleedingfrom the lungs, consumption and excessive menstruation. The leaves have been applied as a poultice tocleanse foul wounds (Chopra et al. 1986). The plant is harvested as flowering begins and can beused fresh or dried, either as an infusion or a tincture. Current uses are predominantly for treatingincreased activity of the thyroid gland and for premenstrual syndrome symptoms such as breast pain(Karalliedde & Gawarammana 2008) (Plants for a future, pfaf.org webpage for Lycopuseuropaeus, accessed 4 May 2020).
Names
The genus name 'Lycopus' is a combination of the Greek 'lycos', 'wolf' and 'pous', 'a foot', meaning'wolf's foot', a translation that suggests little or nothing in relation to the plant and its properties(Gilbert-Carter 1964). There is a fanciful suggestion, however, that it somehow refers to the shape ofthe rhizome (Hyam & Pankhurst 1995). The Latin specific epithet 'europaeus' simply translates as'European'.
There is a range of English common names including 'Gypsywort', 'Green Archangel', 'Marsh Hoarhound','Water Hoarhound', 'European Bugleweed', 'European Water Horehound' and 'Egyptian's Herb'. The'Egyptians' were another group like the gypsies who were imagined to use the plant to dye their skincolour (Grieve 1931). 'Hoarhound' is a bit of a mystery as there is nothing white or hoary about theplant, it not even being particularly hairy. The root 'har' means 'hoar', 'grey' or 'pubescent' (Grigson1974) and the current author (RSF) believes this simply does not fit the plant either.
Threats
None.
Native, locally frequent. Circumpolar boreo-temperate, widely naturalised in both hemispheres.
1892; Praeger, R.Ll.; banks of Ballycassidy River.
July to October.
Growth form and preferred habitats
The mints are a rather difficult plant group to identify, due to their great variability, the ease withwhich they hybridise and the fact that numerous forms are grown in gardens and may escape or bediscarded and then may cross with wild populations. The degree of hairiness, calyx shape, and stamen andpetiole length are all significant distinguishing factors (Garrard & Streeter 1983).
M. arvensis is a very variable species and is usually considered a perennial, although it canbehave as an annual in some situations. It is a rather slender, more or less hairy, sweetly or sicklyscented mint (Clapham et al. 1987), depending on preferences and individual sense of smell! Itproduces simple or branched stems up 10-60 cm tall, and spreading branches issuing from a suckering,stoloniferous rootstock (Clapham et al. 1987). The creeping stolons grow very freely and takefirm hold of ground it occupies, so the plant becomes very difficult to eradicate once it is wellestablished (Grieve 1931).
When in flower, it is readily and easily distinguished from M. aquatica (Water Mint), since itsflowers are all axillary, ie they are associated with leaves in the upper part of the plant. The erectstem is topped by a pair of opposite leaves, rather than by a terminal cluster of flowers as in WaterMint. Non-flowering plants are, however, a different matter, there being no reliable physical vegetativedistinguishing characters to separate these two mint species, although in comparison, M. aquaticadoes have the more pungent odour of the two. Unfortunately, a good sense of smell and the ability todistinguish species by odour alone is not within everyone's capability.
The established strategy of M. arvensis is categorised as CR, ie it is a competitive ruderal thatpreviously was widespread in B & I in arable fields of corn and other cultivated crops, especiallyon wetter ground (Grime et al. 1988, 2007). However, while it can be a vigorous coloniser, M.arvensis tends to be a rather poor competitor (Sinker et al. 1985).
In other parts of B & I, M. arvensis is a frequent plant of woodland rides, paths andclearings, marshy grassy waysides, disturbed waste places and margins of arable fields on damp to wet,intermittently flooded, acidic to neutral soils. It is essentially a lowland species, reaching analtitude of only around 370 m. M. arvensis frequently overlaps with M. aquatica and thetwo regularly hybridise. However, where water levels fluctuate markedly, or ground remains drier forrelatively long periods, Corn Mint typically replaces Water Mint (K. Walker, in: Preston et al.2002).
Fossil history
The hard nutlets of Mentha species make good macrofossils, being able to survive secondarydeposition in glacial and periglacial deposits. As a result, they are almost exclusively used to recordthe members of the genus (Godwin 1975). Fossil records for M. aquatic, or for M. aquaticaand M. arvensis together, are very similar and show these mints were present in everyinterglacial from the Pastonian onwards, proving the native status of both these species. However,fossils from the current Flandrian interglacial are sparse although they do increase later in time, andinclude finds from both Roman and Mediaeval sites (Godwin 1975).
Flowering reproduction
M. arvensis flowers from May to October, the small, 4 mm diameter flowers being borne in distant,compact, leafy whorls on the short, erect flowering branches. The bracts associated with theinflorescence are leaf-like, gradually decreasing in size upwards but always much longer than theflowers. Pedicels (flower stalks) are hairy, the calyx teeth, triangular, acute and much shorter thanthe bell-shaped, hairy calyx tube. The corolla is usually blue-lilac, but it can vary to pinkish orwhite (Clapham et al. 1987). The corolla is hairy inside and the four stamens are usually exertedfrom it. The flowers are pollinated by a variety of insect visitors (Garrard & Streeter 1983). Thefour fruit nutlets, 1.0 × 0.5 mm, are ellipsoid, slightly keeled, finely marked and pale yellow incolour (Butcher 1961).
Nutlets are shaken out of the calyx by wind or simply fall from the plant in autumn. If they drop intowater, or are washed away in rainwater, they may float or drift away from the parent plant. The outerwall of the fruit nutlet of Mentha species contains a dry fleshy substance composed of air-cellswith thin walls that allow the propagule to float. In the case of M. arvensis, floatationpersists for only 36 hours or so (Praeger 1913, quoted in Ridley 1930, p. 222). Otherwise, the seed hasno specialised dispersal mechanism, but they can survive burial and persist in the soil seed bank forfive years or longer (Thompson et al. 1997).
Vegetative reproduction
The fact that, like other members of the Lamiaceae family, M. arvensis possesses a rhizome orrootstock that develops horizontally spreading, low-growing quadrangular stolons that can root at nodesalong their length, confers additional reproductive potential to the species by means of this asexualgrowth process. Stoloniferous growth enables the plant to form clonal clumps or carpets, which enablethe individual to compete more strongly than otherwise would be the case, allowing a greater level ofpersistence in the particular site occupied. The relative importance of seed production versus stolongrowth to the colonising ability and stability of species populations remains unknown.
Fermanagh occurrence
M. arvensis is certainly the second most frequently recorded mint in the Fermanagh flora by a longmargin. Apparently, this is an extremely common species in damp fields, lakeshores and roadsides aroundUpper Lough Erne. However, the superabundance of sites in this area is a reflection of the vast amountof data from unnecessarily detailed recording carried out in this particular region by the EHS HabitatSurvey Team between 1986-7. Undoubtedly, this over sampling skews the distribution picture of manylakeshore species in Fermanagh and the current author (RSF) and RHN believe this is particularly so inthis case. It appears rather doubtful that M. arvensis really is quite so frequent around UpperLough Erne and so comparatively rare elsewhere. The tetrad map almost certainly reflects the extent ofthis unfortunate recorder bias, and as a result, is something of an artefact.
M. arvensis has been recorded in 73 Fermanagh tetrads, 13.8% of the total in the VC. There are nopost-1975 records from five of the earlier tetrads recorded, losses very probably associated with thealmost total move from arable to pasture agriculture in the survey area that has continued to the timeof writing (2022).
As the common names applied to the plant 'Corn Mint' and 'Field Mint' indicate, to a large extent this isa species of tilled fields and damp, disturbed sandy ground. The modern scarcity of such habitats inFermanagh suggests it would be more sparingly distributed than appears the case from the databaserecords. On the other hand, M. arvensis certainly does occur on intermittently floodedlakeshores, especially in limestone districts, plus in open areas within woodland and on marshy groundalong roadsides. Thus while some of its local habitats have been lost since the 1950s due to changes inagricultural practices, others suitable sites remain available.
British and Irish occurrence
Previously M. arvensis was very widespread throughout B & I as a cornfield weed, but hasbecome increasingly scarce in N Scotland and entirely absent from Orkney and Shetland. The species hassuffered a marked decline across B & I, especially from the English Midlands northwards since the1950s. This appears to most probably reflect changes in agricultural practice, including drainage,cleaner crop seed, herbicides and a hugely significant shift from arable tillage towards pasture farmingin the N & W (Braithwaite et al. 2006).
In Ireland, although it remains quite frequent, M. arvensis has become more scattered since the1950s and has declined most in the centre of the island (Parnell & Curtis 2012).
European and world occurrence
M. arvensis is a polymorphic species containing several subspecies. It is widespread in most ofEurope, N Asia and the Himalaya. A N Europe type is referred to as subsp. lapponica (Wahlenb.)Neum. The one area where the species is absent from is Iceland and Greenland (Hultén & Fries 1986Map 1614). The form in E Asia is referred to as subsp. piperascens (Malinv.) Hara. Corn Mint iswidely introduced as a weed elsewhere, including in N & C America, Java and New Zealand. M.arvensis s.l. belongs to the circumpolar boreo-temperate phytogeographic element (Hultén &Fries 1986, Map 1614).
Uses
Unlike Peppermint (M. × piperita) and Spear Mint (M.spicata), M. arvensis has not been used in herbal medicine, nor in the kitchen.Indeed, it can be a nuisance weed in Peppermint plantations where its strong odour would taintthe valuable peppermint oil, spoiling its quality and value (Grieve 1931). The minty fragrance ofPeppermint is replaced in M. arvensis by a smell described by Grigson (1955, 1987) as, "wet,mouldy gorgonzola". In 1798, William Sole published a survey of British mints in which he included25 kinds. He made vivid attempts to pin down and describe their characteristic mint-like scents and hereferred to Corn Mint having, "a strong fulsome mixed smell of mellow apples and gingerbread".
Names
The genus name 'Mentha' is the Latin name of a plant in Pliny after the Greek 'Minthe' andrefers to a myth by Ovid about a nymph of that name. It is one of the oldest plant names still in use,going back perhaps 4,000 years (Gilbert-Carter 1964; Stearn 1992). The Latin specific epithet 'arvensis'means 'growing or pertaining to cultivated fields' (Stearn 1992).
Despite its strong, unpleasant smell, apart from 'Corn Mint', M. arvensis has been given severaladditional English common names, including 'Apple Mint' and 'Lamb's Tongue' (Grigson 1955, 1987). TheEnglish name 'Mint' is derived or evolved from the Old English 'minte', in its various forms a namecommon to the west Germanic languages and ultimately derived from the Latin 'menta', from the Greek'minthe' (Grigson 1974).
The name 'Corn Mint' is widely known and used, the 'corn' element being a collective noun for all kinds,or fields, of cereal. The Old English 'corn' was already used to refer to the seed, not the standingcrop. With the related Germanic words, 'corn' goes back to an Indo-european base meaning 'to wear down'or 'to wear away'. This implies that 'corn' was used in the same meaning as we now use the cognate word'grain', eg 'a 'corn' of salt or of sand, equivalent to a 'grain' of salt or of sand. Cereals, peppersand other plants are said to produce corns or grains (Grigson 1974).
Threats
None.
Native, occasional.
1806; Scott, Prof. R.; Co Fermanagh.
July to November.
Apart from M. arvensis (Corn Mint), the less common of its parent species, this extremelyvariable, usually sterile hybrid is the most commonly grown garden mint in Co Fermanagh, rather thanM. spicata (Spear Mint). Whorled Mint grows to 30-90 cm tall and is very variable in its leafsize, shape, dentation and hairiness (Sell & Murrell 2009). When it flowers, from August to October,it can be distinguished from its parents by its tubular (not campanulate) calyx, 3.0-3.5 mm, the tube ofwhich is twice as long as wide, and by the form of the inflorescence. The inflorescence is of distantwhorls below or crowded ones in the upper part of the flowering stem. The whorls are often stalked. Thebracts on the inflorescence are like the leaves, often decreasing in size, the upper ones sometimesshorter than the flowers. The pedicels of the flowers are eglandular hairy (ie simple hairs withoutglands) (Sell & Murrell 2009).
In common with both its parents, it is a rhizomatous perennial, spontaneous hybrids arising where the twospecies overlap, there being no genetic barrier between them (Stace et al. 2015). Although therewill be a 'weeding out' at the seedling stage, hybrid plants often establish in the wild mint populationand, with their characteristic vigour and well-developed rhizomatous growth, they can persist and evenspread clonally to colonise new sites in the absence of their parents (Stace et al. 2015).
M. × verticillata is usually sterile, but occasionally highly fertileplants do occur, probably as a result of backcrossing. Interestingly, Whorled Mint, the most commonlyoccurring mint hybrid in B & I, is derived exclusively and spontaneously from native wildpopulations. Other mint hybrids also tend to originate from garden material, probably dumped in waysideor disturbed ground sites, but this does not occur in the case of Whorled Mint. However, since it iswidely cultivated for culinary use, M. × verticillata canoccasionally be found as a surviving relic of garden cultivation around old, abandoned homesteads (Staceet al. 2015).
M. × verticillata mainly grows in damp to wet ground in the presenceof its parents and only very occasionally without them. It is found in grasslands, alongside tracks andnear water margins in ditches, streams, slow rivers, lakes and ponds. Like both its parents, it istolerant of winter or wet-weather flooding.
Fermanagh occurrence
M. × verticillata has been recorded in 34 Fermanagh tetrads, 6.4% ofthe VC total. As the tetrad map of its distribution shows, like M. arvensis this hybrid is morefrequently found around the Lough Erne basin. Beyond the lough, it is only rather thinly scattered tothe NE of the VC and even more sparsely so to the SW.
British and Irish occurrence
This hybrid mint is by quite a long margin the most frequent and widespread of the total of nine hybridsof the genus known in Britain (two of them have no Irish records and one (M. spicata × M. longifolia (= M. ×villosonervata)) is represented by a solitary record near Belfast) (Stace et al. 2015).It is recorded throughout lowland Britain from S to N, becoming scarcer towards the N & W ofScotland. In Ireland, this hybrid is most frequently recorded in the N and the far S, but is muchscarcer and thinly scattered in the W and the centre of the island (Stace et al. 2015).
There is evidence of a decline in the populations of this hybrid across B & I from the hectad datadisplayed in the BSBI New Atlas (Preston et al. 2002), a high proportion of the recordsdating from the pre-1970 period.
European and world occurrence
M. × verticillata is described as, "very widespread in mainlandEurope" (R.M. Harley, in: Stace et al. 2015). While frequent in Europe, it is, however, rarein the Mediterranean area (Sell & Murrell 2009).
Introduction, very rare.
1990; NI Lakes Survey; unnamed Lough, Tullyvocady Td.
Fermanagh occurrence
The Fermanagh Flora Database contains a solitary record of this spontaneous hybrid between a native mint(M. arvensis (Corn Mint)) and an alien, introduced mint (M. spicata (Spear Mint)), whichwas discovered by the NI Lakes Survey in 1990 on the damp shore of a small, unnamed lake. A tallperennial, stems 30-90 cm, with a creeping rhizome, this is an extremely variable plant and therefore adifficult hybrid to identify with certainty. It is most like its M. arvensis parent, differing inits very short, narrow calyx teeth and its high degree of sterility. The herbarium label in BELindicates that it was identified by its pungent odour when fresh. Plants of this hybrid arisingspontaneously in the wild are hairy (pubescent), while those of garden origin are glabrous or almost soand arise in the wild as escapes or garden discards (K. Walker, in: Preston et al. 2002).
While it is sometimes grown in gardens, this hybrid is unfamiliar to many recorders in Ireland at leastand, therefore, could very easily be overlooked or mistaken for the more frequently occurring M.× verticillata (Whorled Mint), resulting in it probably beingunder-recorded.
Irish occurrence
The New Atlas hectad map and that in The Hybrid Flora of the British Isles (Stace etal. 2015) both show that M. × gracilis is only sparselyrecorded in Ireland, although O'Mahony (1975, 1995) reports it widespread in Co Cork (H3-H5),particularly along rivers. There have been rare recordings from a total of 18 Irish VCs over the years,many of which are regarded as garden escapes or discards, but apart from Co Cork, very few reports ofthis plant have appeared in recent years (Cat Alien Pl Ir).
The hectad map in Stace et al. (2015) displays data for Ireland from a total of 43 hectads, fivewhere both parents occur, 23 where the hybrid and M. arvensis overlap, none where the hybrid andM. spicata overlap and 15 squares where the hybrid is independent of both parents.
British occurrence
In Britain, in comparison with Ireland, M. × gracilis is very muchmore frequent and widespread in lowland areas, although it is rather scarce in the English EastMidlands. In Scotland, the hybrid becomes rare and ± coastal northwards (Stace et al. 2015). Forcomparison with Ireland, the equivalent hectad statistics from the distribution map in The HybridFlora of the British Isles are: a total of 645 hectads, 468 where both parents occur, 112 wherethe hybrid and M. arvensis overlap, 39 where the hybrid and M. spicata overlap and 26where the hybrid occurs independent of both parents (R.M. Harley, in: Stace et al. 2015).
European occurrence
M. × gracilis is widespread over most of Europe (R.M. Harley, in:Stace et al. 2015).
Native, common and locally abundant. European temperate, widely naturalised in both hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
February to December.
Growth form and preferred habitats
M. aquatica is a very variable, creeping, rhizomatous, aromatic perennial of wetland habitats.Stems grow 15-90 cm tall, 4-angled, ± erect, simple or branched, hairless to densely tomentose. The stemhairs can vary from rather soft, long and spreading, to short, downwardly pointing, white.
The leaves release a powerful minty smell when pressed, which is generally perceived by the majority ofpeople as pleasant and refreshing. It is sometimes alternatively described as 'orange-like' (Genders1971). Perception of odours by the human nose is very variable, however, and finding words to describesmells can be extremely difficult and subjective. Other descriptions of M. aquatica suggest thesmell of the plant varies from a fetid pungent odour to a pleasant camphor-like perfume (Melderis &Bangerter 1955).
Water Mint most typically occurs in permanently wet ground around the margins of open water in marshyground, or in streams and ditches where it is regularly submerged. It is also very common, however, ontemporarily flooded grasslands and, to a much lesser extent, in moderate broadleaf shade in wet, muddywoods and scrub.
M. aquatica forms extensive clonal patches by means of its creeping rhizome, but it is restrictedto situations where competition from taller wetland dominants is reduced. The outcome of thiscompetition may involve some degree of either soil infertility, or disturbance, limiting the growth ofthe dominant species, eg grazing, trampling or active management involving the cutting of the tallervegetation. Having said that, M. aquatica tends to be absent from unproductive soils of reallylow fertility (ie markedly acidic ones) and also from heavily disturbed sites (Grime et al. 1988,2007).
Differences between aerial and submerged stems
Emergent plants of M. aquatica are well furnished with white hairs, but when the stem and foliageare submerged they are quite hairless. Submerged leaves are generally thinner and limper than aerialones; the amount of chlorophyll is often decreased, but non-photosynthetic pigments, especiallywater-soluble anthocyanins and oil-based anthoxanthins, often appear in the vacuolar cell-sap of theouter tissues of the epidermis, turning the plant a dark crimson-purple colour (Sculthorpe 1967). Underthese circ*mstances, the production of the anthocyanin pigment is a reaction to nutrient stress, mostlikely the result of an imbalance in the C/N ratio in the plant's amino-acid (R.O. Mackender, pers.comm., June 2002). Phosphate deficiency might also sometimes be involved (Hopkins 1995).
Anthoxanthins are plastid-based pigments involved in the dissipation of excess light energy absorbedduring photosynthesis, diverting and breaking down excess excitation energy and converting it to heat.The anthoxanthins impart a yellowish-purple colour to tissues (Taiz & Zeiger 1998).
Anaerobic stress
As a semi-aquatic species, M. aquatica has only a limited ability to tolerate anaerobic stress,although in experimental measurements on detached rhizomes, Barclay & Crawford (1982) found itsurvived for four days immersion without any loss of regenerative power, a result similar to other marshemergents such as Carex rostrata (Bottle Sedge), Juncus conglomeratus (Compact Rush) andJ. effusus (Soft-rush). Anaerobic tolerance can vary with season and the state of theplant tissues. The two common Juncus species mentioned are both killed by as little as five daysanoxia, as they possess a relatively shallow root system. These two Rush species really are oxygenstress avoiders, rather than tolerators and the same may well apply to M. aquatica. The roots ofboth Mentha and Juncus species produce relatively open aerenchyma tissue in their roots inwet or flooded soil. This provides large, continuous air spaces within the tissue that allows oxygen todiffuse down to roots penetrating below the water-table. Normally anoxia tolerance is greatest in springand becomes lowered in summer when carbohydrate reserves are reduced (Crawford 1989).
Fermanagh occurrence
The most common and abundant mint in Fermanagh as elsewhere in B & I (present in every VC), M.aquatica frequents a wide range of wetland habitats in both sun and shade and occurs in 333tetrads, 63.1% of the VC total. Water Mint is the 16th most frequently recorded plant in the FermanaghFlora Database, a reflection of the extent of wetland in the survey area. As the tetrad map shows, M.aquatica is common on wet ground everywhere in Fermanagh and it is absent only from the mostacidic bogs and the better drained lowland farmland in the east of the county.
Flowering reproduction
Flowering takes place from July to September, the small lilac flowers being clustered in a dense terminalhead on the tip of the unbranched, erect stem, plus in axillary whorls around the next 1-3 lower nodeson the stem. The protandrous flowers have a short tube and protruding stamens and the stigma is of the'brush type' (Faegri & van der Pijl 1971, p.241). The flowers attract flies, including those of thecrane fly and mosquito families, as pollinators, plus occasional butterflies, such as the Small Skipper,Thymelicus sylvestris (Proctor et al. 1996). If insects fail to visit, the self-compatibleflowers set seed regardless (Harley & Brighton 1977). It would be entirely normal and expected forthe number of selfed seed set to be considerably fewer than those following cross-pollination. Seed isset from August to October, after which the aerial shoots die down. Seed dispersal is mainly byflotation, the nutlets having a thin corky outer coat with rather large air-filled intercellular spacesin the pericarp, providing prolonged buoyancy (Ridley 1930, p. 222; Sculthorpe 1967, p. 328).
Dormancy is broken after chilling, seeds either germinating in the spring in vegetation gaps, or becomingadded to the persistent soil seed bank.
Toxicity and herbivory
Mint species contain irritant oils that can cause inflammation of the skin, cramps and other unpleasantsymptoms if eaten in large quantity (Cooper & Johnson 1998, p. 126). However, M. aquatica issuppressed to some extent in grazed marshes, which suggests that it is browsed by stock animals.
Vegetative reproduction
Against the above mentioned suppression of the species populations by browsing stock, fragmentation ofthe rhizome due to trampling of wet, waterside ground by heavy animals enables dispersion of plant partsby flotation, increasing existing populations and extending distribution at least downstream.
In common with most non-weed plant species, no statistics are available on the frequency of M.aquatica seed regeneration. Seedlings are rarely observed, however, which suggests establishmentand spread probably quite frequently involves vegetative reproduction.
Fossil record
As with all Mentha species, the fossil record for M. aquatica is very much based on thehard fruit nutlets and they are regularly inseparable from M. arvensis. They occur in the CromerForest Bed series and in every interglacial since. Godwin (1975) reckons M. aquatica has beenpresent in B & I at least from the opening of the current Flandrian interglacial (in Ireland knownas the 'Littletonian'), if perhaps not during the previous glacial period.
British and Irish occurrence
M. aquatica is common, widespread and often locally abundant throughout lowland areas of Britainand Ireland and there has been no obvious change in its presence, measured at the hectad level ofmapping, in the two BSBI survey atlases (Perring & Walters 1962; Preston et al. 2002).
European and world occurrence
M. aquatica s.l. belongs to the European temperate element and occurs throughout Europeexcept the extreme north, although it is present in Iceland. It is also present and considerednative in SW Asia, N Africa, The Azores and Madeira. It is introduced (somehow) in N & S America, SAfrica and New Zealand (Hultén & Fries 1986, Map 1615).
Medicinal uses
The dried Water Mint herb yields about 4% essential oil that has an odour of Pennyroyal (M.pulegium). Russian Spearmint oil is also derived from a form of M. aquatica (Grieve1931). The medicinal action of the plant is said to be emetic, stimulant and astringent. In herbalmedicine, it is used to treat diarrhoea and as an emmenagogue (ie a substance that stimulates orincreases menstrual flow). A mint tea made with this plant is recommended for colds and influenza or inany complaint where setting up perspiration is necessary and in all inflammatory cases, internal orexternal, it is regarded as a helpful treatment. A strong infusion is inclined to be emetic. A decoctionof the plant prepared with vinegar is recommended to stop blood vomiting (Grieve 1931). A secondaryfocus of M. aquatica medicinal use is in treating indigestion, constipation, stomach ache andother digestive system ailments (Allen & Hatfield 2004).
Names
The genus name 'Mentha', in Latin was first used by Pliny, the myth associated with it appearing in Ovid,but it goes back further than this suggests. In Greek, the name is 'Minthe', the name of a beautifulnymph, and it is said to have been associated with the mint plant for up to 4,000 years, making it oneof the oldest plant names still in use (Gilbert-Carter 1964; Stearn 1992). The Latin specific epithet'aquatica' means 'growing in water'.
Threats
Probably under pressure from increasing eutrophication of wetland sites.
Introduction, naturalised garden escape, very rare, but possibly or probably overlooked to some extent.
1934; Praeger, R.Ll.; shore of Lower Lough Erne near Enniskillen.
Growth form and preferred habitats
M. × piperita is a very variable hybrid ± intermediate betweenits parents. Like Mentha arvensis × M. spicata (M. × gracilis) (Bushy Mint), it is the sterile product of interbreedingbetween a native and an introduced mint species and, therefore, it might occur both spontaneously (veryrarely) and more often as a garden escape or discard (R.M. Harley, in: Stace et al. 2015). Leavesgive off a pungent smell and taste of peppermint or lemon when crushed. Glabrous plants are generallybelieved to be of garden origin, while pubescent ones are considered more likely to be spontaneoushybrids arising in the wild. These two forms are often not distinguished by field recorders and the truepicture is, therefore, unknown (K. Walker, in: Preston et al. 2002).
Peppermint usually grows in marshes and wet grassland on the margins of lakes, ponds, rivers and streams,as well as by artificial habitats such as ditches, reservoirs and canals. It is also found by roadsides,near habitation and by abandoned buildings on waste ground and in places where garden waste material isdumped. Well established populations may be large and long persistent, since although the plants aresterile, they have vigorous hybrid growth (heterosis) and strong colonising ability due to theircreeping rhizomes (R.M. Harley, in: Stace et al. 2015).
Variation
Sell & Murrell (2009) distinguish four varieties on the basis of leaves with or without crispedmargins, inflorescence shape (oblong spike versus globose head) and hairiness of pedicels and calyx.
Fermanagh occurrence
There are only four records for this familiar, but very variable garden hybrid mint in the FermanaghFlora Database.
With so very few records in the current Fermanagh survey, it is very possible that M. × piperita is being overlooked and under-recorded to some extent.Peppermint has not been recorded in Fermanagh by anyone since 1975.
The local habitats for this vigorous, rhizomatous perennial are perfectly typical of elsewhere in theseislands – in damp to wet waste ground near habitation, in ditches and on lakeshores. The details of theother three Fermanagh records are: S of Lisnaskea, Upper Lough Erne, 1950, MCM & D; near CorrasloughPoint, by Derryvrane L.O.L. (Loyal Orange Lodge) Hall, Upper Lough Erne, 1950, MCM & D; River Erneat Enniskillen, 18 August 1975, Mrs C.R. Hackney & P. Hackney, voucher in BEL.
Irish occurrence
Elsewhere in Ireland, M. × piperita is an occasional garden escape,growing and sometimes persisting for long periods in damp places on river and canal banks, roadsides andnear houses. The Cen Cat Fl Ir 2 and Cat Alien Pl Ir together indicate that thishybrid has been found at least once in 39 of the 40 Irish VCs, the exception being Co Meath (H22). TheNew Atlas hectad map coverage, on the other hand, shows that Peppermint is scarce across most ofS Ireland, with widely scattered recent records only, but it is somewhat more frequently andconsistently recorded in NI.
British occurrence
As the New Atlas hectad map shows, M. × piperita iswidely scattered throughout lowland Britain from S to N, including a presence in both Orkney (VC 111)and Shetland (VC 112) (Preston et al. 2002). The high proportion of pre-1970 records in theNew Atlas from E England suggests the hybrid has declined in that area, probably reflecting theloss of casual or naturalised populations that arose from the previous (now gone) commercial scalecultivation of Peppermint that took place in that part of the country (K. Walker, in: Preston etal. 2002).
European and world occurrence
Peppermint is widespread in temperate Europe and the Middle East, both as a spontaneous hybrid and as agarden escape or discard. It has been very widely introduced and cultivated worldwide for its culinaryand many medicinal uses (Sell & Murrell 2009). Most commercial Peppermint oil production takes placenow in The United States of America and in Morocco (R.M. Harley, in: Stace et al. 2015).
Uses
M. × piperata is one of the most important sources of the volatile oilof Peppermint distilled from the plant and it has been cultivated as such for many centuries for use asa flavouring and therapeutic agent. According to Pliny, Peppermint was used by the ancient Greeks andRomans to flavour both their food and wine. While there is evidence of ancient medicinal use in Egyptand in Iceland in the 13th century (!), Peppermint only came into general use in western medicine aroundthe mid-18th century and then was first used, and commercially cultivated, in England (Grieve 1931).
Peppermint is regarded as the most important essential oil available to herbal medicine, the chiefconstituent of which is menthol, although it also contains menthyl acetate (an ester that gives the oilmost of its characteristic minty odour), isovalerate, menthone, cineol and limonene. The oil iscontained in numerous brands of proprietary medicines such as tinctures, salves, essences, inhalants,herbal pills, toothpaste and tea-mixtures (Launert 1981).
Peppermint oil is more anti-spasmodic than any other volatile oil and adds greatly to its power torelieve pains arising in the alimentary canal. Its stimulating, stomachic and carminative properties arevaluable in treating dyspepsia, being mostly used for flatulence and colic. Oil of Peppermint helpstreat sickness and nausea and it is also used to disguise unpalatable drugs. Peppermint tea is useful incombating the early stage of colds, sometimes effecting a cure. Menthol is used to relieve the pain ofrheumatism, neuralgia, throat affections and toothache. It acts as a powerful local anaesthetic,vascular stimulant and disinfectant. It is inhaled for chest complaints and nasal catarrh, laryngitisand bronchitis. It is even used to ease or prevent sea-sickness (Grieve 1931).
Threats
None.
Introduction, archaeophyte, garden escape, very rare. European temperate.
1953; MCM & D; waste ground in Enniskillen Town.
Growth form and preferred habitats
Mints are undoubtedly the best known culinary crop plants to the public of Britain and Ireland. Theseculinary favourites are a mixture of archaeophytes, such as this species, and neophytes, such as AppleMint (M. × villosa), Peppermint (M. ×piperita) and Eau de Cologne Mint (M. × piperata var.citrata) (Stace & Crawley 2015). Spear Mint, sometimes referred to as 'Common Spear Mint', isa rather stout, scented, almost glabrous perennial with branched flowering stems, 20-40 cm tall,produced during August and September, growing from an extensively creeping and branching rhizome. Itslanceolate, acute, glabrous leaves have sharply toothed margins, unwrinkled surfaces and are sessile orsub-sessile. The leaves are also opposite, oblong or oblong-lanceolate, rounded (almost cordate) at thebase, triangular pointed and are at first spreading, soon becoming decurved. The inflorescence is along, slender, somewhat interrupted, catkin-like spike, of closely arranged whorls of 3 mm diameter,blue-lilac flowers. These spikes are often arranged into a loose panicle, either continuous or slightlyinterrupted with occasionally a small lateral spike at the base (Butcher 1961; Hutchinson 1972).
M. spicata grows in lowland areas in a variety of damp to wet places, often in ditches, riverbanks, on rough or waste ground near habitation. Unfortunately, it also occurs in places where gardenrubbish is fly-tipped (K. Walker, in: Preston et al. 2002).
Garden mints of all types, possess vigorous, or even rampant, rhizomatous rootstocks and, in the case ofM. spicata, the plant is fertile and sets seed. Thus, with both seed and vigorous vegetativereproduction, they tend to quite rapidly outgrow their allocated garden plot. Reflecting thisreproductive ability and vigorous growth, the established strategy of M. spicata is categorisedas C/CR, meaning it is intermediate between a Competitor and a Competitive Ruderal (Grime et al.1988, 2007). Gardeners are often recommended to grow mint species or hybrids in buckets or in sunkencontainers, rather than in beds in open ground to prevent or restrict the invasive spread of the plants.This spreading property leads to garden mint plants frequently being divided and portions donated tofriends or dumped, and hence the propensity of various mints to appear on waste ground or tips nearhabitation in towns and villages, where, as in this instance, they can easily become naturalised and mayprove long persistent (Crawley 2005).
Hybrid origin
The exact origin of M. spicata is unknown, but it is an archaeophyte and is believed to havearisen in cultivation, possibly in C or S Europe (Grieve 1931; Garrard & Streeter 1983). Grieve(1931) suggests it came from the Mediterranean basin and that it was introduced to Britain by the Romanswho were great promoters of the species. It also appears to be of hybrid origin, probably derived from across between the diploids M. suaveolens (Round-leaved Mint) 2n=24 and M. longifolia(Long-leaved Mint) 2n=24, followed by chromosome doubling. Evidence comes from morphological,cytological and chemical data (Harley 1963; Hendriks 1974). The tetraploid result of this speciationprocess (2n=48) is a segmental allopolyploid that occasionally segregates parental characters in itprogeny, giving rise to the morphological identification difficulties in separating M. spicatafrom its hybrids (R.M. Harley, in: Stace et al. 2015).
Fermanagh occurrence
Spear Mint is a commonly grown garden plant and it is therefore a little surprising that it was firstfound outside gardens in Fermanagh as late as 1953, naturalised on waste ground below the old workhousein Enniskillen. A second find was made by RHN in July 2001 beside a farm track in a remote part of theWestern Plateau and the third, and only other station, was discovered by I. McNeill near the Lough ErneYacht Club, Goblusk Bay, Lower Lough Erne, in October 2002. At the latter site, RHN observed a largepatch of this mint covering an area 10 m × 10 m near the bridge on the lane to the Club on 10 September2010.
It is peculiar that there are no other Fermanagh records for Spear Mint, which is such a widely grownrhizomatous perennial, easily identified by its familiar and distinctive aroma when bruised.
Irish occurrence
In comparison with the Fermanagh situation, the editors of the FNEI 3 account regarded this mintas a casual in the three most north-eastern Irish VCs. They catalogued very few records for Co Down(H38) and Co Antrim (H39) and none at all for Co Londonderry (H40). The Cen Cat Fl Ir 2lists 16 Irish VCs in which Spear Mint has been previously recorded and the Cat Alien Pl Ir addsa further seven VCs to this listing (not including Fermanagh, H33). The current author (RSF) reckonsthat this leaves 16 Irish VCs out of the 40 total where M. spicata has never been recorded atall.
The New Atlas hectad map shows M. spicata is extremely thinly scattered across Ireland andnot at all concentrated on the larger conurbations as one might expect (New Atlas). With regardto areas of human population density, the Urban Flora of Belfast records Spear Mint in seven 1-kmsquares in the W of the city, while in Co Dublin (H21), the Flora of Co Dublin listed sixpost-1984 records for the species in five of their eight flora districts, four of which are coastal.
British occurrence
M. spicata is very widely cultivated as a pot or plot herb for the kitchen and often outgrows itswelcome in a garden setting and is sufficiently vigorous to be able to thrive and become naturalised inthe wild when it escapes or is discarded; it is not surprising that it is very commonly recorded insuitable lowland situations throughout Britain, from the Channel Isles to Orkney and Shetland (NewAtlas). It is, however, very variable (especially with regard to degree of hairiness) and isvery difficult to distinguished from its hybrids with other mints (Crawley 2005). Another problem isthat hybrids involving M. spicata as one of the parent species, nearly all possess thecharacteristic spearmint scent (R.M. Harley, in: Stace et al. 2015). Thus it is very possible foridentification errors to occur and M. spicata is probably over-recorded to some unknown extent.
European and world occurrence
M. spicata is also widely naturalised in W, E & S Europe and is introduced in S Africa, NAmerica, S Australia and New Zealand (Hultén & Fries 1986, Map 1618). Being of garden cultivatedorigin, it is probably true to say that it is not indigenous anywhere, although some authors suggest itcomes from the Mediterranean or the mountains of C Europe (eg Grigson 1974).
Uses
M. spicata is mostly used for culinary purposes and was greatly appreciated by the Romans. Pliny(according to Gerard (1597)) said of it, "The smell of Mint does stir up the minde and the taste toa greedy desire of meate." Ancient Greeks and Romans believed mint (Spear Mint) would prevent milkcoagulating and its acid fermentation. They also used it to perfume rooms, baths and as a restorativelike smelling salts are used today. Gerard (1597) considered it, "good against watering eies andall manner of breaking out on the head and sores. It is applied with salt to the bitings of maddogs." It was also believed good for treating wasp and bee stings. Culpeper (1653) lists 40maladies for which he thinks mint is, "particularly good".
Spear Mint has been used for whitening teeth and its distilled oil is still used to flavour toothpasteand chewing gum and to perfume soap. A strong decoction is said to cure chapped hands (Grieve 1931).
The properties of Spear Mint oil are similar to those of Peppermint, being stimulant, carminative andantispasmodic. Its effects are, however, less powerful and it is less used than Peppermint although itis better adapted for use on children's maladies. Its taste is more pleasant and less strong thanPeppermint (Grieve 1931).
Names
Turner (1566) referred to garden mint of his time as 'Spere Mynte'. He ascribes the name on account ofthe leaf shape, not the spiked inflorescence. This accords with the 16th century botanists' Latin nameof the plant 'mentha angustifolia spicata', 'narrow-leaved spiked mint' and 'mentha acuta','sharp-pointed mint' (Grigson 1974). However, the Latin specific epithet 'spicata' does refer to anelongated inflorescence of sessile flowers, ie 'a spike' (Gledhill 1985).
Under the synonym M. viridis (L.) L. (R.M. Harley, in Tutin et al. 1972), there are fiveEnglish common names listed by Britten & Holland (1886), given as
'Brown Mint' (Gerard 1597); 'Garden Mint' (Lyte 1578); 'Mackarel Mint' (Gerard 1597); 'Our Lady's Mint'(Gerard 1597) and 'Spearmint'.
Threats
None.
Introduction, neophyte, very rare.
1997; Wolfe-Murphy, S.A.; Colebrooke River, unspecified site.
Growth form and preferred habitats
M. × villosa is an extremely variable, rhizomatous, perennial, completelysterile hybrid between two widely cultivated mints, the introduced Spear Mint (M. spicata) andthe perhaps partially native Round-leaved Mint (M. suaveolens). [See the species account on thiswebpage.] There is a history of taxonomic confusion associated with this hybrid thanks to its extremevariability, the extent of which is demonstrated by the fact that Crawley (2005) provides a synonymywith no less than seven alternative names. M. × villosa is so veryvariable, four varieties of it are recognised and named in the flora of B & I. Of these four, M.× villosa var. alopecuroides, which is definitely of gardenorigin, appears to be, and probably is, the most widespread form (Sell & Murrell 2009; R.M. Harley,in: Stace et al. 2015).
Unlike the other three taxonomic varieties, var. alopecuroides is a uniform hybrid, probablyrepresented in B & I by a single clone. It has been known in Britain since at least the 18thcentury, but is frequently mistaken for M. suaveolens (R.M. Harley, in: Stace et al.2015). In this variety, the stem has middle stem leaves 4-8 cm long, broadly ovate, sub-rotund ororbicular, softly hairy, margins serrate and with teeth large, patent (ie directed outwards, ± at 90°).There are punctate glands on both leaf surfaces that give the variety its sweet fragrance similar toSpear Mint. The inflorescence is robust, calyx 2 mm and the corolla is pink.
M. × villosa var. alopecuroides is, however, frequentlyconfused with other forms of round-leaved mint, including M. spicata (Spear Mint) itself and,unsurprisingly, also with M. × rotundifolia (False Apple-mint), whichis the hybrid between M. longifolia (Horse Mint), a European species not recorded in Britain(Stace 2019) and M. suaveolens.
Other forms of M. × villosa are usually more narrow-leaved than var.alopecuroides, with a generally oblong or oblong-ovate to lanceolate outline.
For this reason, and because the whole Mentha genus is difficult to unravel, largely due to anabsence of genetic barriers to hybridisation, M. × villosa var.alopecuroides is certainly unevenly recorded throughout B & I and is veryprobably under-recorded in the wild.
Fermanagh occurrence
The solitary record of this mint hybrid listed above is from the typical 'rough grassland near water'habitat that so regularly provides the necessary conditions for sustained survival of garden escapeswithin this genus. This hybrid often forms small patches near water, but can also occur in drier sites,where it is generally associated with fly-tipping of garden waste or excess.
Irish occurrence
The Cat Alien Pl Ir indicates that this hybrid has been recorded in ten other Irish VCs (notincluding Fermanagh), seven of them with post-1980 records. It may be that Garden Apple-mint is becomingmore popular with gardeners and cooks and could therefore perhaps be making more appearances in the wildthan before, but the present state of knowledge does not support this contention. It merely indicates orreflects the need for more active recording of this rather difficult group of plants.
British occurrence
The hectad map in Stace et al. (2015) shows that M. × villosais a very widespread hybrid throughout lowland Britain, and since var. alopecuroides is verycommonly grown in gardens, it is by far the most frequently recorded variety of the four in the wild.The map shows M. × villosa throughout Britain from S to N, from theChannel Isles to Shetland. In Scotland, northwards of the Glasgow-Edinburgh conurbations, it quitequickly becomes somewhat confined to coastal areas. The map in Stace et al. (2015) containsrecords of M. × villosa from a total of 1,222 hectads in Britain. In Ireland, bycomparison, the same map displays symbols from just 24 hectads.
As with other garden mints, this sterile hybrid is very vigorous, spreading rapidly in fertile cultivatedground by underground rhizomes and above ground stolons to form large colonies. It can thus quitequickly become invasive and outgrow the space allocated to it, resulting in excess material being dugout and dumped, sometimes illegally in waste ground or along roadsides or on riverbanks (K. Walker, in:Preston et al. 2002; R.M. Harley, in: Stace et al. 2015).
European and world occurrence
This hybrid mint is widely naturalised in mainland Europe.
Introduction, garden escape, very rare. Submediterranean-subatlantic.
1939; Praeger, R.Ll.; Belcoo, Lough Macnean.
Growth form and preferred habitats
Another rhizomatous, aromatic, vaguely apple-scented mint of damp places, M. suaveolens is arather stout or sturdy perennial with erect stems, 40-100 cm tall, pale green in colour, usually quiteheavily branched above the middle. The stems are very variably hairy, ranging from sparsely to denselyclothed with white, simple eglandular hairs. The leaves, 3.0-4.5 cm, ovate-oblong to sub-rotund, arestrongly rugose (sage-like), pubescent, margins serrate, grey-or white-tomentose beneath. Floweringoccurs in August and September, the inflorescence of numerous congested whorls, forming a terminalspike, interrupted below and usually branched. The calyx is campanulate, hairy and the corolla whitishor pink and hairy outside (Sell & Murrell 2009). Although it is sometimes referred to as'Apple-scented Mint', the aroma or flavour is described by Grieve (1931) as rather like Spear Mint(M. spicata), but stronger. Other descriptions consider it has a 'fruity' mint flavour.
Growth of the plant is rapid and its rhizomatous spread enables it to quickly form extensive clonalpatches that may prove excessively invasive in a garden setting.
Typical habitats are either coastal ditches, or naturalised as garden escapes or discards on damp,lowland roadsides and waste places inland (K. Walker, in: Preston et al. 2002). It is alsoassociated with churchyards and the sites of ruined monasteries, monks having cultivated it formedicinal as well as for culinary use (Grieve 1931; Chater 2010).
British and Irish status
M. suaveolens is regarded by some as a native species in coastal parts of Britain, but probably itis only so in parts of SW England and Wales (K. Walker, in: Preston et al. 2002). The currentauthor (RSF) cannot locate any convincing proof of native status for this species: there does not appearto be any fossil evidence (Godwin 1975) and since the herb has a very long history of garden cultivationand culinary use, reliance on findings dating from John Ray's excursions in the 1660s really do notprove indigenous origin (Pearman 2017).
In his Flora of Cardiganshire, Chater (2010) quotes Salter (1935) on this mint, saying,"Occurs usually, but not always, in the neighbourhood of cottages, having been grown, along withother species of Mint, by the old herbalists." Salter gave a dozen localities. Chater then refersto more recent records, "six other records since 1970 from waste ground, road verges andchurchyards" and, later, "Whether it is native in any of these sites is uncertain…". Hethen maps them all as aliens. The Flora of Cornwall, on the other hand, treats M.suaveolens as native without any reservations, describing it as, "scattered across Cornwallalongside roads and streams and on waste ground" (French 2020).
Elsewhere in Britain, and everywhere in Ireland, M. suaveolens is very clearly accepted as beingan introduced alien and a garden escape or discard (K. Walker, in: Preston et al. 2002).
Fermanagh occurrence
There are only three very old records for this garden mint in the Fermanagh Flora Database, and none atall from the post-1975 period when so much more recording effort has taken place in the VC. Thissuggests to the current author (RSF) and RHN, that this mint really is rare, but it could also reflecthesitancy on the part of field recorders to distinguish this species. It is not easy to separate thisplant from its hybrid with M. spicata (M. × villosa(Apple-mint)), although there is in fact only one 1997 record of this hybrid in Fermanagh. M.suaveolens is a smaller plant than M. × villosa, its leavesmore finely toothed and the flowering spikes are narrower, less than 10 mm wide (Halliday 1997).
The Fermanagh habitats of Round-leaved Mint are listed in the Revised Typescript Flora as,"waste ground, roadsides and a canal bank". The details of the other two records for thespecies are: near Lisnarrick, 1949, R. Mackecknie; and by the Old Ulster Canal near Gortnacarrow Bridge,1950, MCM & D.
Irish occurrence
For a local comparison, the FNEI 3 also lists relatively few records of M. suaveolens,ie around 16 in the post-1938 period in total for the three NE Irish VCs (H38-H40).
The Cat Alien Pl Ir describes M. suaveolens as an occasional garden escape, persistent inabandoned gardens and in a few situations along river banks. It lists occurrences in 34 of the 40 IrishVCs, the exceptions being Cos Kildare (H19), Westmeath (H23), Leitrim (H29), Monaghan (H32) and Donegal(H34, H35). The New Atlas hectad map displays the fact that recent Irish records are veryscattered and sparse, the frequency and pattern being similar to that in C & N England.
British occurrence
In Britain, M. suaveolens has been recorded from the Channel Isles and the Scilly Isles (VC 1a) toOrkney (VC 111) in the far north. The New Atlas hectad map shows it is most concentrated in theSW of England and Wales, but quickly becomes scattered and scarce northwards and eastwards until inScotland it is very rare and almost entirely coastal.
There is also evidence from hectad data in the two BSBI Atlas surveys (Perring & Walters 1962;Preston et al. 2002), that occurrences of M. suaveolens have been in decline for at least50 years. However, some of these apparent losses may have arisen through early records of M.suaveolens representing over-recording, due to errors in the identification of M. × villosa and, in particular, with a variety of the latter referred toas nv. alopecuroides (Hull) Briq. (Brewis et al. 1996; K. Walker, in: Preston etal. 2002).
European and world occurrence
This variable species was originally restricted to Europe and the adjacent part of N Africa but, being aculinary herb, its distribution is insufficiently known since it has been spread by garden cultivationfrom its unknown indigenous origin. It has been introduced to N & S America and to New Zealand(Hultén & Fries 1986, Map 1616).
Uses
Apple-scented Mint (sometimes mistakenly referred to as 'Apple Mint' which is more accurately M. × villosa nv. alopecuroides) or 'Round-leaved Mint' is oftenregarded as a substitute for, or is even used in preference to, M. spicata, Spear Mint, primarilybecause it has a sweeter flavour than the latter, but secondly on account of its greater ability toresist the pathogen, Mint Rust (Webb et al. 1988).
Round-leafed Mint has been frequently used as a domestic herbal remedy, like other mint species beingvalued mainly for its antiseptic properties and its beneficial effect on digestion. Again, as with othercultivated mints, a tea made from the leaves has traditionally been used in the treatment of fevers,headaches, digestive disorders and various minor ailments (Grieve 1931; Foster & Duke 1990).
Names
The Latin specific epithet 'suaveolens' is a combination of 'suāvis' meaning 'sweet' or 'agreeable' and'oleo', 'to emit a smell', and thus 'sweet-scented' (Gilbert-Carter 1964; Gledhill 1985).
English common names include 'Apple-scented Mint', 'Apple Mint', 'Pineapple Mint', 'Round-leaved Mint','Woolly Mint' and 'Horse Mint' (Britten & Holland 1886).
Threats
None.
Introduction, neophyte, garden escape, very rare.
1952; MCM & D; Castle Coole Estate.
Growth form and preferred habitats
This absolutely prostrate, Mediterranean mint with tiny, thyme-like, oval leaves not more than 6 mm long,forms low-growing, dense perennial mats by means of its distinctive, slender, thread-like, creepingstems that root at intervals. It is endemic to Corsica, Sardinia and the Italian island of Montecristoand was first introduced to decorative alpine and rock gardens in B & I in 1829 (K. Walker, in:Preston et al. 2002). It is now frequently cultivated for its pleasant, pungent peppermint-likescent. Gardeners nowadays tend to grow it in cracks in paving, where trampling releases its intensearoma. The plant produces small, 1.6 mm diameter, lavender or pink flowers in 2- to 6-flowered whorls orclusters in June and July. It spreads quickly but avoids dry soils. M. requienii seeds itself andoccasionally escapes onto damp paths, relatively bare grassy ground and rocky places (Farrer 1930;Grey-Wilson 1989; Clement & Foster 1994).
First British and Irish records in the wild
Corsican Mint was first recorded in the wild in Castletownsend, Co Cork (H3), outside the wall of LordBantry's demesne (More 1872), where it still persists (Cat Alien Pl Ir). In England, it firstappeared ex-garden in Cumbria in 1890 (VC70)(Halliday 1997).
Fermanagh occurrence
M. requienii has only been recorded once in a non-garden setting in Fermanagh, when it wasdescribed as being, "abundant on shaded paths", on the Castle Coole Estate in 1952 (RevisedTypescript Flora).
Irish occurrence
M. requienii is very rare in Ireland, but it has been recorded at some time at least once in fourother Irish VCs listed in Cat Alien Pl Ir, namely W Cork (H3), W Galway (H16), Tyrone (H36) andCo Armagh (H37). The New Atlas map for Ireland shows a total of just eight hectads scatteredacross the whole island. Habitats include on a peaty bank at the head of Killary Harbour in W Galway(H16) and on the north slope of Slieve Gullion, Co Armagh (H37) (Clement & Foster 1994).
British occurrence
M. requienii is also very rare in ex-garden settings in Britain, but the New Atlashectad map shows it extends from Jersey and the Scilly Isles to as far north as Angus in NE Scotland (VC90). There are a total of 43 hectads with records in Britain, including three in the Isle of Man (VC 71)(K. Walker, in: Preston et al. 2002). It is established in scattered localities across Britain,including in a few woodlands in the south.
Native, occasional, locally abundant. Circumpolar boreo-temperate, considered native also in Chile.
1900; Praeger, R.Ll.; Co Fermanagh.
April to October.
Growth form and preferred habitats
The genus Hippuris is often (but not always) considered to contain just a single species that iswidespread over the whole world. Other botanists recognise two or more closely related species andseveral varieties have also been described (Preston & Croft 1997). At the same time, H.vulgaris is regarded as one of the most highly evolved species in the flora of B & I, itsextremely minute flowers displaying a remarkable degree of biological reduction and its adaptations tothe aquatic environment being so extremely successful, they have been repeated in several totallyunrelated flowering plant families, plus in ancient cryptogamic genera such as Equisetum andChara. H. vulgaris is such a successful species in reproductive and ecological terms, ithas achieved circumpolar distribution in the northern hemisphere, plus a limited presence in thesouthern. H. vulgaris is a rhizomatous, glabrous, heteromorphic perennial, having two growthforms, submerged aquatic and aerial emergent. The latter is unbranched, jointed and has many crowdedwhorls of leaves giving it a very similar appearance to a developing Horsetail (Equisetum) andthus, by comparison, it has been given the English common name, 'Mare's-tail'.
The emergent plant is most often observed and is very easily recognised by its erect, fleshy but rigid,hollow stems, rising 15-30 cm above the water surface, bearing the minute flowers and leaves that are10-20 mm long, in symmetrical whorls of six to twelve. The submerged or floating form of the plantproduces flaccid, translucent stems up to a metre long, bearing very much longer whorled leaves, 30-40mm, than does the emergent form (Sell & Murrell 2009). The submerged leaves have wavy margins and asoft tip, while the aerial emergent ones are entire-margined and are hard tipped. The hollow stems havenumerous well-developed air spaces that transport oxygen down to the submerged parts of the plant(Melderis & Bangerter 1955).
In B & I, H. vulgaris is occasional to locally frequent or abundant in shallow, sheltered,still or slow-moving mesotrophic to eutrophic, nutrient-rich, waters of lowland lakes, ponds,slow-flowing streams and ditches. It is most frequent in lime-rich waters and is usually rooted in finemineral or muddy organic substrates (Garrard & Streeter 1983; Preston & Croft 1997).
Fermanagh occurrence
H. vulgaris has been recorded in 54 Fermanagh tetrads (10.2% of the total), 48 of them containingpost-1975 finds. As the tetrad map indicates, the species is mainly confined to southern parts of LowerLough Erne, plus Upper Lough Erne and the River Finn catchment. Elsewhere in the VC, it is much morescattered in isolated, shallow water, lowland situations.
Mare's-tail is abundant in some lowland lakes, particularly in and around marl or limestone shores,including the Green Lough turloughs (ie vanishing lakes in limestone country linked to subterraneancaves), and along the marly soils of the River Finn. It is also occasional to rare on more acidicterrain, as in flushes on Sphagnum bog, and in recently created temporary pools in disusedquarries and sand pits.
Flowering reproduction
The flowers are borne singly in the leaf axils of the emergent portion of stem in June and July. AsHutchinson (1948) commented, "… the calyx of the bisexual flowers is very minute around the top ofthe inferior ovary, there are no petals, there is only one stamen, one subulate style (ie with a fine,abrupt, sharp point), one pendulous ovule, and of course only one seed in the small, nut-like fruit.Reduction could hardly go any farther in a bisexual flower!" The extremely reduced, wind-pollinatedflowers can be bisexual or unisexual on the same plant. The tiny flowers go through two distinct stages:firstly the female stage in which the tapering, pointed style is prominent and the red anther is sessileand still unopened, and secondly the male phase, with the style withered and the anther now elevated ona slender filament, open, exposing the pollen to the wind. The flowers set seed efficiently and dispersethem very effectively (Proctor et al., 1996; Preston & Croft 1997).
After fertilisation, the inferior ovary and its pendulous ovule develop into a very small, ellipsoid nutcontaining a single seed. In terms of seed dispersal, Ridley (1930) quotes Praeger (1913) who found theseeds were capable of floating for over 15 months and remained viable throughout. Ridley (1930) alsolists various ducks which eat and transport the (presumably undigested) nuts internally, thus enablingthem to reach and colonise isolated so-called temporary 'dew ponds' in quarries and in sand pits remotefrom connected water systems. H. vulgaris is also tolerant of saline conditions in damp to wetcoastal habitats. The established strategy of H. vulgaris is categorised by Grime et al.(1988, 2007) as CR, meaning Competitive Ruderal, which the current author (RSF) believes, describes theecological behaviour of this often extensive colony-forming aquatic species very well.
Fossil record
While fossil pollen of H. vulgaris is very rare, its mericarp fruits are amongst the mostcharacteristic and, in older deposits, the commonest of Pleistocene plant remains recorded in B & I(Godwin 1975). The fossil record shows H. vulgaris was present in these islands in all of thelast five interglacial periods and it has definitely been present throughout all zones of the currentFlandrian warm period (in Ireland, called the Littletonian). Its fruits have also been recognised fromall the last four glacial stages and, in the latest, the Weichselian, the records are extremelyabundant, occurring in association with a rich periglacial (ie near-to-glacier) flora and fauna. Fromthis evidence, it seems highly likely that H. vulgaris has persisted in Britain throughout thewhole Pleistocene era (Godwin 1975).
Irish occurrence
While H. vulgaris has been recorded at least once in every Irish VC (Cen Cat Fl Ir 2), theimpression this creates is quite misleading. The New Atlas hectad map shows that the species isvery much more consistently found in lowland VCs with permeable aquifers associated with fissured rocks.Generally speaking, these conditions apply across the Central Plain of Ireland and in much of Ulster(Haughton et al. 1979) and they literally underlie the main rivers and lakes from Coleraine onthe NE coast to Limerick in the SW, where the great, long River Shannon enters the Atlantic.
Elsewhere in Ireland, Mare's-tail is only very thinly scattered across the province of Munster and thesouthern half of Leinster, where the rocks are more impervious and the rivers are much less impressive.
British occurrence
H. vulgaris is widely distributed and sometimes abundant in lowland parts of England andWales, especially in limestone regions. It is represented across all latitudes from the Channel Isles tothe northern tip of Shetland (VC 112). It is, however, less frequent in W Scotland, Wales and the SW ofEngland. It ascends to around 900 m in Scotland, but is rare above 400 m (C.D. Preston, in: Prestonet al. 2002). It is reported to be declining in parts of Essex and Sussex (VCs 13, 14 and 18,19), but increasing in other areas such as Gloucestershire (VCs 33, 34). As it is frequently used todecorate and oxygenate garden lakes and ponds, and is readily available in the horticultural trade,deliberately planted material may escape into the wild, or perhaps H. vulgaris is sometimesdiscarded when it becomes too vigorous and abundant for the particular garden setting (Preston &Croft 1997).
European and world occurrence
H. vulgaris is described as being ± ubiquitous in Scandinavia and Hultén & Fries 1986 (Map1376) show it as very frequent and circumpolar in the northern hemisphere in arctic, boreal andtemperate zones, although it becomes scarce to rare and widely scattered towards the Mediterraneanbasin. It is also present as a native species in Chile from 50° S, and southwards to Tierra del Fuego.The Chilean plant appears identical with that of the northern hemisphere (Hultén 1974). While H.vulgaris is present throughout the subarctic and lower arctic regions, and reaches anorthernmost locality at 76° 49' N in E Greenland, in the Canadian E Arctic, however, it is rare northof the Arctic Circle (Godwin 1975).
Names
The genus name 'Hippuris' is from the Greek 'hippos', 'horse' and 'oura', 'a tail', the stems with theircrowded hair-like leaves supposed to resemble a horse's tail and hence 'Horse-tail'. This appliedequally to Equisetum species and botanists call them by that name. Hippurisvulgaris being a flowering plant, rather than a cryptogam, botanists distinguish it as it were afemale form of 'Horse-tail', ie 'Mare's-tail', following the ancient classical example, whereEquisetum species were called by the medieval Latin name 'cauda equina' (also translating as'Horse-tail') and H. vulgaris in relation to this was 'cauda equina femina' (Turner 1538; Gerard1597; Grigson 1974). The Latin specific epithet 'vulgaris' simply translates as 'common' (Johnson &Smith 1946).
The name 'Horse-tail' or 'Horsetail' has never been in common English folk useage for Equisetumspp., or for Hippuris, according to Grigson (1955, 1987). Thus 'Horse-tail' and 'Mare's-tail' aretwo perfect examples of English 'book names', invented by a botanical writer and subsequently taken up,repeated and promoted by others, but not used elsewhere. Traditional local English folk names listed forH. vulgaris include 'Cat's-tail' (the commonest name), 'Fox-tail', 'Bottle-brush', 'Joint-grass','Paddock's Pipes' (ie 'Toad's Pipes') and 'Witch's Milk'. Similar names listed for Equisetumspecies include 'Fairies' Spindles', 'Trowie Spindles' ('trow' is a reference to a goblin or devil),'Snake-pipes', 'Toad's Pipes' and 'Paddock's Pipes'. Both Equisetum and Hippuris have farand wide longstanding folk associations with fairies, goblins, toads, snakes, witches and the Devil (OldMan's Beard) (Grigson 1955, 1987).
Threats
None.
Native, common. Circumpolar boreal-montane.
1980; Weyl, R.S.; Clonshannagh Lough.
Throughout the year.
Recorded at the genus level from 286 tetrads, 54.2% of those in the VC, in a very wide range of wetlandhabitats, these small, annual or perennial floating or submerged aquatic herbs mainly grow in lakes,ditches and streams, but they do also occur in bogs and in turloughs or vanishing lakes in limestonedistricts. Most Callitriche species are capable of growing as both aquatic and terrestrial forms(Lansdown 2008) but, much more rarely than in truly aquatic settings, Callitriche spp. arerecorded from wet terrestrial situations, including disturbed ground such as in gravel pits and, in thewet climate of Fermanagh, even from churchyards.
Lansdown (2008) points out that there is no rigorous means of defining the terms 'aquatic' and'terrestrial' in relation to the growth forms of Callitriche species, because they are simplyextremes on a continuous moisture gradient, rather than being really and truly discrete. Aquatic plantsare those growing in and supported by water, while terrestrial plants are those not growing in water(unless they have recently been submerged). Terrestrial Callitriche are usually obviouslyappressed to the substrate and they form small creeping mats or rosettes. All leaves of terrestrialplants possess stomatal pores and the leaves are 10 × 5 mm. Plants that are found growing out of waterbut not showing the features mentioned, may have been recently exposed to the air by declining waterlevels, and they cannot be considered either aquatic or terrestrial (Lansdown 2008).
The genus is taxonomically difficult, the species being very variable in vegetative form, so thatsometimes fruit characters are essential for discrimination (Preston & Croft 1997). There is anexcellent and helpful treatment of identification problems summarised in the BSBI Plant Crib1998, and although it appeared after the fieldwork for this study, the BSBI Handbook No11 by R.V. Lansdown (2008) is now the definitive account of the genus Callitriche for thewhole of Europe. Prior to the availability of the latter reference, the fact that field botanists havebeen rather wary of the genus helps explain the large number of records at generic level compared withrelatively few for each of the six species recorded in Fermanagh.
As the tetrad map indicates, the genus is widely and evenly scattered in Fermanagh in aquatic and wetterrestrial habitats, but it is not possible from existing data to say much about the distribution ofthe individual species, other than that some are more local than others throughout the VC. A very usefulaccount of the genus and the biology and ecology of the species in B & I is provided by Preston& Croft (1997).
Native, occasional. Circumpolar boreal-montane.
October 1906; Tetley, W.N.; Blaney Bay, Lower Lough Erne.
June to October.
Fermanagh occurrence
The Fermanagh Flora database contains record details of this species from 27 tetrads (5.1%), 24 of themwith post-1975 records. It is a rather localised, morphologically variable, annual species, which thetetrad map indicates was previously confined to Lower Lough Erne (Revised Typescript Flora). Itis now also known from five lake and pond sites on or around Upper Lough Erne, and also from the remotewestern lakelet of Legalough above Marlbank, where it was found in September 1999 by RHN and HJN. TheseUpper Lough Erne sites are Killymackan Lough (an ASSI); the shore at Kilmore South Td, N of RabbitIsland; the shore at Crom Td, Lough Corby; and, finally, Lough Nacallagh. Vouchers exist for three ofthe five Upper Lough Erne sites that emanate from the 1986-7 EHS Habitat Survey of Upper Lough Erne, the1988-91 NI Lakes Survey and an ENSIS new lake survey carried out in 2006. Greater confidence exists ofthe correct identification of these particular records than to other un-vouchered records in theFermanagh Flora Database.
The distribution of C. hermaphroditica appears to closely mirror that of Zannichelliapalustris (Horned Pondweed), another submerged species of lowland aquatic habitats with siltysubstrates that range from mesotrophic to eutrophic in terms of their productivity. The New Atlashectad map shows that C. hermaphroditica has an essentially northern distribution in B & I.It is certainly much better represented (or better surveyed) in NI than in the RoI, although it doeshave a thin scatter of occurrences, including remote outlying stations in the far SW in Kerry (H1) and WCork (H3) and in the SE in Wicklow (H20) (New Atlas).
Threats
Insufficient information exists to determine whether Callitriche species are vulnerable or not.
Native, frequent. Circumpolar boreal-montane.
1881-2; Barrington, R.M.; Lower Lough Erne.
January to November.
Fermanagh occurrence
As the New Atlas map indicates, this is the most common species of Callitriche in B & Igenerally and this is also the case in Fermanagh. It is also regarded as being the most common,widespread and locally abundant member of the genus in Europe, and can occur in a very wide range of wetto damp habitats, but it occurs less frequently in deep or fast-flowing water, than in still orslow-flowing water (Lansdown 2008). C. stagnalis has been recorded in 113 tetrads, 21.4% of thosein the VC. As the Fermanagh tetrad map shows, Common Water-starwort is widely distributed over the wholeof the area, but it appears somewhat more frequently around Upper Lough Erne. As usual, however, it isimportant to remember and recognise that thanks to the NI Lake Survey, Upper Lough Erne has been moreextensively surveyed in comparison with the remainder of the county.
Preferred habitats
C. stagnalis occurs as a shallow, floating and submerged water aquatic over the whole range ofnutrient status from oligotrophic to eutrophic, avoiding only the most acidic, nutrient-starved peatypools, but including seasonally flooded basins like Roosky turlough. It frequently also behaves as aterrestrial annual of seasonally flooded or regularly damp ground, both in shade and in more opensituations. This behaviour appears to be especially the case the further west one goes in B & I(Preston & Croft 1997). C. stagnalis generally occurs with a mixture of terrestrial andamphibious species, such as Agrostis stolonifera (Creeping Bent), Apium nodiflorum(Fool’s-water-cress), Epilobium hirsutum (Great Willowherb), Juncus articulatus (JointedRush), J. bulbosus (Bulbous Rush) and J. effusus (Soft-rush). It rarely occurs along withtruly aquatic plant species, but it frequently co-habits with Callitriche palustris(Narrow-fruited Water-starwort) and C. brutia var. brutia (Pedunculate Water-starwort) inephemeral wetlands and on the exposed muddy margins of lakes and pools with fluctuating water levels(Lansdown 2008).
Local examples of terrestrial habitats of C. stagnalis occur in ephemeral pools in damp lakeshorewoods, such as Reilly Wood and Silles Wood, but it has been recorded also in the much more upland valleywood of the Correl Glen NR, plus on steep sloping wooded scarps at Pollaphuca (also known as'Poulaphouca') and Hanging Rock NR. It has been found in the marsh beside Doagh Lough, in hollows andruts in Lackboy meadows and on Skreen Hill, Derrin Mountain and Legacurragh above Florencecourt. Thelatter represents more calcareous terrain than would be normal for this species, but the ground isoverlain in places by fairly shallow, somewhat disturbed peaty soil. C. stagnalis has also beenrarely recorded in ditches along roadsides.
British and Irish occurrence
Lansdown (2008) warns that while C. stagnalis is recorded as widespread and locally abundantthroughout B & I, it is probably over-recorded for C. platycarpa (Various-leavedWater-starwort) as the two are not easily and reliably separable, the only really definite distinctioninvolving the uniformly spherical pollen grains of C. stagnalis. In addition, Preston& Croft (1997) suggest that at least in SE England some records of C. stagnalis may perhapsbe referable to C. obtusangula (Blunt-fruited Water-starwort).
European and world occurrence
C. stagnalis s.s. occurs throughout Europe from the Azores to Iceland and east to Belarus and WRussia. It is an introduced alien in N America, Australia, New Caledonia and New Zealand (Lansdown2008).
Threats
Probably none, but insufficient accurate data to be certain.
Native, rare. European temperate.
1881; Stewart, S.A.; Carrick Td, on the Western Plateau uplands.
Throughout the year.
Fermanagh occurrence
A taxonomically difficult and, locally, apparently rare perennial species with just three voucherspecimens and a total of 26 records in the Fermanagh Flora Database. It would therefore be foolish topretend to know much about the real local occurrence of this species which can easily be confusedwith the definitely much more common C. stagnalis (Common Water-starwort). Having said that,C. platycarpa has been recorded in 18 Fermanagh tetrads, 16 of them with post-1975 records.
The presence of stem scales and clearly winged fruits that are as wide as long, separates C.platycarpa from all other Callitriche species except C. stagnalis, from which itcan only reliably be separated by the pollen shape. The fruit wing is generally much wider in C.stagnalis, but there is some overlap with C. platycarpa. The bluntly triangular pollengrains of C. platycarpa can only be properly seen under 100-400 × magnification (Lansdown 2008).
Preferred habitats
The typical habitats are in shallow, still, slow-flowing or sheltered waters, or in muddy ground bylowland lakes, rivers and ditches. C. platycarpa appears sparsely scattered across Fermanagh inboth acid and base-rich areas. It might be slightly more prevalent around Lough Erne, but thedistribution data do not really support this contention, since its presence here could merely reflectthe recording bias created by excessively detailed lake survey recording carried out in this area by theEHS Habitat Survey Team.
Preston & Croft (1997) indicate that, like the morphologically similar C. stagnalis, thisspecies has both aquatic-perennial and terrestrial-annual life forms and, likewise, it too has a verywide ecological amplitude in terms of both water/soil chemistry and shade/open sunlight. Plants of C.platycarpa can be shy of flowering or apparently totally sterile, thus denying botanists thefruits necessary to confirm the species identification.
British and Irish occurrence
Preston & Croft (1997) believe C. platycarpa is grossly under-recorded in B & I, but withthe identification problems being significant (see the accounts by R.V. Lansdown in Rich & Jermy(1988) and in the BSBI Handbook 11, Water Starworts, Callitriche of Europe(Lansdown (2008)), it is hardly likely that they can really tell this for certain. It is perhaps more ofa hunch, but Lansdown (2008) is of the opinion that C. stagnalis is over-recorded at the expenseof C. platycarpa.
Visual comparison of the hectad maps in Preston & Croft's (1997) book, Aquatic plants in Britainand Ireland, with those in the New Atlas (2002) displays a considerable recent increaseof this species, especially in parts of C & S England. Although the distribution pattern stillremains rather patchy, becoming more coastal in Scotland north of the Glasgow-Edinburgh conurbations, itis felt that the species increase might be real, rather than reflecting recorder effort. The apparentincreased presence may result from widespread habitat eutrophication since the 1970s, which this speciescan tolerate better than many other aquatic species, since a similar circ*mstance has previously beenobserved in southern Sweden where C. platycarpa appears to be a recent arrival (R.V. Lansdown,in: Preston et al. 2002; Lansdown 2008).
European occurrence
On the continental mainland, C. platycarpa occurs mainly in W Europe, throughout France extendingnorth into S Sweden and east to the Czech Republic. It has also been reported from a few sites in theIberian Peninsula. It appears to be extending its range to the N & E (Lansdown 2008).
Native, very rare. Suboceanic boreo-temperate.
1946-54; MCM & D; drain at Inver Lough, Tattygormican Td, 2 km SW of Rosslea.
June to August.
Fermanagh occurrence
There are just six Fermanagh sites for this apparently rare perennial Water-starwort recorded in fivetetrads as the accompanying map shows, three around Upper Lough Erne and two in isolated lakelets on theeastern boundary of the VC. Only one of the seven records has a voucher in BEL and, rathersurprisingly, C. obtusangula was not observed at all in the VC by the very thorough and extensiveNI Lakes Survey carried out between 1988 and 1991.
Locally, it is very rarely recorded in still or slow-moving water in lowland lakes, streams and ditches.The details of the other Fermanagh records are: N shore of Moorlough Lake, 1946-53, MCM & D; wetditch, Inishroosk, Upper Lough Erne, June 1974, P. Hackney, voucher in BEL; Rabbit Island, UpperLough Erne, 5 August 1986, S.J. Leach & A.S. McMullin; Colebrooke River estuary, Kilmore Td, UpperLough Erne, S.J. Leach & A.S. McMullin (two sample areas recorded); and Rose Lough, Mullycanvan Td,20 July 1988, RHN & J.S. Faulkner.
Habitat preferences
As with all Callitriche spp., the species is taxonomically difficult and C. obtusangulaoccurs in still to slow-flowing, mesotrophic and eutrophic water, perhaps most abundantly overcalcareous soils, but also over peaty substrata. It is strictly a lowland species. In coastal areas, itcan grow in brackish water, typically in ditches in grazing marshes. Terrestrial forms grow on damp mudat the water's edge and are always annuals (Preston & Croft 1997).
The aquatic form of C. obtusangula is a perennial with a life-form similar to C. platycarpa(Various-leaved Water-starwort) and the two species frequently occur together – plants overwintering assubmerged shoots with linear leaves. In the spring and summer, the stems grow to reach the water surfacewhere they develop floating rosettes. Plants in still or quieter, slow-flowing waters can develop large,dense beds under favourable conditions and flower and fruit freely, while those in faster-flowingconditions generally remain submerged and may fail to fruit (David 1958; Lansdown 2008).
British and Irish occurrence
C. obtusangula is widespread in England and
Wales south of a line between Lancaster and Hull, but it quickly becomes rare and widely scatterednorthwards and it is very rare or absent in most of Scotland. In Ireland, it is scattered down both theW & E coasts, and apart from a few hectads along the River Shannon, decidedly rare or absent in thecentre of the island.
Preston & Croft (1997) stated that in their opinion, while C. obtusangula is a fairlydistinctive species, it is almost certainly under-recorded in B & I, being most likely confused withC. stagnalis (Common Water-starwort). However, with so few records here in Fermanagh and, indeed,similarly elsewhere in most of B & I, more careful, systematic recording is required to determinethe true picture of its occurrence and distribution.
European occurrence
C. obtusangula belongs to the suboceanic southern-temperate phytogeographical elementand, on the continental mainland, it occurs from the Netherlands to Austria and the formerYugoslavia in the east, and from there, westwards along the Mediterranean coast, including on theislands of Corsica and Sardinia, into Spain and Portugal. It also occurs in N Africa, in both Moroccoand Tunisia (Lansdown 2008; Sell & Murrell 2009).
Native, scarce but locally abundant. Possible mis-identification. Mediterranean-Atlantic.
1986; McMullin, A.S.; Drummoy Lough.
May to October.
Growth forms and preferred habitats
Sometimes the terrestrial form of this annual or perennial is abundant on exposed mud of lakeshoresduring dry weather, but it is capable of growing in more ephemeral pools and in damp exposed ground,including beside ditches, puddles and in poached or rutted muddy ground in fields or by tracks. Afloating aquatic form of this strictly calcifuge species also grows in more permanent shallow, still orslow-moving water of acid, nutrient-poor status (Preston & Croft 1997). The species is aself-pollinated annual, pollination being submerged or aerial (Lansdown 2008). The fact that it isfrequently recorded in a variety of artificial man-made or disturbed habitats, such as sand- and gravelpits or rutted tracks, somewhat remote from other populations, indicates that sexual reproduction andseed dispersal are effective and sufficiently efficient to allow this to happen (Preston & Croft1997).
C. brutia has only fairly recently been split off from C. hamulata s.l. (IntermediateWater-starwort) at species level, and it differs from that taxon by its asymmetric, notched leaf-tipsand smaller fruits, which in the terrestrial form of the plant are distinctly (at least 2 mm) stalked(ie they are pedunculate). In reality, the separation of the two species is a very difficult one,especially in the field, due to considerable variation in both of them in their aquatic and, to a lesserextent, in their terrestrial growth forms. Subsequent to the Fermanagh Flora survey featuring inthis website, Lansdown (2008) downgraded and reclassified C. hamulata as a variety of C.brutia. In this new Lansdown interpretation, var. hamulata appears to be more frequent inpermanent, swift-flowing water, while var. brutia typically occurs in ephemeral water bodies andat the margins of lakes and streams. Also, plants from the southern part of the C. brutiaspecies range in Europe tend to be var. brutia, with var. hamulata appearing the morefrequent form in the north (Lansdown 2008).
In the Callitriche Handbook, Lansdown (2008) points out that, "C. brutia [includingC. hamulata as a variety] can be distinguished from all other European Callitriche taxa bythe strongly reflexed style, the remains of which are appressed to the side of the fruit, combined withthe lack of a strongly ornamented exine [outer wall] on the pollen grains".
During the summer, terrestrial plants generally fruit abundantly, but non-fruiting plants, aquatic orterrestrial, often cannot be identified confidently and certainly not in the field.
Fermanagh occurrence
Field records of Pedunculate Water-starwort have been made in Fermanagh a total of 41 times in 21 tetrads(4%). Twenty-nine records were made in August and September 1986 by members of the EHS Habitat SurveyTeam when working around the southern portion of Upper Lough Erne (14 tetrads). Apart from this, only afurther eight more remote records exist, most of which were made by RHN. The details of the latter are:swamp at Derryclawan, 2 km NNW of Lisbellaw, 30 May 1992; lakeshore at Mullaghfad, Gubbaroe Td, LowerLough Erne, 30 October 1992; Lough Naman (B), Conagher Upper Td, 2 September 2009; Lough Formal, 19August 2010, Lake Ecology Team, also 11 October 2010, RHN; by stream opposite old house, Tullygarry,near Meenatully Hill, 4 October 2010, RHN & HJN; gravel pit N of Derrin Mountain, 26 October 2010,RHN & HJN; and Lough A Waddy, 26 October 2010, RHN & HJN. The NI Lakes Survey made no records ofthis species.
As none of these records have vouchers, they are not fully acceptable as valid, but are included here andto some extent given the benefit of the doubt.
British and Irish occurrence
When it comes to considering mapped B & I distribution, C. brutia (in the Lansdown sense,including var. hamulata) equates with C. hamulata s.l. in the Preston & Croft (1997)and New Atlas sense. The map in the two latter references shows the relative abundance of C.hamulata s.l. in areas where acidic waters predominate, primarily in the N & W of bothislands, and its scarcity elsewhere, ie in E England and C Ireland. There does not appear to besufficient accurate data on C. brutia or C. hamulata s.l. to determine whether there hasbeen any historical decline or increase (Preston & Croft 1997; R.V. Lansdown, in: Preston etal. 2002; Lansdown 2008).
European and world occurrence
C. brutia (including var. hamulata) or C. hamulata s.l. is a widespread speciesthroughout N, W & C Europe, extending from Iceland and the Faeroes to Scandinavia southwards to thePyrenees, Poland, the Czech Republic, N Italy, Greece and Romania. Beyond Europe, it has been recordedfrom Greenland, Morocco, Iran and the Caucasus. Var. brutia has been introduced to Australia andnorth island, New Zealand. Var. hamulata has been found to be widespread on the W coast of NAmerica, where again, it is almost certainly an introduction (Lansdown 2008).
Callitriche hamulata Kütz. ex W.D.J. Koch s.s., Intermediate Water-starwort
Native, occasional and local. Suboceanic boreo-temperate.
1881; Stewart, S.A.; Carrick Lough, Dresternan Td.
June to September.
This is another taxonomically difficult species, previously regarded as a subspecies of C.intermedia and closely related to C. brutia (Pedunculate Water-starwort – see the C.brutia account on this website), itself once regarded as a variant of C. hamulata.Lansdown (2008) reversed the status of these two taxa, re-classifying C. hamulata as a variety ofC. brutia!
Whatever its rank, this taxon is found in a wide variety of clear, acid, nutrient-poor waters at alllevels.
Fermanagh occurrence
C. hamulata (or C. brutia var. hamulata) has been recorded in 37 Fermanagh tetrads(7%), 30 of them with post-1975 records. As the tetrad distribution map shows, this perennial appears tobe thinly and widely scattered across the VC, typical habitats being lakes, muddy pools, streams and bogdrains. In shallower, more temporary water bodies, the species (or variety) is very probably forced tobehave as an annual. In contrast to the situation with C. brutia, in the Fermanagh Flora Database29 of the 51 records of C. hamulata s.s. were located by the 1988-91 NI Lakes Survey. It isimportant to recognise and emphasise that knowledge of Callitriche species in Fermanagh andelsewhere is very incomplete and that the present details are only given for the guidance of futurerecording.
One of the best distinguishing features of this species or variety is the abruptly expanded, spanner-liketip of submerged leaves of the aquatic form of the plant (ie a broad, regular, apical notch). A veryhelpful set of three identification keys to this and the other species of Callitriche waspublished by R.V. Lansdown in the BSBI Plant Crib 1998, and now that a well-illustratedBSBI Handbook exists, covering all the Callitriche taxa in Europe (Lansdown 2008), it is hopedthat future Water-starwort recording throughout B & I will be more securely founded than haspreviously been the case.
Native, very rare. Eurosiberian wide-boreal.
1947; MCM & D; on a clay bank by NE shore of Ross Lough.
April to July.
Growth form and preferred habitats
P. maritima is a rosette-forming perennial with a branched, woody rootstock and long, narrow andrather thick, fleshy leaves, rounded on the back and channelled on their upper surfaces. The greenishflowers are borne on a slender cylindrical stem in a narrow spike rather similar to that of P.major (Greater Plantain). Each plant rosette usually forms numerous flowering stems. When itflowers, from June to September, the yellow anthers on long filaments are very conspicuous and the plantis wind pollinated (Hepburn 1952; Clapham et al. 1987).
Like other Plantago species, Sea Plantain has no powers of increase or spreading other than byseed. Again, in common with all maritime species, Sea Plantain is a salt-tolerant (ie halophyte) plantof middle and upper reaches of salt-marshes, in the middle and upper zones of salt-marshes and in open,low-growing coastal turf, often alongside Armeria maritima (Sea Pink or Thrift). The leaves ofP. maritima are relished by sheep and, like Thrift, it stands up well to natural grazing in asalt-marsh setting (Hepburn 1952). P. maritima can also grow in shingle at the top of beaches, inrock crevices and on short turf on sand near the sea. It is also frequent in sea cliff grassland, whereit sometimes becomes dominant or co-dominant with other plantains or Thrift, especially if sheep grazingis taking place or other stress features including sea spray and wind exposure help reduce interspecificcompetition from grasses and other somewhat taller-growing or more aggressive plant species (Chapman1976). Sometimes this form of plantain sward vegetation is extremely dwarf, being just a few cms inheight (Hepburn 1952). Like Sea Aster (Aster tripolium), Sea Plantain can also grow in brackishsituations in salinities of, "a quarter of sea water or less" according to Proctor (2013, p.428).
P. maritima is a phenotypically very plastic species that can tolerate an extremely wide varietyof soil physical and chemical conditions, ranging from clay-rich muds to sandy gravel marshes andpastures. However, it cannot tolerate much plant competition (Ranwell 1972). Nevertheless, Sea Plantaindoes survive considerable pressure from other forms of stress, enabling it to colonise, thrive andsometimes even dominate salt-laden, weather-exposed and sheep or rabbit grazed habitats, always insituations that minimise direct species competition (Hepburn 1952).
In addition to its common and very widespread coastal occurrences, inland populations of P.maritima occur beside streams or around wet rocks and in stony flushes in the higher mountainsin the English Pennines, N Wales, NW Scotland and the N of Ireland, and on limestone in western parts ofIreland, sometimes at low altitude (Botanist in Ireland, paragraph 64; Godwin 1975; Garrard &Streeter 1983; Sell & Murrell 2009). It tends to frequent both acidic and limestone soils wherethere is some degree of flushing water moving through silty soil on cliff rock ledges and screes(Tansley 1939; Graham 1988; Halliday 1997; G.M. Kay, in: Preston et al. 2002).
In recent decades, and especially since the advent of motorways in B & I, P. maritima has beenspreading inland along salted roads in many places and doing so particularly commonly in Scotland (G.M.Kay, in: Preston et al. 2002).
Variation
Various attempts have been made to subdivide this rather variable species, but the taxa proposed haveproved difficult to define morphologically.
Plants from Arctic Europe have been named subsp. juncoides (Lam.) Hultén, which is believed todiffer from the widespread form subsp. maritima in having wide bracts, a more ovid to globosecapsule shape, the scapes not exceeding the leaves and more numerous seeds per capsule. However, thesecriteria for discrimination have not proved very satisfactory.
Another variant named subsp. serpentina (All.) Arcangeli is described as having long-acuminatebracts and the posterior sepal with the keel often narrowly winged. It occurs in S Europe, especially onmountains above 2,000 m. However, it is probably only an ecological variant and therefore not worthy ofsubspecific rank (D.M. Moore, in: Tutin et al. 1976).
A more complicated set of suggestions for subdivision of what is considered a polymorphic species tosubspecific or varietal rank on a worldwide basis has been made by Hultén (1974) and applied to a map inHultén & Fries (1986, Map 1726), without being any more convincing to the current author (RSF).Apart from the questionable taxonomy, the Hultén (1974) nomenclature is also confusing.
The more recently published British Floras do not recognise any taxonomic subdivisions in P.maritima (Sell & Murrell 2009; Stace 2019).
Fossil history
Fossil evidence proves that P. maritima has been present in B & I in every glacial andinterglacial sub-stage or zone from the very ancient late Anglian to the present day (Godwin 1975). Thusthe inland mountain and salt-marsh sites in B & I represent very long-established populations ofthis fleshy perennial. On the basis of fossil pollen and macrofossil remains (including in the case ofP. maritima, seeds), this species, and a limited number of others like it, are recognised asrelict survivors from Late-glacial vegetation populations in these isles that were established undertotally different climate and soil conditions from those pertaining today (Godwin 1975). P.maritima and its previously widespread Full- and Late-glacial plant associates were forced toretreat to a range of tree-free coastal and high mountain habitats in order to avoid the shade, closedcover and severe competition of taller-growing, better adapted, immigrant colonising plant species thatarrived when temperatures rose and glaciers melted 10,000 years BP. In later post-glacial stages andclimates, these Late-glacial flora relicts also had to survive the development and expansion of acidpeat deposits (Raven 1956).
The fact that some of these Late-glacial species populations survive as relicts today is quite amazing,and it is not surprising that after persistence of thousands of years, a number of them represent therarest and most thinly scattered species in B & I. While inland mountain and salt-marsh populationsof P. maritima belong to this group of rarities, the species is very common, widespread andindeed often super-abundant in coastal maritime sites.
Fermanagh occurrence
In land-locked Fermanagh, Sea Plantain is locally very rare, but it has been recorded in four tetrads inthe VC over a 50 year period. The Fermanagh sites where this species has been recorded are unexpectedand varied, including both relatively upland calcareous lakeshores around or close to Monawilkin Loughand Ross Lough (the VC has no high mountains) and in lowland grassland in Knockmore, Tonlisderrit andTirmacspird townlands. At Ross Lough, it appears confined to the more or less steep slope of the old,superseded shoreline, particularly where cattle trample and poach the ground, keeping it open so thatbare soil is always present. There is a danger that nearby gorse bushes may spread into this site.
The additional local record details are: around Monawilkin Lough, 10 July 1985, S.J. Leach & S.A.Wolfe-Murphy; W end of Monawilkin Lough (to N of track under cliffs), 1994 & 1996, I. Rippey;Tonlisderritt, 1 km SE of Cashel, 28 April 1996, RHN & HJN; lowland acid grassland, Tirmacspird Td,NW of Lack, 7 June 1997, RHN; and also at Meikle and co-workers' old site, the clay bank by NE shore ofRoss Lough, 25 July 2000, RHN & RSF; also here 17 September 2006 & 19 September 2009.
Other normally maritime species which 'misbehave' in this ecological manner by occurring in land-lockedFermanagh include Carex distans (Distant Sedge), Asplenium marinum (Sea Spleenwort) andViola tricolor subsp. curtisii (a seashore form of Wild Pansy,Seaside Pansy).
British and Irish occurrence
P. maritima is a very common and widespread species around almost the entire maritime coastline ofB & I. While of a very much rarer occurrence, it is also found in upland species-rich pastures andin rocky mountain ground in the Pennines and scattered across the N & W of B & I. It is rarestin inland salt-marshes, which are themselves rare habitats, but it has more recently appeared along somestretches of salt-treated motorways, chiefly in Scotland (G.M. Kay, in: Preston et al. 2002; Sell& Murrell 2009).
European and world occurrence
On the European mainland, P. maritima extends in coastal habitats from the North Cape toSpain and it also occurs rarely in inland situations (Godwin 1975). It is rarer in the south of Europeand extends into N Africa. It is also present in Greenland (Böcher et al. 1966), Iceland (Löve1983), N America and southern S America (Sell & Murrell 2009).
Uses
Apart from the fact that it was appreciated to be nutritious for sheep and pigs to graze, there does notappear to be much mention of folklore uses for the plant (Grieve 1931). It is mentioned in passing byAllen & Hatfield (2004) as having a reputation in the Cork region of Ireland as a general stauncherof wounds, in common with other plantains.
Names
The genus name 'Plantago' is from the Latin 'planta', 'the sole of the foot',referring to the broad, flat leaves that are often pressed to the ground and from the femininetermination 'āgo' (Gilbert-Carter 1964). The Latin specific epithet 'maritima' means 'belonging to thesea', or 'growing on the sea coast' ('mare') (Gilbert-Carter 1964).
English common names include 'Buck's-horn', Buck's-horn Plantain', 'Sea Plantain' and 'Gibbals Kemps':the first two of these is nowadays transferred to Plantago coronopus. "'Gibbals' is or was aname applied to P. maritima around Rommy marshes, about 2 miles [3.2 km] from Cardiff". Amanuscript account of a botanical excursion in 1775 by Lightfoot recounts how hogs were encouraged tograze on the roots that they were reputed to be fond of, that were grubbed up for them, and on whichthey grew fat (Britten & Holland 1886). 'Kemps' ('kemp' is to fight in northern dialect and was moreoften applied to P. lanceolata (Grigson 1955, 1987). It is not known how it came to be applied toP. maritima.
Native, very common, locally abundant. Eurasian wide-temperate, but widely naturalised in bothhemispheres and now circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
March to December.
Growth form and preferred habitats
A large and long-persistent soil seed bank enables this all-too-familiar, yet phenotypically extremelyvariable, weedy, rosette-forming perennial (or annual or biennial), to germinate at any time during thegrowing season (with a peak flush in late May) (Hawthorn 1974). Irrespective of seasonality, it can veryquickly colonise open, short-turf grassland, or bare, disturbed, compacted, wet or dry, very fertile,heavy soil types. Germinating seedlings show effective root penetration in compacted soil (Blom 1979)and the species tends to be especially frequent in heavily trampled ground, including around gatewaysand along lanes and roadside verges. Experimental work has found that the primary roots from smallseeded Plantago species, such as P. coronopus (Buck's-horn Plantain) and P. major subsp. major, are more capable of penetrating the surface of thesubstrate than those of relatively large seeds such as P. lanceolata (Ribwort Plantain) and P.media (Hoary Plantain) (Blom 1978).
P. major is capable of forming almost pure stands occupying the zone closest to maximumdisturbance, although it often associates with Matricaria discoidea (Pineappleweed) and Poaannua (Annual Meadow-grass). Even in closed turf, if P. major manages to colonise anygaps, it will quickly establish and become very persistent as gardeners know to their cost. It canbecome a real nuisance in lawns (Sagar & Harper 1964).
The primary roots of the seedling are replaced early by a fibrous, adventitious root system froma very short (1 cm), woody taproot or caudex that eventually represents the overwintering food storageorgan of the mature, established plant. In P. major, some of the later roots are contractile andhelp to keep the growing point at the soil surface where it is protected (Holm et al. 1977). Inthe spring, established plants quickly develop a rosette of spirally arranged, broad, ovate basalleaves. The broad leaves are 5-10 cm long, have an entire or obscurely toothed margin and have 5-7parallel ribs converging into a rather long petiole. Rapid growth and long leaf petioles allow the plantto produce its characteristic wide-spreading leaf rosettes that establish and occupy valuable surfacespace. The individual plant can then tenaciously hold its ground against almost all other species,forming small or larger aggregations of offspring around it (Holm et al. 1977).
P. major is generally deep-rooted and can tolerate a range of soil reaction from moderately acidto calcareous (Grime et al. 1988, 2007). The mature root system of P. major may exceed 80cm in depth and 80 cm in diameter (Kutschera 1960), which enables the species to survive droughtconditions. Being resistant to both water-logging and drought and tolerant of half-shade, GreaterPlantain appears in a wide variety of lakeshore habitats and even to a limited extent in open areas inwoodland in Fermanagh. Generally, it avoids only extremely acid peat or mineral soils, mountaingrassland, deep shade, or sites that are continually wet during the growing season (Hawthorn 1974).
The established strategy of Greater Plantain is categorised as intermediate between Ruderal and abalanced C-S-R species (ie Competitor-Stress-tolerator and Ruderal). It is essentially a lowland speciesof heavily disturbed, fertile, productive vegetation, but it can occur and be widespread up to around450 m (Grime et al. 1988, 2007).
When growing in exposed places, P. major overwinters as a bud on the woody rootstock at or nearground level. On the other hand, when growing in herbage, an above ground leaf rosette is maintained inall but the most severe winters (Bates 1935).
As Sagar & Harper (1960) point out, many aspects of the life-cycle of P. major suit it forlife as an annual species, for example, its self-compatibility, rapid establishment from seed, quickflowering and equally quick maturation of seed. Nevertheless, P. major is capable of persistingas a true and potentially long-lived perennial.
Flowering reproduction
Flowering begins in early June and continues for about four months with a peak in July. Each rosetteproduces one to many short, leafless scapes bearing the characteristic, cylindrical spike inflorescencethat can vary between 1-30 cm long (Holm et al. 1977). The flowers are greatly reduced,wind-pollinated and capable of full self-fertilization. The individual flowers are insignificant inappearance, scentless and without nectar. Seed set is rapid after fertilization, some fruits maturing2-3 weeks after flowering (anthesis). The species normally produces abundant seed each year and theseed, which is always viable, is small, hard and flattened (Sagar & Harper 1964; Hawthorn 1974).
While some early formed capsules shed seed in autumn, frequently capsules do not disperse their seeduntil the following spring or summer. Although the capsule splits in the circ*mscissile mannercharacteristic of the genus, ie opening around its equator, there is no active dehiscence of seed fromthe capsule after it splits open. Thus seeds must be knocked off the plant by animals, wind or water. Ifthe seeds are not taken off, they remain attached to the spike until it rots at the base and falls tothe ground the next year (Holm et al. 1977).
The number of seeds per capsule varies from 3-28 and the estimated seed production for a large plant isreckoned at c 14,000 per year (Kerner 1896). The seed shape varies with the number contained inindividual capsules, a clear indication of the degree of variability within the species, in that acharacter usually so very conservative can be modified in P. major.
While the main period for seed germination is in spring (peak in May), seeds also germinateintermittently throughout the growing season (Hawthorn 1974) from a large, persistent soil seed bank(Thompson et al. 1997). Seeds of P. major gave 10% germination after burial for 40 years(Crocker 1938).
Despite the purple colour of the anthers, the spikes in full bloom are much less conspicuous than theyare in fruit. Since the seeds are attractive to birds, which aid their dispersal, the species bearsseveral English common names that emphasise this property, such as 'Birdseed' and 'Bird's Meat'(Salisbury 1964). In water, an adhesive layer of mucilage on the seed coat (testa) of P. majorswells to increase the seed size by about 40%. The ability to imbibe water increases the stickiness ofthe seeds to soil particles, feathers and fur and greatly enhances the potential for dispersal (Hawthorn1974).
Variation
Plants vary greatly in size and appearance according to the environmental conditions, ie it isphenotypically very plastic (Salisbury 1964). Infraspecific variation is extensive in P. majorand has resulted in a number of lengthy subdivisions into subspecies, varieties and forms (Hawthorn1974). Sagar & Harper (1964) described Greater Plantain as, "certainly one of the most variablespecies in the British flora". Sell & Murrell (2007) subdivide P. major into twosubspecies, subsp. major and subsp. intermedia, and list four varieties of the latter.
Although subspecific variation has not been sought out in the VC survey, the Fermanagh plant isundoubtedly subsp. major which has cordate leaves with 5-9 veins. In comparison, the leaves ofsubsp. intermedia have 3-5 veins and the leaf blade narrows to the petiole (Garrard &Streeter 1983).
Hybrids
No hybrids have ever been reported and attempts to produce them artificially have failed (Sagar &Harper 1964).
Vegetative reproduction
Unlike P. lanceolata (Ribwort Plantain) and P. media (Hoary Plantain), P. major doesnot reproduce freely by vegetative means. While reproduction is primarily by seed, about 20% of plantsin trampled laneways and in recently disturbed soil also produce ramets or offsets from buds on thepersistent woody crown, undoubtedly in response to damaged tissues. No plants in pasture produce ramets.The ramet remains attached to the parent plant for two to three years, after which the bond breaks andthe ramet initiates root development. The daughter plants produced in this way are immediately capableof flowering and seed production (Hawthorn 1974).
Economic importance as a weed
As with P. lanceolata, the success of P. major has been very closely linked to man'sactivities and especially with agriculture, for a very long time (Holm et al. 1977). In localsituations, P. major can reduce the early growth of arable cereal crops such as wheat and oats.While originally P. major and P. lanceolata were temperate zone plants, they have bothentered the warmest regions of the world to become pests of almost all tropical crops. They are uniquein that more than 50 countries report them as weeds in 26 crops that are as different as alfalfa,coffee, temperate orchards, rice, pastures, cereals, onions, cotton and small fruits. They are mostserious competitors in cereals and are common in arable fields of sugar producing areas. Their seedsfrequently contaminate small-seeded legume seed-stocks (Holm et al. 1977).
In pasture (and in lawns), P. major plants benefit from occasional mowing as the tall vegetationof competing species is temporarily removed. As expected, P. major grows best in hay pasturewhere grazing and trampling are prevalent (Hawthorn 1974). The most important factor governing thedistribution of P. major around the world is the disturbance of an area by man or animals (Holmet al. 1977).
P. major also hosts a number of viral, fungal and insect pathogens (Hawthorn 1974).
Fossil record
Continuous pollen and macrofossil records from the Weichselian and Flandrian periods prove that P.major is a species of ancient native status in Britain at least (Godwin 1975).
Fermanagh occurrence
Very common throughout Fermanagh on disturbed ground including heavily grazed and/or thoroughly trampledgrassland, margins of cultivated ground, lawns, gateways, farmyards, car parks, tracksides, roadsidesand waste ground. P. major has been recorded in 447 Fermanagh tetrads, 84.7% of the VC total,which means that it ranks 28th most widespread vascular plant in terms of tetrad frequency.
British and Irish occurrence
In suitably disturbed sites, P. major is extremely widespread and virtually ubiquitousthroughout both B & I; in the New Atlas map there are very few hectads where it has not beenrecorded, most of them probably representing high ground above 630 m (G.M. Kay, in: Preston etal. 2002).
European and world occurrence
The species belongs to the Eurasian Wide-temperate phytogeographical element and is very widespreadacross Europe, N Africa and N & C Asia. It has been very widely spread by man across bothhemispheres, eg to N America where it occurs parallel to indigenous related taxa (Hultén & Fries1986, Map 1725).
Uses
The fact that plantains frequently grow on paths and around gateways where they get trampled upon,crushed and damaged suggested to herbalists that the plant was so powerful, tough and elastic itmust heal crushing, tearing and bruising wounds in humans (Grigson 1955, 1987). Dioscorides wrotethat Plantain was a vulnerary (wound-healing herb) and was good in treating ulcers, sores and otherforms of wounds or blemishes, including burns. In time, Plantain (probably both P. major andP. lanceolata) became herbs of medico-magical power and divination, the root worn around theneck, for example, as an amulet against the King's Evil.
Grieve (1931) describes P. major as being refrigerant, diuretic, deobstruent and somewhatastringent. It has been used in treating skin inflammation, malignant ulcers, intermittent fever and asa vulnerary. The fresh leaves were also used to staunch bleeding, applied externally as a poultice(Grieve 1931). Additionally, the leaves were used to treat piles, varicose veins, festering wounds andto draw out pus from infected wounds or swellings. It was also considered useful for relieving pain, asin stings. In Ireland, plantain juice was used for coughs and the plant was put to very many othermedicinal uses including gout and for treating sore eyes (Allen & Hatfield 2004).
Names
No less than 31 English common names are listed by Grigson (1955, 1987), at least six of which refer tobirds, since they are reputed to help disperse the seeds, eg 'Canary Flower', 'Canary Food', 'LarkSeed'. Other names refer to the supposed healing powers of the herb, eg 'Healing Blade', the Scottish'Slanlis' and the Irish equivalent 'Slanlus', meaning 'healing herb' (Grigson 1955, 1987).
Threats
None.
Native, very common. Eurosiberian southern-temperate, but very widely naturalised in both hemispheres andnow circumpolar.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
P. lanceolata forms a rosette plant from a short, thick, erect rhizome. The root system iscommonly represented by a few deep roots capable of descending over a metre and a mass of surfacefibrous roots 5-10 cm in length suggesting physiological differentiation into water andnutrient-absorbing systems. The leaves are simple, 10-15(-45) cm long, 2-3.5 cm wide, linear-lanceolateto elliptic-spathulate, margins usually entire and with 3-5-longitudinal ribbed veins (raised on theleaf underside). It is the conspicuous ribs on the leaves that give the plant its English common name'Ribwort Plantain'. When it has more linear leaves, the blade narrows gradually into the leaf base, thepetiole length apparently depending on the height of the surrounding vegetation (Sagar & Harper1964).
As with P. major (Greater Plantain), there is a great amount of phenotypic variation in manyRibwort Plantain characters and undoubtedly the species embraces a number of genetically differentstrains. Generally, the plant follows a perennial life history and reproductive strategy, but rarely (asin cultivation experiments), it can behave as either an annual or a biennial. Overwintering rosetteshave wider leaves than spring and summer ones and those growing in exposed sites are more prostrate, sothey can sometimes be confused with P. major.
P. lanceolata occurs in a very wide range of habitats and although it is largely absent fromdeeply shaded woodland, wetlands and the most acidic ground, it does commonly occur on their margins.The species is most commonly found in pastures, rough grassland and disturbed ground, including ingardens, lawns, roadsides and waste places. P. lanceolata is frost-tolerant and also very droughtresistant. It competes best in drier environments and in hard-packed neutral to basic soils. It is alsowell able to cope with shade and can germinate and establish even when sited in closed turf vegetationamong tall grasses (Blom 1978; Cavers et al. 1980).
Ribwort Plantain is fast growing and very competitive, being quickly able to exploit increased fertilityeither as a result of additional water or nutrients or both of these factors. It is also extremelysuccessful in exploiting gaps in turf by both vegetative expansion and seedling recruitment from apersistent buried soil seed bank (Thompson et al. 1997). The deep taproot allows the plant to behighly drought-resistant and to survive on very dry soils, even in crevices in rocks and on walls. It isone of the world's most successful (or worst) weeds (Holm et al. 1977) and is ranked in the topdozen most successful colonising plant species in B & I. The established strategy is categorised as'C-S-R', meaning it achieves a balance between or involving all three ecological strategies, Competitor,Stress-tolerator and Ruderal (Grime et al. 1988, 2007).
Variation
It is well recognised that P. lanceolata is extremely variable, much of the variation beingwith respect to the particular environment in which it grows (ie it is phenotypically plastic). Howeverthere is a clinal aspect to the variation also, reflecting genetic change across its geographical range.In the past, the variation within the species has frequently been split into a series of subspecies andvarieties. As an example of this, Druce (1928) distinguished as many as twelve varieties, and Pilger(1937) set up just two varieties, var. communis Schlechtend and var. dubia Wahlenberg.However, the first of these he spilt further into five sub-varieties, and in the second variety hedistinguished two more sub-varieties (Sagar & Harper 1964).
In the account in Flora Europaea 4, D.M. Moore came to the conclusion that formalrecognition of any of the suggested variants as subspecies was impractical (Tutin et al. 1976).Having said this, in their critical Flora of Great Britain and Ireland 3, Sell &Murrell (2009) distinguish five varieties, based largely on habitat or geography, but recognised bydifferences in leaf shape, size and hairiness together with inflorescence shape. None of these formshave been looked for in Fermanagh.
Palatability and benefits to grazing animals
P. lanceolata is nutritionally valuable for stock animals, providing a particularly good source ofessential minerals, including calcium, iron, sodium, chlorine, nitrogen, potassium, phosphate andcobalt. It also provides small amounts of magnesium but is poor in manganese (Thomas & Thompson1948; Wilman & Derrick 1994). Ribwort Plantain has in the past been sown in herb strips along withley grass and legume mixtures in order to provide forage variety to stock and to supplement theirmineral nutrition. In Germany, P. lanceolata has been a recommended constituent of pasturemixtures because it is known to improve animal health and milk flavour (Heeger 1949). In 2021, RibwortPlantain is again being introduced to cattle and sheep grazing experimental multi-species herb andlegume swards in Co Down (NI) and elsewhere in B & I as part of an ongoing government sponsoredEuropean Innovation Partnership Project. This experiment is investigating both meat production andadditional benefits to soil and animals from a reduced need for fertiliser nitrogen, extension of thegrowing season, drought resistance, reduced parasite burden and enhanced biodiversity above and belowground.
In dry pastures, the plant is palatable to livestock (especially sheep) and provides, "fair qualityfodder, although inferior to grasses" (Clark & Fletcher 1906). An animal grazing study byMilton (1933) found it, "definitely palatable and may even be sought after". Sheep are so keenon it that they, "will actually chisel the crowns out of the ground, using the lower incisors"(Sagar & Harper 1964). Grazing by cattle in short herbage is less important than by sheep as theplantain plants react to being browsed by developing a ± appressed rosette habit and the cow's tonguecannot then pull the leaves off (Sagar & Harper 1964). In grassland communities, P.lanceolata is reduced by grazing more than by mowing (Tamm 1956). Unsurprisingly, P.lanceolata is, therefore, less common in pastures than in ungrazed, mown meadows. It is alsoless tolerant of trampling than P. major, although like it, the small seed does colonisecompacted dry soil very well (Cavers et al. 1980).
Cattle and sheep will remove flowering spikes of P. lanceolata and P. media (HoaryPlantain), but apparently not of P. major (Sagar & Harper 1964), obviously negativelyaffecting their seed production. A new crop of leaves is produced by plants in the autumn and the plantis wintergreen and frost resistant. Since these leaves have appreciable nutritive value, they serve asuseful winter fodder (Salisbury 1964).
Flowering reproduction
From April to September, or until the first frost or competition prevents anthesis. The rosette plantproduces one or more leafless flowering stems or peduncles (scapes) longer than the leaves, 10-60(-100)cm tall, strongly ribbed or furrowed and appressed-hairy. The terminal spike inflorescence is dense,ovoid to cylindrical in shape, 0.7-5.0 cm, lengthening up to 12 cm in fruit. The much reduced, denselypacked flowers are sessile and are subtended by dark green-black, ovate-acuminate bracts with membranouspoints, the lower bracts larger than the upper ones. The calyx of four sepals is c 4 mm in diameter,brownish-white or yellow with a prominent green or brownish midrib reaching the tip on each sepal. Thecorolla has a tube 1.5-3.0 mm long and is black, with lobes broad, lanceolate and acute. The fivestamens are yellowish-white, with slender filaments equalling or exceeding the corolla tube and theanthers are yellowish. The style is c 2 mm, the stigma filiform and feathery.
The flowers are protogynous, wind pollinated and are described as being 99% self-incompatible. They arevisited by insects collecting pollen, including the Honey Bee (Apis mellifera). The insects donot pollinate the flowers as they do not touch the stigmas of flowers that have no anthers exerted. Malesterility is not infrequent.
The fruit capsule is a dehiscent pyxidium, ie it splits around the equator (circ*mscissile) and openslike a box with a lid. It is 3-6 mm long and contains one or two seeds only, each 1.2 × 0.5 mm,half-spherical (Butcher 1961; Sagar & Harper 1964; Grime et al. 1988, 2007; Sell &Murrell 2009).
Seed production in natural habitats in Europe for most plants is rather small. In pastures grazed bysheep and cattle, seed production will usually be seriously curtailed by animals removing the floweringstems as well as the leaves. Salisbury (1964) estimated the normal level of annual production per plantwas probably around 2,500 seeds, but large plants developing more than 30 spikes could generate over10,000 seeds.
As in P. major, the seeds of P. lanceolata are not actively dispersed after the capsulesplits. They are coated with mucilage that swells when wet, making them adhesive and helping to dispersethem on soil particles and by attachment to hair, feathers and skin, again as happens in P. major(Sagar & Harper 1964). Cattle and birds, including pigeons, sparrows, bullfinch and greenfinch, alsofeed on the seed and probably are important dispersal agents (Salisbury 1964; Sagar & Harper 1964).Animals brushing against a ripe spike may scatter seeds from the open capsule, or scapes may rot at thebase and fall to the ground, shedding the seed. Some seed may be retained in open capsules for eight ormore months.
Seed matures rapidly after fertilisation takes place, in some situations after two or three weeks,depending upon the weather (Sagar & Harper 1964). As mentioned above, seed is long-persistent in thesoil seed bank, evidence suggesting up to 60 years (Holm et al. 1977). Germination normallyranges from 60 to 90% and seeds that have passed through birds have displayed 100% germination(Salisbury 1964). Germination occurs both in the autumn of production and in the following spring,contrasting with P. major and P. media, both of which germinate only in spring (Sagar& Harper 1964).
Vegetative reproduction
Individual plants can survive for twelve or more years and older plants tend to develop a thick rhizomewhich can sometimes produce side buds that develop new aerial shoots in the field leading to clumping ofthe species. In experimental studies, early autumn (September) planted P. lanceolata was able toregenerate buds from root fragments, but not from November or February plantings. It is not clear towhat extent vegetative reproduction occurs under field conditions, but it is probably of limited andminor importance in comparison with colonisation by seed (Sagar & Harper 1964).
Fossil history
Pollen grains are identifiable to species level and prove that P. lanceolata has been present inBritain since every zone of the Late Weichselian glacial onwards to the present day (Godwin 1975). Inlate glacial times, it may have been restricted to arctic-alpine grasslands, but as an indicator speciesof open ground, it has provided science with a dramatic picture of the extent and speed of deforestationfollowing the arrival of Neolithic man clearing the landscape for farming. P. lanceolata spreadwith the grass, hay and the herds at that time in European history, and it did the same when the NewWorld and other colonies were invaded and farming practices, fodder and seed-stocks exported (Holm etal. 1977).
Fermanagh occurrence
This is one of the most common and widespread species in the county; Ribwort Plantain occurs in 496Fermanagh tetrads, 94% of the VC total and it might even then be ignored and overlooked elsewherethrough it being considered by some recorders as an excessively common, weedy, perennial. Despite thisreservation, the species ranks as 5th equal along with Crataegus monogyna (Hawthorn) in terms oftetrad frequency in the VC.
British and Irish occurrence
P. lanceolata is common and ± ubiquitous throughout B & I, being absent only from the mostacidic soils, deep shade and wetlands. It achieves its greatest altitude (845 m) at Great Dun Fell inWestmorland (G.M. Kay, in: Preston et al. 2002).
European and world occurrence
While this very common species is now ubiquitous across the whole of Europe except the extreme north andis also considered native in N Africa and N & C Asia, its centre of origin remains unknown, althoughit may probably have been somewhere in C E & S Europe. P. lanceolata has been introduced totemperate areas across the whole world and may sensibly be referred to as 'cosmopolitan'.
Holm et al. (1977) stated in their book entitled The world's worst weeds, "On aworld basis, P. lanceolata is said to be one of the 12 most successful non-cultivated colonisingspecies." P. lanceolata and P. major are such successful worldwide weeds, since indisturbed agricultural areas where they enter a microsite, become established, spread out and form arosette to occupy valuable space, they can then manage to hold it against almost all other species. Inthis fashion, they have invaded many of the world's agricultural fields and have been reported as weedsfrom more than 50 countries in 26 crops that are as different as alfalfa, coffee, rice, cereals, onions,cotton, small fruits as well as temperate orchards and pastures. The most serious competition takesplace in cereals and in sugar beet fields and plantain seeds frequently contaminate small-seeded legumeseed-stocks. These two plantains are thus a constant source of difficulty and the farmer must spend agreat deal of time, energy and resources attempting to hold them back. However, Holm et al.(1977) concluded, "On account of their individual limited size, they are seldom reported as aprincipal weed for any particular crop."
Ribwort Plantain belongs to the southern temperate phytogeographical element, but is now circumpolarsouthern temperate in its distribution, although there is a wide gap between C Siberia and Alaska(Hultén & Fries 1986, Map 1724; Sell & Murrell 2009).
Medicinal and other uses
As mentioned under P. major (see this website), plantains, including P. lanceolata have areputation as vulnerary herbs in quickly staunching bleeding wounds. Usually it is found sufficient fora fresh leaf to be crushed and applied (or chewed) for enough of the healing chemicals to be released,but for severe wounds the whole plant has been boiled. Alternatively, an ointment has been manufacturedand applied. The herb has also been used to treat skin rashes, friction damage, burns, stings, varicoseveins and piles, and for drawing out pus from infected wounds. In Ireland, plantain juice was used forcoughs and the plant was used for very many other medicinal treatments including for gout, livertrouble, jaundice and even for treating sore eyes (Allen & Hatfield 2004).
Another quite different farm use made of Ribwort Plantain was as a means of testing the dryness of hay.The leaves of P. lanceolata dry more slowly than those of grass, so when making hay it wasimportant to make sure that the plantain leaves were dry, or else the resulting hay stack might overheatand perhaps even catch fire. This gave rise to English common names such as 'Fire grass' and 'Fireweed'(Vickery 2019).
Weed control
As with P. major, cutting off the leaf rosette of P. lanceolata at ground level merelystimulates the formation of a ring of lateral buds that re-establish the plant. Both species can be besteliminated from areas where they are unwanted by spraying with an herbicide, such as 2-4-D (Salisbury1964).
Names
No fewer than 46 English common names are listed by Grigson (1955, 1987). Vickery (2019) manages to beatthis total with his 52 names. Twelve of these names include or infer 'black' as a word element, aprobable reference to the black colour of the immature spike inflorescence or the more mature spike witha dusting of white from the stamens. Children, from time immemorial, have matched one 'Black Man', one'Soldier', one 'Hardhead', one 'Fighting co*ck', swinging its long peduncle against another until theweaker one loses its head. This game, which is widely played across Europe, has given rise to at least15 of the English common names listed by Grigson and Vickery, including 'Conkers', 'Fighting co*cks','Fox and geese', 'Hard Heads', 'Knights' and 'Kings'.
Threats
None.
Native, common and locally abundant. Suboceanic temperate.
1882; Stewart, S.A.; eastern end of Carrick Lough.
Throughout the year.
Growth form and preferred habitats
L. uniflora is a type of plant called an 'isoetid', a category named after the Quillwort genusIsoetes although, of course, while there is a superficial similarity in general growth form, iethe plant is a small, slow-growing, evergreen, amphibious aquatic with thick, stiff leaves or stems thatform basal rosettes and have a proportionally large below ground biomass. The plants of this groupingare not closely related in their taxonomy (eg Isoetes belongs to the spore-bearing Lycopsida).Incidentally, in relation to the mention of the genus Isoetes and the isoetids, the name isderived from the Greek 'isos', meaning 'equal' and 'etos', 'a year' or 'the year', combined to mean'equalling one year', but really referring to 'evergreen' or 'green throughout' (Johnson & Smith1946; Gilbert-Carter 1964; Gledhill 1985).
In the case of Littorella uniflora, the plant has a short, white, erect rootstock that sends offlong white roots. The leaves are bright green, 2.0-10.0 cm long, nearly round in section and incross-section show numerous lacunae. L. uniflora is readily distinguished from othersimilar-looking aquatics such as Isoetes lacustris (Quillwort), Lobeliadortmanna (Water Lobelia) and Subularia aquatica (Awlwort) by its developingstoloniferous, carpet-forming colonies, rather than possessing separate, discrete rosettes as do theseother isoetid genera that occur in similar lakeshore and other aquatic habitats including rivers,streams, reservoirs, ponds and dune-slacks (Preston & Croft 1997).
The terrestrial form of Shoreweed is produced by germination from a large, long-persistent seed-bankwhenever water bodies are drawn down sufficiently in suitably dry summers to expose a broad expanse ofbare, often muddy shore available for colonisation. In addition to this annual form of growth, however,Shoreweed is a perennial species and has great powers of vegetative spread, its abundant stolons,rooting and producing rosettes of leaves at nodes enabling it to form dense, turf-like colonies,especially when it grows permanently submerged in relatively shallow, oligotrophic or mesotrophicwaters. L. uniflora can grow on a very wide range of waterside or bottom substrates includingstones, gravel, sand, peat, marsh and soft silty mud (Preston & Croft 1997; Sell & Murrell2009). The established strategy of the species is categorised as 'SR/CSR', ie intermediate between aStress tolerant Ruderal and a balance of all three strategies, Competitor, Stress tolerator and Ruderal(Grime et al. 1988, 2007).
In Britain, some of its sites, including winter-flooded depressions on heaths and sea-cliffs, dry outcompletely in summer. In deeper water, in sites that remain wet all year, L. uniflora grows inopen swampy conditions dominated or characterised by Eleocharis palustris (Common Spike-rush) andEquisetum fluviatile (Water Horsetail). Here it appears both on exposed shorelines, and belowthis it forms a carpet on lake bottoms down to depths of around 3 to 4 m, where it is often accompaniedby Isoetes lacustris and Lobelia dortmanna (Preston & Croft 1997).
There are interesting variations in tolerances across B & I. In Scotland and Ireland, L.uniflora is most frequent in regions with base-poor acidic geology and waters, but it can alsogrow in definite base-rich habitats such as clear limestone lakes, streams and turloughs (ie vanishinglimestone lakes that drain vertically into subterranean rivers and streams). In the Burren, Co Clare(H9), it can even grow in pools perched on limestone pavement, as for example is the case on the islandof Inishmore (Webb & Scannell 1983). In contrast, in SE England, L. uniflora is completelyconfined to acidic habitats (Preston & Croft 1997).
Flowering reproduction
Shoreweed flowers from June to August and usually does so only when in dry seasons it becomes emersed, iethe flowering stem protrudes above the water surface. Totally submerged plants always fail to flower.The species is monoecious (ie separate male and female flowers are borne on the same plant). Both kindsof flower are very simple, greatly reduced and are borne together on a flowering stem, c 10 cm,generally shorter than the leaves. The flowers are very definitely wind-pollinated, the solitaryterminal male flower being stalked and the one, two or a few female subsessile ones, arranged below atthe base of the male scape, hidden by the surrounding leaf bases. One or more bracts separate the malefrom the female flowers. Each plant can produce three, four or more flowering stems.
The flowers consist of four green sepals with scarious margins and four white petals, also scarious ontheir margins; the males have four stamens with very extended white filaments 10-20 mm long; the fewfemale flowers are uni-ovulate and have one thread-like style, c 10 mm long, bearing a terminal, linearstigma. The fruit is a single-seeded, ovoid or cylindrical, indehiscent (non-splitting) capsule or nut,c 3 mm, its surface keeled, honeycombed and black (Hutchinson 1945; Melderis & Bangerter 1955;Butcher 1961; Sell & Murrell 2009).
There is convincing evidence from isolated ponds and marshes that fruit dispersal in L. uniflorainvolves transport overland by ducks (Stelfox 1922; Ridley 1930, p. 547). Most likely the fruit, whichis not particularly large or bulky, has been conveyed attached in mud on the feet or feathers of ducks.L. uniflora has similarly reached remote oceanic island groups, such as the Azores (Ridley 1930,p. 550). Some seed is not or is hardly dispersed at all: Cook (1987 (b)) reported that inLittorella (no particular species stipulated, but read in context, this could refer to allspecies of the genus), "the seeds germinate while still attached to the mother plant".
Most published ecological treatments suggest that seed of L. uniflora is long persistent in thesoil, ie more than five years and perhaps for decades (eg Grime et al. 1988, 2007; M.J.Wiggington, in: Preston et al. 2002), although some doubt on this was indicated in the latterreference. However, the published Survey of soil seed banks in NW Europe contained just oneestimate of seed longevity, which concluded it was transient, ie seed persisting for less than one year(Thompson et al. 1997).
Vegetative reproduction
It is very likely that L. uniflora depends for local survival on its very effective vegetativereproduction more than on seed production, which only occurs whenever water levels drop and marginalground becomes exposed for a sufficient length of time to allow anthesis, pollination and fertilisation.However, transport of seed and vegetative fragments are both likely to be very significant in achievingspecies dispersal to fresh habitats, with seed being especially so when jump dispersal to remote orunconnected water bodies is concerned (Ridley 1930).
Fossil record
The pollen of L. uniflora is as recognisable as the one-seeded, hard, indehiscent 'bony' fruitsand fossils of both contribute substantially to the historical evidence. The fossil record in B & Igoes right back to the Cromer Forest Bed series and there are records from every sub-stage of thesubsequent Hoxnian interglacial. In the Ipswichian, it is only recorded from sub-stage IV, but in allthe Flandrian zones there are pollen records with a thin scatter of fruit records. There are alsoabundant records from some of the glacial stages, many of the fruit records being from Ireland. It isevident that L. uniflora was prevalent in the shallow lakes of the Weichselian glacial period andthat it has persisted in suitable situations ever since then (Godwin 1975).
Fermanagh occurrence
On the shores of many lakes, especially around mesotrophic to more eutrophic Lough Erne, this perennialgrows abundantly in shallow water and produces its greatly reduced unisexual flowers when stretches ofbare gravelly, sandy, muddy or peaty lakeshores become exposed in summer. Despite its frequency onlakeshores and especially so around the larger loughs in the VC, rather surprisingly L. unifloradoes not appear to occur on any river sites in Fermanagh. It has been recorded in 132 tetrads, 25% ofthose in the VC. As the tetrad map shows, it is most frequently found around the larger lakes and israther rare beyond them.
British and Irish occurrence
L. uniflora has a predominantly northern and western distribution in B & I, where it is oftenabundant in suitable wetland habitats (New Atlas). In SE England, it is rare and confined toacidic habitats, but in Ireland and Scotland, where it is much more frequent, widespread and locallyabundant, it displays a much wider ecological range. The ability of L. uniflora to toleratemesotrophic water, and even some degree of eutrophication, has allowed it to survive in areas whereother isoetids have declined.
In S England and S Ireland, L. uniflora lost many suitable habitats due to drainage operationsbefore 1930 and this progressive loss has continued to the present in S & E England where thespecies is now rare and widely scattered. Apart from drainage, some losses appear to have resulted fromhabitat change due to overgrowth by tall, rank vegetation (Preston & Croft 1997; M.J. Wigginton, in:Preston et al. 2002).
European and world occurrence
L. uniflora belongs to the suboceanic temperate phytogeographical element and occurs widelyscattered in W, C & N Europe, with diminishing frequency eastwards and northwards, although itreaches 68°N in the Arctic. It is also present on Iceland, the Azores and on the tip of N Africa.Presence becomes more scattered in the south of Europe, although it reaches W Spain and Portugal. Thespecies is sparse or absent from most of the Mediterranean basin, but is present in N Italy and inSardinia (Godwin 1975; Clapham et al. 1987; Hultén & Fries 1986, Map 1729; Sell & Murrell2009).
Names
The genus name 'Littorella' is derived from the Latin 'litus' or 'littus' meaning 'shore' or 'of theshore', referring to the habitat where it is most visible (Johnson & Smith 1946; Gilbert-Carter1964). The Latin specific epithet 'uniflora' means 'one flowered', a reference to the solitary terminalmale flower which is so conspicuous (Johnson & Smith 1946).
The English common name 'Shoreweed', or older and better 'Plantain Shoreweed' (Prior 1879) is anotherperfect example of an invented 'book-name', of no folk significance, but in this case, a useful reminderof the habitat (Britten & Holland 1886).
Threats
Experience in Fermanagh shows L. uniflora can tolerate a degree of eutrophication, provided it isnot excessive. Acidification of upland lakes due to forestry run-off could see it decline locally.
Introduced, established garden escape, only occasional, but very under-recorded.
July 1987; Northridge, R.H.; along embankment from West Bridge, Enniskillen Town to the Scillies River.
February and July.
There are only 21 records in the Fermanagh Flora Database for this large, very commonly grown, deciduousgarden shrub, a native of China (E & W Sichuan and W Hubei), that grows in its native territory onshingle by rivers and streams, on roadside cliffs and in scrub at 1,300-2,600 m (Phillips & Rix1989). It was introduced to gardens in B & I in the 1890s and very quickly became a populardecorative subject on account of its ease of growth (it thrives in almost any soil and revels in fullsun), range of inflorescence colour variation (from a strong red to rose-purple to violet-purple orwhite) and flowering period (July to September), but especially for the honey fragrance of its longracemes that regularly attract large numbers of butterflies in late summer (Anon. 1981). However, inaddition, thanks to its large size (2.5 m or more in both height and width), prolific flowering andproduction of abundant windborne seeds, Butterfly-bush frequently jumps the wall or fence, escapes intonon-garden settings and is now commonly naturalised in a wide range of habitats. It is very resilient,being hardy to -15°C and can still achieve flowering in one growth season even if cut to the ground byfrost (Phillips & Rix 1989).
Fermanagh occurrence
B. davidii has been found in twelve scattered Fermanagh tetrads located mainly in or nearEnniskillen, which is by far the largest conurbation in the county. However, the low number of recordslargely reflects a lack of recorder interest in the occurrence of the plant, rather than any degree oflocal extreme rarity! Until the 1980s, the common attitude of plant recorders in Ireland had been formany decades that native species are the major priority and that garden escapes, unless they becomeinvasive, are of little or no significance and therefore not worthy of much notice.
All 21 Fermanagh records were made by RHN, HJN and I. McNeill since 1987, 17 of them since 2000. Inreality, here in Fermanagh, B. davidii is just as frequent as anywhere else in these islands.Seedlings, saplings and mature plants of Butterfly-bush are now common all over towns and cities acrossmost of B & I. It grows in dry, disturbed, often stony conditions in waste ground, disused quarries(where dumping of garden waste sometimes takes place), along lakeshore embankments, around derelictcottages, on old stone or brick walls, in neglected gutters and in chimneys – usually in more urbansettings but also in villages such as Derrygonnelly, Clabby and Tamlaght.
The ballast beds of railway tracks and areas around sidings are one of the more recent habitats thatB. davidii has successfully colonised and overrun. Old railway photographs taken in England showthat the shrub did not become really abundant in this type of habitat until the 1980s, and the fulleconomic effects of its rampant colonisation have yet to be felt and measured since the damage inflictedby the roots of Butterfly-bush, growing through and destroying brickwork, has not been widelyappreciated. Herbicide control is very expensive and spraying needs to be regularly repeated to get ontop of even a moderate scale of infestation by this prolifically fecund, highly resilient shrub.
Should any strain of B. davidii evolve to become herbicide resistant, there would be very serioustrouble controlling this extremely vigorous woody weed (Stace & Crawley 2015).
British and Irish occurrence
Introduced in late Victorian times to B & I gardens, B. davidii was appearing naturalisedbeyond garden confines by 1922 in Merioneth (VC 48) and by 1927 in Middlesex (VC 21). The 1962 BSBIAtlas showed it was locally quite well naturalised in S England and coastal Wales, but only verythinly scattered elsewhere across N England, Scotland and Ireland (Perring & Walters 1962).
The New Atlas hectad map indicates just how rapidly and much more widely it has spread insubsequent decades, becoming common throughout all of lowland England and Wales as far north as a linebetween Lancaster and Hull, and scattered north to reappear in quantity around the three major Scottishconurbations of Glasgow, Edinburgh and Inverness. In Ireland, it has also greatly increased itspresence, especially around Belfast and Dublin, but thinly and widely scattered right across the islandto Galway and most west coast VCs (Preston et al. 2002).
Names
The genus name 'Buddleja' is named in honour of the Reverend Adam Buddle (1660-1715), English botanistand vicar of Farmbridge in Essex. The Latinised specific epithet 'davidii' is a name honouring theFrench missionary and plant explorer who worked in China, Father Armand David (Stearn 1992).
Native, common, locally abundant. European temperate.
1884; Barrington, R.M.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Ash is the only member of the Oleaceae, the Olive family, which penetrates this far north in Europe andit is the only wind-pollinated species in the family, all the other members being insect-pollinated(Milner 1992). Ash is very prevalent in semi-natural woods, timber plantations (chiefly for poles orstaves), secondary scrub and managed hedgerows. It is also a pioneer colonist on cliffs, riverbanks,lakeshores, roadside and field hedgerows and as a weed in old quarries. It occurs on all types of soilin Fermanagh except the most acid and peaty, the permanently water-logged, or the extremely shallow orvery steeply sloping and unstable ones. Wardle (1961) reckoned Ash is usually absent where the pH of thesurface soil goes lower than 4.2.
Ash trees are never very tall in Fermanagh, often only around 10 m, occasionally up to 15 m in height andonly rarely in more nearly optimal growing conditions are they taller than this. Despite this lack ofstature, Ash still forms the dominant tree in old valley woods over limestone, eg around the Knockmorearea. It definitely prefers and is most abundant in areas with calcareous rocks and near neutral soils.Having said this, young plants in particular often produce pale yellow foliage and appear to suffer fromlime chlorosis when growing on basic soils.
In areas of more acid geology and soils, Ash is more frequent or almost restricted to lower slopes ofvalleys, hollows and stream banks, where percolating ground water is sufficiently enriched with nutrientbases to support the tree and allow it to compete. Previously, on the better loam soils in uplandsheltered valleys and on lake islands, Ash would have shared the canopy of mixed deciduous woodland withQuercus petraea (Sessile Oak). Now, however, apart from the odd inaccessible scrap of such woodsand on a few protected estate and NR examples, almost all of these semi-natural woodland stands havebeen felled and destroyed to create additional sheep pasture.
On cliffs, rocks, or on screes that are sufficiently stable to permit the growth of any woodyspecies, the young soils present in crevices are generally able to supply the Ash tree's needsirrespective of the rock chemistry (Wardle 1961). On limestone scree, Ash seedlings can withstandlimited down-slope movement better than Hazel (Corylus avellana), but the saplings and eventualtrees often end up with very strongly curved trunks!
F. excelsior also competes very well on winter-wet or seasonally water-logged, often clay soilsaround many lake shores, as well as on better drained soils subject to intermittent flooding alongriverbanks. This is because the young seedling requires only quite a shallow depth of well-drained soilon which to establish itself in wet ground. Subsequent growth in water-logged habitats is poor, but thespecies will respond rapidly to any improvement in drainage. It has been suggested that the poor growthof Ash on water-logged sites may paradoxically be due to periodic water shortage in the plant, since itsroots are unable to penetrate deeply in such soils and suffer when even a brief drought occurs (Wardle1961).
Fossil record and early Ash woodland colonisation
The fossil pollen record of Fraxinus excelsior is virtually restricted to earlier interglacialsgoing back to the Pastonian, plus current Flandrian periods (Godwin 1975). As the tree is recognised asbeing frost-sensitive, it is not really surprising that it is totally absent from all the interveningglacial stages. The reappearance of Ash pollen and macrofossil wood in the Flandrian interglacial was,in Godwin's opinion, rather tardy. It did not appear at all in B & I until zone V, when just tworecords were found. Ash fossils make a vast leap forward in frequency and abundance in zone VIIa (theAtlantic period) 8,000-5,000 BP, the Holocene climatic optimum for both warmth and moisture. Temperaturethen is reckoned to have been warmer than today. Dates quoted for this period of climatic optimum varyin almost every reference examined, some making the period a thousand years later and shorter (ie7,000-5,000 BP). The current author (RSF) is merely a lay reporter, not a palaeo-climatologist and,therefore, cannot comment further. The preceding period, known as the Boreal, had an overall temperaturesimilar to that of today.
The increases that F. excelsior made in zone VIIa became extended and sustained in zones VIIb (theSub-boreal) and VIII, the expansion in site numbers being accompanied by a substantial increase in theproportion of total tree pollen contributed by Ash. The low pollen productivity characteristic of Ashmeans that in effect abundance of the tree increased even more than initially appears the case throughthese time periods (Godwin 1975). Having said this, the fossil pollen figures imply that Ash waswidespread in the Atlantic period (Holocene climatic optimum) wildwood, but it was only locally commonat that time (Rackham 1980, p. 204).
The fossil evidence strongly suggests that the expansion of Ash in the later stages of the Flandrian(called the 'Littletonian' in Ireland), primarily reflects human clearance of the pre-existing climaxforest (which included Elm (Ulmus spp.)). The Ash trees' strong invasive response to clearancesand deforestation ± coincided with, or shortly followed, the major decline of Elm pollen, which probablyinvolved the effects of both plant disease and human felling operations, that opened up growingconditions favourable to the Ash.
Why it should have been Ash that profited from the clearing of primary forest is not clear. It issomewhat unlikely that Ash was deliberately favoured by early farmers, but the species is a goodcoloniser and often forms secondary woodland, especially on chalk and limestone soils. Since it ispalatable to stock, it is also possible that it was used as an alternative fodder in hedges andwood-pasture to replace the missing Elm, the branches of which, prior to its decline, were an importantsource of animal food (Rackham 1980). Hay as storage fodder made from grass did not exist in agricultureuntil long after this prehistoric period.
Fermanagh occurrence
Definitely the most frequent and often the dominant tree in limestone districts, Ash ranks second only toAlder (Alnus glutinosa) among the most commonly recorded tree species in Fermanagh – provided weset aside the Hawthorn's claim to the ranking on account of it being a hedgerow shrub, ie defined asmulti-stemmed and lacking a definite bole (Mitchell 1974). Either way, Ash, which after all can bepresent in shrub form too, is present in 478 Fermanagh tetrads, 90.5% of the VC total. It is thus one ofour top two or three tall woody plants (we must not totally forget about the heathers!) in terms of bothabundance and distribution. In Britain, by comparison, Ash is considered the fourth commonest woodlandtree species (Milner 1992).
The most impressive Ash woodland in Fermanagh is at the bottom of Hanging Rock, a 50 m high limestonecliff below the grazed limestone uplands of the Marlbank plateau near Florencecourt. The Hanging RockNature Reserve sits between the cliff base and a narrow band of agricultural fields above Lower LoughMacnean. The Ash woodland is damp in the oceanic climate of this part of the world and it has a richground flora and an extensive list of lichens, both of which indicate it is ancient woodland. Thelimestone soil is shallow and rocky. The wood is characterised by numerous old, fallen trees withsaplings growing up through them. The secondary canopy is composed of Hazel, Holly (Ilexaquifolium) and Bramble (Rubus spp.).
Seed production, dispersal and habitat colonisation
Winged seeds are produced most, but not every year, often appearing in large numbers (100,000 ormore/tree), enabling Ash to disperse very effectively. Thus it frequently invades derelict sites, eg inold quarries or on waste-ground as a pioneer scrub species – it is virtually a woody weed in such sites.Ash is planted or arrives self-sown in many, possibly most hedgerows in Fermanagh, both along fieldboundaries and on roadsides. As a pioneer, F. excelsior seedlings are usually in competition withgrasses and they are only able to establish where there are gaps in the sward (Wardle 1961). Theestablished strategy of the species is categorised as C/SC, ie intermediate between Competitor andStress-tolerant Competitor (Grime et al. 1988, 2007).
Seedling growth is rather slow at first, but once the individual is well established it becomes vigorous.In good growing conditions of full light or semi-shade, moisture and fertile soil young plants canachieve elongation growth of 50-100 cm or more per year from the second year onwards, emulating suckersof Elm (Ulmus spp.) and Wild Cherry (Prunus avium) (Oliver 1998). Young seedlings andnewly opened buds, both floral and foliage, are susceptible to late frosts and are often killed in moreexposed sites in April and May.
Stress tolerances and growth requirements
Growth and reproductive success of F. excelsior is also very dependent upon moisture relations,the shoots being very sensitive to any inadequacy of water supply due to drought or drying winds,particularly during the months of May and June when growth and reproduction are restarting. Rathersurprisingly, this even applies to plants in sites that are water-logged in the winter months. While Ashrootlets are always concentrated in the top 5 cm of soil, lateral roots may be sparsely distributed togreat depths, reaching down as far as the permanent water table. Nevertheless, larger plants displaystrong powers of vegetative recovery from the negative and exhausting effects of frost and drought(Wardle 1961). Ash plants therefore need some shade, shelter and relatively moist, but not too wet soilconditions in their early years but, thereafter, the plants are very much more those of alight-demanding species (Hadfield 1957).
In the full sun conditions typical of a managed linear hedgerow setting, Ash grows very much more rapidlythan it does in woodland shade. It is said to have the highest nutrient uptake of any native tree(Milner 1992, p. 10). The forester and tree identification expert, Alan Mitchell, who had a poor opinionof the species, referred to it disparagingly as, "a gross feeder", ie behaving like StingingNettle (Urtica dioica) with respect to nutrient demand (Mitchell 1996). On the other hand, Ash isvery tolerant of poor soils and although often severely stunted by nutrient stress, it can survive andgrow where few other tree species can, including in cracks and ledges on rocky crags. Occasionally iteven manages to grow on, or through, old stone or brick walls, where conditions must obviously be veryunfavourable yet clearly lie within the wide tolerances of the species.
The efficient mineral uptake by F. excelsior is reflected in the very rapid decay of its leaflitter, which in damp woods disappears within nine months, ie by the following June (Wardle 1961). Inthe field layer of woodland, Ash seedlings are very shade-tolerant and can persist in almost suspendedanimation for years under heavy canopy shade, effective increase in their height only becoming possiblewhenever they experience conditions of almost full sun. When a gap eventually appears in the woodlandcanopy, they are released from domination and shoot up in height, competing fiercely until one or moremanages to fill the ecological space temporarily available (Wardle 1961; Rackham 2014).
Canopy opening, leaf fall and their incorporation in soil
While the familiar folk saying has it, 'Oak before Ash, splash, splash, splash. Ash before Oak, soak,soak, soak', in reality F. excelsior is almost always the last native tree to come into leaf,usually in mid- to late-May, but occasionally some trees in a cool season are still bare-branched, withtheir distinctive black buds intact in early June, eg in 2002 (RSF, personal observation). The delay inleafing is undoubtedly due to the vulnerability of the leaves and new shoot growth to attack by lateseason frost. The natural distribution of F. excelsior in W Europe is undoubtedly defined by itsintolerance of winter cold, late spring frosts and any drought in May and June when the tree is mostactively producing its new season's growth and beginning reproduction.
The leaf canopy of Ash casts a relatively light shade when compared with Oak (Quercus spp.), oreven with Rowan (Sorbus acuparia), or other secondary canopy species of the typical mixeddeciduous woodland in B & I and, as a result of this, a rich field and ground flora is able to growbeneath it (Kelly & Kirby 1982). The light degree of shading by Ash is partly due to the fact thatthe species has opposite leaves and the inevitable opposite branching pattern which leads to thedevelopment of a remarkably open crown (Edlin 1964).
Ash is also one of the first native trees to drop its leaves, which either turn yellow or fall green,typically in early October. Although the leaves are pinnately-lobed and are composed of up to 13leaflets, they detach and fall in one piece. In the past the leaf litter attracted earthworms whichdragged them down into their burrows leaving the stalks in view, sticking out of the ground at oddangles. The current author (RSF) has not seen evidence of this happening for many years since earthwormshave become a rarity, possibly due to the arrival in NI of predatory New Zealand Flatworms, but perhapsalso as a consequence of pollution associated with the excessive use of agrochemicals and farm slurry.
Flowering reproduction
Flowering generally begins when the tree is between 30 and 40 years old (Wardle 1961). The flowers aresimple, reduced structures which open in mid-April just before the leaf canopy does. The denselyclustered flowers may be either separate male and female or perfect (ie bisexual or hermaphrodite). Thesame tree can bear flowers of all three kinds, although in general the species is dioecious, havingseparate male and female trees and, as such, it is best described as partially dioecious (Edlin 1964;Grime et al. 1988, 2007). Although whole trees tend towards one sex or the other, the sex ofindividual tree is not clear cut and they can change the sex balance from year to year (Rackham 2014).
The flowers are as simple as flowers can be: each consists of two stamens and/or an ovary, without sepalsor petals, and producing no nectar. They do tend to be purplish in colour and are borne in axillarypanicles of more than 100. Pollination is mainly by wind, but for a few days sufficient pollen isproduced to attract honey bees and this almost certainly results in some degree of cross-pollinationbetween flowers and trees (Rackham 2014). The fruit is an elongated, winged, samara, referred to in bulkas 'Ash keys', each usually containing just a single seed and borne in dense bunches from the beginningof July onwards.
Viable seed samara are blown from the tree from September onwards, but many continue to remain on theparent tree until the following spring. Dispersal distances of up to 200 m or more by winter storm windscarrying samaras have been recorded (Ridley 1930; Edlin 1964; Rackham 2014). However, in conditions ofwoodland shelter, obviously most keys simply drop immediately beneath the parent tree.
When released from the tree in its samara fruit, the embryo seed is immature and requires a prolongedperiod of dormancy in order to mature and become capable of germination. Ash seed also has a chillingrequirement and needs a stratification of two winter's duration. Together these dual delays mean thatgermination usually takes place in the spring of the second year after shedding (Wardle 1961; Gardner1977). During this long delay, a great many dormant seeds are eaten by mice, voles, birds andcaterpillars (Milner 1992). Under natural conditions, a small number of seedlings can be expected toappear every spring and some seed may remain viable and hidden from predators for up to six years.Seedlings are also subject to similar animal predation, including by rabbits; damping off is a verycommon fate if the adjacent vegetation and the soil are too moist. Saplings that suffer the deleteriouseffects of browsing, however, often demonstrate strong powers of recovery (Wardle 1961).
Vegetative reproduction
Although the plant responds superbly well to coppice cutting by re-sprouting from the basal crown, therenormally is not any really effective mode of vegetative reproduction in F. excelsior andtherefore no clonal growth. However, when the basal root crown of a tree becomes decadent due to old ageor unfavourable changes in habitat conditions, epicormic shoots often spring from the lower part of thetree trunk. True suckering has been reported, but it must be of rare occurrence (C.D. Pigott, in: Wardle1961).
Toxicity and herbivory
Sheep and goats are fond of browsing the tree foliage and in past times young shoots of coppice Ash werecollected and used as animal fodder (Milner 1992). The tree contains poisonous principles, however,including a glycoside called 'aesculin' which also occurs in Aesculus species. Cattle feeding onrecently fallen green leaves may suffer from an impaction of the rumen, a condition formerly known inthe English Midlands as 'wood evil', the symptoms of which include drowsiness, abdominal pain andconstipation. The condition also results in reduced milk yields. There are no reports of humanpoisoning, but handling the foliage can cause dermatitis (Cooper & Johnson 1998).
A local study in England found young, fast-growing sapling Ash with pale-green leaflets were oftenheavily grazed by deer, hares and especially rabbits, that ate every leaf and gnawed the bark, sometimesring-barking and killing plants. Slower-growing varieties in the same area, with more leathery, darkerleaflets were less often savaged by herbivores and also seemed to be less susceptible to being burnt andblackened by late spring and early autumn frosts. These observations suggest that penalties for vigourmay include both greater palatability and frost-tenderness (Oliver 1998).
Considering that Ash is a long-resident native tree, only a moderate number of invertebrates feed on it,a total of 68 species, approximately half of them macro-Lepidoptera and Heteroptera (Kennedy &Southwood 1984). Of these, 29 are specific to Ash (Rackham 2014). The database of the Biological RecordsCentre has a list of eleven insect and mite species for which Ash is the obligate food-plant. Presumablythe relatively low number of feeding invertebrates is a direct consequence of the chemical defences ofthe species.
Epiphytes
On the other hand, Ash bark can be either acidic or alkaline and a total of 230 species of epiphyticlichens have been recorded on the tree, making it second only to oak in this respect (F. Rose, in:Morris & Perring 1974, p. 256, Table 3; Gilbert 2000).
Uses
Reproduction in Ash is virtually entirely by seed but the mature tree also displays remarkable powers ofvegetative regrowth when cut to the ground. Ash responds well to being cut as coppice in this way,rapidly re-sprouting numerous straight poles from the cut crown that traditionally have been used aswalking sticks and as staves for all kinds of building and support functions. "Throughout historyAsh has generally been the second commonest recorded timber tree." (Rackham 1980). The uses go backwell into pre-history, eg Ash and Hazel were much used in Neolithic trackways in the Somerset Levels(Rackham 1980).
Top quality Ash timber has great impact strength and elasticity which in past times made staves bothsuitable as weapons themselves, or as shafts for spears. This use is reflected in the English commonname, 'Ash', which is derived from the Anglo-Saxon 'aesc', a poetic word meaning, 'a spear' (Milner1992). The Anglo-Saxon 'aesc' could also mean, however, 'a vessel', meaning a boat, since Ash timber wasalso much used in boat building, although it is not suitable for the outer planking as it rots quicklyin water (Prior 1879; Rackham 1980).
Ash wood was also used for the handles of more peaceful woodworking and gardening tools, and for theframes and shafts of vehicles. This latter use continued from early chariots to the Morris Travellercar, the Morgan sports-car, and even to aircraft construction.
Ash has the best burning qualities of any native tree timber, even when it is green or wet and Milner(1992, p. 101) quotes an interesting poem to the effect. There is a substantial body of folklore,superstition, divination and magic associated with Ash. It was one of the Irish 8th century, 'Nobles ofthe wood', and several authors go into the matter in more detail than there is room to include here (egGrigson 1955, 1987; Milner 1992; Nelson & Walsh 1993; Vickery 1995).
Ash was also held to have herbal medicinal qualities and these and numerous other uses of the timber aredescribed in detail by Grieve (1931).
Lifespan and senescence
Longevity of the standard (ie un-pollarded) Ash tree stretches to around 250 years at a maximum, withmost trees less than 200 years (Rackham 1980; Nelson & Walsh 1993; Mitchell 1996; Thomas 2000).However, observation regularly confirms that rather stunted individuals are frequently overcome byevergreen Ivy (Hedera helix), which the light open canopy of the Ash tree is unable to shade out.These festooned trees readily fall victim to wind throw, probably when only about half their potentialage or less. Old, senescent trees tend to drop major boughs as well as the perfectly normal small branchshedding from the crown that continues throughout the life of the individual specimen. These older treesthen often begin to form basal epicormic shoots (ie dormant buds on the trunk producing new shoots,literally, 'upon the bark'). These vigorous shoots are almost, but not quite, suckering, invariablyattached right at the base of the original tree, rather than spreading out around it underground liketrue suckers.
On the other hand, Ash stools and pollards, formed by regularly cutback either to ground level or trunkheight of around 2 m, are capable of living indefinitely, being rejuvenated at every three or four yearcut-down. Very old stools in ancient English woods can measure up to 3 m in diameter and most stools inthis type of woodland are 1.5 m in diameter (Rackham 1980).
Declining tree size
In the not quite so distant past, there were some enormous historic elderly Ash trees in Ireland,including the famous, 'Great Ash of Leix', the circumference of which was measured in 1792 at one foot(30 cm) from the ground to be 40 feet 6 inches (equivalent to 12.46 m) (Nelson & Walsh 1993).Mitchell (1996) lists modern so called 'champion trees', but the largest extant Irish Ash (theBallynatray Ash, Youghal, Co Cork) has a girth of only 8.5 m (26 ft). It is very misshapen, has split ordecayed and is now represented by two rugged, hollow trucks. It does not appear to have been pollarded,which can prolong the lifespan (eg the Glen Lyon Ash in Perthshire, Scotland (Stokes & Rodger 2004))and is estimated to be perhaps 400 years old. Two hundred years is a very good age for an Ash, while 300years would be exceptional (Fennell 2013).
This observation of surviving old tress supports something also noticed with other species, such asAlder, that suggests, for unknown reasons, trees nowadays do not seem to achieve anything like thedimensions they previously did in historical or pre-historic times. Evidence for this arises from bothold published records (which could be erroneous) and from preserved tree trunks recovered from bogs thatunequivocally prove the point.
British and Irish status and occurrence
In the British Isles, F. excelsior is native and common throughout England, Wales and Ireland,although it becomes scarce or absent in more exposed coastal sites in W Ireland. In Scotland, it isregarded as native on limestone but here, as elsewhere, it is widely planted and it becomes rarer as onetravels northwards (Preston et al. 2002).
Ash disease
Ash dieback first came to public notice in B & I in October 2012 when it was reckoned to be both'new' and 'terrible' (Rackham 2014). The disease was not new; it had been making its way across Europefor the previous 20 years and should have been noted by those in charge of tree health attendingInternational conferences on the subject. However, B & I representatives did not attend thesemeetings and they did not take heed of the impending arrival of the disease organism on these shores.The 'new' Ash disease is caused by a microscopic fungus called Chalara fraxinea. It invades theleaves and damages the tree by manufacturing a chemical called 'viridol' that is extremely toxic. Insummer, the fungus in the leaves is in an asexual phase of its life-cycle and produces spores calledconidia which form sticky masses and that spread with rain to infect the leaves of adjacent trees.
Like many other fungi, the organism has an alternation of generations and in autumn it turns into itssexual phase, a white cup-fungus fruit body about 3 mm across, called Hymenoscyphus pseudoalbidusor H. fraxineus that appears on the midrib of the ash leaves. This midrib or rachis fallsseparately from the leaflets that it normally bears. When it reaches the woodland floor, the cups of thefungus puff out little clouds of 'ascospores'; it is a type of fungus called an 'Ascomycete'. A singlemidrib may bear 20 cup-fungi each releasing 1,500 per hour for up to two weeks, ie several millionspores from each Ash leaf! The spores are extremely light and float on the slightest breeze. They cantravel to start infections on distant trees and when new leaves appear in the spring, the fungus revertsto the infectious Chalara phase, penetrates the tissues and continues the cycle within the tree.
Experience in continental Europe, being replicated in the UK in 2012, indicates that 'Chalara Ashdieback', as it is now referred to, can kill young and coppiced ash trees quite quickly. However, oldertrees are able to resist it for some time until prolonged exposure, or another pest or pathogen such asArmillaria (Honey Fungus), attacking them in their weakened state, eventually causes them tosuccumb (Rackham 2014). Needs a reference?
Infected trees can be distinguished from a distance by the thinning leaves on the crown and sidebranches, and bare, dead or dying, defoliated uppermost branches. However, shoot death and dieback canhave other causes apart from Chalara attack and fruit hanging on the tree in autumn and wintercan be mistaken for the blackened, dead leaves characteristic of the disease. Closer inspection showsfour characteristics of the Chalara dieback: wilted, rigid, blackish-brown leaves from July toSeptember; a blackish strip of cankerous bark (lesions) spreading up and down the twig; long, narrowdark cankers (lesions) where twigs join a larger branch or trunk with a dead twig in the middle; long,re-growth (epicormic) shoots springing from the branches, leading to bunched foliage, a phenomenon knownas 'proliferation', giving a 'witches'-broom' appearance to the tree foliage (Rackham 2014;https://www.forestresearch.gov.uk/tools-and-resources/fthr/pest-and-disease-resources/ash-dieback-hymenoscyphus-fraxineus/,accessed 12 August 2022).
Chalara does not always kill infected trees. There is some genetically based resistance to thedisease. Studies in Finland and Czechoslovakia suggest that some infected trees can recover, whileothers get worse from year to year and eventually die (Rackham 2014, p. 123). The dying process may takemore than 20 years, depending on the age of the tree when it is infected. Ash disease was first noticedin or near Latvia around 1990 and has apparently spread across Europe from there. H.pseudoalbidus could be an introduction to Europe from E Asia, possibly from China or Japan, orit could be a newly arisen mutation or hybridisation of an existing fungus (Rackham 2014). Does thatneed a reference or is it covered by Rackham 2014?
There is currently no cure for Chalara Ash dieback, and no clear method for stopping its spread.Therefore, the aim of management, as outlined in the National Chalara Management Plan, should beto slow the spread,minimise the impact of the disease and preserve as many Chalara-tolerantash trees as possible.
Trees in areas with high levels of public access need to be monitored carefully for risks to publicsafety, and some felling or pruning of dead or dying trees is advisable if risk assessments show theyare a hazard.When assessing trees' health, it is important to look for signs oflesions(cankers)or Honey Fungus (Armillaria) near the base of the trunks sincethese canweaken the trunks andmake trees more prone to fall.
European and world occurrence
F. excelsior is indigenous and widespread in W & C Europe from the Cantabrian coast to beyond63ºN up the coastal fringe of Norway. From 61ºN on the Baltic coast, it stretches SE through C Russia(excepting the steppes), to Turkey, the Caucasus and parts of the Mediterranean – although there itmingles with other Fraxinus species (especially F. angustifolia) and the distribution ofthe species becomes unclear (Wardle 1961; Hultén & Fries 1986, Map 1489).
Names
The genus name 'Fraxinus' is the ancient classical Latin name of F. excelsior as used by Virgil,Ovid and others of the period (Gilbert-Carter 1964). The Latin specific epithet 'excelsior' means 'verytall' (Gledhill 1985). As with Beech (fa*gus sylvatica), Ash contains relatively little variation,but it has produced a number of abnormal forms, some of which are regarded as of horticultural merit.
Threats
The dieback disease has been killing many young trees in recent years and there is little evidence of itbeing brought under control, or of any real hope of doing so. Some trees will prove resistant, however.
Introduced, neophyte, garden escape. Rare, but probably under-recorded.
7 August 1993; McNeill, I.; roadside hedgerow, Cloghoge Td, W of Clabby.
May to August.
Growth form and preferred habitats
Syringa vulgaris is a deciduous, decorative, garden shrub or small tree 4-7 m in height withopposite, ovate leaves 4-12 cm long. It flowers in April and May, the 'wild' or original introduced formof the plant having mauve or rarely white, terminal, pyramidal, panicles 10-20 cm long.
The plant prefers moist, well-drained, moderately fertile soils of alkaline or neutral reaction. It doeswell on chalk and limestone, but cannot tolerate very acid, very dry, very wet or very shady conditions.It is not a very aggressive coloniser or competitor, but once it is established, which may requireseveral years' growth, S. vulgaris is long-persistent, its vigorous suckering habit enabling itto form small clones over a period of decades (Clapham et al. 1987; Phillips & Rix 1989).
Introduction and spread
S. vulgaris has been in cultivation in B & I for more than four centuries having beenintroduced from SE Europe pre-Gerarde's Herball, ie probably sometime just before the publicationof the book in 1597 that first describes its cultivation in England. The species is a Balkan native ofrocky hillsides from N & C Romania, Yugoslavia, Bulgaria, C Albania and NE Greece (Phillip & Rix1989). It now seems that many gardeners have over the subsequent centuries developed a love/haterelationship to the plant. The flowers of Lilac cultivars are beautiful and fragrant for a few weeks inearly summer, but after this the blossoms die disgracefully, hanging in ugly, brown, persistent paniclesand all that otherwise remains for the rest of the growing season is month after month of rather dismalfoliage taking up garden space. In late summer, lilacs are frequently attacked by a powdery mildewcalled Erysiphe syringae, there is no autumn leaf colour display and the seed clusters also failto present any aesthetic attraction (Clapham et al. 1987; Buczacki 2007).
The heyday of the species greatest popularity was the 19th century and most of the numerous, more than150, single- and double-flowered cultivars were selected and developed from then, or else from the earlyyears of the 20th century (Griffiths 1994; Sell & Murrell 2007). It became common practice to graftthe newly selected cultivars onto the original S. vulgaris rootstock which has the property ofsuckering freely (Buczacki 2007).
S. vulgaris was first recorded in Britain beyond the garden confines around 1879. It is largely bythe power of its vegetative reproduction that the different cultivated forms of Lilac have left gardensand become so familiar a feature of the wider environment into which the species continues to spread atthe present time. The increase of S. vulgaris records since the 1962 BSBI Atlas (Perring& Walters 1962), led T.D. Dines to describe the rate of change as, "astonishing",attributing it most probably to better recording of aliens in general, plus a genuine increase of thespecies presence (Preston et al. 2002).
Flowering reproduction
The many small, bisexual flowers that make up the inflorescence borne in April and May are 15-20 mm indiameter, strongly perfumed, often double. The calyx is yellowish green, divided one-third of the way tothe base into four lobes. The corolla varies in colour from pale to dark lilac, blue, purple, pink orwhite, the tube 8-12 mm, the four lobes 7-8 mm, obovate and rounded at the apex. There are two stamens,adnate to, and enclosed within, the corolla. The style is solitary, green and stays within the corollatube. The flowers attract insect visitors (mainly bees) that pollinate them and the fruit is a dry,smooth, hard, grey, ovoid capsule, 8-12 mm long that splits into two cells to release fourpapery-winged, wind-dispersed seeds. Presumably these wind-carried seeds are the primary means of thespecies' escape from gardens (Ridley 1930; Edlin 1964; Sell & Murrell 2007).
The current author (RSF) has not been able to locate information on some aspects of the basic biology ofthe plant, such as for instance whether or not it is self-compatible, the level of seed production, seedviability and longevity in soil, germination requirements, and other similar information and statistics.
Fermanagh occurrence
This familiar flowering garden shrub was previously ignored and un-recorded in Fermanagh. It appears rareto occasional in the just eleven tetrads so far recorded, scattered across the east of the VC. The findsare all in roadside hedgerows, where it appears as isolated individuals or as short stretches ofsuckering shoots. S. vulgaris can occur in situations quite remote from habitation. This suggeststo RHN and the current author (RSF) that it most likely originates locally as self-sown garden escapes,rather than as deliberately planted individuals, or dumped garden material. Apart from its initial rolein 'jumping the garden wall', seed reproduction appears of less significance in the wild in maintainingthe species than the observed active vegetative spread.
British and Irish occurrence
Elsewhere in Ireland, Reynolds (2002) suggests Lilac is deliberately planted in hedgerows. However,usually it is rare, often solitary and forms a patchy distribution in hedgerows, appearing as a relictin old, neglected gardens, on the margins of woods and on waste ground. Generally, it occurs thinly andirregularly distributed near habitation in the lowlands.
Reported instances of plants established in quarries, on cliffs, amongst sand-dunes, by roadsides,shrubberies and thickets in other parts of B & I suggest an origin involving either self-sown seedor discarded garden waste (or both), although it at first might appear unlikely that such a decorativeand attractively scented shrub or small tree could be a garden outcast on the scale required to matchits present day distribution and frequent occurrence.
Another frequently mentioned habitat is alongside railway lines, where the winged seed from gardensbacked onto the railway provided a source of propagules that, being caught in the slipstream of traffic,are swept along to colonise fresh sites, sometimes remote from habitation. In some situations, eg in WBerks, it has become locally abundant by the railway a good distance from any houses (Crawley 2005). Insome areas of England, as in Ireland, it probably remains under-recorded, possibly due to lingeringrecorder prejudice against non-native species (Halliday 1997; Greenwood 2012).
Locally under-recorded in Fermanagh
The presence of this unmistakable and familiar garden shrub was only realised relatively late on in theFermanagh Flora survey, so it is very probably more frequent in non-garden habitats than has been so farrecorded. In adjacent Co Tyrone (H36), for example, S. vulgaris has been recorded from no lessthan 60 5-km squares, strikingly more than in Fermanagh, and clearly the result of a greater awarenessof garden escapes in that VC (McNeill 2010).
The New Atlas editors have commented on the much wider occurrence of S. vulgaris throughoutB & I when compared with the 1962 BSBI Atlas. This suggests that Lilac might be activelyspreading in more recent years.
The details of the ten other Fermanagh hedgerow records are: Druminiskill, 3 km SE of Derrylin, 27 May1995, RHN & HJN; Carrignabrook, 2 km E of Glen Bridge, 3 August 1995, RHN & RSF; Knocks Td, 1 kmNE of Magheraveely, 4 August 1995, RHN & RSF; Knocks Td, ENE of Lisnaskea, 7 August 1995, RHN &RSF; Drummaw, 1.5 km WSW of Magheraveely, 2 May 1997, RHN; Eshnadeelada Td, 7 km NNE of Lisnaskea, 15July 1997, RHN; Tullynageeran Td, 1987-99, I. & D. McNeill; near Killyreagh House, Tamlaght, 17 May2005, RHN; Cloncoohy, 3 km SSW of Teemore, 11 September 2010, RHN & HJN; and E of Summerhill Lough,15 September 2010, RHN & HJN.
Threats
None.
Naturalised introduction, common. European temperate, widely naturalised including in N Europe, N Americaand New Zealand.
1884; Barrington, R.M.; Co Fermanagh.
Throughout the year.
Growth form, flowering reproduction and preferred habitats
A rapid-growing, deciduous, or in milder areas semi-evergreen, spreading, bushy shrub, 3-5 m tall, Wildor Common Privet bears lanceolate, dark green, opposite leaves, 6 cm long and panicles of small, whiteflowers at the stem tips in early summer. The over-wintering leaves tend to be broader at their apex andoften develop a bronze colouring (Lang 1987). Flowering occurs from mid-June to late August, the blossomis very fragrant, but it is considered by many to have a rather unpleasant, sickly-sweet scent. Eachflower is 3-4 mm long, with a tubular four-toothed calyx and a tubular corolla with four petal lobes, aslong as the tube. The flowers are insect pollinated. The fertilised flowers develop into shiny, black,rounded berry fruits, 6-8 mm in diameter, carried in dense erect clusters. The fruits vary in sizedepending on the number of seeds (one to four) they contain, the usual number per berry being two orthree. While the berries begin to form from late July onwards, it usually takes until the end of Octoberfor them to be fully ripe, and they then attract birds (Snow & Snow 1988). In mild winters, theberries often remain on the bushes until the end of the following February as birds appear to preferother available food. The seeds are large, 5 × 4 mm, shiny and black with a pitted surface and with flatsurfaces where they abut (Lang 1987). The berries are eaten and dispersed by Blackbirds and othermembers of the Thrush family, plus Robins and Magpies, while Blue tit* are occasional pulp predators(Snow & Snow 1988).
L. vulgare commonly grows in woods, scrub and hedgerows throughout B & I. The status of thespecies is questionable and problematic due to a scant, tentative and decidedly inadequate fossil record(Godwin 1975). Even granting some possibility of native status, it still remains difficult todistinguish indigenous from relic introduced stations since it is well known that the shrub has forcenturies been frequently planted in hedges and woodland, and it can spread further by self-seeding frombird droppings, typically giving rise to isolated individuals (Harron 1986; T.D. Dines, in: Prestonet al. 2002).
However ambiguous its status may be, L. vulgare is quite widely and traditionally regarded as anative plant in S Britain at least. The editors of the BSBI New Atlas hectad map gofurther and plot the species as native ± throughout England and Wales, although in the view of thecurrent author (RSF), the evidence to support this is shaky or absent. It is also acknowledged that theNew Atlas map may contain some records for the related evergreen species, L. ovalifolium(Garden Privet) due to recorder errors (T.D. Dines, in: Preston et al. 2002).
Wild Privet prefers well-drained, calcareous or base-rich soils that are moderately fertile, somewhatsheltered and in half-shade or full sun (Sinker et al. 1985). While growth is favoured bylimestone soils, L. vulgare is certainly not confined to them. It can tolerate most soil typesprovided they are not wet and soggy and is capable of surviving both drought and pollution, making itvery suitable hedging material. Although it avoids soggy ground, it regularly occurs on wooded riverbanks (Green 2008). Its drought resistance is exemplified by the fact that it can occasionally be foundbird-sown growing on walls (Reynolds 2013), presumably in part encouraged or allowed to survive onaccount of the lime in the mortar.
Irish occurrence
Commonly naturalised in hedgerows and spreading into derelict ground and the margins of woodland, WildPrivet is considered by some botanists to be a very rare native species in Co Waterford (H6), NTipperary (H10), SE Galway (H15) and Co Dublin (H21) where, apart from in hedges, it is confined tocliffs and rocky places (The Botanist in Ireland, paragraph 518; Cen Cat Ir Fl 2;An Irish Flora 1996). Praeger (1934d) seems to be the main authority for this status claim, buthe was partisan and therefore not always reliable in such matters. Reynolds (2002) in her Catalogueof alien plants in Ireland, recognises the long history of L. vulgare in planted hedges.
Fermanagh occurrence
Elsewhere in Ireland, including in Fermanagh, the shrub is in the opinion of RHN and the current author(RSF) sensibly regarded as a naturalised introduction, planted in hedgerows and in gardens and spreadinginto adjacent scrub and the margins of woods, through both naturally bird-sown seed and as clippingsdiscarded along with other garden waste.
As the Fermanagh species tetrad distribution map shows, L. vulgare is very widespread in the VCbeing recorded in 227 squares (43%), but it is more common in the southern and eastern lowlands wherethe agriculturally better soils are located.
Toxicity
As with L. ovalifolium (Garden Privet) (see account on this website), all parts of the plant aredangerously toxic to both stock animals and humans, although animals know to avoid it and poisoning isvery rare. Children are most at risk and the toxicity should not be under-estimated (Cooper &Johnson 1998).
British and Irish occurrence
Very common and widespread throughout lowland B & I, there can be no doubt that this species hasspread widely beyond its natural distribution, the limits of which are now obscure, or indeed impossibleto discern (T.D. Dines, in: Preston et al. 2002). The New Atlas hectad map plots L.vulgare as being an introduced alien throughout Scotland and Ireland, although in the latter itis sometimes considered to have native status in a few mainly coastal parts of the south and west (seeabove).
European and world occurrence
L. vulgare is a plant of wood margins and scrub in Europe, preferring calcareous soils. It isrestricted to S, W & C Europe, north to 59°N, and in a small area of SE Norway and SW Sweden (Lang1987). It is probably only native in regions south and eastwards of Germany towards the Mediterraneanbasin (Hultén & Fries 1986, Map 1490). As a native, it is confined to Europe, but is quite widelyintroduced in eastern parts of N America and has also been recorded in New Zealand.
Names
The genus name 'Ligustrum' is the name of an unknown plant in the ancient Romanwritings of Virgil that had white flowers (Gilbert-Carter 1964). It has clearly been'borrowed' to fit the privet species. Another suggestion is that the name is derived from the Latin'ligo', meaning 'to bind', the twigs having been used for tying objects (Johnson & Smith 1946). TheLatin specific epithet 'vulgare' means 'common', which in this case fits the occurrence rather well!
No fewer than 13 English common names are listed by Grigson (1955, 1987), of which the two earliest,mentioned by Turner (1548), were 'Privet' and 'Primprint'. However, the origin of such words is obscureand they also exist in other variant names as 'Pevit', 'Pevot', 'Prim', 'Privy' and 'Privy Saugh' (iePrivy Willow), leaving us none the wiser. It is very possible that 'privet' and its variousmodifications arose from the Old English 'pryfet', which is found in place-names such as Privett inHampshire, and minor names of the Middle Ages such as Prevetmore or Privetheye, although 'privet' on itsown is not recorded until 1542. Thus the name remains of unknown derivation, although the suffix '-et'indicates, as in other words from trees and shrubs, a collective, such as a 'thicket' of the plant namedin the first syllable of 'privet' (Grigson 1974).
Prior (1879, p. 192) offers another suggestion for the derivation of 'privet', from a mention by the poetand farmer Thomas Tusser (1524-1580), most famous for his instructional poem Five hundred points ofgood husbandry, first published in 1557, of another name for the plant, namely 'Privy'. This wassaid to be altered from 'Prymet', an abbreviation of 'Primprint', or 'Prim', the Primrose, through aconfusion of this flower and the shrub, from the application to both of them by medieval writers of theLatin name 'Ligustrum'. The names 'Primprint' or 'Prim' are nowadays transferred to the Privet shrub,but formerly belonged to the Primrose, from the French 'prime printemps', meaning 'first spring' andexactly corresponding to the modern French name for the flower, 'primevére' (Prior 1879, p. 190).
Threats
None.
Naturalised introduction, both deliberately planted and a garden escape. Apparently rare or very rare,but possibly mis-identified as L. vulgare and therefore under-recorded.
1 July 1986; Wolfe-Murphy, S.A.; S shore of Lusty More Td, Upper Lough Erne.
April to September.
Growth form, reproduction and preferred habitats
This semi-evergreen or evergreen monoecious hedging shrub, 3 m tall, is rarely recorded inFermanagh, but it may possibly be overlooked to some extent in favour of L. vulgare (WildPrivet). Garden Privet is a Japanese hedging species similar to Wild Privet that was introduced tocultivation around 1842 and first appeared in the wild in Britain in 1939 (T.D. Dines, in: Preston etal. 2002). Like L. vulgare, it also possesses shiny black berry fruits in late summer.However, L. ovalifolium can be distinguished through having broader, glossy, dark green ±evergreen leaves, hairless young twigs and bisexual flowers in which the fused corolla tube is longerthan the spreading petal lobes. The leaves are described as '± evergreen' since considerable quantitiesof them are occasionally shed in prolonged cold weather.
Garden Privet forms a dense and effective screening hedge, but it is neither ornamental nor decorative,except in its less vigorous golden- or silver-leaved cultivar forms. As hedging, the species recommendsitself by tolerating dry, impoverished soils where little else could survive, plus coping well with highlevels of atmospheric pollution in towns and cities. It is even more stress tolerant when it comes tomanagement, being able to survive hard clipping back at any time of year (Harris & Harris 1981;Buczacki 2007).
L. ovalifolium flowers in June and July and is pollinated by various insects. It subsequentlyproduces fleshy drupe fruits, 6-10 mm, that turn shiny, black as they mature. Reproduction beyond thegarden setting is probably mainly by seed, but hedge clippings and deliberate cuttings also root easily.In gardens, privet hedges are clipped so regularly, flowering and fruiting are generally preventedaltogether.
The plant remains abundantly planted as garden hedging and it persists as naturalised relicts in oldgardens. It is also common in parks, estates, amenity areas and waste places, in some of which itrepresents deliberated planted hedges, or rather often as established isolated bushes derived fromunofficial, fly-tipped garden discards (Buczacki 2007). Solitary plants may also arise more occasionallyas bird-sown seedlings, extremely rarely observed growing on walls (Green 2008; McNeill 2010).
Yellow- and silver-variegated leaved forms of L. ovalifolium are also popular garden subjects andare widely used in decorative gardening as well as for hedging. L. ovalifolium is generallypreferred over L. vulgare for hedging purposes as it is the more evergreen of the two (Sell &Murrell 2007).
Fermanagh occurrence
There are a total of only eight hedgerow records in the Fermanagh Flora Database, occurring across seventetrads. Except for the plant(s) found at Inver Bog by RHN, all are sited around the shores of UpperLough Erne and they were identified by members of the EHS Habitat Survey Team. The remaining recorddetails are: E shore of Lough Digh, 1 August 1986, S.J. Leach; S shore of Derryvore Td, 11 September1986, P. Corbett & A. Farr; N shore of Derryvore Td, 11 September 1986, S.A. Wolfe-Murphy & L.W.Austin; shore of Formil Td, 12 September 1986, S.A. Wolfe-Murphy & A. Farr; hedge near Inver Bog, 9April 1999, RHN; Drumskinny, 30 August 2003, I. McNeill; Tedd Crossroads, Raw Td, 9 September 2010, RHN& HJN.
British and Irish occurrence
From its mid-19th century introduction onwards, L. ovalifolium has been popular and widely andabundantly planted in gardens throughout the whole of lowland B & I. However, since the 1960s ittended to fall out of favour as hedging material and was replaced to a considerable extent by conifers,which have now also ± run the course of their popularity. L. ovalifolium is frequent andwidespread across England and Wales as far north as Carlisle and Newcastle-upon-Tyne; it has a similarpresence in Scotland across the central belt urban areas of Glasgow and Edinburgh with a thinningpresence elsewhere and becoming increasingly coastal northwards in Scotland (New Atlas).
In Ireland, Garden Privet is rather more thinly and widely scattered and appears less obviously urbanthan it does in Britain. In its Irish sites, it is nearly always deliberately planted and is only veryoccasionally believed to be bird-sown (Cat Alien Pl Ir). However, it is recognised that isolatedplants must either be bird-sown or the result of discarded garden hedge material or trimmings.
Toxicity
All the aerial parts of the species, but especially the berries, contain the poisonous glycoside,ligustrin. This plant toxin is so powerful it can kill a horse and should always be regarded aspotentially dangerous. This adequately explains why such a common garden hedging species is only veryoccasionally found along rural field margins. Privet can also cause dermatitis and some unfortunateindividuals can suffer severe skin reactions when clipping hedges (Cooper & Johnson 1998).
Names
See the account on this website for Ligustrum vulgare for derivation of names. The Latin specificepithet 'ovalifolium' means 'oval leaved', which does not adequately distinguish between the two privetspecies in the opinion of the current author (RSF).
Threats
None.
Introduction, occasional and only casual. Eurosiberian temperate, widely naturalised including in N &S America and New Zealand.
1902; Abraham, J.T. & McCullagh, F.R.; on old walls, Enniskillen Town.
August to November.
Growth form and preferred habitats
This large, deep tap-rooted, conspicuous, monocarpic, biennial or rarely annual plant produces a lowvegetative rosette up to 60 cm in diameter that overwinters and in the succeeding summer it develops astrikingly stout, erect, flowering stem, 0.3-2.0 m tall, densely covered with soft, white hairs. Infact, the whole plant, stem and leaves, is covered with a dense, greyish or whitish, felt-like, woollytomentum of long, soft, rarely shortly branched, hairs (Ross-Craig 1966, Part xxii, plate 1). Theflowering stem is also longitudinally ridged by the decurrent bases of the stem leaves. The lower,radical, rosette leaves are alternate, oblong or oblanceolate, 10-40 cm long, their bases againdecurrent on the internode below.
The tall flowering stem with its tiered decurrent leaves, their wing-like margins running down the stem,provides a means of channelling rainwater from above down to the roots. The upper leaves droop at theirtips and discharge rainwater into the channelled midribs of the leaves below them that then act likegutters, so that water eventually trickles its way down the entire height of the plant using thedecurrent leaf-bases, until it reaches the roots. This is an important adaptation as the species mostlyoccupies open, ± disturbed, ungrazed ground with well-drained or dry soils (Melderis & Bangerter1955).
In Britain, Great Mullein is a frequent plant of rough grassland, hedge banks, roadsides, near quarries,railway sides and waste ground, preferring warm, dry, open, recently disturbed or bare, sandy, stony,rocky or calcareous soils that are moderately fertile and ungrazed. It can sometimes also grow on walls(Sinker et al. 1985).
The dense coat of felted leaf hairs makes V. thapsus foliage unpalatable to stock animals such ascattle, the hairs setting off intense irritation in the mucous membranes of the grazing animal, so theyquickly learn to avoid the plant. The hairs also protect the plant from grazing by some phytophagousinsects (Grieve 1931; Gross & Werner 1978). When preparing the useful Mullein Tea, a remedy of thegreatest antiquity for coughs and colds, it must always be strained through fine muslin to remove anyhairs that may be floating in the hot water that has been poured over the flowers, or leaves, asotherwise they cause intolerable itching in the mouth (Grieve 1931).
The felted hairs also protect the species from aqueous solutions of the herbicide 2, 4-D as the waterdroplets cannot penetrate the leaf surface (Gross & Werner 1978).
The established strategy of V. thapsus is categorised as SR/CSR, meaning it is intermediatebetween a Stress-tolerant Ruderal and a more balanced mix of all three strategies, Competitor,Stress-tolerator and Ruderal (Grime et al. 1988, 2007).
Variation
V. thapsus is described as a polymorphic species by Hultén & Fries (1986) and they mention theease with which it becomes involved in hybridisation. Stace et al. (2015) list a total of sevenrare hybrids involving V. thapsus, none of which has ever been recorded in Ireland.
The variation in Europe is sufficient for three subspecies to be recognised in Flora Europaea3, these being: subsp. thapsus, throughout the range of the species; subsp.crassifolium (Lam.) Murb., mountains of S & C Europe, from Portugal to the E Alps; and subsp.giganteum (Willk.) Nyman, mountains of S & SE Spain (I.K. Ferguson, in: Tutin et al.1972).
There are no subspecies presently recorded in B & I (Sell & Murrell 2007).
Fossil record and the question of native status
There is no fossil pollen record of V. thapsus at all and Godwin (1975) mentions only two seedoccurrences, the first in the very early Cromer Forest Beds and the second in the Ipswichianinterglacial which long predates the current Flandrian (or in Ireland, Littletonian) interglacial.Therefore, there is no relevant, direct evidence of fossil support for native status in B & I andother circ*mstantial evidence, either way native/introduction, must be sought to resolve or illuminatethe question (Webb 1985).
The fact that V. thapus is a weedy, colonising species of disturbed ground, very long persistentin the soil seed bank and has been widely used for many years – throughout recorded history and inClassical times – in herbal medicine for both man and beast, and cultivated for this reason in Irish andE & SE England cottage gardens, sometimes on a considerable commercial scale (Allen & Hatfield2004), suggests that man has had an important role in its introduction, increase and spread acrossEurope and B & I.
The genus has many species, some more suitable for garden cultivation than others, but all areacknowledged as, "Stately plants, mostly of biennial duration, but the best are so handsome andlong flowering as to be quite essential in the garden." (Robinson 1909). The current author (RSF)is not suggesting that V. thapsus has been grown for decorative reasons, although this isnot an impossibility since it is a very notable, sizeable, striking plant, but it certainly has beencultivated for medicinal use, probably until the late 19th century at the very least.
Phenology and flowering reproduction
V. thapsus germinates in early spring and forms a leaf rosette that continues to grow into lateautumn and then overwinters. After a cold vernalisation period, in the following spring the plantproduces a tall flower stalk. Flowering begins in late June and peaks in early August. The inflorescenceis a very dense, spike-like raceme, 20-50 × 3 cm. Rarely, additional axillary racemes develop from theupper leaves.
Flowers mature on the inflorescence stalk from the bottom to the top in successive spirals, so that atany one point in time they appear in a loose spiral pattern up the length of the raceme. As theflowering season progresses, each successive spiral of blossoms begins slightly higher on the stem, butit overlaps for most of its length with the earlier spirals. Growth of the inflorescence stalk isindeterminate and since the duration of the flowering period is a function of stalk height, tallerstalks can continue flowering into the autumn (late September or early October).
The flowers are large, sessile, bisexual and only slightly irregular. Each flower is subtended by aprominent green bract. The calyx is deeply 5-lobed, the lobes narrowly lanceolate. The corolla is'regular' rotate (ie symmetrical, not irregular), with a very short tube and five broad, spreadinglobes, 20-35 mm in diameter, pale yellow, or rarely white and with five irregular stamens directlyattached to it (ie they are epipetalous). The three upper filaments are shorter, clothed with dense,yellowish or white-woolly hairs and their anthers are reniform (ie kidney-shaped) and medifixed, whilethe lower two stamens have longer, glabrous filaments. The lower pair of anthers are large, elongate,coloured, obliquely attached and produce a copious supply of pollen. The flowers produce very little orno nectar. The superior ovary is 2-celled and is surmounted by a solitary style, topped by aclub-shaped, capitate stigma (I.K. Ferguson, in: Tutin et al. 1972; Hickey & King 1981; Staceet al. 2015).
The individual flowers of V. thapsus are ephemeral, opening before dawn and closing aroundmid-afternoon of the same day. They are protogynous, the style maturing first before the anthers andthen bending downwards once the anthers appear. The two anterior stamen filaments are naked and theyproduce most of the pollen that leads to fertilisation. The woolly hairs on the filaments of theposterior three stamens appear to provide a useful platform foothold for insect visitors such ashoverflies and bees that gather pollen from the posterior anthers while their abdomens are dusted withpollen from the anterior pair. Although visited by a wide variety of insects, including butterflies,moths and flies (Fitter 1987), it is said that only long- and short-tongued bees are effective vectorscarrying out cross-pollination (Pennell 1935). Since there is very little or no nectar in the flowers,this latter claim appears open to question and the current author (RSF) expresses his doubt on thematter.
Verbascum is an unusual genus in that the stigma is not merely passive but, rather, is active whenit comes to pollen capture. In Verbascum, and also in Acanthus, the stigma is like avalve, opening only when it is receptive and closing again shortly after pollen is deposited upon it(Percival 1965).
Towards the end of the day (ie usually mid-afternoon), if cross-pollination has not occurred, the stylereturns to its original position and the corolla closes, pushing the still receptive stigma against theanthers and effecting self-pollination (ie autogamy takes place) (Kerner von Marilann 1895).
The fruit is a capsule, 3-6 mm long, ovoid, longer than the calyx and very hard. It splits into twovalves at maturity to release the numerous minute seeds which are columnar, truncate, light to darkbrown, 0.5-1.0 mm long, their lateral surfaces angular, six-sided and rugose with blunt tubercles invertical rows (Butcher 1961; Hutchinson 1972).
Seed production, dispersal and longevity
Seed production can be very large, but it is also extremely variable, ultimately depending to agreat extent upon the scale of the plant and size of the inflorescence. Salisbury (1942) reports themean number of capsules per plant as 226 ± 42, with an average of 596 ± 30 seeds per capsule. This givesan approximate average of 136,000 seeds per plant. The number of seeds for a population from a threeyear old Michigan field in August 1976 ranged from 0 to 749 per capsule, with a mean of 208 ± 200 for atotal of 175,000 per plant. The extreme variation in this latter example was caused by weevil predation(Gross & Werner 1978).
Seeds possess no specialised morphological adaptations for dispersal by wind or animals, but they aretiny, very hard and produced in abundance. The fruit capsule splits open along its long axis when matureand any slight movement of the tall flowering stalk by wind or a large animal is all that is required torelease the seeds from the parent (Mclean & Ivimey-Cook 1956).
Seeds are dispersed as far as 11 m, although 93% of them fall within 5 m and 75% of them fall within 1 mof the parent plant (Salisbury 1964; Gross & Werner 1978).
Seed of V. thapsus is capable of immediate germination upon release, there being no embryoafter-ripening requirement (Gardner 1921). Buried dormant seed survival in soil is long-term persistent,viable seed having germinated after periods of 38 years or much longer. Seed in Denmark from soil levelsarchaeologically dated to 1300 AD were still viable (Oosting & Humphreys 1940; Ødum 1965; Thompsonet al. 1997).
Fermanagh occurrence
An occasional casual in Fermanagh, Great Mullein is only sporadically found in gardens, waste ground,walls and disturbed soil. Although the plentifully produced wind-dispersed seed is long-persistent inthe soil seed bank (Salisbury 1942, pp. 126-7), it is hardly persistent in any site within the VC. V.thapsus has been recorded in 19 tetrads and, as the map indicates, they are thinly scattered inthe centre and southern half of Fermanagh, generally in sites near habitation. Seventeen of the tetradshave post-1975 records.
Irish occurrence
Rare and local in Ireland, Great Mullein is listed in the Cen Cat Fl Ir 2 asoccurring at least once in every Irish VC except W Donegal (H35). In the past, it has been presumednative throughout the island. It is a locally frequent colonist in stony or sandy banks, seashores andbare waste places in the southern half of Ireland, but is very much rarer in the north of the island.The rare, sporadic, casual occurrence in Fermanagh and elsewhere in NE Ireland (FNEI 3), leadsthe current author (RSF) to believe it is much more likely to represent an alien here. The occasional,somewhat more clumped occurrence in adjacent Co Tyrone (H36), mostly on waste ground and dumps in the SEand NW of the VC, is not dissimilar to the Fermanagh experience and, again, McNeill (2010) considersGreat Mullein an introduction.
British occurrence
There are six British species of Verbascum and natural hybrids between most of them are notuncommon. V. thapsus is widely distributed throughout England and Wales but it becomes scarcenorthwards and it is rare and local in Scotland. It should also be noted that it is absent from theOuter Hebrides (VC 110) and Shetland (VC 112) and is recognised as an alien introduction in both Orkney(VC 111) and the Isle of Man (VC 71) (A. Horsfall, in: Preston et al. 2002).
European and world occurrence
V. thapsus belongs to the Eurosiberian temperate phytogeographical element. It is distributedacross most of Europe except the extreme north and much of the Balkan Peninsula. It is also present inAsia south to the Caucasus and Himalaya and eastwards to W China. It is introduced and widelynaturalised across N America and is present in Argentina, Patagonia and the islands of New Zealand,Hawaii and Réunion (Hultén & Fries 1986, Map 1629).
Uses
It is probably the case that all the various Mullein species found in Britain possess similar medicinalproperties. Stace (2019) lists 13 species plus 20 hybrids, most of which being garden escapes would beof very limited distribution (New Atlas; Stace et al. 2015). Of the more common andsignificantly available and useful species, V. thapsus is by far the one most employed (Grieve1931). For medicinal purposes, it is generally collected from the wild, although in Ireland where it ismuch more scarce in comparison with England and Wales, it was carefully cultivated in gardens because ofa steady demand for the plant by sufferers from pulmonary consumption (Grieve 1931).
The leaves and flowers are the parts used medicinally. The leaves are nearly odourless and are of amucilaginous and bitter taste while the flowers are slightly more agreeable in flavour. The plant hasvery marked demulcent, emollient and astringent properties that make them useful in herbal medicine forthe treatment of both pectoral complaints and bleeding of the lungs and bowels. It was greatly valuedfor the treatment of consumption. The herb was also used to treat the pain and irritation ofhaemorrhoids and an infusion was given to treat diarrhoea and bleeding of the bowels (Grieve 1931).
Use in cattle included treatment for strangury (difficulty in passing urine), for which finely choppedMullein leaves were mixed with bran and water fed to the animals (Vickery 2019).
In continental Europe, a sweetened infusion of the flowers, strained in order to remove the rough hairs,was frequently used as a domestic remedy in mild catarrhs and colic. A conserve of the flowers was alsoemployed against ringworm and a distilled water of the flowers was long reputed to be a cure for burns.These, and many other herbal medicinal uses, plus reports of various other non-medicinal uses, forinstance as a yellow dye, sometimes used as a hair dye, and as a fish intoxicant, the seeds beingslightly narcotic, have been documented by Grieve (1931)! Vickery (2019) mentions finds of V.thapsus at Mount Grace Priory in N Yorkshire after an archaeological dig and the possibilitythat monks based there would have used the dried flower stalks of Mullein, which were dipped in tallowto make wicks for processional candles. This religious establishment was dissolved in 1539, but Vickerywarns that the idea that the Mount Grace plants grew from 16th century seed is probably no more thanwishful thinking.
Names
The genus name 'Verbascum' is the classical Latin name of an unknown plant in the Roman writer Pliny'sworks borrowed and reused for this group of plants (Gilbert-Carter 1964; Gledhill 1985). Anothersuggested origin is that it might be a corruption of the Latin 'barbascum', meaning 'a hairy plant',from 'barba', 'a beard', many of the members of the genus being characterised by the possession of downyfoliage (Johnson & Smith 1946). The Latin specific epithet 'thapsus' is after the town of that namein ancient Africa (now Tunisia), or after the Greek island Thapsos (Johnson & Smith 1946; Gledhill1985).
No less than 50 English common names are listed by Grigson (1955, 1987) and 43 by Vickery (2019), many ofthem containing the word elements 'flannel' or 'blanket', clear references to the soft, woolly hairsthat cover the plant. Names that suggest everything that 'soft' include references to Rabbit's ears,Donkey's ears, velvet, rag paper and flannel. Other names contain elements such as 'candle' and 'taper'which refer to the use of the dried plant as kindling, or as lighting when dipped in suet or wax.
The most frequently used name 'Mullein' dates from the 15th century from the French 'moleine', possiblyfrom 'mol' ('mou'), meaning 'soft' (from the Latin 'mollis'), an obvious reference to the conspicuouslysoft, felted leaves (Grigson 1974). The French 'moleine', from the Old French 'malen' and the Latin'malandrium', refers to the scab disease in cattle and to the 'malanders' or leprosy. 'Malandre' alsorefers to other diseases in cattle, including lung problems (Prior 1879). The name 'Mullein' was speltvariously by early herbal writers, eg 'Moleyne' (Grete Herball, Anonymous (1526)), 'Mollen'(Turner (1568) A New Herbal), 'Mullen' (Turner (1548) Names of herbs), 'Mulleyne'(Lyte (1578) A Niewe Herball, or Historie of Plantes). Gerard also has 'Mullein' or rather'Woolen', a reference to the generally woolly appearance of the plant (Britten & Holland 1886).
Threats
None.
Native, frequent to locally common. Eurosiberian temperate, introduced in eastern N America.
1881; Stewart, S.A.; Derrygonnelly.
Throughout the year.
Growth form and preferred habitats
A stout, erect, unbranched perennial, growing 60-120 cm tall, from a rhizome or rootstock that istuberous and covered with numerous 'knotted' nodules – hence one of its English common names 'KnottedFigwort', and its Latin specific epithet 'nodosa', meaning 'swollen joints'. This underground organ,which today we usually are forbidden to excavate and examine for conservation reasons, is very wellillustrated by Ross-Craig (1966, Part XXII, plate 16). The glabrous stem is sharply four-angled, dullgreen and finely speckled with purple. The opposite leaves are stalked, glabrous, 5-12 cm long, dark,slightly bluish-green on the upper surface, paler beneath, ± triangular ovate, acute at apex. The leafmargins are coarsely and unevenly or doubly toothed, with larger teeth occurring towards the base(Hutchinson 1972; Sell & Murrell 2007).
S. nodosa differs from S. auriculata and S. umbrosa in the sharply four-angled, butun-winged stem and the very narrow scarious border of the sepals (Garrard & Streeter 1983).
S. nodosa is a quite frequent but non-gregarious, often local perennial of shaded or semi-shaded,moist to damp, but well-drained, fertile soils in a wide variety of lowland sites, some of which aredisturbed, eg along roadsides, ditches, hedgerows, waste ground and quarries (Garrard & Streeter1983; Sinker et al. 1985). It is chiefly found as more or less isolated plants in woods alongtheir margins and rides, and in scrub, often on lakeshores, streamsides or river banks, but it is alsoless frequently found in drier situations on screes and along lanes, gravel driveways and on wasteground. While it tolerates a considerable degree of shade (Halliday 1997), it can also grow on walls atheights of 3-4 m above the ground (Ridley 1930).
S. nodosa was included by Salisbury (1942) with a list of 16 other, "woodland typescharacteristic of intermittently open habitats" that for instance included Chamerionangustifolium (Rosebay Willowherb), Cirsium palustre (Marsh Thistle), Digitalispurpurea (Foxglove), Galeopsis tetrahit (Common Hemp-nettle), Gnaphaliumsylvaticum (Heath Cudweed), Hypericum perforatum (Perforate St John's-wort), Seneciosylvaticus (Heath Groundsel) and Verbascum thapsus (Great Mullein). The 17 species onSalisbury's list all tend to be common features of coppiced areas or clearings in recently felledwoodland and they share the characteristic of sometimes occupying such temporary open areas in amazingabundance, almost certainly due to the presence of their dormant seed in the woodland soils. Speciesthat occupy habitats that are only intermittently available for colonisation tend to have much largerseed productivity, for instance, 38 of these species produced approximately 28,200 seeds per plant,compared to 38 species of permanently open habitats that produced just over 3,000 seeds per plant(Salisbury 1942).
A successful, competitive perennial species that holds its ground well, Common Figwort features in thetall herb community of damp ground, but it is intolerant of both grazing and mowing (Sinker etal. 1985). However, the leaves are bitter-tasting so that browsing animals normally avoid itentirely. "Cattle, as a rule, refuse to eat the leaves as they are bitter, acrid and nauseating,producing purging and vomiting if chewed."(Grieve 1931).
The established strategy of S. nodosa is characterised by Grime et al. (1988, 2007) asC/CR, meaning it is intermediate between a straight Competitor species and a Competitive Ruderal.
Flowering reproduction
S. nodosa flowers in summer and early autumn, from July to October. The flowers are borne inloose, pyramidal panicles composed of small opposite to alternate cymes, the oldest flowers being in themiddle of each cyme. The branches of the inflorescence are covered with short gland-tipped hairs.
The flowers are bisexual, 6-7 × 3-4 mm, the calyx saucer-shaped, 5-lobed, the lobes ovate with shortteeth on the margins. The sepal margins have a very narrow, often scarcely visible, scarious margin. Thecorolla is irregular, 2-lipped, 7-10 mm, the tube greenish and the upper lip purplish-brown, 2-lobed.The lower lip of the corolla is greenish, 3-lobed and shorter than the upper lip. There are four fertilestamens plus a solitary sterile staminode that is broader than long and is emarginate (ie slightlynotched). The fertile stamens are at first curled downwards, away from the style which matures first andoccupies the mouth of the corolla (ie the flower is protogynous). The ovary, which is seated on a large,fleshy, nectar-secreting receptacle disk, is 2-locular, with numerous ovules on the two fleshy axileplacentas (Butcher 1961; Hutchinson 1972; Sell & Murrell 2007).
Pollination
Cross-pollination is effected primarily by wasps of the genus Vespula, and by flies, thebrownish-purple petal colour being typical of 'wasp-flowers'. The insects are attracted by thecarrion-like smell of the flowers. The stigmas are receptive before the anthers open and they remain sofor 48 hours. After this, the stamen filaments straighten, the pollinated stigma withers and the stylebends down over the lower lip of the corolla, its place being taken at the mouth of the corolla by thedischarging anthers. Insect visitors alight on the upper, younger flowers that are in the female phaseand work their way downwards to the older flowers that have moved to the male phase, thus carrying outcross-pollination.
The fruit is a broadly ovoid, pointed capsule containing many rugose (rough textured), brown seeds 1.2 ×0.6 mm, that are half ovoid, marked with deep vertical grooves and horizontal striae. The capsule opensby splitting into four valves and the small seeds are shaken out by wind or by passing animals movingthe plant (Butcher 1961; Hutchinson 1972).
Seed production
In terms of fruit and seed production, Salisbury (1942) examined 61 plants and found the number of fruitsranged from 42 to 956 per plant. The total number of capsules counted was 14,354, giving a mean of 235 ±16 capsules per plant. Sixteen capsules were examined for seed content and the number of seeds percapsule ranged from 105-272. The mean number of seeds in the 16 capsules was 204 ± 8.6. Thus the meanseed output per plant per year was between 43,000 and 54,000 seeds. Germination in the light is around60%, so the calculated mean reproductive capacity would be around 28,000 potential offspring each year(Salisbury 1942).
Salisbury (1942) does mention that another researcher, Eklund (1929) in Scandinavia, produced data forsix capsules of S. nodosa which yielded a much lower average, namely 105 seeds per capsule.Eklund also gave the numbers of fruits for two individuals as 60 and 76. As Salisbury (1942) commented,these data would, if typical, suggest a much lower productivity in Scandinavia than in his own studiesin Britain.
Seed dispersal and longevity in soil
There is no specialised seed dispersal mechanism, but the tall plants provide a favourable starting pointfor wind to catch and carry the small, lightweight propagules at least a few metres from the parentfruit capsule. As mentioned above, S. nodosa has often been observed growing on walls in elevatedpositions (up to 3-4 m) (Ridley 1930, p. 28), clearly carried aloft by wind.
The seed is also long-term persistent in the soil seed bank, some of it remaining viable for at leastfive years (Thompson et al. 1997).
Vegetative reproduction
There is evidence of vegetative reproduction in S. nodosa from an examination of the swollenunderground rhizome or tuber and its nodules. Salisbury (1942) contrasted the rapid vegetative extensiongrowth of Ranunculus repens (Creeping Buttercup) runners with what he referred to as, "theclosed type of extension growth of S. nodosa" involving very small annual increments in thespread of the underground tuber over an eleven year period of study (Salisbury 1942, pp. 225-6, Fig.35). This 'closed extension of growth' is described by him as, "owing to the swollen character ofthe short stolons and their perennial character, almost complete exploitation within the periphery isassured."
Fossil history
There are no fossil pollen records for S. nodosa and Godwin (1975) reports a solitary seedidentification by Reid from the Hoxnian interglacial. Thus there are no fossil records of any sort fromthe present interglacial in either Britain or Ireland that the current author (RSF) has informationabout. This does not decide native or introduced status, but it means that all and any availableevidence regarding status will be circ*mstantial, and needs to be carefully analysed.
Fermanagh occurrence
Although seldom if ever plentiful, as the tetrad map shows, S. nodosa is fairly frequent and verywidespread in Fermanagh, having been recorded in 209 tetrads, 39.6% of the VC total.
British and Irish occurrence
Widely distributed throughout B & I, except for N Scotland, the Scottish Highlands and the NW ofIreland where the soils are probably too acid and peaty for it (New Atlas). There is littleevidence of any significant change in the distribution or frequency of S. nodosa from BSBIsurveys covering the period since the 1960s (Braithwaite et al. 2006).
European and world occurrence
Eurosiberian temperate in its phytogeography, widespread in Europe and W Asia, but noticeably thinningtowards both north and south in Scandinavia and the Mediterranean (Hultén & Fries 1986, Map 1631).It is rarely recorded in eastern N America, where it is an introduction, and it has also been introducedto South Island, New Zealand where it is very rare (Webb et al. 1988).
Uses
Common (or Knotted) Figwort has a long history of herbal and folk medicine uses, the 'fig' in the namereferring to piles. This brown-flowered and very smelly herb has knobbly, tuberous roots (hence thealternative common name), which by the 'doctrine of signs' suggested its use for treating the diseasecalled 'scrophula' (or 'scrofula'), that was characterised by glandular swellings and probably was aform of tuberculosis. The disease was formerly also referred to as 'the King's Evil', characterised byswollen tubercular glands on the neck (Grigson 1955, 1987). S. nodosa was called 'the ScrofulaPlant' on account of its value in treating all forms of cutaneous eruptions, abscesses and wounds and itwas formerly considered a specific for this purpose (Grieve 1931; Stearn 1992). The flowers also havesmall glands on the corolla, which being far more visible than the tuber, might also suggest thespecies' connection with Scrophula or Scrofula disease (Gledhill 1985).
In herbal medicine, the plant is considered to possess diuretic and anodyne properties, ie it helpsproduce urine, or assists its flow and is painkilling. The whole herb, collected fresh and dried in Juneand July, is made into a decoction for external use, and the fresh leaves are also made into an ointment(Grieve 1931). In Ireland, the dried root or tuber was ground to a powder and mixed with lard to make anointment used for treating piles and all kinds of skin troubles. The roots and leaves, and sometimes theseeds, were boiled and the liquid drunk as a kind of tonic, "held to clear the blood of impurities,including boils and rashes". The forerunning examples and numerous other Irish herbal medicinalapplications are detailed by Allen & Hatfield (2004).
British use of this herb has been very slight in comparison with Irish use and apparently has beenrestricted to the leaves alone, for poulticing skin eruptions, abscesses, ulcers and wounds (Allen &Hatfield 2004).
Of all the different species of Figwort, S. nodosa is the most used, principally as a fomentationfor sprains, swellings, inflammation, wounds and diseased parts, especially in scrofulous sores andgangrene (Grieve 1931).
Names
The genus name 'Scrophularia' is from the Latin 'scrofule', translating as 'scrophula', the tuberculardisease it was supposed to cure. The Latin specific epithet 'nodosa' means 'having conspicuous joints ornodes', or 'knotty', ie referring to the root tuber and its nodules (Gilbert-Carter 1964; Gledhill 1985;Stearn 1992).
Grigson (1955, 1987) reported that in Ireland, Figwort was considered the 'Queen of Herbs' and Foxglove'the King of Herbs'. He lists a total of 15 additional English common names for the species, several ofwhich refer to the greenish-brown tint of the leaves and the brown colour on the petals, such as'Brown-net', 'Brownwort', 'Brunnet' and 'Brennet'. A few of the names refer to the medicinal propertiesof the herb, such as 'Poor Man's Salve' and 'Cut-finger'. Several names refer to the unpleasant smell ofthe plant, such as 'Stinking Christopher' and 'Stinking Roger' (Grigson 1955, 1987).
Threats
None.
Native, frequent. Suboceanic southern-temperate.
1902; Abraham, J.T. & McCullagh, F.T.; Castle Caldwell.
February to December.
Growth form and preferred habitats
This glabrous, pale, yellowish-green stemmed perennial, 50-100 cm tall, grows on somewhat wetter lowlandground than S. nodosa (Common Figwort) and was for many years known as S. aquatica. Whilethese two rather unpleasant smelling Figwort species overlap, they do not hybridize. Typicalwaterside habitats of S. auriculata include the sides of ditches, river- and stream-banks and wetareas on lakeshores. It can also occur in wet woods and damp meadows.
The established strategy of S. auriculata is characterised by Grime et al. (1988, 2007) asC/CR, meaning the species (in parallel with S. nodosa) is intermediate between a straightCompetitor species and a Competitive Ruderal.
Fermanagh occurrence
In the FNEI 3, Hackney et al. describe S. auriculata as being, "very rare andlocal" in NE Ireland, but it is really quite frequent in Fermanagh, occurring in 63 tetrads, 11.9%of those in the VC. While it is quite widely scattered across the VC, the high presence of Water Figwortaround Lower Lough Erne in particular is rather surprising. However, having a more obviously calcareousshore the Lower Lough supplies the moderately fertile, neutral, lowland soils that S. auriculataprefers. The species is also known to occur occasionally on drier conditions in calcareous soils and itdoes so especially in western parts of B & I (Garrard & Streeter 1983; Flora of Connemara andthe Burren).
Differences between S. nodosa and S. auriculata: S.auriculata differs from the more common S. nodosa in being a somewhat taller plant,rather less branched, with winged angles on the square stems and obtuse, kidney-shaped, bluntly toothedor crenate, ovate to elliptical leaves, obtuse at the tip. Sometimes the leaf blades have one or twolobes at the base (hence the name 'auriculata' (see below)) and the petiole is pale green, winged andglabrous. The leaves are dark green on the upper surface, paler beneath. It also has a fleshy, short,simple rhizome and a fibrous root system, rather than the nodular rhizome or tuber characteristic ofCommon or Knotted Figwort (Sell & Murrell 2007).
Flowering reproduction
S. auriculata flowers from July to September and has narrow linear bracts below each pair of cymesin the inflorescence. The two-lipped flowers, 6-9 × 3-5 mm, are borne in a narrow, cylindrical panicle.The calyx is five-lobed, each lobe ovate-obtuse with a broad, conspicuous scarious margin. The corollais 5-9 mm, tube greenish, globose; the two-lobed upper lip is brownish-purple, longer than the green,three-lobed lower lip. Occasionally the flower is pure white. The flower is otherwise very similar toS. nodosa, with four fertile stamens, with a staminode that is sub-orbicular and entire. Thestyle is solitary, 3.5-4.0 mm, and the stigma capitate (Melderis & Bangerter 1955; Butcher 1961;Hutchinson 1972; Sell & Murrell 2007).
Pollination
Pollination is by wasps, the flowers being favoured by Vespidae, especially Vespula vulgaris andV. germanica and less frequently visited by other insects including honeybees and various otherbees. The insects are attracted by the stinking odour of carrion the whole plant emits and the flowerfoods. Being typical of 'wasp-flowers', S. auriculata offers insects readily accessible nectar,but releases relatively little pollen from the four fertile epipetalous stamens held close to the lowerlip of the corolla. As in S. nodosa, the flowers are protogynous, the female phase being primaryand lasting just two days. The previously bent back stamens then straighten and bring the anthers to themouth of the flower above the now withered stigma (Proctor et al. 1996).
There does not appear to be any satisfactory explanation as to the function of the sterile staminode inthe upper part of the flower. It is a distinctly uninteresting flap of tissue attached to the petals,unlikely to divert or delay the insect visitor inside the flower in its all-engrossing search for a foodreward (Proctor et al. 1996; Sell & Murrell 2007).
Fruit and seed behaviour
The fruit is a 4-6 mm subglobose capsule with an apiculate apex (ie possessing a short, abrupt point).The numerous small seeds are pale brown to black in colour, 0.8-1.0 mm long, oblong and rugose withwavy, vertical ribs (Butcher 1961; Sell & Murrell 2007). As in S. nodosa, the capsule splitsinto four valves and the small, lightweight seeds are shaken out by wind or passing animals shaking thetall fruiting stem.
Since S. auriculata typically grows beside water, it is obvious that some seed will fall intowater directly, or soon be washed into it following rainfall. Ridley (1930) observed and was puzzled bythe fact that seeds of this river-plant sink immediately in water. He investigated and found that seedsput in water sank and remained at the bottom for about a week. The seed then germinated under water andemitted the radicle (ie the first root), but did not float for a further two days – ie when thecotyledons expanded. This experiment took place in the month of August and the seedlings involved werestill floating on 13th October. Ridley concluded that it is undoubtedly due to the floating seedlingsthat S. auriculata disperses along the edges of rivers and other water bodies and eventuallysettles and roots. Another related genus that also does this is Mimulus (Ridley 1930) and otherspecies that disperse in this way include Baldellia ranunculoides (Lesser Water-plantain),Lythrum salicaria (Purple-loosestrife) and some species of Juncus (Rush spp.) (Sculthorpe1971).
The seeds of Water Figwort are also short- to long-term persistent in the soil seed bank (ie surviving1-5 years or more burial), although there are only three such estimates listed in the soil seed banksurvey of NW Europe (Thompson et al. 1997).
Toxicity
Like Common Figwort, the S. auriculata plant has an unpleasant odour and taste and it containstoxic glycosides that reduce milk yield if cattle graze it. This has caused at least one reportedpoisoning of young calves in B & I (Cooper & Johnson 1998).
British and Irish occurrence
A common plant of wet, waterside growing conditions throughout lowland Britain, the distribution plottedin the New Atlas hectad map is broadly similar to that presented in the earlier BSBI Atlas(Perring & Walters 1962, 1976), although there are considerably more records towards the north ofthe contiguous British range of the species (the N limit approximately represented by a line on the mapbetween Liverpool and Durham). Most probably this expansion is due to a better, more thoroughexamination of previously less studied areas of England. In Scotland, S. auriculata is decidedlyrare or very rare and in some sites it is considered an introduction. The species very likely reachesits world NW limit in S Scotland. While frequent in the Channel Isles, it is rare in the Isle of Man.Many records in Scotland have had to be discounted, being regarded as likely errors for S. nodosa(A. Horsfall, in: Preston et al. 2002).
In Ireland, S. auriculata is quite widely distributed (New Atlas), but is much morefrequent in the S & W and is only occasional elsewhere (Parnell & Curtis 2012).
European and world occurrence
In phytogeographical terms, it is described as suboceanic southern-temperate and is present in W Europenorthwards from Spain, Portugal and France to the Netherlands, Germany, then SE to Switzerland andextending further southeast to the Balearics, Italy, Sicily, Corsica, Sardinia and Crete. It is alsorepresented in N Africa (Morocco and Tunisia) and the Azores (Tutin et al. 1972; Pignatti 1997;Sell & Murrell 2007). Quite how it manages to travel so widely to colonise isolated islands, such asthe Azores, is a bit of a mystery, unless it is assisted by human activities.
Uses
As with Common Figwort, S. auriculata has been used in folk medicine for centuries (Grieve 1931;Allen & Hatfield 2004). See the current author's account of S. nodosa on this website formore details of typical use.
Names
The genus name 'Scrophularia' is from the Latin 'scrofule', translating as 'scrophula', the tuberculardisease it was supposed to cure. The Latin specific epithet 'auriculata' means 'lobed like an ear' or'with ear-like lobes', a reference to the one or two basal leaf lobes, despite the fact that they arenot always present (Gledhill 1985).
Twenty-one English common names are listed by Grigson (1955, 1987) and 25 by Vickery (2019). Twelve ofthe names are unique to S. auriculata, and the others are shared with S. nodosa. AsGrigson points out, similarity of names given to plants in folklore and herbalism was often due to thesimilarity of parts rather than the whole of different species, ie the emphasis was on the parts used inpharmacy. Thus leaves of S. auriculata resemble leaves of Betony (Stachys officinalis), sothe herbalist, apothecary or patient grouped the two species and distinguished them as 'Water Betony'and 'Wood Betony'. Water Betony was another stinking herb sympathetically (following the 'Doctrine ofSigns') clapped on to stinking sores, ulcers, wounds and bruises (Grigson 1955, 1987).
Another interesting name for S. auriculata was 'Crowdy-kit'. 'Crowdy' referred to a fiddle and thesqueaky noise some people make when playing it. Children make a squeaking, fiddler's noise by scrapingone dry stem of the plant across another and hence the connection and label for the species. As withS. nodosa, many of the names refer to the rather brownish colour of the plant, or to its strong,unpleasant smell, eg 'Black doctor', 'Brown net', 'Stinking Roger' and 'Scaw-dower' or 'Scaw dowr' inthe Cornish tongue, meaning 'Water Elder', a reference to the smell (Vickery 2019).
Threats
None.
Native, very rare.
1939; Praeger, R.Ll.; wooded lakeshore, Davy's Island (also known as Inish More), at the E end of LowerLough Erne.
This Figwort hybrid is extremely rare in Ireland, having only ever been recorded twice on theisland: by the River Liffey in H21 (Co Dublin) in 1937, and here in Fermanagh at Davy's Island on LowerLough Erne in 1939. Both Irish sites were discovered by Praeger, growing where plants of the parentsclosely coincide and in what has been described in Stace (1975) as, "the two main localities ofS. umbrosa in Ireland". There is no specimen from the Lough Erne site in the DublinHerbarium (DUB) (Wilcox 2013).
Hybrid plants have been identified in sites where the two parents grow close together. The hybrid isintermediate in a whole range of characters, including the staminode, so one would need to bevery familiar with the species to recognise the hybrid (C.A. Stace, in: Hybridization,pp. 367-8). With this in mind, it is not quite so surprising that this hybrid has never beenrefound in either of its reported Irish stations in recent years, and that nobody has increased thedistribution with additional records (Flora of Co Dublin).
Stace (1975) pointed out that, plants from the banks of the River Shalbourne, S Wiltshire (VC 8)identified as this hybrid by Druce (1916), were later said to be diseased-affected (mildewed) examplesof S. umbrosa with malformed inflorescences and abnormal staminodes. Accounts of these fourspecimens appear to differ: either no flowers at all, or only a few normal, un-diseased flowers werepresent on the Wiltshire specimens, but all four are said to bear well-formed fruit and seeds (Stace1975; Wilcox 2013).
Subsequently, when the four specimens (two pairs of specimens) in the Oxford herbarium, Druce(OXF), originally collected in October 1915 by C.P. Hurst, were re-examined by Wilcox in 2012,none of them proved to be hybrids, but rather, two turned out to be S. umbrosa, and two wereidentified as S. auriculata (Wilcox 2013).
The Co Dublin voucher for Praeger's (1951) plant in DBN was also examined by Wilcox, who foundthat it had fertile capsules. On that basis he re-determined it as S. auriculata. Thus, as aconsequence of these investigations and considerations, Stace et al. (2015) gave it as theiropinion that, "hybrid figworts do not deserve a place in our flora".
Artificial hybrids between the above two species have been successfully synthesised, being mentioned onthe annotations added to the specimens by Goddijn & Goethart (1931 in sched.), by Vaarama& Hiirsalmi (1967), and by Wilcox (2013). The latter author reports that, "The hybrid plantsare said to be intermediate, but generally resemble S. auriculata in the leaf type (ie withoccasional 'auricles' on the petioles). However, the staminode shape is only relatively intermediate andmore closely resembles S. umbrosa in shape, although very slightly larger (deeper = height) thaneither parent".
Wilcox (2013) provides a figure comparing the staminodes of the parent species and the artificial hybridhe created. He is of the opinion that, "fertile hybrids would be almost impossible to detect in thewild without careful searching, but plants with similar staminodes to the artificial hybrid would beworth collecting." In this short paper Wilcox encourages collection of potential hybrid materialand indicates a willingness to examine any specimens sent to him (Wilcox 2013).
Introduction, neophyte, very rare, but possibly overlooked and under-recorded to some extent.Eurosiberian temperate, with a distinctly continental distribution in W Europe.
1939; Praeger, R.Ll.; Davy's Island (also called Inish More), at the E end of Lower Lough Erne.
Growth form, identification and preferred habitats
S. umbrosa is a stout, erect, perennial, 60-100 cm tall, with a simple, compact, fleshy rhizomethat like other Scrophularia species tends to grow on fertile lowland soils at the sides ofrivers and streams and in damp woodland, in both open and shaded situations (A. Horsfall, in: Prestonet al. 2002).
The square stems of S. umbrosa are even more obviously 4-winged than those of S. auriculataand both have a distinct scarious margin to the sepals that is up to 1 mm wide. One of the alternativeEnglish common names draws attention to the winged stem, calling the plant 'Green-winged Figwort'(Butcher 1961) and the current author (RSF) recommends that this name be taken up generally. 'GreenFigwort' does not convey anything useful. The two Figworts differ in that S. auriculata hasleaves that are bluntly toothed and sometimes have small, leaf-like lobes at their base. Also thestaminode (a sterile stamen situated under the upper lip of the corolla) in S. auriculata isrounded or kidney- shaped. The leaves of S. umbrosa, on the other hand, are lanceolate, brightgreen, sharply toothed and with no leaf basal lobes, and the staminode is distinctly two-lobed(Garrard & Streeter 1983; Parnell & Curtis 2012; Wilcox 2013, Fig 3).
Green-winged Figwort individuals tend to be scattered amongst other more common waterside plants such asCaltha palustris (Marsh Marigold), Iris pseudacorus (Yellow Flag), Oenanthe crocata(Hemlock Water-dropwort), Phalaris arundinacea (Reed Canary-grass), Schoenoplectuslacustris (Common Club-rush) and Scirpus sylvaticus (Wood Club-rush), or else it may formsmall, pure stands.
There is some degree of negative association between S. umbrosa and S. auriculata which mayreplace it, at least in some more open situations in southern England where the latter is quite common(M.E. Braithwaite, in: Stewart et al. 1994; A. Horsfall, in: Preston et al. 2002).
S. umbrosa reproduces by seed which mainly germinates in the spring. Unlike S. nodosa andS. auriculata, it does not form a persistent soil seed bank
(M.E. Braithwaite, in: Stewart et al. 1994).
Fermanagh occurrence
There are just three records for S. umbrosa from two widely separated areas in Fermanagh, and R.H.Northridge and the current author (RSF) consider one of them very, or even extremely, doubtful. In anyevent, S. umbrosa is very rare and local on damp lakeshore woods and possibly also on woodedriver banks in the Cladagh River Glen. The two Fermanagh records that are more acceptable are bothPraeger's and date from 1939. The first Praeger site is the above one from Davy's Island, which lies offthe Castle Archdale estate on the E shore of Lower Lough Erne. Here Praeger, who would have previouslyseen the plant on the River Liffey, described the S. umbrosa representation as, "veryfine". The second Praeger discovery was on the closely adjacent larger White Island, where herecorded it on the eastern shore of the wooded island.
The solitary post-1975 record in the Fermanagh Flora Database was from the Marble Arch or Cladagh RiverGlen woods, and RHN & RSF would greatly prefer to have a voucher of it to confirm its correctidentification. The record was made in February 1993 by a visiting English bryologist working on asurvey commissioned by the EHS. RHN & RSF very much doubt that he could identify this rare andcritical figwort in February, and without a voucher are loathe to accept his record.
Irish occurrence
Elsewhere in Ireland, S. umbrosa has been only very rarely recorded from just five widely spacedVCs (Irish Red Data Book). It is most abundant on the banks of the River Liffey in Co Dublin(H21), where its Irish discovery was made in 1895. The only other recent record is from north CoLondonderry (H40), on the River Bann just S of Coleraine (FNEI 3). S. umbrosa would appearto be declining, but it is quite probably under-recorded, being mistaken for or not distinguished fromS. auriculata (Irish Red Data Book).
British occurrence and status
S. umbrosa is rare and very local, scattered throughout England, Wales (especially the Welshborders) and Scotland as far N as the Moray Firth (New Atlas). The patchy pattern of occurrencein B & I and the late dates of its discovery around both islands make RHN and the current authorcertain that this species is a modern introduction: not native, but a neophyte (New Atlas; Sell& Murrell 2007).
In Britain, while it remains scarce, S. umbrosa has certainly increased quite markedly inabundance during the 20th century. Despite its similarity with S. auriculata, previous confusionbetween the two species does not appear to explain the perceived increase of S. umbrosa. Itappears to be steadily becoming more frequent and is spreading beyond its previous main centres, perhapsbeing dispersed from them by wildfowl (M.E. Braithwaite, in: Stewart et al. 1994).
European and world occurrence
Belonging to the Eurosiberian temperate phytogeographical element, S. umbrosa has a chiefly C& E European distribution that stretches discontinuously northwards to Denmark, S Sweden and Latvia,and southwards to Corsica, Sardinia, Italy, Sicily and the Peloponnese. It appears to be absent or rarein most of W Europe. It also occurs in W & C Asia to Tibet and south to Palestine. In outlying areasbeyond the stations mentioned, it is a casual species (Hultén & Fries 1986, Map 1632; Sell &Murrell 2007).
Names
The genus name 'Scrophularia' is from the Latin 'scrofule', translating as 'scrophula', the tuberculardisease it was supposed to cure. The Latin specific epithet 'umbrosa' is from 'umbra' meaning shade andtranslates as 'growing in shady places' or 'shade-loving' (Gilbert-Carter 1964; Gledhill 1985). Asmentioned above, in the opinion of the current author (RSF), the BSBI recommended English common name'Green Figwort' (Dony et al. 1974) is not as meaningful or informative as 'Green-winged Figwort'.
Threats
None.
Introduction, neophyte. A mis-identification or a transcription error.
8 October 1996; Waterman, Mr T.; Tullynanny Lough.
Since it has never been recorded anywhere else in Ireland, the Fermanagh record of this perennial isregarded as a definite error. A long-lived, ruderal neophyte in Britain, S. scorodonia is mainlyconfined to coastal situations in SW and SE England and the Channel Isles, where it is reckoned to be anaccidental and only occasional introduction, although in some areas it is long-established (A. Horsfall,in: Preston et al. 2002).
Introduction, established neophyte of garden origin, occasional to locally frequent.
1892; Praeger, R.Ll.; Inish Dacharne peninsula, Lower Lough Erne.
Throughout the year.
This established western N American alien is a 19th century garden escape, now naturalised in or nearstreams or other wet, muddy lowland situations throughout most of the British Isles. A vigorous,stoloniferous, patch-forming aquatic, up to 30 cm tall, it both seeds itself abundantly and spreadsvegetatively in autumn to form creeping or clumping monocultural patches (Sell & Murrell 2007). Thespecies has very successfully dispersed since it first appeared in the wild in Abergavenny, S Wales in1824 and now extends in a scattered manner across Britain from the Channel Isles to Shetland (Ellis1993; A. Horsfall, in: Preston et al. 2002).
It has been recorded in 61 Fermanagh tetrads (11.6% of the total in the VC), 55 of them with post-1975records. As the tetrad distribution map indicates, Monkeyflower is widespread on muddy stream-sides andlakeshores across the VC, but is most frequently found in the east of the county.
Elsewhere in Ireland, M. guttatus has been recorded in at least 31 of the 40 VCs (Cen Cat Fl Ir2), including all those in the north of the island. The New Atlas hectad map, however,displays recent records from only 16 Irish VCs. The atlas is very clearly in error since this count doesNOT include Fermanagh – where the records actually fall into as many as 19 hectads!
M. guttatus has numerous blood red spots on the bright yellow lower lip of the open, two-lippedflowers, which are solitary in the axils of upper leaves. A number of other closely related anddifficult to distinguish American Mimulus species and hybrids of garden origin have also escapedand become naturalised in wet, lowland places.
Threats
This alien is very probably still spreading.
Mimulus guttatus × M. luteus (M. × robertsii Silverside), Hybrid Monkeyflower
Introduction, garden escape. Very rare, but probably overlooked and under-recorded.
4 July 1997; McNeill, I.; Tempo River at Tonyglaskan Bridge.
All apparently in July.
This hybrid, rhizomatous or stoloniferous perennial Monkeyflower of garden origin was first recognised inBritain in 1964 and only properly named in 1990. It is an established garden escape that has alreadybeen recorded scattered throughout B & I, but with a predominance in NW England, S Scotland and NIreland (Ellis 1993; Clement & Foster 1994; New Atlas). At the moment, the mappeddistribution as displayed in the New Atlas in the opinion of the current author (RSF) reflectsonly the home territory of the interested recorders of this taxon.
Two recorders have been looking out for this plant in Fermanagh and the seven records in the NE of thecounty made by Ian and David McNeill are very probably just the beginning of the story for this taxon.RHN and the current author (RSF) are sure it has been overlooked elsewhere, or been mistaken for theundoubtedly much more frequent M. guttatus (Monkeyflower) – a situation very commonly foundthroughout these islands (A. Horsfall, in: Preston et al. 2002).
Like M. guttatus, which is one of the parent species, the hybrid grows in shaded, wet, muddy orshingly soils by flowing water, or in flushes on marshy or fen-rimmed lakeshores. It reproduces not onlyvegetatively by fragmentation of the short, brittle stolons, but also by seed, since it has formed ahexaploid amphidiploid form by carrying out spontaneous chromosome doubling (Mimulus peregrinusVallejo-Marin (New Monkeyflower)), that has restored sexual fertility to the previously sterile triploidhybrid (Sell & Murrell 2007; Stace et al. 2015).
The McNeills have recorded sufficient M. × robertsii in Tyrone (H36)to regard it as being quite common there (McNeill 2010). This hybrid has previously been recorded in CoAntrim (H39) and in at least five other Irish VCs (Cen Cat Fl Ir 2; FNEI 3; Cat AlienPl Ir). The most accurate picture of Mimulus distribution is presented in the hectad mappublished in Stace et al. (2015).
The remaining Fermanagh record details are (some dates vague): Killerbran Bridge, Colebrooke River,1987-99, I. & D. McNeill; stream at Glengesh, 7 km NE of Tempo, 1987-99, I. McNeill; Tempo River atTonyglaskan Td, 1987-99, I. McNeill; stream at Jamestown House, Magheracross, 2000, I. McNeill; PollboyBridge, Colebrooke River, 16 July 2000, I. McNeill; and Derryloman, Colebrooke River, 16 July 2002, I.McNeill.
Threats
Probably spreading along with the M. gutattus parent and perhaps replacing it, especially onhigher ground.
Native. Very rare. Circumpolar boreo-temperate.
19 July 1957; MCM & D; Corraslough Point, near the entrance of the Swanlinbar River to Upper LoughErne.
August and September.
Growth form and preferred habitats
Mudwort (or Common Mudwort) is a rather inconspicuous, semi-aquatic, loosely tufted, emergent dwarfannual with slender, creeping stems 10-15 cm long and leaf rosettes only 2-7 cm high, composed of roots,leaves and numerous very reduced flowers produced at each node. It does not have an upright stem, butit* slender leafless runners or stolons begin their growth erect before becoming horizontal andspreading. The stolons are short-lived and are produced during optimal vegetative activity, theirprimary function undoubtedly being as organs of reproduction, rather than perennation (Sculthorpe 1967).
The leaves are radical (ie all basal) and can be either emergent, floating or submerged. The floatingleaves have blades entire, elliptic, obtuse, spoon-shaped, glabrous, apex obtuse to rounded with aslender petiole that is often reddish and nearly twice as long as the blade (ie 2-7 cm long). Insubmerged plants, the leaf petiole can be elongated to as much as ten times that of the emergent leavesand the leaf blade is almost non-existent, the leaf being subulate or ± filiform (Sculthorpe 1967;Meikle 1985).
Mudwort, occasionally also referred to as 'Water mudwort', is found on exposed mud of temporary pools andstream banks, from 0-300 m altitude. As a semi-aquatic plant, it grows in moist or wet ground habitatsincluding damp hollows in meadows, in mud and wet sand next to water and partly submersed or floating inshallow depths around pools, lakes, streams and ditches.
As Salisbury (1970) pointed out, the great majority of the 50 or so B & I species characteristic ofexposed muds are actually of local, sometimes extremely local, occurrence, which suggests their need foran infrequent combination of environmental conditions.
In B & I, L. aquatica is a typical member of the flora of drying mud, but it is very local andgenerally is regarded as a rare annual (Salisbury 1967). It grows on and is confined to the stretch ofmud exposed on the wet margins of mainly lowland lakes, ponds, reservoirs, rivers, streams and ditchesand other periodically flooded places as water levels shrink in hot, dry summer weather. As Salisbury(1970) points out, "Where mud borders the larger, more permanent, areas of inundation a graduallowering of the water-table may only enhance the vigour and centrifugally extend the zone of the'reed-swamp' species, or other shallow-water vegetation, so that, normally, a severe summer drought (orartificially abrupt lowering of the water-level in a lake or reservoir) will alone provide an expanse ofbare mud where pioneer species can colonize and flourish."
Plants may be partly or completely submerged as a result of renewed rainfall and a rising water table.L. aquatica is a very rapidly germinating ephemeral that colonises bare mud when it first becomesexposed to the air in summer, or later in the season and the small plant completes its short life cyclebefore autumn or early winter rains re-flood the occupied ground.
The species is described as erratic as well as ephemeral, L. aquatica often reappearing at siteswhere it has been absent, or has not been seen, for many years. Population sizes can vary enormouslyfrom year to year, depending on the extent of the muddy shore exposed; the number of individuals in alocality, when it sporadically reappears, may be astonishingly large (Salisbury 1967). The explanationfor this appears to be that seed can remain dormant and viable in submerged or regularly wetted mud formany years.
L. aquatica does not appear to be a very competitive species, so that winter submergence ofits seed, as well as summer desiccation providing abruptly available expanses of bare mud, may havetheir main significance in diminishing competitive pressure from other species, rather than throughconferring a direct benefit on Mudwort (Salisbury 1970).
While in Fermanagh and its few other sites in Ireland, including in the karst limestones of the Burren,Co Clare, L. aquatica appears calcicole. Elsewhere in B & I, there is a feeling that thespecies is somewhat (maybe only slightly) calcifuge, preferring mildly acidic soils, which arenutrient-enriched by manure from grazing animals and, when sufficient water is present, by the droppingsof flocks of feeding water fowl.
Flowering reproduction
Plants flower from June onwards, although the normal season generally stretches from July to September(Salisbury 1967). The numerous, tiny, bisexual, regular (actinomorphic) flowers, 2-3 mm in diameter aresolitary on rather short peduncles, 1-2 cm long (shorter even than the leaf petioles) in the axils ofbracts. The peduncles become spreading (decurved) when in fruit. Flowers have a 5-lobed calyx thatpersists and becomes membranous in fruit. The corolla consists of five white or pale pink petals fusedtogether into a short basal tube. The flowers are pollinated by insects and, if this fails, theyself-pollinate (Fitter 1987). In a number of mud species, cleistogamy and self-fertilization is thenorm, at least in submerged plants, so seed set is automatic and very large soil seed bank numbers havebeen reported in some instances.
After fertilisation a bluntly ellipsoid capsule, usually c 3.0 x 1.5 mm, the largest fruits rarelyexceeding 4.0 × 3.0 mm, develop, each containing a surprisingly large number of very small seeds(Salisbury 1967).
Seed production
Salisbury (1942, 1967) examined a total of 178 plants over three seasons and discovered the mean numberof capsules per plant was 31, and the mean number of seeds per capsule was 117, a high figure for such asmall fruit. The mean seed productivity is therefore calculated to be around 3,600 per plant (Salisbury1967).
It has to be emphasized that the size attained by individual L. aquatica plants varies enormouslyowing to the marked diversity in the density of the Mudwort sward on the one hand and, on the other, tothe seasonal timing of the plants' development associated with the onset of drought conditions and mudexposure. If a sunny summer occurs after mud is exposed, seeds germinate rapidly and large plants maydevelop producing secondary fruiting rosettes. If the seedling sward is very dense, however, rosetteformation will be reduced or even prevented, resulting in fewer fruits per plant and possibly smallercapsules containing fewer seed.
Having said this, clearly potential seed production of L. aquatica is remarkably large for aspecies with such a small photosynthetic assimilatory surface. A large seed output in relation to sizeappears characteristic of a number of species that specialise in the colonisation of bare mud. A strongparallel is afforded by Elatine hexandra (Six-stamened Waterwort), an even smaller plant whichalso produces thousands of seeds and which regularly co-habits with Mudwort. Environmental conditionsfor successful reproduction by these ephemeral species recur at infrequent intervals making themsporadic in their occurrence and natural selection has made them opportunistic, giving them rapidgermination and colonising ability when conditions are favourable, enabling them to generate enormouspopulations on bare mud (Salisbury 1967).
Fully developed specimens of L. aquatica are quite easy to identify by the shape of the leaves andthe small white flower, although often plants fail to develop fully.
Seed dispersal
Seeds are often released underwater when the capsules open and may thus be readily dispersed within thewater catchment in flowing streams. While the seeds are very small and lightweight and could easily becarried by wind blowing across open capsules, wetland birds probably are significant vectors also,carrying seed of the plant in attached mud on their feet and feathers to fresh sites of a more remotenature (C.D. Preston, in: Stewart et al. 1994).
Variation
A second very rare species of the genus occurs in Britain only very locally in Wales, and is otherwisetotally absent in Europe. Elsewhere it occurs in eastern N America, and is more widely distributed inthe southern hemisphere and the N Pacific. It is L. australis R.Br. (syn. L. subulata E.Ives), Welsh Mudwort, which differs in having all its leaves needle-like (ie subulate), and its calyx isshorter than the corolla tube. A plant of wet sandy mud at the margins of pools where water has stoodfor some time, in Wales L. australis is very local in Glamorganshire (VC 41), Merionethshire (VC48) and Caernarvonshire (VC 49). This usually annual species produces stolons or rhizomes both above andbelow the soil surface, and these organs can occasionally perennate (Hutchinson 1972; Sell &Murrell 2007).
Fermanagh occurrence
The mouth of the Swanlinbar (or Claddagh) River as it enters Upper Lough Erne and Ross Lough (includingthe mouth of the Sillees River) are the only two sites where L. aquatica has ever been seen inNI. When Meikle and co-workers discovered the plant in 1957 it was abundant and some was growing up to10 cm tall on soft muddy ground under 5–30 cm of water along with Eleocharis acicularis (NeedleSpike-rush). The L. aquatica plants varied greatly in size and luxuriance, some withfloating leaves closely resembling Baldellia ranunculoides (Lesser Water-plantain), whileothers plants were minute (Carrothers et al. 1958).
From 1991 onwards this little, ephemeral, sporadically appearing annual has been seen at Ross Loughgrowing on mud exposed after periods of summer drought. In September 1991, it covered a strip ofshoreline 100 m long × 5 m wide. In August 1994, it formed an elongated mat 60 m long × 3 m wide here,positioned several metres out from the normal shoreline. On these two occasions, the plants were sodensely packed they resembled a grassy sward. Accompanying species on the exposed mud were Alismaplantago-aquatica (Water-plantain), Bidens cernua (Nodding Bur-marigold),Callitriche agg. (Water-starworts), Persicaria hydropiper (Wall-pepper), Rorippaamphibia (Great Yellow-cress) and R. palustris (Marsh Yellow-cress).
At the entrance of the Sillees River to Ross Lough in 1992, 1994 and 2002, the species grew verysparingly on exposed mud, but only as scattered individual plants. In September 2002, very scatteredL. aquatica grew along an extremely muddy track trampled by cattle about a metre above theforeshore of Ross Lough, rather than on the shore itself where it had been recorded on earlieroccasions. In October 2005 and 2006, it was very scattered along approximately 150 m of muddy shore,which had been trampled by cattle. The plant had to be searched for on the latter occasion. In July2010, it was found growing in its usual place, just above the normal high water mark, scattered along200 m of the muddy lakeshore.
Ross Lough is next to the limestone hills at Boho and the local habitat for the plants here hassimilarities with turlough sites, ie vanishing lakes sited above limestone cave systems in Cos Clare andGalway, where drainage is essentially vertical and where the habitat is grazed by cattle during thesummer months (Curtis et al. 1985). At Fisherstreet, a coastal area of the Burren, Co Clare (H9),the plant has rarely and very exceptionally been found growing in small, extremely shallow solution cupsetched into the horizontal surface of clints, the flat surface slabs of limestone pavement (Curtis etal. 1987). This could hardly be a more ephemeral habitat for a semi-aquatic plant.
British and Irish occurrence
Most of the 50 or so 'mud-species' that occur in Britain have greatly diminished in their frequency, theaverage loss in the period around the late 1960s being probably about 40% of their former locations(Salisbury 1970). Since then, the decline has continued: an analysis of the total number of hectads in B& I with records in the period 1987-99 displayed in the New Atlas, showed just 80 squareswith L. aquatica present. A further 20 hectad squares had records dating from 1970-1986 and therewere pre-1970 records known from another 136 squares, making the rate of decline on both islands veryobvious.
In all or almost all its stations in B & I, L. aquatica is sporadic and very variable inabundance from year to year. As has just been discussed, it has been recorded from more than one hundredstations scattered over England, in a considerable proportion of which it no longer occurs, and it isalso known from only a very small number of sites in Scotland and Ireland (Perring & Walters 1962;Preston et al. 2002). The species is very strikingly intermittent in its occurrence, sometimesreappearing after many years of apparent absence (missing for up to eight decades), yet the number ofindividuals in a locality when it reappears may be astonishingly large (Salisbury 1967; C.D. Preston,in: Stewart et al. 1994). Mudwort is thus scarce or rare, extremely local and thinly scatteredacross Britain from Hampshire northwards to NE Aberdeenshire.
There has been a remarkable increase in records of this species in Scotland since the 1950s (Leach etal. 1984), probably due to better recording of an easily overlooked, inconspicuous plant, butperhaps indicating some degree of northward spread. Losses in England and Wales appear to be associatedwith pools and a range of smaller water bodies that previously represented the most characteristichabitat of the plant, while by comparison the species has survived better around larger waters (C.D.Preston, in: Stewart et al. 1994).
In Ireland, sites again are very rare, widely scattered and confined to a few sites in Co Clare (H9), CoCork (H3 & H4) along the River Lee and its reservoirs, Co Fermanagh (H33) and SE Co Galway (H15)(Curtis & Ryan 1985; O'Mahony 1986; Curtis & McGough 1988).
The fact that Mudwort has become such a very rare and rather widely disjunct species in Ireland and ascarce, erratic and declining local one in Britain, suggests that losses of suitable, very specificmuddy habitats may have reached a critical threshold. This is an especially serious situation for asporadic species which only erratically produces a very large, but perhaps a not always very long-livedsoil seed bank.
European and world occurrence
L. aquatica is widespread in most of Europe except the Mediterranean region, although it ispresent in N Italy and Cyprus and has scattered colonies in the Alps. It is often rare and local. It isalso present in Egypt and in arctic and temperate Asia eastwards to India and Japan. The species has acircumpolar boreo-temperate distribution that includes Greenland and N America from Labrador to theNorth West Territory and south in the mountains to Colorado and California (Meikle 1985; Hultén &Fries 1986, Map 1626; Sell & Murrell 2007).
Names
The genus name 'Limosella' is a diminutive, derived from 'limosus', meaning 'muddy'. The Latin specificepithet 'aquatica', translates as 'aquatic'.
The English common name 'Mudwort' or 'Water Mudwort' delimits the habitat concisely.
Threats
On account of its rarity in NI and the vulnerability of its habitat, L. aquatica has been listedunder the Wildlife (NI) Order, 1985 as one of 56 plant species given special conservation protection inSchedule 8, part 1.
Introduction, neophyte, garden escape, very rare.
8 August 1996; Northridge, R.H. & Forbes, R.S.; waste ground on N bank of River Erne at Belleekvillage.
Growth form and preferred habitats
A prolifically seeding, familiar, colourful garden annual, biennial or short-lived perennial of endemicPyrenean and mountains of SC France and Mallorca origin, A. majus was probably introduced to B& I in the 16th century (Grigson 1974). Snapdragon (or better, Common Snapdragon), grows 30-80 cmtall from a strong taproot and produces flowers 40-45 mm long in short terminal racemes of up to 20blossoms, usually with purple, pink or yellowish-white corollas. The form that is grown in gardens isdeveloped from subsp. majus, but in the Mediterranean basin, subsp. tortuosum (Bosc) Rouyis also planted and naturalised widely (D.A. Webb, in: Tutin et al. 1972). While it can beperennial in milder areas of the country, and if given winter protection elsewhere, A. majus isusually grown in British gardens as a half-hardy annual. It flowers from July to September and ispollinated by large, weighty insects like bumble-bees that can open the closed corolla mouth, orself-pollinated (Garrard & Streeter 1983).
Plants are very stress-tolerant, make few habitat demands and appear as sporadic garden escapes(sometimes long-persistent), usually growing on old walls, in pavement crevices, or on waste ground nearhabitation (Hackney et al. 1992).
Fermanagh occurrence
This species has only ever been recorded once in the VC, on waste ground on the riverbank at the town ofBelleek. The belated appearance of the species in Fermanagh and at least five other Irish VCs, maysimply be the result of deliberate neglect by earlier recorders who considered it beneath them to recordsuch obviously non-native garden material in artificial or disturbed habitats of no conservation value.
Irish occurrence
Elsewhere in Ireland, Common Snapdragon is a fairly regular escapee, especially in the eastern half ofthe country where the gardening population is most densely congregated. By the current author's (RSF's)reckoning, A. majus has been recorded at least once in 19 of the 40 Irish VCs, including thewestern Cos Mayo and Galway (Clapham et al. 1987; Hackney et al. 1992; Cat Alien PlIr). Reynolds (2002) and Hackney et al. (1992) also note that plants that establishthemselves on old walls and which can survive for over 50 years in such situations, tend to be thepredominant reddish colour form, whereas those in more disturbed ground display a wider range of bothflower colour and plant form.
British occurrence
A. majus has been a popular garden subject in Britain since the time of Elizabeth I. It was firstrecorded in the wild in 1762 and remains abundantly planted, seeds profusely and frequently escapes, sothat it is widely recorded across lowland Britain, especially near gardens. In England, it is mostfrequent and widespread south of a line between Lancaster and Hull, becoming much more scarce andscattered north of this limit. In Scotland, it is frequently recorded around and between the two majorconurbations of Edinburgh and Glasgow and again appears further north around Perth and Inverness.
Inspection of the hectad maps in the two BSBI atlases shows the frequency and distribution of CommonSnapdragon has greatly increased in the 40 years since the 1962 Atlas, probably reflecting both agenuine greater spread of the plant, as well as better recording of aliens in the flora (Perring &Walters 1962, 1976; A. Horsfall, in: Preston et al. 2002).
European occurrence
Endemic in SW Europe and introduced more widely to gardens and escaped and naturalised widely across theMediterranean, so it occurs at least as far east as Cyprus (D.A. Webb, in: Tutin et al. 1972;Meikle 1985)
Names
The genus name 'Antirrhinum' is a name in Dioscorides from the Greek 'anti', meaning 'resembling' and'rhis' or 'rhinos', a 'nose' or 'snout' (supposedly of a dragon), alluding to the shape of the flower.An older generic name for the plant was 'Asarina' or 'Asarrhina', which carried the probable meaning'Gummy-snouted' (Johnson & Smith 1946).
The English common name 'Snapdragon' was first used in 1573 (reference not quoted by Grigson 1974, but isprobably Thomas Tusser, Five Hundred Pointes of Good Husbandrie (1573)), the jaw-like flowers,being supposed to suggest a dragon's head, can be easily opened by a gentle squeeze, and will snap backinto their natural posture when released (Grigson 1974). The name 'Snapdragon' has been used fornumerous related and unrelated plants, including Antirrhinum majus, Aquilegia vulgaris,Digitalis purpurea, Fumaria officinalis and Linaria vulgaris (Britten & Holland1886).
Threats
None.
Introduction, archaeophyte, very rare and sparse. European temperate but widely naturalised in thenorthern hemisphere.
1934; Praeger, R.Ll.; railway at Belleek.
July to September.
Growth form and preferred habitats
Previously a familiar weed of ancient introduced origin, C. minus is now found in very open, stonysites along railways and on disturbed, lowland waste ground, quarries, walls, short permanent grasslandand in stony or rocky areas of both agricultural and garden cultivation. Familiar and regarded as acommon arable weed by Colgan & Scully (1898) (known then as Linaria minor (L.) Desf.), sincethe advent of herbicides in the 1940s, this often very diminutive, 5-30 cm tall, glandular, viscid,branched, spring-germinating, summer annual has declined very markedly. It now only occurs very rarelyon dry, or well-drained, bare gravel or stony soils, often of a calcareous nature and never below pH4.5. The established strategy is categorized as R/SR, meaning the species is intermediate between a pureRuderal and a Stress-tolerant Ruderal (Grime et al. 1988, 2007).
Small Toadflax appears in Fermanagh, as elsewhere in most of Ireland, almost entirely associated with thesites of derelict old railway stations and on the previous permanent way, track bed ballast, somestretches of which have been developed, since the closure of the railways, into car-parks or cattletracks to and from fields (FNEI 3; An Irish Flora 1996).
Flowering takes place from June to September, the shortly-spurred purple and red flowers being small(6-10 mm long), solitary and axillary on short stalks (peduncles) at the base of the upper leaves. Thetwo-lipped corolla in C. minus is ± open at the mouth (ie ajar), unlike in Linaria, whichin botanical terminology is described as 'personate', meaning 'masked' or 'hidden', the throat of theflower being closed by a pair of bulges on the lower lip. Pollination is probably by selfing and theresultant fruit capsule, 3-6 mm, is ovoid and shorter than the calyx (Clapham et al. 1987). Thecapsule opens to release the small black seeds by means of three irregular pores per cell (ie 3 pores ×2 capsule cells) at its apex (Butcher 1961; Sell & Murrell 2007).
Seed production by this often very small plant is relatively large, Salisbury (1942) calculating the meanproduction of an average-sized individual at around 2,200 seeds per plant. Even though the plants aresmall, they are erect and the seeds being small and light are readily wind dispersed. Howeverpersistence in the soil seed bank is rather variable: sometimes prolonged (more than five years),sometimes short-term persistent (1-5 years), and occasionally transient (less than one year) (Thompsonet al. 1997), so regular disturbance is essential in order to keep the habitat sufficiently opento permit germination, establishment and repeated successful seed production (Grime et al. 1988,2007).
Fossil record
Seeds of C. minus have been identified from the end of the Weichselian glacial period in Kent, SEngland, but there are no fossil records from the current interglacial (Godwin 1975). This has led tothe species being designated as an archaeophyte, ie an ancient introduction, rather than it having anyclaim to native status.
Fermanagh occurrence
As the tetrad distribution map shows, C. minus has been recorded from a total of just ninescattered squares in Fermanagh, only four of which have post-1975 records. In some sites, it is onlyrepresented by around ten individuals, eg on gravel at Newtownbutler Football Ground car park near theold railway station, where it has persisted for over 60 years.
Irish occurrence
C. minus had been recorded up until the advent of the herbicide era in the late 1940s or early1950s in almost all Irish VCs, but since then it has declined to greater and greater rarity. Althoughthe New Atlas hectad map shows the species still widely scattered across the island, there arerelatively few modern records in comparison with the situation in England and Wales. In the north ofIreland, and especially where cross-border transport was concerned, this species decline also in partreflects the widespread dismantling of the railway system that took place in the 1940s and early 1960s,since the slipstream of trains was a significant seed vector.
British occurrence
C. minus is widespread in Britain N to the Tay, commonest in S England, but absent from much of N& C Scotland. It has very much declined elsewhere and has made a major retreat from farmland acrossthe whole of Britain due to agricultural intensification involving commercial seed cleaning andwidespread use of herbicide. The principal habitats Small Toadflax now maintains are disturbed sites onwaysides, in quarries and especially along railway lines and in station yards (A. Horsfall, in: Prestonet al. 2002).
European and world occurrence
C. minus is a European temperate species, widespread in Europe apart from the Mediterraneanislands (though it has been recorded in Sardinia and in Crete). It is absent also from Russia, Icelandand the Faeroes. It does, however, occur in parts of N Africa, and in W Asia as far east as the Punjab(Clapham et al. 1987). It is introduced in N America and is quite widespread across NE & Cparts, becoming rare towards the W coast (Hultén & Fries 1986, Map 1634; Grime et al. 1988,2007).
Names
The genus name 'Chaenorhinum', sometimes spelt with a double 'r' as 'Chaenorrhinum', is a word formed inanalogy with 'Antirrhinum', from the Greek 'chaeno' or 'chaino', meaning 'splitting' or 'gaping',referring to the open throat of the corolla which distinguishes the genus from both Antirrhinumand Linaria (Gilbert-Carter 1964; Stearn 1992). The Latin specific epithet 'minus' translates as'small', 'smaller' or 'lesser' (Gilbert-Carter 1964; Gledhill 1985).
Threats
None.
Introduced, very rare. Eurosiberian southern-temperate, but widely spread with agriculture in bothhemispheres.
25 August 1984; Northridge, R.H.; disturbed ground (a newly sown lawn) in Enniskillen Town.
Growth form, status and preferred habitats
A variable, erect, 8-60 cm tall, bright pink flowered, summer annual that germinates in May or June,M. orontium (previously known as Antirrhinum orontium L.) is covered with short stickyglandular hairs, at least in its upper half, and develops a tap-root and fibrous side-roots. The plantis simple or branched, sometimes right from the base, and leafy, the leaves opposite below, alternateabove. The leaves are linear or narrowly elliptical, the blade 30-70 × 1.5-5.0 mm, obtuse at apex. Theleaf margin is entire and recurved, the midrib and fine veins prominent below (Garrard & Streeter1983; Sell & Murrell 2007).
M. orontium featured on the list of 41 species previously assumed to be native in Britain whichWebb (1985) considered more probably introduced by man and their status requiring further study. Theeditors of the New Atlas subsequently agreed with Webb's assessment and it is now regarded as anarchaeophyte, an ancient introduction, very probably of Mediterranean origin (Hultén & Fries 1986).
A weedy plant of arable fields on light, sandy, well-drained soils, M. orontium also frequentsroadsides and waste ground (Garrard & Streeter 1983).
The species has declined sharply as an agricultural weed throughout Britain during the last 50 years orso and it has almost, but not quite, disappeared from Ireland altogether where it has always beencasual.
Flowering reproduction
The plant flowers from July to October, producing a terminal raceme inflorescence of subsessile,solitary, bisexual flowers in the axils of bracts similar to the leaves but smaller, exceeding theflowers. The flower pedicels are 2-3 mm long, sepals narrow-ovate to linear, 10-17 mm long. The tubularcorolla, 10-14 mm long along the upper side, equalling or shorter than the calyx, reddish pink or rarelywhite, is two-lipped and broadly saccate or pouched at the base. The mouth of the corolla tube istightly closed by the palate swelling on the 3-lobed lower lip. Nectar is secreted by a swelling at thebase of the ovary and the flower is pollinated by bees or self-pollinates (Hutchinson 1972; Garrard& Streeter 1983; Sell & Murrell 2007).
The fruit is an obliquely ovoid-ellipsoid capsule, 9-10 mm long, thin-walled, dark brown, covered by amixture of short and long glandular hairs. It opens at the apex by two or three small pores per cell torelease the numerous small black seeds (Butcher 1961).
The seeds, 0.9-1.1 mm long, are somewhat flattened, with one face smooth, keeled and produced into anarrow wing: the other face is finely tuberculate or pitted and it has a wide, raised, sinuate border(Sell & Murrell 2007).
The average number of seeds per plant is around 124, although the plants vary greatly in size and in thenumber of fruits they produce. Salisbury (1964) found that individuals producing but a single capsulewere frequent, but a large plant could produce as many as 76. In his study, the average for over 200specimens was twelve capsules and the mean reproductive capacity of the plants was calculated to liebetween 800 and 1,200 seeds. Seeds remain dormant overwinter and germinate in late spring (May or June).A wet spring can prevent germination and seriously affect local survival of the species since it doesnot appear to be long-persistent (Salisbury 1964).
Fermanagh occurrence
A single Fermanagh record exists for this alien casual species. In this instance, it was almost certainlyaccidentally introduced with recently sown garden lawn seed.
Irish occurrence
Previously, M. oronticum was a rare, casual weed of arable agriculture in five Irish VCs in the S& SE of the island (Cen Cat Fl Ir 2). It is now established only in Co Cork (H3-H5),but even there it is regarded as rare and declining. Elsewhere in SW Ireland, it is a rare, or anextremely rare casual (An Irish Flora 1996).
British occurrence
In the past, Weasel's-snout was most frequent on the sandy or gravelly soils of East Anglia, Surrey (VC17), Hampshire (VC 11), Dorset (VC 9), SW England and SW Wales. Although it may remain locally frequentin these areas, it has declined sharply over the last 50 years, very probably as the result ofwidespread use of broad spectrum herbicides to which it is susceptible, the advent of stubble burningand the autumn sowing of crops such as winter wheat and oil seed rape (Garrard & Streeter 1983; A.Horsfall, in: Preston et al. 2002). It survives mainly in the far south of England and in coastalareas of Wales, having become rare, scattered and casual, if not actually extinct, elsewhere.
European and world occurrence
This variable species probably originated in the Mediterranean region where it is very abundant. It is aprevalent weed throughout Europe and SE Asia, extending eastwards to the Himalaya. It also occurs inMadeira and the Canary Isles. M. oronticum has also been spread with agriculture as a crop seedcontaminant to many parts of the world, including C & S Africa, N America (not very widespread),Australia and New Zealand (Hultén & Fries 1986, Map 1633).
Names
The derivation or origin of the genus name 'Misopates' is a tricky question that all books that thecurrent author (RSF) has consulted on plant names appear to avoid. Looking for Greek word elements for'miso-' there is a choice between 'miskos', 'a stem', or 'misein', 'to hate', ie antagonistic tosomething. For '-pates' there is the Greek 'pathos', 'suffering' or 'feeling', Greek 'patos', 'bottom',or 'path', or Greek 'pater', 'father'. In the opinion of the current author, none of these make muchsense when combined, nor do they convey any information regarding the plant in question. The Latinspecific epithet 'orontium' is a bit easier, since there is a river of that name in Syria that gave itsname to a genus of plant from the area, which is mentioned in Galen (Gilbert-Carter 1964; Gledhill1985).
English common names include 'Weasel's snout' and 'Calf's snout' or 'Calves' snout',referring to the elongated blunt shape of the somewhat hairy fruit capsule. Other names are 'CornSnapdragon' and 'Lesser Snapdragon', referring to the quite obviously similar flower shape toAntirrhinum majus (Snapdragon) and the weedy nature of the plant.
Threats
None, since the species is not persistent.
Introduced, neophyte, garden escape, occasional, locally frequent.
1900; Praeger, R.Ll.; Co Fermanagh.
March to December.
Growth form and preferred habitats
A slender-stemmed, hairless or almost so perennial with fibrous roots, the stems of C. muralis canvary from 10-80 cm long, are pale green in colour, often suffused with purple and mainly creep andtrail, rooting at intervals, in and along old walls with lime mortar. The species also occurs much morerarely on limestone rocks and screes in natural habitats, as it does locally in Fermanagh at KnockninnyHill and on the cliffs and screes above Doagh Lough.
An early 17th century garden introduction from the Mediterranean to these isles (possibly dating from1602), it was probably deliberately planted for decoration in walled gardens for its small, butplentifully produced flowers over a long season from May to November, although the main period is fromMay to September. The irregular toadflax-like flowers are pale mauve in colour, marked with a white andlemon-yellow swollen (personnate) lower lip. The flowers, which are borne singly in the axils of theleaves, also each bear a short, curved spur. The leaves are pale yellowish-green, sometimes suffusedpurple. They are thick in texture, usually 5-lobed, ivy-shaped and borne on long stalks. The leavesusually are alternate and their undersurfaces are a reddish-purple colour (Grieve 1931; Salisbury 1964;Ellis 1993; Sell & Murrell 2007).
After its escape from cultivation, which is believed to have first taken place around 1640, C.muralis spread widely in lowland B & I and is sometimes abundantly present, or as Salisbury(1964) so beautifully describes it, [it] "not infrequently exhibits an undesirable luxuriance towhich the popular names of Roving Jenny and Mother-of-thousands pay recognition.". Clearly fromthis, C. muralis is an aggressive coloniser of suitable old lime-rich walls, pavement crevices,rocks, screes and quarries, habitats where it can then readily establish and may persist indefinitely(Grigson 1987; Ellis 1993). The established strategy of C. muralis is categorized as R/CSRmeaning intermediate between Ruderal and a more balanced mix of all three strategies, Competitor,Stress-tolerator and Ruderal (Grime et al. 1988, 2007).
The spread of the species once it has arrived on a wall or rock is easily understood as the fruit-bearingstalks bend away from light and thus position the capsule, containing up to 40 very small seeds, in darkcrevices (Salisbury 1964).
It is remarkable in being one of the very few species which plant physiologists categorise as behaving ina 'negatively phototropic' manner, since almost all other plants react to directional light by bendingtowards it (ie they are positive in their 'tropic' response to light).
Pollination, fruit, seed and dispersal
The flowers are pollinated by Hymenoptera, mainly bees and if this does not occur, they self-pollinate(Fitter 1987; Sell & Murrell 2007). The ripe, globose, fruit capsule opens by two side pores, themargins of which usually split into six teeth: the three teeth on the lower side are more prominent andthe three upper ones are smaller (Clapham et al. 1987). Each capsule usually contains 15-40 dullblack, rounded seeds that bear thick curved (flexuous) ridges on their surfaces and are 0.6-0.8 mm indiameter (Salisbury 1964; Clapham et al. 1987).
Dispersal of C. muralis to new, distant sites is presumably by wind on account of the small sizeand lightweight nature of the seeds, though Salisbury (1964) suggests birds probably also effectcarriage. Having said this, Ellis (1993) notes that Ivy-leaved Toadflax is one of those alien invadersthat, while common and widespread throughout B & I, has maintained a close relationship with man andis rarely found any great distance from human habitation.
Germination takes place in both spring and autumn and the percentage success is high. Seed is the normalmethod of reproduction but, in addition, plant fragments can root readily under favourable conditionsand achieve some measure of vegetative reproduction (Salisbury 1964).
Variation
There is sufficient natural variation within the species for two subspecies to be recognised by Stace(1997) and Sell & Murrell (2007), differing with respect to hairiness. However, since subsp.visianii (plant hairy in ± all of its parts) has so far been recorded in only a handful of sitesincluding Wisley in Surrey (VC 17) (Clement & Foster 1994), the plant has not yet beendifferentiated beyond the species level in Fermanagh!
Albino forms with no red pigment in the stems, leaves and flowers occur and are also said to be notuncommon (Ellis 1993).
Fermanagh occurrence
C. muralis has been recorded in 48 widely scattered tetrads in the VC (9.1%), although notunexpectedly, these are mainly in the lowlands near habitation and only much more rarely is it found innatural or semi-natural limestone habitats.
British and Irish occurrence
Ivy-leaved Toadflax is very common, widespread and well-established on suitably dry or well-drained,calcareous rocky, stony, wall or pavement habitats throughout lowland B & I.
European and world occurrence
C. muralis is native in the southern Alps, W Yugoslavia, C & S Italy and Sicily. It has beencultivated and is now naturalised on walls and more rarely on stony ground or shingle, across most of S,W & C Europe from Holland southwards (Grieve 1931; D.A. Webb, in: Tutin et al. 1972; Claphamet al. 1987; Sell & Murrell 2007).
Uses
Ivy-leaved toadflax has anti-scorbutic properties and being acrid and pungent has been eaten like cressin salads in southern Europe (Grieve 1931).
Names
The genus name 'Cymbalaria' is said to be from Greek 'kymbalon', or Latin 'cymbalum', meaning 'cymbal' or'cymbal-like', supposedly from the form (shape) of the leaves (Gilbert-Carter 1964; Gledhill 1985;Stearn 1992). As the leaves of C. muralis are definitely and quite obviously 'ivy-like', thecurrent author does not follow this argument at all, but it is possible that the leaves of some of theother 14 species of the genus are cymbal-like. The Latin specific epithet 'muralis', 'growing on walls',fits perfectly (Stearn 1992).
Some of the many common names the species has accumulated in the last 400 years indicate its fecundity,for example, 'Mother of thousands' and 'Mother of millions', or even, 'Fleas and lice' (Grigson 1955,1987)! In Italy, it is the 'Plant of the Madonna', which again may reflect fertility, as perhaps doesanother name 'Travelling sailor’, which at the same time refers to its considerable dispersal ability(Mabey 1996)! Grigson (1955, 1987) lists a total of 28 English common names, many of them includingword-elements such as 'creeping', 'climbing', 'rambling', 'roving' and 'wandering'.
Threats
None.
Introduction, neophyte, garden escape, very rare.
July 1990; Northridge, R.H.; on a wall in Pettigo village.
Growth form and preferred habitats
An erect, tufted, colourful, attractive perennial, native of C & S Italy and Sicily, Purple Toadflaxhas been commonly grown in gardens in B & I from sometime before 1648 and remains a populardecorative subject with several named cultivars, including a common pink flowered one, cv. 'Canon J.Went' and a rarer white one, cv. 'Springside White' (Griffiths 1994; Buczacki 2007). It was firstrecorded naturalised outside a garden setting in the wild in Middlesex around 1830 and its frequency anddistribution in Britain, at least, has greatly increased in the 40 years between the two BSBIAtlases (Perring & Walters 1962; Preston et al. 2002).
The plant has a glabrous stem, 40-120 cm tall, often branched above, bearing numerous, glabrous,linear-lanceolate leaves up to 4.5 cm long, the lower ones whorled. The nectar spur on the flowers ofboth L. purpurea and L. repens (Pale Toadflax) is curved. L. repens differs fromL. purpurea in having whitish, purple-striped flowers and a less dense inflorescence (Garrard& Streeter 1983).
Purple Toadflax most commonly grows on old walls, dry hedge banks and stony waste ground along roads andin urban pavement and less disturbed ground around railways and quarries (Garrard & Streeter 1983;A. Horsfall, in: Preston et al. 2002). In all its habitat types, it demonstrates both greatability to tolerate stress and an obvious avoidance of biological competition.
Flowering reproduction
Flowering takes place from June to August, the flowers, 6-8 mm long, being borne in long, dense,leafless, spire-like racemes with bracts equalling the flower stalks. On each flower, the five calyxteeth are linear-lanceolate, acute and longer than the tube. The corolla is lilac-purple topurplish-violet, irregular, 2-lipped, personate, and the long, curved spur is about 5 mm long, as longas or longer than the petal limb. The stamens are four in number and are included within the corollatube. The pale purplish style on the globular, superior ovary is single and has a small, green, capitatestigma. The flowers are very attractive to bees that pollinate them and they are self-incompatible (Sell& Murrell 2007; Stace & Crawley 2015). When fertilised, the ovoid capsule is much longer thanthe sepals and it opens with 3-triangular teeth on each of the two cells. The ovoid seeds are 1.3 × 1.0mm, angled, deeply honeycombed and brown in colour (Butcher 1961; Streeter et al. 2009).
L. purpurea seeds freely and prolifically in a garden setting, spreading into paving, gravel,walls and any open ground, so that it can soon become an over-abundant, weedy pest (Stace & Crawley2015).
Fermanagh occurrence
There are just three records in the Fermanagh Flora Database, all found by RHN from 1990 onwards. Theyare from perfectly typical habitats of the species in these islands, ie walls and dry stony waste groundwhere they are definite garden escapes or discards. The remaining record details are: stony ground atLisbellaw village, July 1994; and on pavement at Windmill Heights, Enniskillen town, May 2001.
British and Irish occurrence
L. purpurea naturalises itself widely in these islands and can become established. Previously ithad been listed as occurring at least once from eleven Irish VCs (Cen Cat Fl Ir 2; FNEI3), but Reynolds (Cat Alien Pl Ir) added a further seven, which together with Fermanaghnow makes a total of 19 of the 40 VCs on the island with records.
The New Atlas hectad map shows that in Ireland and Scotland, L. purpurea is much morescattered, occasional and urban in its distribution than is the case in lowland England and Wales,although Beesley & Wilde (1997) reported it widespread and quite common in urban Belfast, beingrecorded in 34 1-km squares in the city. Similarly, in their survey of urban inner Dublin, Wyse Jacksonet al. (1984) reported L. purpurea locally frequent, mainly S of the River Liffey.
In England, L. purpurea is recorded at the hectad level as being almost omnipresent in the area SEof a line drawn on the hectad map between Bristol and Bridlington and generously scattered in thelowlands northwards to beyond Inverness, chiefly on the eastern side of the island. It is also wellrepresented in Wales, especially in coastal and urban areas of the country (A. Horsfall, in: Prestonet al. 2002).
The considerable increase in the species British presence in the 40 years between the two BSBIAtlas surveys probably reflects both a genuine spread of the plant and a better recording effortdirected at introduced species, a feature that is now noticeable with most garden plants (Clement &Foster 1994).
Threats
None.
Native, common and widespread. Suboceanic southern-temperate, but very widely naturalised in bothhemispheres.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
A large, usually biennial, rarely perennial, monoecious, tap-rooted, pubescent to lanate, rarelysubglabrous, rosette-forming herb up to 200 cm tall, D. purpurea is winter-green and occasionallyflowers in its first, rather than second, year of growth (Grime et al. 1988, 2007; Sell &Murrell 2007). The stem is simple or sparsely branched, round in section and grows erect from a rosetteof oblong-lanceolate, acute, crenate-toothed, softly pubescent, grey-green leaves. The radical leaveshave a long-winged petiole, while the stem leaves are sessile.
It is widespread and common to abundant on dry or well-drained acid soils, particularly after they aredisturbed, such as by woodland or scrub clearing, fire, or more intermittently as the result of animalscrapes or trampling. Foxglove also occurs, although much more rarely, in limestone districts. Typicalhabitats include wood, open heathland, scrub margins and clearings, hedgebanks, roadside verges, rough,occasionally mown or grazed grasslands and waste ground, especially when it is near habitation. D.purpurea is tolerant of considerable levels of shade but grows most rapidly in fully-litconditions. It can also survive and develop to the flowering stage on stony, rocky, rather dry, shallowsoil conditions, including on mountain- and sea-cliffs and on walls. It is widespread and common onground up to around 885 m in B & I (Sell & Murrell 2007).
The established strategy of D. purpurea is categorised by Grime et al. (1988, 2007) asSR/CSR, ie it is intermediate between a Stress-tolerant Ruderal and a more balanced mixture ofCompetitive, Stress-tolerant and Ruderal strategies altogether.
Variation
This extremely variable, polymorphic species originated in the W Mediterranean and taxonomically is atit* most complex in the SW part of that basin, there being up to five different subspecies named, plus anumber of local variants based largely on plant height, leaf size and shape, degree of hairiness(indumentum), calyx, corolla colour and spotting. Only the most widespread form, subsp. purpurea,occurs in B & I.
Some local populations demonstrate combinations of these variable characters, but the pattern ofvariation is too complex and changeable (fluctuating) to allow them to be regarded as subspecies. Inaddition, dwarf alpine ecotypes occur and regional differences exist between populations on the Iberianpeninsula and the rest of Europe (V.H. Heywood, in: Tutin et al. 1972; Hultén & Fries 1986).
Garden cultivars 'Alba', 'Apricot', 'Campanula' and 'Dwarf Sensations' occur, together with a range ofnamed garden hybrids, including 'Excelsior Hybrids' and 'Foxy Hybrids' (Griffiths 1994).
Fermanagh occurrence
When precipitation is heavy and frequent, as is very much the case in Fermanagh, lime is graduallyleached from soil surface layers and they become increasingly acidic and peaty, even when borne directlyover limestone rock. Thus, D. purpurea is not localised in Fermanagh, but rather it is scatteredwidely across 238 tetrads, representing 45.1% of the VC total.
Flowering and vegetative reproduction
Reproduction is almost exclusively by seed although damaged plants can asexually develop daughterrosettes. Flowering takes place from June to September. The average plant bears a terminal, unilateral,spike-like raceme inflorescence with around 60 irregular, downward-directed (pendulous), pale to deeppinkish-purple, bell-shaped flowers with deeper purple spots on a ciliate white ground inside. Theflowers, with a corolla 40-55 mm long, produce nectar and attract bumble bees that pollinate them, butif this fails, then the flowers are self-compatible and fertilise themselves (Hickey & King 1981).
Seed production and dispersal
An average plant of D. purpurea is capable of producing around 85,000 tiny seeds to add to thesoil seed bank and large plants can generate between 100,00-500,000 (Salisbury 1942; Richards 1997b).The fruit capsule, 11-13 mm in length and longer than the calyx, contains a mean of around 1,030 seedsand germination in light is 95% (Salisbury 1942). Seed dispersal is by wind. "The capsule, at firstconic, splits into halves, each half dehiscing again on the inner face, but remaining so closelyconnected with the other half at the base that the seeds collect near the bottom and are shaken out ofthe cup so formed in small quantities at each puff of wind." (Ridley 1930).
Seed longevity
The seed of the species has a reputation of being long-persistent in soil (Grime et al. 1988,2007), but the survey of soil seed banks of NW Europe found a range of values: three estimates suggestedthe seed was transient (surviving less than one year), eight studies considered it short-term persistent(less than five years), four reckoned it was long-term persistent (more than five years) and threeestimates could not assign their samples to one of the three seed bank types (Thompson et al.1997).
A complicated reproductive strategy
Being monocarpic, the parent plant dies after flowering and although often described as a biennial, inreality in many less-favourable habitats D. purpurea may take more than two years growth to reacha size and have accumulated the photosynthetic energy resources to trigger successful flowering andfruiting. Perhaps it is best considered as a short-lived, once-flowering (ie technically speaking'hapaxanthic') perennial [from the Greek 'hapax' = 'once' and 'anthos' = 'flower'] (Holmes 1979).
Even this, however, does not fully cover the potential reproductive strategy of D. purpurea, sincevery occasionally individual Digitalis plants (particularly those regularly monitored in acultivated garden setting) are known to 'forget to die' after their first seeding. These individuals maycontinue to grow and flower one or two additional times (Cragg-Barber 1996). One such specimen in Bergenflowered three times and it also developed a complicated caudex (ie a thickened, woody, truck-likeappendage) at the base of the stem, which bore many adventitious leaf rosettes, apparently making itcapable of continuing life indefinitely (Faegri 1980). Faegri (1980) even went as far as suggesting thatthe early demise of hapaxanthic specimens of species like D. purpurea might be more due toexternal competition than to internal factors.
Empirical observations indicate that mechanical damage to the flowering stem, sufficient to prevent itfrom setting seed, can lead in about 20% of instances, to specimens producing basal side rosettes andoverwintering to flower the following year. This is particularly the case when considering well-grownplants in sheltered, rich, damp soil situations, Richards (1997b), who made these observations, agreeswith Faegri that the capacity for perennation in D. purpurea certainly seems to be underenvironmental control. However, further observation of related European species suggests thatperennation may be under genetic control as well (Richards 1997b).
Toxicity
D. purpurea is a very poisonous plant, all parts of it being toxic, especially the seeds(Hutchinson 1972). The active toxic principles, digitoxin, digoxin and gitoxin (plus many otherbiochemically similarly substances with names derived from the botanical name of the genus (Cooper &Johnson 1998, p. 197)), are cardiac glycosides that remain widely prescribed in orthodox medicine asheart muscle stimulants.
The toxicity of the plant is unaffected by drying, storage or boiling and, as with Seneciojacobaea (Common Ragwort), stock fed hay containing Foxglove may be poisoned. Animals usuallyknow to avoid the plant, but if alternative grazing is scarce, they may eat it and can develop a cravingfor the plant after being poisoned (Cooper & Johnson 1998, p. 63).
British and Irish occurrence
D. purpurea is common and widespread, and locally abundant on suitable soils throughout most ofBritain and most of Ireland, although it is purely an introduction on Shetland (VC 112). In Ireland, itappears scarce, rare or absent in a band across the island from Dublin to Co Clare (H21-H9), whichrepresents the underlying region of most limestone geology, although many large local peat bogs alsooccur in this area of the country (New Atlas).
In B & I, decorative forms of D. purpurea are also widely cultivated in gardens, sometimesinvolving imported foreign seeds from major horticultural suppliers; some of these forms are believed toescape and may become naturalised (Reynolds 2002).
European and world occurrence
Native in W, SW & WC Europe, D. purpurea is also found in Morocco. It belongs to thesuboceanic southern-temperate phytogeographical element and has been widely introduced and cultivatedelsewhere for ornamental and medicinal drug purposes (Clapham et al. 1987).
Subsp. purpurea covers most of the European range of the species; subsp. mariana (Boiss.)Rivas Goday occurs in S Spain, subsp. dubia (Rodr.) on the Balearic Islands and subsp.mauretanica Maire & Emb. in the Atlas Mountains.
As a medicinal and ornamental plant, D. purpurea has been spread well beyond its original range inW Europe and is now naturalised in parts of C & N Europe, N & S America, S Australia and NewZealand (Hultén & Fries 1986, Map 1638).
Medicinal uses
Foxglove was used by old herbalists for various purposes, most of them entirely remote from the valuableproperties used in modern scientific medicine. Gerard (1597, 1633) recommended it for falls from highplaces, and the expressed juice of the leaves was used in the form of an ointment to treat scrofulousswellings. Bruised leaves were applied to clean old sores and ulcers. The plant was also boiled in wineand used as an expectorant (Dodoens 1554). The leaves alone are used to manufacture the drugs for whichthe plant is now used in the treatment of heart conditions.
The plant contains important glucosides, three of which are cardiac stimulants. The most powerful,digitoxin (also referred to as 'digitalis'), is an extremely poisonous and cumulative drug, insoluble inwater; digitalin is crystalline and also insoluble in water; and digitalein, which is amorphous butreadily water soluble and therefore can be administered subcutaneously in very minute doses. Dosage isextremely important in this latter case, as too little of the drug is ineffective, and a slight excesscan prove fatal. Another ingredient, digitonin, is a cardiac depressant ± identical to saponin, thechief constituent of the Senga root (Grieve 1931). Digitoxin is used as a medication for heart failure.Its clinical use was pioneered by William Withering who in his Account of the foxglove andsome of its medicinal uses etc. (1785) recognized that it reduced dropsy, increased urine flowand had a powerful effect on the heart.
Names
There are 91 English common names listed for D. purpurea by Grigson (1955, 1987), ranging from thefamiliar 'Foxglove' to 'Dead Man's Bells', and 'Hedge Poppy' to 'Poppers' and 'Scotch Mercury' to'Witch's Thimble'. Many of the names are variations on the idea of the flower corolla being like afinger-stall and they were often playthings of children in this way, despite the extremely poisonousnature of the species. The fact that some names referred to 'Deadmen's fingers' and Deadmen's Thimbles'suggests an awareness of the toxic content of the plant tissues.
There are numerous references to flowers that 'pop', such as 'Flop-a-dock', 'Flop Poppy' and 'Flops'. Theconnection with foxes is also frequent, including 'Fox-and-leaves', 'Fox Docken', 'Fox Fingers','Foxter' and 'Foxy', possibly derived from the notion that the plant grew from fox droppings (Grigson1955, 1987).
Foxglove was also considered a fairy plant, or a goblin's plant with magical connections that had to berespected. There were also supposed to be dangers that had to be evaded, including avoiding cutting theplant, and connections with changlings and missing children.
Threats
None.
Native, common and widespread, locally abundant. Circumpolar boreo-temperate, also widely naturalised inboth hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This small, creeping perennial has hairy stems, 5-15 cm long, that lie prostrate or decumbent, rooting atthe nodes, but branches are ascending, forming dense, leafy tufts of growth. Leaves are opposite,shortly stalked, blades 5-20 × 4-15 mm, leathery, broadly ovate or oblong-ovate, rounded at their apexand also at the base, margins entire or shallowly crenate, glabrous or almost so. Flowers are rathernumerous in terminal leafy racemes, the subtending leaves gradually decreasing in size upwards andbecoming alternate. Typically, the lower flowers are in fruit before the upper have finished (or evenbegun) blooming (Hutchinson 1972; Sell & Murrell 2007).
It generally appears sparse to abundant in a very wide range of open, more or less disturbed, damp, acidhabitats. These include poorer grassland, grazed water meadows, fens, heaths, damp garden lawns,clearings in woods or their margins, plus any bare, damp, disturbed soil along tracks and on wasteground. The plant is tolerant of soil chemistry, moisture status, medium shade, grazing pressure andtrampling. In mesic conditions, about the only thing it cannot cope with is competition from tall orvigorous, tight turf-forming grasses and herbs, since it requires more open, 'gappy' conditions thanthese plant associates allow (Sinker et al. 1985).
The established strategy of V. serpyllifolia is categorised as R/CSR, meaning it is intermediatebetween a straight Ruderal species and a more balanced mix of all three strategies, Competitor,Stress-tolerator and Ruderal (Grime et al. 1988, 2007).
Flowering reproduction
The species has a very long flowering period, almost all year round, although the main period is fromMarch to October, and the peak lies between May and July. The small bisexual flowers, 5-10 mm indiameter, borne in lax leafy racemes, have calyx lobes linear, obtuse, glabrous, and the 4-lobedcorolla, divided almost to the base is white, pale blue or bright blue, striped with darker lines (Sell& Murrell 2007). The two anthers lie close together either side of the small, capitate stigma, sothat insect visitors (usually flies) may effect cross- or self-pollination (Hutchinson 1972; Garrard& Streeter 1983). Insects visiting Veronica species grasp the bases of the two stamens forsupport, pulling them together and, in so doing, they dust themselves and the stigma standing betweenthem with pollen (Proctor & Yeo 1973; Hyam & Pankhurst 1995).
The fruit is a flat, somewhat rounded, notched capsule, very shortly glandular-hairy and tipped by theslender persistent style. The ripe capsule dehisces into two valves, usually apically and the flat,ovoid seeds, 0.7-0.9 mm, are pale straw-coloured (Sell & Murrell 2007).
Apart from being attached and transported in mud, seed is known to be dispersed internally by browsinganimals, including cattle, horses and goats (Salisbury 1964, p. 103).
There are a total of 37 estimates of seed longevity in the soil seed bank survey of NW Europe, of whicheight consider it transient (surviving one year or less), ten reckon it is short-term persistent(survives less than five years), eleven regard it as long-term persistent (surviving at least fiveyears) and eight found seed present but could not assign it to one of the three seed bank types(Thompson et al. 1997).
Identification
The species has glabrous, almost completely untoothed leaves, prostrate, spreading stems that root at thenodes and branches bearing terminal racemes of pale blue flowers with styles 2.5 mm long that, takentogether, distinguish V. serpyllifolia from both V. arvensis (Wall Speedwell) and V.officinalis (Heath Speedwell). The bracts of the flowers are longer than the pedicels, which inturn are longer than the calyx (Garrard & Streeter 1983).
Variation
V. serpyllifolia is a rather variable plant of which at least 20 variations have been described(Hultén 1974). Two subspecies occur in B & I, but as elsewhere in Ireland, only the more commonform, V. serpyllifolia subsp. serpyllifolia, has been found in Fermanagh, and mostrecorders do not work at the subspecific rank. The other subspecies, subsp. humifusa (Dicks.)Syme is an arctic-montane form that belongs to the discontinuous circumpolar plants (Hultén 1974, p. 260& Map 252). It has rooting stems and broader, almost entire leaves and larger, bluer flowers. InBritain, it is a scarce plant of damp places in mountains (Garrard & Streeter 1983).
Fermanagh occurrence
V. serpyllifolia is very common and widespread, occurring in 346 Fermanagh tetrads, 65.5% of thosein the VC.
British and Irish occurrence
Widespread and common throughout B & I, the New Atlas hectad map showing V.serpyllifolia is almost omnipresent in lowland Britain and at altitudes of up to around 825 m.The distribution is stable, but thins somewhat to the N & W in Scotland where wetter, more acidicpeatlands predominate. In Ireland, the picture is similar, the distribution almost complete at thehectad level of discrimination, but thinning in the Central Plain and in the western coastal region,where very wet, strongly acidic peatlands become the dominant vegetation under a hyper-oceanic climateregime (A. Horsfall, in: Preston et al. 2002).
European and world occurrence
V. serpyllifolia subsp. serpyllifolia originated in the temperatepart of Eurasia, from where it has spread northwards as a weed throughout almost all of Europe, fromIceland to Scandinavia (71°N) and south to C Spain and Portugal, Sicily, Albania and Thrace. It is alsopresent in the mountains of N Africa and has been spread as a weed to many areas around the globe,including N & S America (both E & W USA), E Asia, Australia, New Zealand, Madeira, the Azoresand Canary Islands, Tristan da Cunha and St Helena, S Greenland and doubtless in other places in itspresent range (Hultén 1974, Map 251; Hultén & Fries 1986, Map 1640; Clapham et al. 1987).
Names
The origin of the genus name 'Veronica' is problematic. It may be named in memory of St Veronica, who themyth has it, wiped the sweat from Christ's face (Gilbert-Carter 1964; Gledhill 1985). Alternatively,some authorities give it as a corruption of 'Betonica', plants of the two genera having some degree ofsimilarity in their leaves. Other suggestions are that the name is a Latin form of the Greek word'Beronike', or else refer it to the Greek 'hiera eicon', meaning 'sacred image', or the Arabic 'viroonikoo', 'beautiful remembrance' (Johnson & Smith 1946). In truth, the origin or derivation isunknown.
The Latin specific epithet 'serpyllifolia' translates as 'Thyme-leaved' (Gledhill 1985), or (better)'with leaves like Thymus serpyllum' (Stearn 1992).
The English common name 'Speedwell' was first recorded for V. officinalis at the end of the 16thcentury by Lyte (1578) but since then its use has been extended to numerous (or all) other species ofthe genus (Britten & Holland 1886). The healing medicinal properties of the genus Veronicawere linked to Wales by Gerard (1597, 1633), where he said it was known by the name 'Fluellen'. He wenton to create confusion by extending this Welsh name to other Veronica species and to two Britishspecies of another genus, Kickxia, as well (Grigson 1974; Allen & Hatfield 2004).The meaningor 'sense' for a medicinal plant means 'prosper well', or 'go on well' or just 'get well soon'. V.officinalis was regarded as a significant strengthening and wound-healing herb, also said to begood for treating coughs (Grigson 1974).
Threats
None.
Native, common and widespread. Eurasian boreo-temperate, widely naturalised in both hemispheres.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This is a low, creeping, branching, rooting and mat-forming, hairy perennial with opposite, elliptical,coarsely-toothed, nearly stalkless leaves. The leaves are wedge-shaped at their base, rounded or with abroad, triangular top and the leaf blades are very variable in size, 5-30 × 3-30 mm. V.officinalis grows only around 15-25 cm tall and its slender, straggling stems are thinlypubescent with rather long hairs all around.
V. officinalis is common and widespread in a varied range of habitats including open woodland,short turf on dry, grassy areas of heaths, stony banks and pastures at all levels, by mountain streamsup to around 900 m, as well as on cliffs, screes and walls on moderately acid, well-drained, sandy,stony or loamy, often leached soils, including by roadsides (Webb & Scannell 1983; Sinker etal. 1985; A. Horsfall, in: Preston et al. 2002). It also regularly occurs on slightlybetter drained raised mounds and on anthills in calcareous grasslands. In fact, despite its common namesuggesting a rather specific heath habitat, it can be found in short turf almost anywhere, except on thevery highest or the wettest ground.
The plant can spread vegetatively and often forms large clonal mats in suitable short turf, where ittolerates both heavy grazing and moderate soil disturbance from trampling (Sinker et al. 1985).The established strategy of V. officinalis is categorised as SR/CSR meaning it is consideredintermediate between a Stress-tolerant Ruderal and a more balanced combination of all three plantstrategies, Competitor, Stress-tolerator and Ruderal (Grime et al. 1988, 2007).
Variation
This is a very variable species with respect to both size and degree of hairiness. Plants with thelargest leaves and the greatest development of hair appear most frequently in open woodland conditions,while plants on the coast, around the mountains and on acid heaths tend to be smaller than average (Sell& Murrell 2007).
Fermanagh occurrence
While as the accompanying tetrad map indicates, Heath Speedwell is common in a wide range of localhabitats liberally scattered across the VC, it is neither as widespread nor as frequent as V.serpyllifolia (Thyme-leaved Speedwell) (see the current author's account on this website). Inthis case, V. officinalis has been recorded in just 204 tetrads, 38.6% of the VC total, incomparison with 346 tetrads, 65.5% of squares for V. serpyllifolia.
Flowering reproduction
V. officinalis produces dense, spike-like, erect, stalked racemes of 15-25 small flowers, 6-7 mmin diameter, with lilac-blue or violet-blue corollas marked with dark blue veins, borne in the axils ofleaves, rather than terminal on the stem (Garrard & Streeter 1983; Sell & Murrell 2007).Characteristically, the pedicels are 1-2 mm long, shorter than both the bracts and the pubescent,4-lobed flower calyx (2.0-2.5 mm). The flowers are borne from May to August and they produce nectar toattract various small insects that pollinate them, or else they self-pollinate as the flower fades, thefilaments of the two stamens twisting inwards and downwards as the corolla withers until the dehiscedanthers touch the stigma with their pollen (Hutchinson 1972).
The fruit capsule is partially flattened, heart-shaped, shortly pubescent and it exceeds the calyx inlength. It dehisces into two valves, opening apically. The seeds, 0.8-1.2 mm, are yellow or light brown,sub-round or elliptical, flat and smooth (Butcher 1961; Hutchinson 1972; Sell & Murrell 2007). Inone English study, the mean seed production of V. officinalis was around 8,700 per plant(Salisbury 1942, p. 228).
There does not appear to be any specific dispersal mechanism mentioned in the literature – a topicawaiting study – but the very small, lightweight seeds are undoubtedly lifted out from the open capsuleby the slightest breeze passing across it. It is also very probable that seeds of plants growing on ornear anthills are distributed by the insects, as is known to occur in other parts of the world (Ridley1930).
British and Irish occurrence
Widely distributed throughout the whole of both islands but, while the overall distribution is littlechanged since the 1962 BSBI Atlas (Perring & Walters 1962), there have been local declinesdue to contraction in range caused by habitat loss, especially in the English Midlands. In Ireland, itis very frequent, locally abundant, but apparently less so in the centre of the country (Webb etal. 1996; New Atlas; Parnell & Curtis 2012).
European and world occurrence
V. officinalis belongs to the European boreo-temperate phytogeographical element and occurs inwoods and heaths almost throughout Europe from Iceland to the Mediterranean basin and the Azores. Italso occurs in Asia Minor and the Caucasus and has been widely introduced, including to eastern NAmerica and to New Zealand, where it is extremely rare or now extinct (Hultén & Fries 1986, Map1645; Webb et al. 1988; Sell & Murrell 2007).
Medicinal uses
This species of Veronica retained a place in herbal medicine until quite recent times due to itspossessing astringent compounds, including a glycoside called aucubine, tannins and several acids, whichgave it diuretic, expectorant and tonic properties. According to Grieve (1931), it was used forhaemorrhages, treatment of wounds, skin complaints and an infusion of the dried plant treated coughs andcatarrh. On the other hand, Launert (1981) describes the effect of V. officinalis as mildlydiuretic, mildly expectorant and anti-diarrhoeal, and an infusion was recommended for chronic skincomplaints, stomach upsets, arthritis and rheumatic conditions. Allen & Hatfield (2004) reviewed theevidence for medical use of this species in B & I and found most or almost all of it unreliable,book derived recommendations with little or no first hand verified accounts of its use other than fortreating colds in Donegal.
Names
The origin of the genus name 'Veronica' is problematic, and as discussed in the current author's accountof V. serpyllifolia on this website, in truth, the origin or derivation is unknown.
The Latin specific epithet 'officinalis' as usual indicates the medicinal use of the herb in the past, itsupposedly being collected by pharmacists and sold in their shops (Stearn 1992). However, there seems tobe only very slight evidence of any real medicinal use and the name may be another example of adiscredited 'book name'.
Threats
None.
Native, common and widespread. Eurosiberian boreo-temperate and widely naturalised.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
This very familiar woodland margin, hedgebank, rough grassland, road- and lane-side, prostrate ordecumbent, slender perennial, up to 30 cm high, Veronica species is immediately recognised by thetwo distinct rows of long whitish hairs along the opposite sides of the stem, which distinguishes itfrom the similar V. officinalis (Heath Speedwell) and V. montana (Wood Speedwell), whichhave hairs all round the stem (Garrard & Streeter 1983). The beautiful and relatively large, blueflowers, with an obvious white 'eye', are reminiscent of blue eyes, a feature which is frequentlyalluded to in many alternative English common names of the plant throughout the British Isles, eg 'BillyBright Eye' and 'Birdseye Speedwell' (Grigson 1955, 1987). The opposite leaves are almost sessile, thevery short petioles margined with long fine hairs, the blades oval and coarsely or crenately,deeply toothed (Butcher 1961; Hutchinson 1972).
This creeping, stoloniferous perennial is the most common and widespread terrestrial or aquatic Speedwellthroughout the whole of B & I. While it is definitely most commonly found in areas of limestonegeology and soils, Germander Speedwell achieves this degree of dispersal and presence through having awide ecological amplitude, allowing it to crop up and persist in many different habitat types. V.chamaedrys grows in most soil types except those featuring extremes of water, nutrients, acidityor disturbance (A. Horsfall, in: Preston et al. 2002). In the Sheffield area of England, Grimeand his co-workers considered the habitats of V. chamaedrys demonstrated an upland bias ataltitudes up to 410 m, and the species occurred exceptionally at up to twice this altitude (Grime etal. 1988, 2007). On account of the restricted altitudinal range that Fermanagh presents, thesame behaviour is not apparent in H33. Here, in Fermanagh, the only preference V. chamaedrysdemonstrates is towards damp, open, base-rich, infertile calcareous conditions with a pH above 6.0.
Vegetative reproduction
The plant colonises gaps in grass-dominated vegetation by means of its prostrate creeping stolons thatroot at their nodes. It is therefore favoured by disturbance and competes best on less fertile soils andin pastures, both situations where its more vigorous neighbours are somewhat curtailed and fail to oustit. The shallow-growing underground parts comprise tangled, tough stem branches that sprout clusters offibrous roots containing a mycorrhizal associate at the nodes along their length. The species regularlyoccurs in woodland shade often in a non-flowering state and the role of seed in its local dispersal maybe secondary to vegetative reproduction in this and in other disturbed ground situations.
Flowering reproduction
In sufficiently open growing conditions, which occasionally can include garden lawns and flowerbeds,plants of V. chamaedrys flower freely, producing blossoms two at a time on the long 10-20flowered raceme from April to early July (Clapham et al. 1962). Each flower, 10-12 mm diameter,lasts only one or two days before shedding its fused petals and stamens. They are visited by flies andthe small mining and ground nesting bee, Andrena labiata, plus small black Halictus bees,honey bees and hoverflies. Despite their name the latter insects, like the others mentioned, alight onthe two prominent diverging epipetalous stamens, or on the corolla, seeking the nectar secreted by adisc at the base of the ovary and protected by erect bristly hairs around the base of the very shortcorolla tube. The two stamens are lateral and divergent and are seized by the insect alighting on theflower to steady its landing. In doing so, it draws the anthers against its body on which pollen isdeposited and carried to the stigma of another flower (Hutchinson 1972; Proctor & Yeo 1973; Knight1999a).
After fertilisation, the two-celled, flattened, heart-shaped fruit capsule, 5 mm, shorter than the calyx,may contain up to ten small, flat seeds, 1.8 × 1.2 mm, ovoid, smooth and brown (Butcher 1961) which areprobably wind and/or ant dispersed (Ridley 1930, p. 31). In pasture situations, the seed can surviveingestion by cattle, which along with vegetative fragmentation due to trampling may thus provide anothermeans of species transport and dispersal (Ridley 1930, p. 361).
The survey of soil seed banks of NW Europe found numerous reports of seed survival varying from transientto long-term persistent (Thompson et al. 1997). Grime et al. (1988, 2007) categorised theseedling establishment strategy as S/CSR, meaning intermediate between a Stress-tolerator and a morebalanced mix of all three plant strategies, Competitor, Stress-tolerator and Ruderal, ie this indicatesor suggests that the vulnerable seedling stage of the species is somewhat better adapted to long-actingenvironmental stress than it is to immediate competition or heavy or frequent disturbance. These sameworkers, however, point out that regeneration of V. chamaedrys from seed is insufficientlystudied to allow definite pronouncements.
Fermanagh occurrence
In Fermanagh, V. chamaedrys is recorded in 439 tetrads, 83.1% of those in the VC, making it the35th most widespread species in the entire county survey. Germander Speedwell is widespread throughoutthe area, except on bogland, aquatic habitats and the higher mountains. The usual habitats includemarginal rough grassland on hedgebanks, roadsides, on the margins of woods and scrub on lakeshores,pastures, rock outcrops including cliffs, screes and limestone pavement, and also on waste ground anddisturbed soil.
British and Irish occurrence
V. chamaedrys is very widely distributed throughout B & I, but in the N & W ScottishIsles it is almost certainly an introduction (ie on the Outer Hebrides, Orkney and Shetland), allrecords there being confined to disturbed ground near habitation (Pankhurst & Mullin 1991; NewAtlas).
European and world occurrence
V. chamaedrys is native and widespread throughout Continental Europe and boreal Asia, but it isalso anthropophilic, being favoured by early man's landscape clearances and subsequent disturbances. Itis a scattered, established introduction in Japan, the eastern states and W coast of N America, Icelandand New Zealand, where it is occasionally cultivated (Hulten & Fries 1986, Map 1646; Webb etal. 1988, p. 1210).
Medicinal uses
Germander Speedwell had a reputation as a cure for jaundice "throughout Ireland", according toVickery (2019), the flowers being boiled in a little milk and the infusion drunk. The same authorrecords that in Norfolk the species was known as 'Sore-eyes' and the flowers were infused to make asoothing eye-bath. Vickery also mentions that a tea made from the species was esteemed a valuable remedyfor indigestion and stomach pains in Guernsey.
Names
The origin of the genus name 'Veronica' is problematic, and as discussed in the current author's accountof V. serpyllifolia on this website, in truth, the origin or derivation is unknown.
The origin of 'Germander' in the English common name, a name which is also applied to several species ofthe genus Teucrium, including T. chamaedrys (Wall Germander), comes as 'chamaedrys'through medieval Latin 'gamandrea', from the Greek 'khamandrua', or 'chamandrua', being a corruption of'khamai' or 'chamai', meaning 'on the ground', and 'drus', meaning 'oak'. The prominently toothed leavesof T. chamaedrys were likened to dwarf or ground-hugging oak leaves, ie the leaves of the pricklyevergreen Mediterranean shrub, Quercus coccifera, the Kermes Oak. The leaves of V.chamaedrys are thought to resemble those of Teucrium chamaedrys in shape, size andtexture and during the 16th century the name became transferred to it, either as straightforward,'Germander', 'English Germander' or 'Wild Germander'. The current 'recommended' English common name'Germander Speedwell' is a book name, dating only from the 19th century (Prior 1870; Grigson 1974).
The origin of the name 'Speedwell', nowadays applied across the genus, really had its origins in thesupposed medicinal properties of one or a few species of Veronica, such as V. officinalis.It meant the equivalent of 'prosper well', or 'go on well' or just 'get well soon', and may have hadother connotations apart from health, perhaps some magical good luck wish or charm, such as 'good speed'or 'God speed', 'Wish-me-well', or 'safe travel' on a journey (Grigson 1955, 1987).
There are 33 alternative English common names listed by Grigson (1955, 1987) and 15 in Vickery (2019),six of which are additional to Grigson's listing. Many of the names make reference to eyes (Angel's-,Blue-, Bobbies'-, Bird-Christ's-, God's-, Devil's-, Deil-, Cat's-, Lark's-, Milkmaid's-), and sight, orlack of it, eg 'Blind-flower', all referring to the beautiful blue and white 'eye' of the individualflower.
Threats
None.
Native, common and widespread. European temperate.
1881-2; Barrington, R.M.; Devenish Island, Lower Lough Erne.
Throughout the year.
Growth form, recognition and preferred habitats
A widespread but scattered and relatively uncommon, wintergreen, low-growing, patch-forming polycarpicperennial with slender, creeping stems up to 50 cm long, adventitiously rooting at the nodes, branchends curving upwards and ascending to 15 cm. It is like V. chamaedrys (Germander Speedwell) andV. officinalis (Heath Speedwell) in both its hairy appearance and its clonal patch-forming habit,but V. montana has hairs all round the creeping, rooting stems (unlike the two lines of hairs onthe stems of V. chamaedrys). It also differs from V. chamaedrys in having clearly stalked(petioles 6-20 mm), larger leaves and fewer, smaller, paler, lilac-blue flowers per axillary raceme(Melderis & Bangerter 1955; Butcher 1961). However, V. chamaedrys can sometimes occur inwoodland and along overgrown rides and trails and, in damp, shaded conditions, the characteristic twolines of hairs may not develop, the stems becoming ± uniformly hairy all round (R.W.M. Corner, in: Rich& Jermy 1998).
V. montana is also more shade-tolerant than V. chamaedrys and it will flower underdeciduous canopy, though not unnaturally it performs better in this respect and others, when growingunder somewhat better lit, half-shade conditions. It usually occupies damp to moist, lowland woods andsheltered, shaded banks. It prefers moist, seepage areas, avoids the most acidic soils and is oftenfound in coppice situations. While it can be found on damp mildly calcareous soils, it shows a definitepreference for mildly acidic ones (Garrard & Streeter 1983; Grime et al. 1988, 2007).
Despite its specific epithet, V. montana is not a mountain, nor even an upland species at all. Itreaches its highest site at 435 m on Pont y Daf, Brecs., Wales (VC 42) (A. Horsfall, in: Preston etal. 2002). Rather than a montane plant, in some parts of B & I, it is reputed to be a goodindicator species of ancient woodland, of oak, beech, ash and alder, usually on nutrient-rich soils(Crawley 2005). It is certainly rare in unshaded and regularly disturbed ground. Having said that, it isprobably not a very competitive species, appearing to frequent habitats offering less-than-ideal growingconditions that will limit the vigour of accompanying species. The established strategy of V.montana is categorised as SR/CSR, meaning intermediate between a Stress-tolerant Ruderal and amore balanced mix of all three plant strategies, Competitor, Stress-tolerator and Ruderal (Grime etal. 1988, 2007). Regeneration is principally vegetative, the species readily forming spreadingclonal mats. In linear habitats such as streamside banks and along woodland rides, the plant shows anability to spread that is very probably partially achieved by plant fragmentation and re-establishment,as well as by seed.
Flowering reproduction
Wood Speedwell flowers from April to July and in doing so, it shows a remarkable tolerance of woodlandshade, flowering over a wide range of light levels that limits many other associated species includinggrasses such as Holcus mollis (Creeping Soft-grass). Having said this, it flowers best inlightly-shaded sites (Grime et al. 1988, 2007). The flowers, 8-10 mm in diameter, are borne inlax, leafless, few-flowered racemes of usually between two and eight lilac-blue flowers. The pedicelsare long and the bracts subulate (ie awl-shaped: narrow and tapering from the base to a fine point). Theflowers are pollinated by small flies and Hymenoptera (Garrard & Streeter 1983).
The fruit capsules are 1.2 × 1.0 cm, flattened, kidney-shaped, longer than the sepals. The seeds are 2.0× 1.8 mm, almost orbicular, flat, smooth and unmarked, yellow (Butcher 1961; Sell & Murrell 2007).Seed is set from July to August, the amount of seed produced often being low. The soil seed bank surveyof NW Europe contains five estimates of V. montana longevity, two of which regard the seed astransient, while the other three suggest it is either short-term persistent (1-5 years) or long term(over five years) (Thompson et al. 1997). Apart from the spread of the species along linearhabitats mentioned above, evidence of seed dispersal has not been observed. The fact that V.montana often appears to be ± restricted to long-persistent habitats, like ancient woodland,suggests seed dispersal is not well developed (Grime et al. 1988, 2007).
Fermanagh occurrence
Wood Speedwell has been recorded in 214 Fermanagh tetrads, 40.5% of those in the VC. It is mostfrequently found on appropriately shaded and sheltered lakeshores and river banks. In Ireland, V.montana is sometimes considered mainly a plant of base-rich soils (An Irish Flora 1996).However, in Fermanagh, while it does quite frequently occur over limestone, it is more commonly found onmildly to moderately acid soils, varying from light sandy to heavy clay in texture.
British and Irish occurrence
V. montana has a widely but thinly scattered to locally frequent distribution throughout most ofEngland and Wales, but it is more scarce in the less wooded parts of East Anglia and the east Midlands.Comparison between the two BSBI atlases shows there is no noticeable change in the species distributionduring the 1962-2002 period (A. Horsfall, in: Preston et al. 2002). In Ireland and Scotland,V. montana is very local and the species is absent from Orkney, Shetland and the Outer Hebrides.
European and world occurrence
V. montana belongs to the European temperate phytogeographical element. It is restricted toC & S Europe and adjacent parts of Africa and Asia. The distribution stretches from N Spain andPortugal to N Denmark and eastwards to Latvia and W Ukraine and the Caucasus. In N Africa, it is foundon the mountains of Algeria and Tunisia (Hultén & Fries 1986, Map 1647; Sell & Murrell 2007).
Threats
None.
Native, frequent. Eurosiberian boreo-temperate, but also in N America and widely naturalised beyond itsnative range.
1881-2; Barrington, R.M.; Devenish Island, Lower Lough Erne.
April to October.
Growth form, recognition and preferred habitats
Marsh Speedwell is a small, slender, creeping, semi-recumbent, straggling, ascending perennial the stemsof which root at the lower nodes. It bears narrow, opposite, lanceolate, stalkless leaves that slightlyclasp the stem. The leaf margins appear almost entire, but closer examination shows they possess veryminute, distantly spaced teeth. Plants are usually hairless, but occasional hairy forms are known tooccur, especially in rather drier habitats, and have been named var. villosa Schumach (Claphamet al. 1987; Sell & Murrell 2007).
Marsh Speedwell produces alternate, very loose, few-flowered raceme inflorescences of small, whitish,pinkish or pale blue flowers borne on long, slender stalks (Perring et al. 1989). It is readilydistinguished from V. catenata (Pink Water-speedwell) and V. anagallis-aquatica (BlueWater-speedwell) with which it might be confused, in having its usually rather few-floweredinflorescences arranged alternately on the stem, ie only a single inflorescence is produced fromeach pair of leaves at a stem node. The linear-lanceolate leaves are often reddish brown to dark bronzedue to the presence of anthocyanin pigments and the petals vary from pale lilac, to pink to almostwhite, striped with darker veins.
Marsh Speedwell is found in a wide range of wetland habitats including in still or slow-moving waters onthe shallow margins of smaller lakes, ponds, pools and ditches, and in moist to damp, poorly drained,often winter-flooded marshy places in wet meadows, fens, peat bogs, wet heaths and hillside flushes,usually on rather infertile, acidic or calcium-deficient soils (Garrard & Streeter 1983; Sinkeret al. 1985; A. Horsfall, in: Preston et al. 2002). It can grow either on bare ground oramong tall herb vegetation and appears tolerant of the grazing pressure and the trampling it receivesfrom cattle in the regularly flooded water meadows around Upper Lough Erne. The species reaches itshighest site in B & I at Cross Fell at 780 m (A. Horsfall, in: Preston et al. 2002).
Climbing ability
V. scutellata creeps at the base, but can ascend to 50 cm or more in height when drawn up bytaller herbaceous vegetation around it. Species like V. scutellata, with very slender stemsascending from a creeping or procumbent base, do not have the photosynthetic reserves and physicalability to stand erect by themselves, so they make use of their better equipped neighbours to propthemselves up and enable them to climb towards the light. The least specialised method of climbing isachieved by plants that weave their leaves and stems in and around the neighbouring plants they areusing as props. At its simplest, this can be achieved by the climbing plant developing rigidlyhorizontal leaves. In Marsh Speedwell, the leaves are initially vertical and so can be pushed throughgaps in the foliage of their neighbouring plants. Later on, the Veronica leaves of this andrelated species are held horizontally, thus anchoring the decidedly weak stems in position at a greaterheight than would otherwise be possible (Fitter 1987, p. 192).
The established strategy of V. scutellata is categorised as R/CSR meaning it is intermediatebetween a straightforward Ruderal and a more balanced mix of the traits of the three main strategies,Competitor, Stress-tolerator and Ruderal (Grime et al. 1988, 2007).
Flowering reproduction
V. scutellata flowers from June to August. The solitary inflorescence at each node on the erectflowering stem bears relatively few, well-spaced bisexual flowers, each small blossom borne on a long,slender, thread-like flower stalk, subtended by a very much shorter linear bract. The flowers, 7-8 mmdiameter, have a corolla that is usually pale blue, striped with purple lines, but they can vary incolour from pale pink to white. They attract flies as pollinators and, if not crossed, the flowers canself-pollinate (Fitter 1987). The fruit capsule, 3.5 × 4 mm, is flat and reniform (kidney-shaped) anddeeply notched, veined and winged (Butcher 1961). It opens by dehiscing into two valves apically. Theseeds 0.8-1.0 mm are broadly elliptical, flat, smooth and pale brown (Butcher 1961; Sell & Murrell2007).
Seed dispersal and longevity
There does not appear to be any specialised seed dispersal mechanism or adaptation, but the seeds arelight and presumably can be wafted out of the open capsule by wind passing over it. Seeds probably canalso be dispersed in flowing water or may be picked up and transported in mud. V. scutellataplants have been listed among many others grazed by Arctic Reindeer feeding on the low vegetation andtransporting seed internally and excreting it with their dung (Ridley 1930, p. 373). It appears likelythat the species would also be transported in this way by cattle and other grazing animals.
There is some doubt as to the persistence of V. scutellata seed in the soil. The survey of soilseed banks in NW Europe contains data from nine studies, three of which considered the seed survivaltransient (less than one year); one study believed it short-term persistent (surviving 1-5 years); onereckoned it long-term persistent (surviving more than five years); and four recognised seed as beingpresent in soil, but could not assign it to one of the three seed bank types (Thompson et al.1997).
Fermanagh occurrence
In Fermanagh, V. scutellata is a frequent to local perennial around the shallow margins of waterbodies and in hollows on adjacent fen water meadows on more or less acidic, poorly drained soilsincluding peat, but unlike V. beccabunga (Brooklime) it is never present in any great quantity.The local headquarters of the plant is undoubtedly around the fen-girdled shores of Upper Lough Erne andin marshy ground around nearby lakelets and rivers. Otherwise, it is scattered and rather local in itsoccurrence, although in reality it has been recorded in 149 Fermanagh tetrads, 28.2% of those in the VC.Essentially, it is a species of shallow water around lakes and rivers, marshy meadows and, very rarely,by bog drains.
British and Irish occurrence
V. scutellata has been recorded from every Irish VC, but it is probably declining in some and, forinstance, in the Flora of Co Dublin, it was regarded as common only in marshes in the mountainoussouthern district and very rare elsewhere (Doogue et al. 1998).
The New Atlas hectad map shows that V. scutellata is very widely distributed throughout B& I, but the distribution is scattered and has a definite predominance in N & W parts of bothislands. In N Ireland, it appears much more consistently recorded than in the RoI, although this maysimply be an artefact of recorder effort visible at the hectad scale. The New Atlas hectad mapshows a definite widespread decline in the species occurrence across B & I, but especially in C& S England and C Ireland. Analysis of the map data indicates these declines have occurred since1950 (A. Horsfall, in: Preston et al. 2002).
European and world occurrence
V. scutellata belongs to the European boreo-temperate phytogeographical element and is of frequentoccurrence across most of Europe, although it becomes rare in the Mediterranean basin and is onlysparsely present in N Africa. It also stretches across into N Asia and eastwards to Kamchatka (Claphamet al. 1987). Hultén & Fries (1986, Map 1648) state that it is considered native in N Americaand they map it as such, although they do mention that it is introduced in S America and in New Zealand.
Names
The Latin specific epithet 'scutellata' is derived from 'scutella', meaning 'a small salver or dish' or'scutellatus', 'shield-shaped' or 'platter-like' and thus refers to something that is a small shield -or saucer-shaped feature, although exactly what this refers to is not obvious to the current author(RSF) (Gilbert-Carter 1964; Gledhill 1985; Stearn 1992).
Threats
None at present.
Native, common, widespread and locally abundant. Eurosiberian temperate, widely but rarely naturalised inAfrica and in N & S America.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
Growth form and preferred habitats
Brooklime is a robust, wintergreen, aquatic perennial readily recognised by its rather thick, fleshy,hairless, purple-brown, hollow, low-growing stems which first creep and root and then ascend, bearingopposite, bluntly rounded, toothed leaves which are shortly stalked. The stalked, rounded leaves aresufficient to distinguish V. beccabunga from the other three water- or marsh-speedwells, V.anagallis-aquatica (Blue Water-speedwell), V. catenata (Pink Water-speedwell) and V.scutellata (Marsh Speedwell) (Garrard & Streeter 1983).
V. beccabunga typically occupies wet, moderately fertile ground around the shallow margins ofstreams, rivers, ditches and lakes. It is also commonly recorded in winter- or intermittently-floodedmarshy grassland and around open flushes and springs with a more or less constant flow of water.
Brooklime appears to thrive in open, disturbed, cattle trampled, somewhat poached waterside situations,perhaps thus avoiding severe competition in more dense vegetation, or at least here being better able toface reduced competitive pressure from taller growing species that are less tolerant than it is ofstressful disturbance. The established strategy of the species is categorised as CR, meaning CompetitiveRuderal (Grime et al. 1988, 2007). V. beccabunga also tolerates quite deep shade infen-carr scrub and other wet woodland situations, so it also displays a degree of stress tolerance(Grime et al. 1988, 2007).
V. beccabunga is equally widespread and common on both acidic and calcareous soils, varying intexture from fine muddy silt to gravel in texture. It avoids very infertile soils, including bog peat,due to its low nutrient status and the extreme acidity of such purely organic soils, but it can occurwhen these acid peats are flushed with suitable oxygen and mineral-enriched ground water. Under suitablegrowing conditions, straddling the boundary between marsh or mire and the aquatic realm, Brooklime oftenbecomes luxuriant and can form extensive clonal mats across shallow flowing water in streams, ditchesand flushes (Preston & Croft 1997).
Flowering reproduction
V. beccabunga flowers freely in most sites it occupies. Erect, lax inflorescences, each of 10-30flowers, are borne in opposite axillary pairs at leaf nodes from May to September. The bisexual,hermaphrodite, 5-7 mm diameter blue-petalled flowers, on 4-7 mm slender stalks, are unusual in the genusin being protogynous, ie they are female first after they open, the male anthers ripening later. Thisseparation of the sexes in the flowers' maturity is clearly designed to facilitate cross-pollination,the flowers mainly attracting flies, and especially hoverflies (Proctor et al., 1996). However,the flowers remain closed or half-closed if weather conditions are unfavourable for insect flight and,in these circ*mstances, they then readily self-pollinate (Knuth 1906-1909; Clapham et al. 1962).Plants set many seeds which germinate readily and a short- or long-term persistent seed bank alsodevelops in the soil (Preston & Croft 1997; Thompson et al. 1997).
Reproduction and dispersal is both vegetative, by means of stem fragmentation, and also by seed. Theseeds are not buoyant, but their seed-coat has a mucilaginous gel layer that becomes adhesive when wet,so they are effectively dispersed by birds and other wetland animals to which they attach (Ridley 1930),making them perfectly capable of reaching land-locked lakelets.
Vegetative reproduction
Brooklime plants are shallow rooted and entire colonies are readily swept away in flood waters: brokenstem fragments also root readily, so that a combination of these two properties enables the species tospread vegetatively and increase distribution within the water body (Haslam 1978; Grime et al.1988, 2007).
Fermanagh occurrence
As the second ranking Veronica species to V. chamaedrys (Germander Speedwell) in Fermanaghboth in terms of frequency and distribution, Brooklime is a very abundant perennial in the VC, as islikewise the case elsewhere in B & I, its occurrence far outstripping that of the other aquaticspecies in the genus (Preston & Croft 1997). V. beccabunga is widespread and commonly foundin 390 Fermanagh tetrads, 73.9% of those in the VC. It is absent only from the highest ground and themost acidic bogs (ie pH below about 5.0).
British and Irish occurrence
V. beccabunga is common, widely distributed and locally abundant throughout most of B & Iexcept for the C Highlands and N & W Scotland where it is rare and ± restricted to the coast. TheNew Atlas hectad map indicates that in the westernmost parts of Ireland, it also becomes somewhatless frequent. Having said this, there does not appear to be any evidence from the New Atlassurvey of a decline, at least when measured at the hectad scale. The only probable area where decline ishappening at present is in built-up and developing urban situations such as around major cities (Preston& Croft 1997; A. Horsfall, in: Preston et al. 2002).
European and world occurrence
V. beccabunga belongs to the Eurosiberian temperate phytogeographical element and is widespread intemperate areas of Europe and adjacent parts of W Asia: it is also native in N Africa (Morocco, Egyptand Ethiopia). In Europe, it extends as far north as 65°N (although absent from Iceland) and southwardsfrom there to S Spain and Portugal, and eastwards through the Mediterranean basin to Sicily, Greece, theBalkans, the Middle East and into W Asia and the W Himalaya (Hultén & Fries 1986, Map 1649).
It is introduced and naturalised in N & S America and parts of Africa, probably transported there inships' ballast. V. beccabunga is replaced by the closely related V. Americana (Raf.)Schwein in temperate N America and parts of Pacific E Asia, the two species taken together having adiscontinuous circumpolar distribution (Hultén 1974, pp. 172 & 378, Map 164).
Names
The Latin specific epithet 'beccabunga' is possibly derived from the German name, 'Bach-bunge', 'bach'meaning a 'brook' and 'bunge' a 'bunch'. The Old Norse 'bekkr' meaning 'brook' may also be involved. Analternative origin is from the Flemish, 'beckpunge', meaning 'mouth smart', alluding to the pungency ofthe leaves, which were once eaten in salads like cress (Grieve 1931).
The English common name 'Brooklime' is thought to have evolved from the Old English 'brōc bleomoc'meaning 'brook' and the name of a plant equivalent to the Old Norse 'bung' which features as a wordelement in the scientific name (Grigson 1955, 1987). Grigson lists an additional 13 English commonnames, several clearly related to 'beccabunga' such as 'Beccy leaves' and 'Bekkabung'. The familiar'Bird's eye' and 'Water Bird's eye' also appear, used for other Veronica species, plus 'Waterpurple' and 'Wellink'. Turner (1548) called the plant 'Brookelm'. Together with 'Watercress', Brooklimewas a salad plant as mentioned above, the young tops and leaves providing not unpleasant eating.
Threats
None.
Native, occasional. Eurasian southern-temperate and widely naturalised in both hemispheres.
1900; Praeger, R.Ll.; Co Fermanagh.
June to October.
Growth form and preferred habitats
This stout, fleshy, short-lived perennial aquatic and semi-aquatic speedwell regularly (probablypredominantly) behaves as an annual, although it can sometimes overwinter. It occurs in similar wetground near water and shallow, still or slow-moving aquatic habitats that V. beccabunga(Brooklime) occupies and sometimes the two species occur together, but it is much less frequent than thelatter across the whole of B & I. Unlike Brooklime, V. anagallis-aquatica often occurs ±submerged as well as emergent in shallow water, as well as a terrestrial herb in ± wetland stands.
Again, like Brooklime, V. anagallis-aquatica has rather thick, fleshy, glabrous, hollow stems thatin this case shortly creep, root and then ascend (Press & Short 1994). Plants are more erect thanBrooklime, more leafy and branched, and grow somewhat taller and more luxuriant, usually between 20 and60 cm in height, but capable of reaching 100 cm or more under optimal growing conditions, eg in Cyprusand New Zealand (Meikle 1985; Webb et al. 1988). The stem can become very stout, 4-angled andsucculent, especially towards the base. The leaves are opposite, the lower ones oval, shortly stalkedand scarcely toothed, the upper ones narrowly lanceolate, serrate towards the apex, stalkless andclasping the stem at their base (Butcher 1961). Upper leaves can vary enormously in size, 1.5-17 cm longby 0.8-3.5 cm wide (Meikle 1985). Submerged leaves are very membranous (Webb et al. 1988).
Blue Water-speedwell grows in shallow water or on bare or trampled broken ground by lakes, ponds, rivers,streams and ditches and in marshy grassland, fens, flooded water meadows and clay- and gravel-pits. Itis mainly lowland, but does reach 380 m at Malham Tarn in the Yorkshire limestones (VC 62) (A. Horsfall,in: Preston et al. 2002). In flowing water, Blue Water-speedwell may persist as submergedvegetative stands, failing to flower but, when emergent, it flowers freely. Plants on fertile mud orsilt by streams and rivers are larger and more vigorous than plants in less favourable, drier, or moreinfertile terrestrial sites.
V. anagallis-aquatica is probably less shade-tolerant and it appears more confined to lowlandsites than V. beccabunga. The latter is able to persist in more shaded, taller vegetation and itoften spreads onto damp ground around upland streams and flushes in a way that Blue Water-speedwellrarely does. Despite its quite frequent luxuriant growth, V. anagallis-aquatica is not a stronglycompetitive species in tall herb vegetation, for instance, lacking the climbing ability of V.scutellata (Marsh Speedwell). The difference in occurrence between V. beccabunga andV. anagallis-aquatica must reflect a greater ecological sensitivity in the latter, which appearsto demonstrate less vigour in its vegetative and reproductive capabilities, both of which vary greatlywith habitat conditions. The competitive and comparative features and adaptations of theseVeronica species are clearly worthy of further detailed study (Preston & Croft 1997). Havingsaid this, the established strategy of V. anagallis-aquatica is categorised as CR, meaning'Competitive Ruderal' by Grime et al. (1988, 2007), that recognises the species does have anumber of ecological traits that enable it to compete and establish successfully under someenvironmental conditions, including rapid germination and growth.
Also, there is an underlying genetic difference in that V. beccabunga is diploid, while both V.anagallis-aquatica and the closely-related V. catenata are tetraploids (Preston &Croft 1997). V. anagallis-aquatica is distinguished from V. beccabunga by its less fleshystems and its stalk-less, oblong-lanceolate upper leaves, while it is readily separated from the veryclosely related V. catenata (Pink Water-speedwell) by its pale blue, rather than pale pinkflowers. Separation of V. anagallis-aquatica, V. catenata and their probably quitecommonly occurring hybrid, V. × lackschewitzii J.B. Keller is notpossible without the presence of flowers and/or fruit (J.H. Burnett, in: Rich & Jermy 1998).
Flowering reproduction
If it is going to flower, V. anagallis-aquatica does so in July and August. The inflorescences arelong, lax, slender, erect, leafless racemes borne in pairs in the axils of the upper leaves. The 15-40(mean of 25) flowers in each raceme, 5-10 mm in diameter, are lilac-blue, pale blue, pinkish-lilac orrarely white, with strongly marked violet lines on the corolla. The flowers are visited by flies thatcross-pollinate them, but self-pollination is also easily performed. The fruit capsule, 3-3.5 mm, ispale brown, orbicular (just longer than broad), inflated, slightly notched and hairless. It dehiscesinto four valves and the numerous seeds, 0.5-0.7 mm, are pale brown, elliptic to globular, flat on oneside, convex on the other and smooth (Blamey & Grey-Wilson 1989; J.H. Burnett, in: Rich & Jermy1998; Sell & Murrell 2007).
Seed dispersal is by wind and flowing water, or in mud transported on animals, including birds(particularly ducks) and on man, enabling colonisation of isolated lakelets, as well as man-made poolsand pits (Ridley 1930, p. 547).
There are just two estimates of seed longevity in the survey of NW Europe soil seed banks, one of whichregards the species as short-term persistent (ie seed surviving between one and five years), while theother recognises seed as being present, but unable to assign it with respect to its transient orlonger-term survival (Thompson et al. 1997).
Vegetative reproduction
Early in their growth, young plants develop stolon-like lateral branches that send out adventitious rootsand spread the plant vegetatively to form clonal mats (Preston & Croft 1997). In common with V.beccabunga, portions of colonies of V. anagallis-aquatica can dislodge or branchesfragment when flowing water is in flood spate and these can root and establish downstream, spreading thespecies vegetatively (Haslam 1978). As reproduction is by both seed and vegetative fragmentation, it isnot clear which in this case is the more ecologically significant method of increase, although seed isobviously required in the longer term in order for the species to be biologically capable of adaptationto changing growing conditions.
Fermanagh occurrence
V. anagallis-aquatica has been recorded in just 72 tetrads in the VC, 13.6% of the Fermanaghtotal, as opposed to over 73.9% for V. beccabunga (Brooklime). Six tetrads contain only pre-1976V. anagallis-aquatica records. The Fermanagh distribution of V. anagallis-aquatica is alsopeculiar in that it is quite frequently found around Lough Erne, but it is scattered and very local torare elsewhere in the county.
British and Irish occurrence
Apart from tolerating a greater range of water depth than V. beccabunga, it is not obvious whyV. anagallis-aquatica and the closely related V. catenata should be so much less commonthan Brooklime in a wetland county like Fermanagh where there would appear to be a wide range ofsuitable sites for all three species. Study of the hectad distribution maps of these three species andthe Water-speedwell hybrid in Preston & Croft (1997) and in the New Atlas, however, showsthat only V. beccabunga is evenly distributed over the whole of B & I (with the exception ofthe Scottish Highlands and the far NW, where it is scarce or absent). The other two Water-speedwellspecies are much more frequent in the climatically most continental portion of Britain that lies southof the familiar line from the Humber to the Severn and they really are only very local elsewhere in bothB & I. This suggests a large-scale climatic factor is affecting the relative success of these twoaquatic Veronica species. Although the distribution of V. anagallis-aquatica, measured atthe hectad level, appears stable, Burnett (1997) is certain that there has been an appreciable declinein abundance of the species in England over a period of around 20-30 years towards the end of the 20thcentury.
European and world occurrence
V. anagallis-aquatica belongs to the Eurasian southern-temperate phytogeographical element and isa native of the temperate and Mediterranean regions of Europe, adjacent Asia and N Africa, extendingfrom Iceland and central Scandinavia into C Siberia and almost entirely around the Mediterranean. Italso discontinuously spreads eastwards from Turkey to N India.
The population in SE Asia is distinguished by some as subsp. divaricata Krösche, which has lax,usually glandular-pubescent racemes and arcuate-ascending pedicels (Hultén & Fries 1986, Map 1650).However, variants like this also occur locally throughout the range of the species and do not thereforemerit subspecific distinction (S.M. Walters & D.A. Webb, in: Tutin et al. 1978, p. 248).
The species has also been introduced in N & S America, C & S Africa and New Zealand (Hultén &Fries 1986, Map 1650).
Names
The Latinised specific epithet 'anagallis-aquatica' of this plant previously was just 'anagallis', whichat least as a genus name, was a label in Dioscorides, probably derived from two Greek roots meaning'without' and 'boast', referring to an unpretentious plant (Gilbert-Carter 1964). An alternativederivation is from the Greek 'anagalao', meaning 'to laugh', the genus Anagallis being fabled tohave the power to remove sadness (Chicheley Plowden 1972; Hyam & Pankhurst 1995). However, this doesnothing to explain how the name 'anagallis-aquatica' evolved and became a label for this aquatic plant,and the current author (RSF) is at a loss as to how to take the quest further.
Threats
None.
Native, rare. Circumpolar temperate, but disjunctly so; introduced in C Africa and S Australia.
1934; Praeger, R.Ll.; Co Fermanagh.
June to September.
Growth form, identification and preferred habitats
Pink Water-speedwell may remain under-recorded, as not everyone in the field yet separates it from thevery similar V. anagallis-aquatica (Blue Water-speedwell), although the two were recognised asdistinct species over 100 years ago (Pennell 1921). These two aquatic, semi-aquatic and terrestrial,usually annual but occasionally short-lived perennial species, obviously differ in flower colour and, inaddition, V. catenata has its pedicels shorter than its bracts at flowering time and they spreadat right-angles after flowering. It also has narrower leaves, purple-tinged stems and fewer and morewidely spaced flowers per inflorescence than V. anagallis-aquatica (An Irish Flora 1996;Burnett 1997). Having said this, the two species cannot reliably be distinguished without flower and/orfruit characters (J.H. Burnett, in: Rich & Jermy 1998). However, of vegetative characters, the mostuseful are the absence of basal petiolate leaves in V. catenata and the more marked, frequent,small serrations of the leaves of V. anagallis-aquatica (Burnett 1997).
When growing in more disturbed, trampled sites, V. catenata produces small plants that behave asannuals and young plants develop basal branches that grow into flowering branches, rather than prostrateor procumbent, leafy spreading stoloniferous stems, like those produced by V. anagallis-aquatica(Preston & Croft 1997; Stace et al. 2015). In more terrestrial, waterside sites, the hollow,glabrous stems, 20-30 cm long, shortly creep, root and then ascend in the same manner as V.anagallis-aquatica (Fitter 1987; Sell & Murrell 2007).
V. catenata appears to have a somewhat wider habitat range than either V.anagallis-aquatica or V. beccabunga (Brooklime), strongly preferring exposed bare mud atthe margin of shallow, standing or running water, although it can also grow in much deeper water thanthese other species, down to 1 m deep in streams, ditches and seasonally flooded ground on the marginsof water bodies. In addition, it can be found growing in flushes, on riverside rocks and in coastalsites, where it is occasionally flooded by brackish water. Having said all this, V. catenata mostfrequently occurs in B & I as a waterside terrestrial plant, preferably in open, disturbed ground(Burnett 1997; Stace et al. 2015).
While it can occur on fertile, moderately acidic soils, V. catenata is more frequently recorded incalcareous surroundings (Preston & Croft 1997). The established strategy of V. catenata iscategorised as R/CR, meaning it is intermediate between a straight Ruderal species and a moreCompetitive-Ruderal one (Grime et al. 1988, 2007).
Hybrids
V. catenata does overlap from time to time with V. beccabunga and with the more closelyrelated V. anagallis-aquatica in both its ecology and occurrence, and it is capable of formingvigorous hybrids with the latter, although these have never been discovered in Fermanagh. The hybridoccurs throughout much of temperate Europe, sometimes forming large clones in streams. The hybridsdevelop long, many-flowered racemes but form very few 1- to 5-seeded capsules and are most easilyrecognised by their high degree of sterility (S.M. Walters & D.A. Webb, in: Tutin et al.1972; Stace et al. 2015).
Flowering reproduction
As is the case with most aquatic or semi-aquatic species, when it is submerged, V. catenata doesnot flower, but remains entirely vegetative. Fortunately, completely submerged plants are unusual andthe species is mostly either terrestrial or emergent and therefore capable of flowering. Flowering takesplace between June and August, the pink flowers with reddish veins attracting flies that pollinate themor, failing this, they self-pollinate (Fitter 1987). Capsules are borne on relatively robust pedicels(stouter than in V. anagallis-aquatica) that may thicken after pollination. The capsule isenclosed within narrowly ovate, obtuse sepals which rarely exceed the orbicular capsule which is usuallybroader than long (3-4 mm broad, 3-3.5 long). It is always deeply notched at the apex in mature,un-dehisced capsules (Burnett 1997).
Presumably, in common with other wetland Speedwell species, seed dispersal would mainly be by windblowing across the open capsule, but some seed may be ingested by grazing animals and become transportedinternally by cattle, horses and birds, especially ducks. Other seed might very possibly be dispersed inflowing water.
There is just one estimate of seed longevity in the soil seed survey of NW Europe, which reckoned thatsurvival in soil is short-term persistent (ie 1-5 years) (Thompson et al. 1997).
Fermanagh occurrence
As mentioned under V. anagallis-aquatica, this is a rare species in B & I apart from in themore continental region of Britain lying south of the Humber-Severn line (Preston & Croft 1997).V. catenata is certainly rare in Fermanagh, occurring on exposed mud by lakeshores and, morerarely, on sheltered shallow stretches of river-banks in just 14 tetrads scattered around the VC.Thirteen of the Fermanagh tetrads have post-1975 records. It has been found almost entirely on lowlandlakeshore sites, plus just a single occurrence on the Colebrooke River.
Irish occurrence
Like these other two wetland speedwells (V. anagallis-aquatica and V. beccabunga),V. catenata has been recorded from almost all Irish VCs (Cen Cat Fl Ir 2), but of thesethree species it is by far the least familiar. On the evidence of the records published in the FNEI3 and in the Flora of Co Dublin, Fermanagh appeared to have more locations for thisspecies in the post-1975 era than anywhere else in Ireland. The New Atlas hectad map now showsthat the species is rather well represented in the Irish Midlands, with numerous scattered stationselsewhere across the island, including near the N coast (A. Horsfall, in: Preston et al. 2002).
British occurrence
Locally frequent in C, E & SE England and Midland Ireland, but scattered, coastal and scarce to rarein Wales, SW & N England, Scotland and the Channel Isles (New Atlas; Sell & Murrell2007). Over the period 1960-97, in the opinion of J.H. Burnett (1997), V. catenata did notdecline in abundance in England as much as V. anagallis-aquatica did, and he raised the question,can it be that these species demonstrate cyclical changes in abundance and distribution? This is aphenomenon, he points out, that botanists have been slow to investigate.
European and world occurrence
V. catenata is widespread but thinly scattered and very probably uncommon across most of Europefrom 58°N southwards and eastwards to C Europe and W Asia. It is, however, absent from Iceland and mostof the Mediterranean region. The species is mapped by Hultén & Fries 1986, Map 1651) as being nativein the Azores and N America, although considered introduced to C Africa (Ethiopia and Kilimandjaro) andS Australia. The fact that the same authors reckon the closely related V. anagallis-aquatica isprobably not native in N America, raises a measure of doubt in the mind of the current author (RSF) thatthis species is indigenous in N America.
Names
The Latin specific epithet comes from the Latin 'catenatus', meaning 'linked', 'chained', or 'chain-like'(Chicheley Plowden 1972), but it is not obvious to the current author (RSF), to which plant feature thismight allude.
Threats
None.
Native, frequent, widespread and locally abundant. European southern-temperate, but widely naturalised inboth hemispheres and now circumpolar southern-temperate.
1881-2; Barrington, R.M.; Devenish Island, Lower Lough Erne.
March to November.
Growth form and preferred habitats
In terms of reproductive strategy and life-history, V. arvensis is a winter- or less commonly, asummer-annual. Wall Speedwell is a very typical example of a stress tolerant, semi-rosette forming,weedy therophyte. The plants are very variable, stems ranging from 5-25(-40) cm in height, erect ordecumbent, often branched from the base and developed on a fibrous root system. The small, ± triangularleaves (up to 15 mm) narrow upwards into the flowering part of the stem. They are downy or thinly hairyand their margins are coarsely and bluntly toothed, features that readily distinguish the plant from thecommon naturalised V. peregrina (American Speedwell) (Perring & Walters 1989; An IrishFlora 1996).
This weedy little annual is frequent and locally abundant in dry, open, sunny, potentially droughted,more or less disturbed, wayside sites where perennial competitors are excluded by the stressful growingconditions. Previously it was a common weed of arable crops and cultivated soil in general includinggardens, but modern herbicides have reduced its presence considerably. It is very frequent on the topsof walls, or on dry, exposed, relatively infertile, bare ground near buildings, car parks, quarries,sand-pits and playing fields. Indeed, it can occur in any disturbed or stony, open, fully illuminatedsite, that offers bare soil of either a calcareous or moderate acidic nature within the near neutral pHrange 6-8 and of almost any texture. In closed turf grasslands, it only colonises open, ± disturbedspots, such as anthills, which are avoided by most other species, or, alternatively, it colonisestransiently available, recently disturbed bare, rocky soil (Grime et al. 1988, 2007).
Flowering reproduction
V. arvensis flowers from March to October or, indeed, in favourable sites, throughout the wholeyear. The numerous flowers are borne in long, but rather crowded, terminal racemes. The tiny, 2-3 mmdiameter flowers, have bright blue or dark sky-blue petals with a whitish tube. The petals are muchshorter than the hairy calyx. The flower pedicels are very short and the associated bracts are entireand longer than the flowers, hiding them to an extent.
The flowers attract insects such as flies and small bees as pollinators, or else they self-pollinate: thetwo anthers and the stigma being at the same level makes this easily happen (Fitter 1987). The fruitcapsule is rounded, about as long as broad, 3.0 × 3.5 mm, notched and fringed with gland-tipped hairs.The seeds are 1.0 × 0.5 mm, flat, with radiating striæ, smooth and light brown (Butcher 1961; Hutchinson1972; Garrard & Streeter 1983; Streeter et al. 2009).
Seed persists in soil for up to five years or so (Thompson et al. 1997), although often it is notproduced in huge numbers. The average capsule contains only 18 seeds and the mean per plant is around1,000 (Salisbury 1942). However, under optimal growing conditions, large plants may produce up to 17,000seeds (Harris & Lovell 1974; Grime et al. 1988, 2007). As the species generally grows inexposed, open sites, dispersal is principally by wind. Although the plant is generally absent fromgrazed grassland sites, should it happen to be ingested, it is possible for seed to pass unharmedthrough the alimentary canal of cattle (Ridley 1930). As it frequently grows on or near anthills, it isvery probable that ants help transport seed. Since it was a familiar weed of arable fields for manycenturies, it is highly likely that dispersal in crop seed and human agency facilitated its worldwidespread.
Fermanagh occurrence
As the tetrad map indicates, Wall Speedwell has been recorded in 117 widely scattered Fermanagh tetrads,22.2% of those in the VC. Sites include on the limestones above Florencecourt and the lower slopes ofCuilcagh mountain.
British and Irish occurrence and status
V. arvensis is widespread and common throughout most of B & I, being the commonest of a groupof annual weedy Speedwells. It has traditionally been regarded as native, although in common with mostother Veronica species, there is no fossil evidence to support this idea (Godwin 1975). Its weedybehaviour, long association with cultivation and spread beyond our shores by man, taken together suggestto the current author (RSF), grounds for suggesting it could be an archaeophyte. The New Atlashectad map shows V. arvensis is very widespread in both B & I, except in the far NW ofScotland and the W of Ireland where unsuitable wet, acid peaty soils are predominant and cultivation isvery rare or almost entirely absent.
European and world occurrence
Very widespread throughout most of Europe, C & W Asia, N Africa and Macronesia. It is regarded ashaving its origin somewhere here, but has spread as a crop weed throughout this area. It has been verywidely introduced by agricultural and trading man in E Asia, S Africa, N & S America, S Australiaand New Zealand, so that the present distribution is discontinuous circumpolar (Hultén & Fries 1986,Map 1654).
Names
The Latin specific epithet 'arvensis' is derived from 'arvum (solum)', meaning 'arable land', suggestingthe species grows mainly on arable land (Gilbert-Carter 1964).
Threats
None.
Introduction, neophyte, rare, but probably slowly increasing.
1902; Abraham, J.T.; Enniskillen Town.
April to November.
Growth form and history of introduction and recording
A small, 5-25 cm tall, branched, erect, annual American immigrant with inconspicuous 2-3 mm diameterflowers, V. peregrina first appeared in Ireland as an abundant garden weed in Baron's Court, CoTyrone (H37) in 1836 (in Cybele Hibernica 1866 referred to as 'Barnescourt'), where there is along established nursery garden. The plant, with its long, lax terminal raceme of tiny, pale blue orwhitish flowers that readily self-pollinate and set fruit freely, is believed to have been accidentlyand unintentionally introduced to garden cultivation in Britain as a 'stowaway' on the soil of adeliberately imported plant, apparently arriving from a European nursery sometime prior to 1680 andunknowingly distributed by the horticultural trade (Crawley 2005). The flowers open for only a few hoursaround midday and they quickly wilt with the petals dropping away. The flowers are replaced by otherones next day (https://www.minnesotawildflowers.info/flower/purslane-speedwell, accessed 24 October2022).
Its slightly succulent, ovate or oblong leaves, entire or weakly toothed, narrowing to a short petiole,serve to distinguish it from V. serpyllifolia. American Speedwell flowers from April to July andgood fruiting material can be found in August, although it is in fruit as early as May. It is a weed ofcultivated ground, particularly fruit and nursery gardens, usually on gravel paths or on ± compactedground (Bangerter 1964; Garrard & Streeter 1983; Crawley 2005).
Soon after its first recognition, and up to the end of the 19th century, records of V.peregrina abounded from N and NW Ireland and it became quite widely established and apparentlynaturalised. First VC records in nearby counties were, E Donegal (H34) 1870, W Donegal (35) 1871,Roscommon (H25) 1872, Sligo (H28) 1872, Londonderry (H40) 1896 and Monaghan (H32) 1900. Hart (1883) saidof it, "In many places from east to west of Donegal it has become the commonest garden weed. Exceptin gardens I have not met with it." In a short note on the species in Ireland, Praeger (1896b)commented that, "It appeared to be thoroughly established in cultivated ground in the north-west ofIreland and the fact is of interest, as, so far as I can find, it is unknown in England, and in Scotlandis recorded from Perth alone." He went on to say, "In the London Catalogue it does notfind a place, being apparently treated as a mere casual, and unworthy of insertion …". At the datePraeger was writing (1896), he pointed out that this American immigrant had also settled down as acolonist in Spain, France, Belgium, Holland, Germany, Italy and other unnamed European counties.
However, at his time of writing, Praeger (1896b) was wrong in his statement about the non-existence ofEnglish V. peregrina records. It had been recorded in S Lancaster (VC 59), being reported byAshfield (1865) as having been discovered by a Miss Becker at Altham, "a few years ago … it has nowspread over the place and taken such possession of the ground that it comes up persistently inever-increasing numbers year after year …". It had also been found in Middlesex (VC 21), at Harrowand at Pinner Hill (Melvill 1864). Another pre-1896 record was at Knutsford Racecourse in Cheshire (VC58) where, "only a plant or two" were known, "for a couple of years" and, "werestill there, 1870" (Warren 1871).
Variation
Two forms of V. peregrina originate from America, north and south, the glabrous species and aglandular-pubescent form, var. xalapensis (Kunth) St John & Warren. The latter form hasappeared in Britain and Europe as a rare wool alien (Stace 1991; Clement & Foster 1994).
Fermanagh occurrence
In Fermanagh, V. peregrina was first found at Enniskillen in 1902, but it remained scarce oralmost unrecorded until RHN started recognising it in 1989. Since then, he has added 36 records, mainlyfrom the grounds of schools and the larger landed estates around the lowlands of the Erne basin, butalso in municipal flowerbeds, urban waste ground and in the disused Brookeborough quarry, where gardenrubbish is fly tipped. It appears to be only casual and sporadic, but has now been recorded in 16Fermanagh tetrads, 3% of the total in the VC.
Irish occurrence
V. peregrina is still present in its original Irish site at Baronscourt nursery, Co Tyrone, andalthough it is still mainly a garden weed, it is occasionally found on roadsides, forest tracks, wasteground, quarries and, in one particular Tyrone site (Augher Lough), it behaves as a lakeshore marshplant (McNeill 2010). It has now been recorded at least once in 15 Irish VCs, but in some of these, asin Fermanagh, it has proved only casual and fleeting (Reynolds 2002). The New Atlas hectad mapplots records in eleven Irish VCs, almost all in the N or NW portion of the island (A. Horsfall, in:Preston et al. 2002).
British occurrence
The expectation of leading botanists in the 19th century that V. peregrina would quickly spreadbeyond garden cultivation and become an invasive and widespread weed has not been fulfilled (Bangerter1964). Despite being abundant in its original localities, it spread from them only slowly, increasingits distribution probably mainly through fresh introductions of seed. It seems to require the optimumsoil conditions of nursery or other cultivated garden settings, where it persists longest and it appearsto have difficulty in surviving for long outside such managed conditions. The gradual increase in thenumber of VCs in which it appears indicates it has established status, but in most non-garden sites itcannot be regarded as completely naturalised (Bangerter 1964; Clement & Foster 1994). In Britain,V. peregrina has become widely but patchily distributed, extending northwards to the far north ofScotland. Apart from cultivated or disturbed moist ground, it occasionally appears in damp waste placesand by streams (Stace 1991; Clement & Foster 1994).
European and world occurrence
Taken together, the present world distribution of the species is disjunct circumpolar and it spans theequator (Hultén & Fries 1986, Map 1656). In Europe, it is quite widely scattered, but in theMediterranean region it is confined to Italy, Corsica, Sardinia and Sicily. It has been introduced to SAustralia.
Names
The Latin specific epithet 'peregrina' is from 'peregrinus' meaning 'foreign' or 'exotic', derived from'pereger' meaning 'abroad' or 'away from home', from 'per' meaning 'through' and 'ager', 'field' or'land' (Gilbert-Carter 1964). There are a number of English common names additional to the book name'American Speedwell'. These include 'Purslane Speedwell', 'Hairy Purslane Speedwell', 'Necklace Weed'and 'Neck Weed'.
Introduction, archaeophyte, rare. European temperate, widely naturalised beyond its native range.
1882; Stewart, S.A.; Co Fermanagh, "Borders of fields, common."
April to November.
Growth form, identification and preferred habitats
A small, 10 cm tall, much-branched, spring germinating, summer annual that flowers throughout the wholeyear (although mainly April to September), V. agrestis was once, but is no longer, a common,locally abundant, ruderal weed of cultivated ground and arable crops across lowland B & I. Nowadays,it is rare and mainly seen in gardens and on waste tips where garden rubbish is discarded. It isespecially characteristic of dry disturbed soils and its established strategy was categorised as R/SR,meaning intermediate between a straight Ruderal and a Stress-tolerant Ruderal (Grime et al. 1988,2007).
Flowers are axillary, solitary in the upper part of the stem and are pollinated by small flies and bees,or else self-pollinated. In dull weather, the flowers open less widely, so that the anthers and stigmalie in contact, automatic pollination being effective (Hutchinson 1972; Garrard & Streeter 1983).
V. agrestis is distinguished from V. persica (Common Field-speedwell) by its smaller flower(only 4-8 mm in diameter) and the lobes of the fruit not diverging. It is separated from V.polita (Grey Field-speedwell) by the lower petal (at least) being milky-white and the corollanot being a uniform dark blue: it also has brighter green foliage, the lower leaves opposite, graduallybecoming alternate upwards (Hutchinson 1972). The fruit capsule is clothed with numerous glandular hairsand normally contains from 8-16 seeds which are about 1.5 mm in their longest dimension, compared with1.3 mm for V. polita. In both V. agrestis and V. polita the seeds are veryremarkable, being boat-shaped with transverse ridges on the convex surface (Salisbury 1964; Garrard& Streeter 1983; Webb et al., 1996). Another description of the seed of V. agrestissays they are, "grub-like, with wrinkled beady margins and scooped out in the middle"(Hutchinson 1972).
Fermanagh occurrence
There are only two pre-1975 records for this weedy annual in Fermanagh and it was not until RHN beganrecognising it in 1990, that 15 subsequent sites were added by him. The two early records were Stewart'sfirst listed above and a 1900 Florencecourt find by Praeger published by him in Irish TopographicalBotany the following year. The flowers are parti-coloured (in shades of white, pale blue orpink), the lower petal or petals being either much paler blue or white. The Fermanagh Flora Databasecontains a total of 18 records from, as the tetrad map indicates, 16 widely scattered tetrads, 15 ofthem with RHN's records.
Irish occurrence
Previously, Green Field-speedwell was a widespread, if not a common weed of arable agriculture andcultivated ground, occurring in every Irish VC (Cen Cat Fl Ir 2). However, it declined to itspresent rarity during the last 50 or more years (FNEI 3; Flora of Co Dublin). The speciesstatus was regarded by the Cen Cat Fl Ir 2 as, "possibly introduced", which is why itis not considered by Reynolds in her Cat Alien Pl Ir. V. agrestis was so common in thelate 19th and early 20th centuries in both NE Ireland and in Co Dublin, that the authors of the countyFloras did not include details of its sites (FNEI 1; Colgan 1904).
British occurrence
There has been a steady, long-term, marked decline in V. agrestis populations to scarcityor near rarity in Britain also, a process that was noted even in the first BSBI Atlas (Perring& Walters 1962). It was previously common and widespread in suitably disturbed cultivated ground inlowland England, but more scattered and local in the north and west, as well as in Ireland.
It is odd that of all the annual weedy species of dry, well-drained, stony or disturbed, moderatelyacidic, cultivated ground, V. agrestis should have declined quite so much. The enormous decreaseof V. agrestis populations may be linked to the rise of the closely allied V. polita, asan example of competitive replacement (Salisbury 1964), or there could be similar interactions involvingother weedy speedwells of similar habitats and habit, eg V. arvensis (Wall Speedwell), V.persica (Common Field-speedwell) and V. hederifolia (Ivy-leaved Speedwell). Detailedstudy would be required to demonstrate the reality of this possibility.
An alternative explanation recognises that there are many other previously frequent weeds of arableground which have also disappeared, or almost so, eg Agrostemma githago (Cornco*ckle),Centaurea cyanus (Cornflower), Papaver dubium (Long-headed Poppy) (including P. dubiumsubsp. lecoqii (Yellow-juiced Poppy)) and Scandixpecten-veneris (Shepherd's-needle). All of these were previously found in Fermanagh andthroughout the country and have been effectively eliminated either by modern seed-cleaning processes andchanges in, or intensification of, agricultural operations involving the heavy use of herbicides andfertilisers (A. Horsfall, in: Preston et al. 2002; Crawley 2005).
Species revised status
V. agrestis, together with V. polita, Centaurea cyanus and Scandixpecten-veneris are among the list of 41 species whose status Webb (1985) suggested requirereassessment and which might well turn out to be best regarded as introductions to B & I. Thecurrent author (RSF) is pleased to see that the editors of the New Atlas have agreed with ProfWebb and now recognise the plant as an ancient introduction to B & I, ie it is an archaeophyte (A.Horsfall, in: Preston et al. 2002).
European and world occurrence
The species has spread from C & S Europe and adjacent parts of N Africa and W Asia to reach N Europeand is now thinly scattered across numerous parts of Asia (including the Far East), N America and NewZealand (Hultén & Fries 1986, Map 1657).
Names
The Latin specific epithet 'agrestis' is derived from 'ager', meaning 'field', and translates as 'offields', 'growing in the fields', or 'wild on arable land' (Gilbert-Carter 1964; Gledhill 1985; Stearn1992).
Introduction, neophyte, rare. Eurosiberian southern-temperate, widely naturalised in N Europe, N Americaand New Zealand.
1934; Praeger, R.Ll.; Co Fermanagh.
April to September.
Growth form, identification and preferred habitats
This is another small, creeping, annual weed, c 10 cm tall, of damp to dry, fertile, cultivated soilfound mainly in lowland gardens and on paths and walls. V. polita is similar to V. persica(Common Field-speedwell) but it has smaller, short-stalked, oval, grey-green leaves that are coarselytoothed, and short-stalked, solitary, axillary flowers, 4-8 mm diameter, that are a uniform dark orbright blue with a prominent white eye (Salisbury 1964). The flowering period stretches from March toNovember and while flowers are visited and pollinated by small flies and other Diptera, they may alsoself-pollinate (Fitter 1987).
V. polita is slightly more xerophytic and drought tolerant than V. persica (CommonField-speedwell) and it can grow in a variety of soils but prefers less acidic, light, sandy or morecalcareous soils than V. agrestis (Green Field-speedwell). Indeed, across its wholedistribution, it occurs mostly on calcareous soils. Like V. agrestis, in past years it too was afrequent or common ruderal, weed species, recorded from every Irish VC (Cen Cat Fl Ir 2). Itcolonised cultivated, disturbed and waste ground and previously also grew on railway tracks. Itsestablished strategy is categorised as R/SR, meaning it is intermediate between a straight Ruderal and aStress-tolerant Ruderal (Grime et al. 1988, 2007).
However, V. polita has been in decline for some time and has now become scarce or a rarity,probably like V. agrestis, due to the widespread use of herbicides and fertilisers associatedwith the intensification of agriculture and its horticultural spinoff (FNEI 3; Flora of CoDublin).
Fermanagh occurrence
The Fermanagh Flora Database contains records from seven tetrads, five of which have post-1975 records.The remaining record details are: Kiltierney Deerpark, 1951, MCM & D; railway near Kilturk Td, 1951,MCM & D; wayside in plantation wood, Florencecourt, 7 April 1990, RHN (and again in September 1992and May 2004); Belleisle Estate, 27 June 1992, RHN; Killreagh House, near Tamlaght, 6 April 1996, RHN;garden at Portora School, 23 April 1996, RHN; and grounds of Carlton Hotel, Erne bridge, Belleek, 12July 2006, RHN.
As the tetrad map shows, the sites are widely scattered across the VC, with a slight concentration aroundEnniskillen, the largest conurbation in the county.
British and Irish occurrence
The New Atlas hectad map shows V. polita widespread in both B & I, but very much lessfrequent and patchier across Ireland. The pattern and spread of recent records makes it very obviousthat V. polita is declining in N England and Scotland as well as in Ireland and it remainsfrequent and widespread only in S & E England and in Wales (A. Horsfall, in: Preston et al.2002).
Species revised status
V. polita, together with V. agrestis, Centaurea cyanus (Cornflower) and Scandixpecten-veneris (Shepherd's-needle), are among the list of 41 species whose status Webb (1985)suggested require reassessment and which might well turn out to be best regarded as introductions to B& I. The current author (RSF) is pleased to see that the editors of the New Atlas have agreedwith Prof Webb, and now recognise V. polita as an ancient introduction to B & I, ie it is anarchaeophyte that was first recorded from the wild in Britain in 1777 (A. Horsfall, in: Preston etal. 2002).
European and world occurrence
The distribution of V. polita is similar to that of V. agrestis. The species probablyoriginated in C & S Europe and adjacent parts of N Africa and W Asia, but it has spread as a weed ofcultivation to N Europe, N America and New Zealand (Hultén & Fries 1986, Map 1658). It has itspresent day centre of variation in the Elburz mountain range in N Iran and was probably native to openplaces in deciduous montane forests, before becoming a weed in Neolithic times and migrating to Europe(Fischer 1987).
Names
The Latin specific epithet 'polita' means 'elegant, polished, refined or neat' from 'polio', 'polish'(Gilbert-Carter 1964; Stearn 1992). It is not clear to the current author (RSF) to which part of theplant this allusion might refer.
Threats
None.
Introduction, neophyte, occasional and local, possibly somewhat under-recorded.
1884; Barrington, R.M.; Crom Castle Estate.
January to November.
Growth form, identification and preferred habitats
This hairy, sprawling winter- or summer-annual with decumbent to erect stems, 10-50 cm long, cangerminate, grow rapidly and flower at almost any time of year, although the main flowering takes placebetween March and September. It branches from the base and bears heart-shaped leaves, the lower onesstalked, upper sessile, margins with few shallow teeth and veins beneath hairy (Butcher 1961). Itresembles V. agrestis (Green Field-speedwell), but differs from it and V. polita (GreyField-speedwell), by being larger all round and especially by its having relatively large, solitary,axillary, pale blue flowers, 8-12 mm in diameter with a white centre, borne on slender pedicels longerthan the leaves that subtend them.
V. persica is a typical example of a ruderal annual weed of lowland arable cultivation, occurring,sometimes abundantly, in fallow areas or under crops in fields, especially cereals, or in gardens or onwaste ground. V. persica also appears sporadically in other disturbed habitats, including spoilheaps, demolition sites and waysides. It prefers or demands regularly disturbed, moist, fertile, loamsoil in the near-neutral, pH range 8.0–6.0 (Grime et al. 1988, 2007).
V. persica is strongly suppressed by shade, however, and its behaviour and restriction tosemi-permanently disturbed ground suggests it displays low competitive ability. However, theabove-ground dry weight and the total seed mass of V. persica is much larger than that of V.agrestis, whereas the seed weight as a percentage of above-ground weight is much lower than thatin V. agrestis. This indicates a comparatively large competitive potential due to its strongvegetative development (Fischer 1987).
Flowering reproduction
As with other small, weedy Veronica species, the flowers of V. persica attract small fliesand other Diptera with nectar and they are either insect pollinated or, if this fails to occur, as itoften does, they self-pollinate (Fitter 1987; Sell & Murrell 2007).
The fruit capsule is distinctive and helps to identify the species, having widely divergent lobes,measuring 5 × 9 mm, its surface ciliate and the lobes distinctly flattened, with a slight keel (Melderis& Bangerter 1955; Salisbury 1964; Garrard & Streeter 1983). The seed is also rather distinctive,being 1.4 × 1.0 mm, cup-like or boat-shaped, coarsely tubercled (rugose) on the convex surface and grey,light brown or yellowish-brown in colour (Butcher 1961; Salisbury 1964; Sell & Murrell 2007).
There are 5-10 seeds in each of the two fruit locules, the average number in a capsule being 13. Anaverage-sized plant produces around 2,000 seeds, but in large ones it can be 5,000-7,000. Moreover,since growth is so rapid, it is perfectly possible for two generations of the species to develop andfruit within a single year (Salisbury 1964). Seeds are said to be dispersed by ants, but the main agentof spread is probably man, transporting the species as impurities in agricultural seeds, in manure andin fodder (Salisbury 1964). Seed is not only prolifically produced and readily dispersed, it is alsolong-persistent in the soil seed bank, surviving for more than five years (Thompson et al. 1997).
Origin and introduction
V. persica is of SE Asian origin and was first recorded in Europe about 1800 and in Britain, fromBerkshire, in 1825 (Garrard & Streeter 1983). After its first appearance, it spread extremelyrapidly in Britain, so that the 8th edition of The British Flora described it as, "Prettyfrequent in England and middle and south Scotland, possibly introduced with clover and otherseeds." (Hooker & Walker-Arnott 1860).
V. persica is a tetraploid species of allopolyploid nature (2n=28), intermediate in characterbetween its probable parents, the diploids V. polita (Grey Field-speedwell) and V.ceratocarpa C.A. Mey. (both 2n=14). The latter species is native to sub-humid or mesophyticdeciduous forests of the Caucasus and Elburz mountains in N Iran, while V. polita also has itscentre of variation in the Elburz range, where it grows as a slightly xerophytic, therophyte (ie aruderal annual of fairly dry habitats) (Fischer 1987).
A characteristic primary habitat, probably for all the mentioned species (V. persica, V. politaand V. ceratocarpa), is pebble and gravel riverside places in mountain forest regions of N Iran.This type of habitat generally favours therophytic species of comparatively high ecological plasticitythat are able to resist drought periods, but are simultaneously capable of benefitting from a good watersupply and which have a broad light intensity tolerance. V. persica probably achieved itscharacteristic wide ecological amplitude by combining the stronger xeromorphic characters of V.polita, with the more mesomorphic features of the other parent, V. ceratocarpa (Fischer1987).
Fermanagh occurrence
V. persica appears to be surprisingly rather local in Fermanagh, for it is generally a verycommon, to abundant, although perhaps sporadic weed of gardens and disturbed waysides elsewhere in B& I (Fischer 1987; New Flora of the BI 1991; An Irish Flora 1996). It has beenrecorded, however, in 51 Fermanagh tetrads (9.7% of the VC total), rather thinly scattered in thelowlands in the typical disturbed habitats it frequents.
While the current author (RSF) and RHN have no direct evidence on the matter, we have a feeling that thisspecies is under-recorded in Fermanagh. It may perhaps be deliberately overlooked, being recognised as awidespread weed and be simply disregarded. Government employed survey workers might well consider it aninsignificant alien of no conservation value and constituting no threat to native species in semi-nativehabitats and, therefore, ignore it. On the other hand, it could be argued that V. persica isprimarily a plant of cultivated ground and the relative scarcity and local nature of the species in theFermanagh might merely reflect the major decline of arable farming to almost total extinctionthat has happened during the past 60 years (Murray et al. 1992).
British and Irish occurrence
The New Atlas hectad map shows that V. persica is very widely distributed across all oflowland B & I (A. Horsfall, in: Preston et al. 2002).
European and world occurrence
V. persica has spread widely over most of Europe and has also been introduced through agricultureto S Africa, E Asia, N America, S Australia and New Zealand (Hultén & Fries 1986, Map 1660).
Names
The Latin specific epithet 'persica' translates as 'Persian', the old name for modern Iran, correctlyindicating the point of origin of the species (Gilbert-Carter 1964). English common names include'Bird'seye Speedwell', 'Common Field-speedwell', 'Persian Speedwell', 'Large Field-speedwell','Bird's-eye', 'Cat's-eyes' and 'Winter Speedwell'.
Threats
None.
Introduction, neophyte, frequent and widespread.
1949; MCM & D; Aghalane Bridge, Woodford River.
Throughout the year.
Growth form and reproduction
A small, downy, prostrate creeping, but rampant mat- or carpet-forming perennial, most commonly found asan aggressive invasive weed of garden lawns, where it can quickly become a problem. Although it flowersfreely from April to June, the solitary, attractive, pale lilac-blue flowers, formed on long,thread-like stalks (pedicels 6-8 times longer than leaves) in the axils of ordinary leaves areself-sterile and do not (or only extremely rarely) form capsules and seeds in B & I (Perring &Walters 1989; Ellis 1993). Salisbury (1962) suggested that the sterility probably arose because all theplants in Britain belong to the same source or mating strain and thus constitute a singleself-incompatible clone. In this connection, it is important to bear in mind that fewself-incompatibility systems are absolute (Stace & Crawley 2015).
It is not known for certain whether more than one mating strain exists in B & I, but the factremains, seedlings of V. filiformis have never been seen in B & I (Bangerter & Kent1957). However, the slender, wiry, creeping stems or runners, up to 50 cm long, spread rapidly andreadily root at their nodes. When it invades lawns, V. filiformis becomes a very close and firmcreeper and is as hard to remove as the native V. serpyllifolia (Thyme-leaved Speedwell). Stemfragments are also extremely successful at regenerating the plant from mown clippings, some of which mayalso disperse considerable distances in strong winds (Ellis 1993). Vegetative material of the species isprobably also transported in contaminated soil, turves used for laying new lawns and in garden plantpots. It could also be a contaminant on contractors' lawn mowers, strimmers and other machinery (Stace& Crawley 2015).
Introduction, spread and preferred habitats
The species was introduced to B & I from Turkey and the Caucasus as a small decorative subject forrock gardens in the early 19th century, perhaps as early as 1800, but certainly by 1808. In its nativearea, V. filiformis is a rare plant with a very restricted range that grows in woods, fields andpastures. Even in its native habitat and environment, the species forms few seeds (Bangerter & Kent1957). It was first reported as having escaped from garden confines in England in the neighbourhood ofColchester in 1838, and the second report was 89 years later by the River Ayr in Scotland in 1927(Bangerter & Kent 1962). In the following 25 years, it spread slowly at first and then, post-1950,quite rapidly across B & I. Unlike other field speedwell species, its ability to colonise is notconfined to open and bare ground. It now is an abundantly naturalised adventive, forming patches ongrassy waysides, stream-side meadows and pastures throughout most of the British Isles. It also remainsa serious park and garden weed and major lawn pest (Bangerter & Kent 1957; Ellis 1993; Stace &Crawley 2015).
Fermanagh occurrence
As the tetrad map indicates, V. filiformis is frequent and widespread across Fermanagh, havingbeen recorded in 129 tetrads, 24.4% of those in the VC. It is found typically in pastures, roadsideverges, disturbed ground, quarries and on riverbanks and lakeshores, but it most commonly occurs as aweed of lawns.
Irish occurrence
V. filiformis was first reported as a garden escape in Ireland from Ballina, W Mayo (H27) around1933, secondly at Wicklow (H20) in 1935 and thirdly on a grassy bank between Sligo town and Ben Bulbenmountain (H28). In 1938, it was found at Greencastle, E Donegal (H34) and in 1946 on sand dunes atStrandhill, Co Sligo (H28) (Bangerter & Kent 1957). By 1957, records had been made from 39 sites andby 1961 this had increased to around 100 (Bangerter & Kent 1957, 1962). It is now, as in Britain, awidespread and common, lowland garden escape, naturalised in lawns and other managed grassland, suburbanand other grassy verges and waste ground. It also appears less frequently in more natural habitats,particularly on the banks of streams and rivers (Reynolds 2002).
The New Atlas hectad map indicates a very much more complete coverage of the VCs in the north ofthe island, but it also shows a greater presence (or more persistent recording effort) around the citiesof Dublin and Cork, while the species appears less prevalent in the far west of the Republic, especiallyin Co Clare (H9), E & W Mayo (H26 & H27) and S Kerry (H1) (A. Horsfall, in: Preston etal. 2002). It will be interesting to compare how the species has been recorded in theforthcoming BSBI Altas 2020 to be published in 2023.
British occurrence
The maps of Slender Speedwell distribution published by Bangerter & Kent in 1957 and 1962 showed thespecies' presence extending from the Channel Isles to the far N of Scotland, a range almost as wide asthat found today (A. Horsfall, in: Preston et al. 2002). However, in the second half of the 20thcentury, the distribution became very greatly consolidated and extended across the sea to Orkney andShetland.
V. filiformis has become an extremely common, widespread and locally abundant invasive perennialacross the whole of lowland Britain in a wide range of habitats from garden lawns and gravel paths tochurchyards, to artificial and semi-natural sites, including roadsides, paths, rubbish heaps, grassybanks (especially near houses), plus along streamsides. It is probably most abundant in S & WEngland, but in parts of S & SW of England in the late 1940s it was described as being dominant overnative vegetation in some sites (Bangerter & Kent 1957).
It has been mooted that anecdotal evidence indicates a decline in the species presence has been noticedsince around 1995 (Stace & Crawley 2015), although the Change in the British Flora 1987-2004survey suggested it is still increasing (Braithwaite et al. 2006).
European and world occurrence
During the 20th century, V. filiformis spread to many areas of N & W Europe and also to both E& W regions of N America and New Zealand, where it was first recorded in 1981 (Hultén & Fries1986, Map 1661; Webb et al. 1988).
Names
The Latin specific epithet 'filiformis' means 'threat-like', from 'filum', 'thread', and 'forma','shape', an obvious reference to the slender, wiry, spreading stems (Gilbert-Carter 1964).
V. filiformis has been given a number of English common names including 'Slender Speedwell','Creeping Speedwell', 'Threadstalk Speedwell', 'Round-leaved Speedwell', 'Caucasian Speedwell' and'Whetzel Weed'.
Threats
An invasive weed of mown or heavily grazed grasslands, in Fermanagh V. filiformis is now alsoactively colonising semi-natural grasslands beside riverbanks and lakeshores and it has arrived in theNR of Hanging Rock.
Introduction, archaeophyte, local but probably under-recorded. European southern-temperate, naturalisedin N America, Japan, S Australia and New Zealand.
1884; Barrington, R.M.; garden at Crom Castle.
April to October.
Growth form and identification
A slender annual of cultivated and open, bare or waste ground, V. hederifolia is an unmistakeableweed with weak, straggling, prostrate or decumbent, thinly and shortly pubescent, hairy stems, 10-60 cmlong. The stems bear nearly all alternate, ivy-shaped leaves, broader than long and rather thick intexture. The leaves are semicircular in outline, 5-lobed and shortly stalked. The cotyledons (quitepersistent) and some of the lowest leaves are opposite. It often forms long side-branches spreading fromthe base.
V. hederifolia is readily distinguished from other creeping, hairy, annual speedwells withsolitary flowers, by its shortly stalked, semi-circular, lobed leaves, at least as broad as long (AnIrish Flora 1996).
When it flowers in spring from April to June, the small, solitary, usually pale-blue bisexual flowers,4-9 mm diameter, are borne on long stalks (pedicels) in the leaf axils, the stalks soon becomingdeflexed in fruit. The fruit capsules are almost spherical, scarcely compressed, hairless, the twochambers of each containing a maximum of just two seeds. The seeds are described as 'grub-like', iecup-shaped, ellipsoid and wavy, almost smooth (Melderis & Bangerter 1955; Perring & Walters1989; Sell & Murrell 2007; Parnell & Curtis 2012).
Variation
There is sufficient variation in V. hederifolia, which comprises a polyploidy complex(basic chromosome number n=9) with three ploidy levels, enabling five taxa to be recognised inEurope, two of which occur in B & I. Two rather weakly differentiated subspecies are recognised in B& I, the archaeophyte nominate subsp. hederifolia (a hexaploid) and the perhaps nativesubsp. lucorum (Klett & Richt.) Hartl. (= V. sublobata M. Fischer) (a tetraploid). Allfive forms are intersterile and there is taxonomic debate or uncertainty as to the rank of their levelof separation (M. Fischer, in: Walters & King 1975).
Subspecies hederifolia has thicker, darker green leaves, 3- to 5-lobed, the middle lobe beingwider than long, and the incisions between the lobes are distinctly deeper than they are in subsp.lucorum. The petals of subsp. hederifolia are usually bright blue with a white eye and theanthers are blue before they release their pollen. The anther colour before the pollen is released isthe least variable, most reliable character to distinguish the two subspecies; in subsp. lucorumthe anthers are whitish rather than blue.
The variability of characters needs to be emphasised, however, since in some plants, one or morecharacters may be less well shown and plants of one subspecies may exhibit a character that is normallyassociated with the other subspecies. Therefore, identification must be determined on a combination ofcharacters (M.J. Wiggington & G.C. Graham, in: Rich & Jermy 1998).
Habitat range
In terms of habitat preference, V. hederifolia is a pioneer weed of bare ground on rich soils,frequently met in a wide range of lowland ruderal situations, from being a garden weed, the edges ofarable fields, colonising shaded banks, plus disturbed and open ground of all types, including sanddunes, railway ballast, disturbed roadside verges, farmyards, compost heaps, rubbish tips and wasteground. It is especially associated with rocks, shingle and silty places by stream sides in woodland(Sinker et al. 1985; Chater 2010). The established strategy of the species is categorised asR/SR, meaning intermediate between a straight Ruderal species and a Stress-tolerant Ruderal (Grime etal. 1988, 2007).
Subsp. hederifolia tends to occur more typically in open, sunny sites such as arable ground,gardens and roadsides, while subsp. lucorum appears confined to more damp, shaded sites such aswoodland and perhaps in more natural or semi-natural habitats than subsp. hederifolia frequents(Green 2008). Subsp. lucorum can, however, also occur as a garden weed (Sinker et al.1985). In Cardiganshire, Wales (VC 46), Chater (2010) reckoned that subsp. lucorum was thecommonest woodland annual in the county, often occurring in vast abundance in spring in the damper andmore fertile woods. However, he went on to warn that this ecological distinction, like the morphologicalones separating the two subspecies, is imperfect (Chater 2010, p. 591).
British and Irish status
Previously a common cornfield and open ground weed and accepted or presumed by all as a native species inBritain and as possibly introduced in Ireland (Scannell & Synnott 1987), this summer annual has onlyrecently been recognised as an archaeophyte introduction throughout these islands (ie an ancient, orpre-1500 BP introduction) (New Atlas). Although Stace (1997) was still treating it as nativethroughout B & I, the species had been considered as a possible or probable introduction in Irelandfrom at least 1950 onwards. In his Flora of County Wicklow, Brunker (1950) regarded V.hederifolia a common colonist of cultivated ground, wall tops and roadsides. In their Floraof Connemara and the Burren, Webb & Scannell (1983) described the species as, "probablyintroduced", and considered it, "a rare plant in western Ireland". However, the authorsof The Flora of County Dublin still reckoned the species, "possibly introduced" (Doogueet al. 1998).
The survey of the flowers of Cork city and county regarded both V. hederifolia subspecies as alienintroductions (O'Mahony 2009), but in their Flora of County Limerick, Reynolds & Reynolds(2013) opt for the best of both worlds, regarding subsp. lucorum as native and subsp.hederifolia as alien. In his Flora of County Tyrone, McNeill (2010) goes against thetrend, considering V. hederifolia, "probably native" and, "rather uncommon".
Fermanagh occurrence
V. hederifolia was first noticed in Fermanagh in a garden at Crom in 1884. Twenty-three of the 29records in the Fermanagh Flora Database have been added by RHN since 1987, so it is very probablyunder-recorded by other workers. V. hederifolia has been found in 20 tetrads, 3.8% of those inthe VC. Therefore it appears thinly and widely scattered, mainly in gardens, estates and near habitationor associated with other forms of disturbance on roadsides or open waste ground.
In Fermanagh, the two subspecies are still in the early days of being distinguished and recorded. Fourrecent records of the 29 recorded for the species in Fermanagh are tentatively identified as subsp.lucorum.
British & Irish occurrence
The difficulty of making field determinations between these two B & I subspecies results inuncertainty regarding accuracy of distribution maps as depicted in the New Atlas. The currentauthor (RSF) has therefore decided not to comment on the subspecies maps in the New Atlas, but toconcentrate on the B & I distribution of the species as a whole.
Irish occurrence
V. hederifolia was recorded as a common weed of cultivation in Ireland during the early 19thcentury (Mackay 1825, 1836). In his Irish Topographical Botany, Praeger (1901) described V.hederifolia as, "often abundant, but rather locally distributed and not common in theCentral Plain". Nowadays, in Ireland, V. hederifolia is considered occasional tofrequent in the eastern half of the country, becoming much rarer west of the River Shannon (NewAtlas; Parnell & Curtis 2012).
British occurrence
The New Atlas hectad map shows V. hederifolia very frequent and widespread throughoutlowland England and Wales, becoming more scarce, more coastal and more eastern in Scotland, althoughpresent in Orkney and Shetland (A. Horsfall, in: Preston et al. 2002).
European and world occurrence
This polymorphic species seems to have originated in C & S Europe and adjacent parts of N Africa andW Asia. It is widespread throughout temperate Europe, subsp. hederifolia and subsp.lucorum spreading northwards into Scandinavia as weeds. The species s.l. has been introduced intoN America, where it has become naturalised, and into Japan, S Australia and New Zealand (Hultén &Fries 1986, Map 1662).
Names
The Latin specific epithet 'hederifolia' translates as 'ivy-leaved'. English common namesinclude 'Ivy-leaved Speedwell', 'Corn Speedwell', 'Mother of Wheat', 'Winterweed' and 'Hen-bit'(Melderis & Bangerter 1955).
Threats
None.
Introduction, neophyte, here a definite mis-identification or recording error. European temperate, spreadbeyond its supposed native distribution by association with agriculture.
10 August 1970; Farrell, Ms L.; quadrat 7, limestone grassland at Monawilkin.
Growth form and preferred habitats
This annual hemi-parasite attacks the roots of grasses, cereals and a wide range of other speciesincluding Salix alba (White Willow). Despite being able to grow to a height of around 60cm when occurring among tall grasses, having an attractive flower with dark pink lips and a terminaltuft of brilliant purple bracts, M. arvense has never been definitely recorded anywhere else inIreland either before or since, and the current author (RSF) and RHN now believe the Fermanagh record isa definite field error or slip of the pen on the record card.
Formerly, M. arvense was an abundant pest weed in cornfields in places including the Isle of Man,the relatively large (5-6 mm × 2.7 mm) black seed discolouring flour. This was probably the case in theearly part of the 19th century, when winnowing with a hand riddle was primitive, hard work and failed toseparate weed seeds from grains of Corn. Following the introduction of more efficient threshing machinestowards the close of the 19th century, M. arvense subsequently became an uncommon or even a rareand local seed-borne weed of arable agriculture in parts of S & SE England and the Isle of Man,especially on chalky soils (Salisbury 1964).
M. arvense survives in a handful of English VCs as a rarity of open, usually calcareous sites ingrassland beside hedges and ditch banks, or in other very open, disturbed or eroding situations where itcan avoid competition from more vigorous perennial species. Seed survives for about two years or perhapslonger in the soil and germination percentages are normally low, around 25% (Salisbury 1964; P.J.Wilson, in: Wigginton et al. 1999).
Fermanagh occurrence
There is a solitary record of Field Cow-wheat as listed above in the Fermanagh Flora Database, but it isalmost certainly an error.
British and Irish status
Until recently, M. arvense was regarded as a diminishing English native cornfield weed, and aBritish Red Book rarity at that with only four scattered sites remaining (P.J. Wilson, in: Wiggintonet al. 1999, p. 243)! It was one of the 41 species previously assumed native in Britain whichWebb (1985) suggested were probably introduced by man and whose reassessment he advocated. It is nowbelieved that M. arvense, which possesses a large black seed like a wheat grain, was originallyintroduced to Britain along with crop seed.
In parts of continental Europe, Field (previously called 'Common') Cow-wheat or 'Poverty Weed', isconsidered an archaeophyte. It was called the latter name because it discoloured the wheat or otherflour and greatly lowered its value to the farmer. However the plant was first discovered in England inNorfolk by John Ray in 1724 (Salisbury 1964, p. 42) and its revised English status is therefore that ofa rare and slowly declining neophyte (Preston et al. 2004).
There is no obvious reason why this crop seed alien could not have been introduced to suitable cerealgrowing areas of Ireland and it might have survived in disturbed marginal sites as has been the case inBritain. Nevertheless, the solitary existing Monawilkin record does not ring true since, in the view ofRSF & RHN this upland, thin-soiled, rocky limestone terrain could not have supported cerealcultivation at any period in history.
Native, occasional, locally frequent. Eurosiberian boreo-temperate.
1882; Stewart, S.A.; Co Fermanagh.
April to October.
Growth form and preferred habitats
This annual, up to 60 cm tall, but often shorter than this and with spreading branches and opposite,entire, narrow leaves is a partial- or hemi-parasite on the roots of an accompanying range of plants,including trees, ericaceous sub-shrubs and Myrica gale (Bog-myrtle). It is not absolutely clearif it can use herbaceous plants, especially Deschampsia flexuosa (Wavy Hairgrass), with which itis often associated, as hosts, but in many situations it exclusively parasitizes woody plants (Smith1963; Masselink 1980). M. pratense is an extremely variable plant, the leaves often bronze-greento chocolate brown and the petal colour varying from pale yellow-white to a rich, bright yellow like abuttercup.
The established strategy of M. pratense is categorised as R/SR, meaning it is intermediate betweena straightforward Ruderal and a Stress-tolerant Ruderal (Grime et al. 1988, 2007). Thesemi-parasitic nature of this annual requires the seedling plant, produced after autumn germination, tosend out sufficient roots of its own in order to locate the roots of a suitable host plant, attach andcreate an effective haustorium linkage. If this aim is achieved, it enables the plant in the spring(around March-time), access to the nutrient support essential to its further growth, development andreproductive survival. Without host plant contact and support, the juvenile M. pratense plantremains undeveloped and it gradually withers away during May and June (Smith 1963). After flowering andfruiting, the plant dies in late summer or autumn (Masselink 1980).
The advantage hemi-parasitism gives M. pratense plants is the ability to grow very rapidly inenvironments with exceedingly low nutrient soils that severely limit the competitive ability of anyother vascular plants present. M. pratense reaches full size and reproductive capability after afew weeks' growth and, in nutrient-deficient habitats, this feat is only possible if a substantialproportion of its nutrients and moisture requirement are obtained from its host plant. M.pratense is a shallow rooted species, but observation in drought conditions shows it is able toobtain a fully adequate water supply from the host plant, making it independent of soil water (Smith1963).
As is often the case elsewhere in B & I, M. pratense usually occurs in small, isolatedpopulations of around 200 plants (Smith 1963). It is locally frequent, but rarely abundant, in astrikingly wide range of habitats in Fermanagh, encompassing acidic (preferred) and calcareous,nutrient-poor, wet but drained or dry soils in both semi-shaded and more open situations.
The species is clearly bimodal with respect to soil reaction, occurring on both acidic soils of pH downto 5.5 and limestone or chalky ones of pH 6.8 or more (Smith 1963). The habitats colonised includewoodland waysides, clearings and margins, lakeshores, peat bogs, upland heath and limestone cliffs. In NScotland and N England, it also occurs in boggy flushes and waterlogged peat in acid bogs (Smith 1963;Garrard & Streeter 1983). M. pratense does seem to be more frequent today in heathlandcommunities managed using fire to regenerate and keep them from senescing (Ingrouille 1995).Interestingly, in the Netherlands, M. pratense does not occur in heath vegetation (except thosewith Vaccinium myrtillus (Bilberry) and/or V. vitis-idaea (Cowberry)), or on peatlands orin conifer woods (Masselink 1980).
Flowering reproduction
Flowering takes place from May to October, the inflorescence consisting of a series of single flowers,15-20 mm long, in the paired axils of the upper opposite, green, leaf-like bracts. However, the nodesare twisted, so the flowers are all turned to the same side to lie alongside one another (ie they aresecund). The bi-lipped petal tube is closed or only very slightly open with a large yellow palate;nectar production is high. The four stamens are positioned with their anthers converging under the upperlip of the corolla forming a 'pollen box' into which the anthers shed their contents. Visitingbumble-bees probing past the anthers for nectar invariably unlock this box and get showered with pollen.This is known as the 'loose-pollen mechanism' (Melderis & Bangerter 1955). In addition to the floralnectary, the bracts at the base of the flowers and leaves close to them are provided withnectar-secreting hairs, acting as extra-floral nectaries that attract ants. The extra-floral nectariesare indistinct depressions on the leaves and bracts of M. pratense (Britton 1943; Hutchinson1972).
If insect visits fail to occur, the flowers are selfed, since they are hom*ogamous and the stigma ispositioned close to the anthers (Clapham et al., 1962). Individual flowers may continue tofunction for up to seven days, which is unusually long in this flower family (Molau 1993).
The fruit capsule is obliquely ovate, compressed and splits along one side to release the maximum of fourlarge, pale- to dark-brown, ellipsoid seeds, 3-5 mm long (Britton 1943; Butcher 1961; Hutchinson 1972).
Seed is shed from July to early September. The seed bears an oil-body (elaiosome) attachment derived fromthe funicle or stalk of the ovule. This, and the extra-floral nectaries that secrete a watery saccharinefluid, provides an attractive food-source that encourages ants and voles to visit and collect the seeds,helping to effect their dispersal (Ridley 1930; Molau 1993). The role of ants must not be exaggerated,however, as in a northern Netherlands study, seeds were not moved more than a few tens of cm, or not atall – small ants merely sucked on the elaiosome and presumably on the extra-floral nectaries too. Thesmall distances the seeds were transported was thought to be due to the ants being hindered by thevegetation density and probably also because the elaiosomes soon lost their attraction by drying up(Masselink 1980).
In a study in the northern Netherlands, the mean number of seeds per flower ranged from 2.2 to 3.1. Oneplant may produce 40-50 flowers throughout the summer and therefore about 100-150 seeds. Countsrevealed, however, that in most populations the number of seeds per plant was much lower, viz. 40-75seeds, of which 20-60 may germinate, the others being viable but dormant. Hence the reproductivecapacity is low. Many seeds fall victim to invertebrate and fungal attack and others are eaten by miceand other small animals. Normally only the most sheltered seeds survive predation after hibernation.Naturally, seeds in open sites will be more easily found by seed eating animals than those in sites withmore dense grassy vegetation (Masselink 1980).
Germination takes place from late September until November (Masselink 1980).
Variation
Two subspecies, subsp. pratense and subsp. commutatum, are now recognised in Britain, butonly subsp. pratense occurs in Ireland. Again, two varieties of subsp. pratense aredistinguished on corolla colour, var. pratense and var. hians, the latter having the wholecorolla a deep golden yellow (Smith 1963; Rich & Jermy 1998). In Fermanagh, the species has not beenexamined in sufficient detail to comment on these forms. As straightforward M. pratense,the species has been recorded in every Irish VC except Meath (H22) (Cen Cat FlIr 2).
Fermanagh occurrence
M. pratense has been recorded in 74 Fermanagh tetrads (14%), 66 of which have post-1975 records.As the distribution map indicates, it is widely scattered in the N and W of the VC, but only rare orabsent from considerable areas of the more intensively farmed east of the county.
British and Irish occurrence
While it is well distributed across the whole of B & I from south to north, although absent from theChannel Isles, M. pratense becomes scarce and local in E Anglia, the east Midlands, SE Scotlandand South Central Ireland. From the lowlands it ascends to 960 m in S Kerry (H1). M. pratense hasbeen in steady decline in both B & I since before the 1930s, but the decline has accelerated sincethen. This is due to a combination of habitat loss and changes in land use involving the felling ofwoods and weedy overgrowth caused by woodland management neglect (F.J. Rumsey, in: Preston et al.2002). In N Scotland, M. pratense appears to be limited to areas of peatland that are too wet toburn, as heaths and bogs are managed by periodic firing to reinvigorate sub-shrub grazing.
European and world occurrence
It belongs to the Eurosiberian boreo-temperate phytogeographical element and is present in most of Europeexcept the south, where it is absent from most of the Mediterranean. It is also in W Asia, east to theYenisei region in Russian Siberia (Clapham et al. 1987; Hultén & Fries 1986, Map 1670).
Toxicity
Cow-wheat, like other members of the Scrophulariaceae, does contain poisonous glycosides which couldcause illness and death in the unlikely event of animals eating a large quantity of them (Cooper &Johnson 1998).
Names
The genus name appears in Theophrastus, referring to a plant growing among wheat, 'Melampyrum' means'black wheat' ('melas'= black; 'pyros'= wheat), the seeds being considered to resemble black grains ofwheat (Gilbert-Carter 1964). The Latin specific epithet 'pratense' means 'growing in meadows'(Gilbert-Carter 1964), which is not quite what we observe today, since the species is much more of awood and heathland species. This objection also applies to the seed colour referred to in the meaning ofthe genus name, which is brown, not black. The current author concludes that both of this species' namesrefer to a black-seeded plant (almost certainly M. cristatum (Crested Cow-wheat)) that grewamongst and probably parasitized cereal grasses. The seeds of M. cristatum are black and those ofM. arvense are brown but go black (Britton 1943; Butcher 1961).
Prior (1879) suggests the common name, 'Cow-wheat' derives from the seed resembling wheat, the 'cow'element indicating their worthless nature as human food.
Grieve (1931) recounts some curious medieval folk-lore which may relate to the prevalence of the speciesin cornfields on land recently cleared of wood, presumably using fire.
Threats
None.
Native, frequent.
1881; Stewart, S.A.; Co Fermanagh.
Throughout the year.
This is a 'catch-all' aggregate for this taxonomically very difficult genus. All seven editions of Webb'sAn Irish Flora have stressed the difficulty of identifying Irish material, mentioning the problemof unspecified differences from species descriptions based on British and Continental material andcomplications involving intermediate hybrids. Hybridisation, regular self-pollination and inbreedingtogether have given rise to stabilised distinct lines of eyebrights and created endemic microspecies(P.F. Yeo, in: Stace 1975; Ingrouille 1995).
As pointed out in the Metherell & Rumsey (2018) BSBI Handbook Eyebrights (Euphrasia) of theUK and Ireland, "Even the arrival of DNA analysis has done little, at this stage, to dispelthe fog of uncertainty surrounding this genus." These authors are also of the opinion that,"Unfortunately Euphrasia is, in taxonomic terms, a very badly behaved genus! It is verydifficult to decide what is a species in Euphrasia." The arrival of this well produced,illustrated handbook, should, it is hoped, stimulate further study of the genus in Fermanagh and in therest of B & I.
Eleven microspecies were keyed and very briefly described in An Irish Flora (1996), and eight ofthese have either small or modest numbers of records in Fermanagh, some of which have been verified bythe referee and British authority on the genus, Dr P. Yeo. The current author (RSF) and RHN do notpretend, however, that the knowledge of Euphrasia occurrence in Fermanagh is other thanrudimentary and we recognise that, as with other critical plant groups, much more focussed work remainsto be done. The treatment and references given by A.J. Silverside in the BSBI Plant Crib,together with Webb's An Irish Flora account, would be a suitable starting point, the former'sapproach being partially based on habitat and the known geographical distribution of the microspecies(BSBI Plant Crib 1998). The BSBI Euphrasia Handbook supersedes these two sources ofidentification information and is certainly the best account, both in terms of descriptions andillustrations that there is of these difficult little facultative hemi-parasitic annuals.
This latter technical term, 'hemi-parasite', means that Euphrasia species and hybrids arefree-living and can produce their own energy through photosynthesis, but they may also attach themselvesby means of underground haustoria, root to root, xylem to xylem, with the living roots of neighbouringplant species (often grasses or legumes). They can then extract water and inorganic nutrients from theirhost plant(s), through their hemi- or semi-parasitic root attachment (Metherell & Rumsey 2018).
As a consequence of the above identification difficulties, most local field recorders including ourselveshave regularly consigned observed or collected eyebright forms into this aggregate, rather than simplyignore them altogether. The majority of the 321 records of this Euphrasia aggregate in theFermanagh Flora Database were recorded from rocky situations in limestone districts, in a wide varietyof sites at all altitudes except the very highest (ie Cuilcagh summit). They are scattered over 164Fermanagh tetrads, 31.1% of those in the VC, but are mainly found on the Western Plateau uplands onscarps and rocky terrain, plus in quarries and sand-pits. A small number are associated with muchwetter, more acidic habitats, including fen, riverside and lakeshore, bog and cutover bog and, had theybeen discriminated, these would probably belong to one of E. nemorosa, E. micrantha orE. scottica.
Native, apparently rare, but probably under-recorded. European boreo-temperate.
1948; MCM & D; damp pastures and grassy roadsides by Lough Melvin near Garrison.
June to September.
Apart from E. salisburgensis (Irish Eyebright), this is possibly the most distinctive microspeciesof the genus known to occur in Fermanagh. E. rostkoviana is an erect plant up to 40 cm tall, butmore usually 20-25 cm in height, branched from near the base. It has large flowers for the genus and theupper part of the plant is grey-hairy with long glandular hairs.
Considering its rather distinctive appearance, it is surprising that E. rostkoviana has so veryfew Fermanagh records and, like every member of the genus, it is very probably under-recorded. E.rostkoviana has been recorded in just nine Fermanagh tetrads, only five of which have post-1975records. It has been found in meadows, pastures and in rough grass on roadside verges. The taxon issometimes referred to as E. officinalis subsp. rostkoviana and,very probably, many of the forms of E. officinalis agg. recorded in damp, river- and lake-sideherb-rich, hay meadows and longer, rougher roadside grassland, really should belong to this microspeciesand be listed here.
The Cen Cat Fl Ir 2 lists 22 VCs in which E. rostkoviana has been recorded at least once,but it fails to include Fermanagh. The four records made by Meikle and co-workers in 1948-50 have beenverified by P. Yeo, but the post-1975 ones have not. The remaining record details are: Upper LoughMacnean, 1950, MCM & D; Lower Lough Macnean, 1950, MCM & D; Mullylusty Td, Lurgan River Glen,1950, MCM & D; grassland by Glasdrummond Bridge, Swanlinbar River, 15 July 1987, RSF & RHN;meadow downstream of Roogagh Bridge – the Cirsium helenioides (Melancholy Thistle) site, 19 June1988, RHN; S of road at Knockennis, 8 September 1994, RHN; Crossmurrin, 25 June 2000, RHN; and meadownear Killy Beg, NE of Garrison, 14 June 2009, RHN & HJN.
It is clear from the New Atlas hectad map and the accompanying brief species account that thistaxon, although not well recorded anywhere in these isles except possibly Wales, is particularlyunder-recorded in Ireland. The authors of the BSBI Euphrasia Handbook (Metherell & Rumsey2018) also recognise the extent of Irish under-recording of this species.
E. rostkoviana forms hybrids with perhaps seven other Euphrasia microspecies, although allof them are rare, extremely rare or very local in B & I. E. rostkoviana × E. angliahas been recorded once in Co Dublin, as has the hybrid with E. arctica, but these are the onlyrecords from Ireland so far (Doogue et al. 1998; Stace et al. 2015; Metherell & Rumsey2018).
Apparently very rare, possibly mis-identified, but probably under-recorded. Oceanic temperate.
1952; MCM & D; sandy lakeshore, at Derreens West, near the outflow of Lower Lough Macnean.
As with E. rostkoviana, this taxon is sometimes referred to as a subspecies of E.officinalis (Silverside 1991). There is some doubt about the identification of this solitaryrecord as in the 1975 Revised Typescript Flora prepared by R.D. Meikle the species name is addedin pencil, prefixed by a question mark.
E. anglica is outlined in An Irish Flora (1996) as being like E. rostkoviana inhaving the upper parts furnished with the same covering of glandular hairs visible to the naked eye as agreyish down, but the plants are less erect, more branched and with a smaller corolla.
A.J. Silverside (in BSBI Plant Crib 1998) lists it as a taxon of damp heath and pastures and F.J.Rumsey (in Preston et al. 2002) adds that it frequents tightly grazed habitats on damp, acidsubstrates. The types of habitat range from old, unimproved pastures, heaths and moors, to disturbedartificial situations in disused quarries. Suitable examples of all these habitats are as widespread inFermanagh as they are elsewhere in Ireland, but forms of this taxon are much more distinctive from E.rostkoviana in SE England and its particular form in Ireland is not readilydistinguished. More work on Irish material is clearly required.
Elsewhere in Ireland, E. anglica has been recorded as occasional in heathy grassland and presentalmost throughout the Burren (H9) and Connemara (H16). Here, the Flora of Connemara and theBurren described the plant as more straggling than E. rostkoviana, with often flexuousstems and slightly smaller flowers. The New Atlas hectad map displays a much more widespreadrepresentation in Connemara than in the Burren, Co Clare (H9), and elsewhere a very sparsely scatteredIrish distribution, especially in the north of the island. The current author (RSF) and RHN concludethat this is a difficult taxon to identify, but it would be worthwhile looking out for it.
E. anglia forms hybrids with six other Euphrasia microspecies (Stace et al. 2015;Metherell & Rumsey 2018) but only one of these (with E. rostkoviana) has ever been recordedin Ireland, in Co Dublin by Doogue et al. (1998).
Euphrasia arctica subsp. borealis (F. Towns.) Yeo,Arctic Eyebright
Native, local. Oceanic boreo-temperate.
1900; Praeger, R.Ll.; Lower Lough Macnean.
July to October.
E. arctica is a very variable, tetraploid annual hemi-parasite that is recognised by its veryshort-stalked glandular hairs, which are visible using a ×10 hand lens and holding the plant against thelight. The plant is robust with large leaves and flowers and it often has a number of long, spreadingbranches (Flora of Connemara and the Burren). The leaves are not fleshy, like in E.tetraquetra (Western Eyebright), nor are the inflorescence bracts crowded and overlapping as inthe latter. The typical habitat is described as damp, disturbed, marginal grassland and the authors ofAn Irish Flora (1996) regard it as, "nearly everywhere the commonest [Euphrasia]species".
There are two subspecies within E. arctica in B & I, but the New Atlas map demonstratesthat while subsp. borealis is without doubt widespread throughout the N and W of these isles,subsp. arctica is much more confined, appearing only definitely in Orkney and Shetland, butpossibly also in other areas of Scotland and the English Lake District (F.J. Rumsey, in: Preston etal. 2002). The BSBI Euphrasia Handbook makes no mention whatsoever of subsp.arctica (Metherell & Rumsey 2018).
In Fermanagh, subsp. borealis has been thinly and widely recorded in 42 tetrads (7.9% of the VCtotal), but only 31 of them have post-1975 records. Locally, it appears to be a plant of disturbedgrassy areas on roadside verges, quarries, lakeshores and any other areas of rough, disturbed,grassland, such as around gateways. There is considerable uncertainty at present associated with recenttaxonomic revision of E. arctica s.l., confusion with E. nemorosa (Common Eyebright) andhybridisation with the latter. Consequently, throughout B & I, E. arctica subsp. borealis is believed to be under-recorded to some degree.
The BSBI Euphrasia Handbook emphasises the very variable nature of this species (referring to itas E. arctica Lange ex Rostrup) and the fact that, as an outbreeding species, it readily formshybrids with a wide range of other Euphrasia microspecies. However, the authors point out that inthe majority of cases, the large flowers and broad leaves of E. arctica should serve to separateit from most other species.
According to the Euphrasia Handbook, E. arctica forms hybrids with no less than 15 otherEuphrasia microspecies, although some of them are very rare and many are extremely local(Metherell & Rumsey 2018).
In the Hybrid Flora, there are six Irish records of a hybrid between E. arctica and E.nemorosa, four with both parents present, one with just E. nemorosa present and one withneither parent present (Stace et al. 2015). The hybrid between E. arctica and E.confusa (Confused Eyebright) has been recorded in three Irish sites around the Dublin area, andthe hybrid with E. micrantha (Slender Eyebright) has been found in another three Irish sites onthe River Shannon and in Connemara. The hybrid with E. scottica (Scottish Eyebright) adds threemore Irish sites, one near Dublin and two in the Connemara region. There is some possibility of anotherhybrid with E. salisburgensis (Irish Eyebright), but apparently there is not yet sufficient hardevidence to substantiate this claim (Stace et al. 2015).
Native, very rare, but a possible mis-identification. Oceanic temperate.
16 August 1992; Northridge, R.H.; scattered scrub woodland at Templenaffrin, near Belcoo.
August.
This annual species or microspecies with erect, purplish stems up to 15 cm is recognised by itsquadrangular flower spike of dense, overlapping bracts and flowers, its rather fleshy leaves and itsshort-stalked, glandular hairs. Overall, the plant has a distinctly four-sided habit. While usuallydistinct, Silverside remarks in his BSBI Plant Crib 1998 account of the genus, thatE. tetraquetra is often confused with condensed, ie compact, dwarf forms of E. nemorosa(Common Eyebright). In N Scotland, in particular, confusion may also occur with dwarf forms of E.arctica (Arctic Eyebright) and E. foulaensis (Foula Eyebright), which can look verysimilar (Metherell & Rumsey 2018).
Although it is occasionally or rarely found inland on grazed limestone pastures, E. tetraquetra isvery much more frequently recorded on short turf on exposed sea cliffs, stabilised sand dunes and ingrassland near the sea and it is widely recorded in such sites especially around the N and W coasts of B& I (Flora of Connemara and the Burren; New Atlas).
RHN recorded this species in scattered scrub woodland on limestone terrain near Belcoo and again the nextday on short grassland near the stony shore of Lower Lough Macnean adjacent to Templenaffrin. However,there is no mention in the Fermanagh Flora Database of a voucher being checked and verified by anexpert. Until this safeguard has been achieved, the possibility that RHN may have come across a dwarfE. nemorosa has to remain a possibility for these two very isolated inland records.
According to the BSBI Euphrasia Handbook, E. tetraquetra forms hybrids with eight otherEyebright microspecies (Metherell & Rumsey 2018). There are two Irish records of the hybrid withE. nemorosa from the Dublin area (H21) and one Irish record of the hybrid with E.micrantha (Slender Eyebright) from Bunduff Dunes, Co Sligo (H28) (Stace et al. 2015).
Native, occasional, locally frequent. European temperate, introduced in N America.
1900; Praeger, R.Ll.; Florencecourt.
June to September.
In Fermanagh, this annual hemi-parasite is locally frequent on thin, heathy pasture over limestone in awide variety of circ*mstances, from lowland lakeshores and sand pits, to screes and around cave mouthsin the upland limestones of the county. There are over 80 records from 22 tetrads in the VC (4.2%),although only eleven of them have post-1975 records. The current author (RSF) and RHN would thereforenot claim this as anything more than an occasional species, although it is certainly under-recorded. Asfar as we can tell, it appears scattered across the limestones lying to the west of Lough Erne.
As is the case with E. arctica (Arctic Eyebright), E. nemorosa is probably one of the mostcommon species of the genus, but like other eyebrights, it is under-recorded due to difficulty inseparating the species. Consequently, in Fermanagh, it is often lumped in the E. officinalisaggregate when recording.
E. nemorosa and E. micrantha (Slender Eyebright) differ from most other species in havingdistinctly mauve flowers (rather than white tinged with mauve), but it can also have both lips white.Given adequate growing conditions, E. nemorosa is a stouter, bushier plant than E.micrantha and it has larger, medium-sized flowers; the lower lip of the corolla, measuring 7-9mm across, is usually deflexed, ie bent down or backwards (Flora of Connemara and the Burren;An Irish Flora 1996; Metherell & Rumsey 2018).
E. nemorosa is a relatively late-flowering species, generally flowering at node ten or above, thecorollas being usually smaller than in the closely related E. arctica (≤7 mm whereas E.arctica can have flowers up to 13 mm). Metherell & Rumsey (2018) point out that, "Thetoothing of the leaves, particularly the lower floral leaves, is [a useful distinguishing character];almost always aristate in E. nemorosa, generally less constantly so in E. arctica(although the basal pair or basal two pairs of teeth are often aristate in that species) and almostnever so in E. confusa (Confused Eyebright)".
Elsewhere in B & I, E. nemorosa can be categorised as a plant of short turf grassland onheaths, downs and dunes, plus in open scrub, woodland rides and upland moorlands. It avoids entirely thecompetitive situation of agriculturally improved land. E. nemorosa is regarded as the most commonand ecologically diverse form of Eyebright in B & I, yet this is qualified by the fact that itbecomes increasingly confined to lowland calcareous soils further north in these islands (F.J. Rumsey,in: Preston et al. 2002).
E. nemorosa can form hybrids with 15 other Euphrasia microspecies (Metherell & Rumsey2018). Irish E. nemorosa hybrid records featured in the Hybrid Flora of the British Islesinclude: two records of the cross with E. pseudokerneri (Large-flowered Eyebright); three recordsof the hybrid with E. confusa; and one record of the cross with E. micrantha, on the ArdsPeninsula, Co Down (H38) (Stace et al. 2015).
Native, very rare, but locally abundant and probably under-recorded. Oceanic boreo-temperate.
1953; MCM & D, det. P. Yeo; side of path at Castle Archdale.
Due to its flowering early in the season, this annual tetraploid species is probably under-recorded.There are a total of ten Fermanagh records from eight tetrads, four of them with post-1975 dates. Thefirst four records were made by Meikle and co-workers between 1953 and 1957, although there are noprecise dates for three of them. On the original card index kept by R.D. Meikle as BSBI VC recorder, thecomment made by the determiner P. Yeo on 11 December 1953 on the first record is written as follows,"The nearest approach to that species that I have seen from Ireland, and similar to W Cornishplants that appear intermediate between E. confusa and E. nemorosa." The site detailsof the other three MCM & D records (all det P. Yeo and presumably with vouchers at K) are:Gubbaroe Point; Cornaleck shore, just SE of Landing Place, Upper Lough Erne; and Derreens West shore,Lower Lough Macnean.
The fourth record of E. confusa was made by RHN (accompanied by HJN on the first outing) on 4 May2000 at the limestone knoll of Knockninny overlooking Upper Lough Erne. The notes made of the privatelykept voucher plant detailed it as being small (4.5 cm), no hairs, flowers largely white and, obviously,it was in flower in early May. Webb & Scannell (Flora of Connemara and the Burren) remarkedthat in their experience in the W of Ireland, plants of E. confusa are usually very dwarf andoften unbranched. Since 2000, RHN has recorded this eyebright on a further five occasions in shortlimestone turf and, as the tetrad map indicates, all the sites were in the south of the VC.
Elsewhere in B & I, E. confusa is an annual of grazed pastures and grassy heaths growing onfree-draining, acid or calcareous soils. It is said to be especially characteristic of heavily grazedhill pastures in N & W Britain, but is occasionally found in open vegetation on sandy soilselsewhere. While it is a widespread and frequent species in Britain, it is much rarer and mainly coastalin Ireland. In the FNEI 3, however, it is described as, "Frequent in the Mourne Mts.Apparently very local elsewhere, but requires further investigation."
As with many other eyebrights, however, E. confusa hybridises prolifically with at least 15 otherEuphrasia microspecies (Metherell & Rumsey 2018), making plants often very difficult toidentify; it is therefore probably under-recorded everywhere, not just in Fermanagh. Up until 2014 or2015, none of the hybrids involving E. confusa have Irish records (Stace et al. 2015).
Native, rarely listed but probably under-recorded. European temperate.
August 1938; Praeger, R.Ll.; Black River.
July and August
This annual, tetraploid species with slender, erect, usually dark purple stems, grows up to 25 cm tall.It has long internodes, the upper ones shorter than or more usually 2-4 times as long as the leaves.Lower floral internodes 1.5-2.5(-3) times as long. The leaves are glossy dark green, often suffusedpurple on both surfaces. The flowers are distinctive, being small and bright purple with a yellow splashon the lower lip (Metherell & Rumsey 2018).
E. micrantha grows on damp to dry places on upland heaths, moors and lakeshore bogland, usuallyamongst Calluna vulgaris (Heather) and other ericaceous sub-shrubs. In other parts of B & I,it is also reported from open clay to sandy substrates in disturbed artificial habitats, includingdisused gravel-pits and quarries and on old airfields (F.J. Rumsey, in: Preston et al. 2002).
It is only very occasionally recognised and recorded in Fermanagh and although there are records from 19tetrads (3.6%) in the VC, only ten of them have post-1975 records. This undoubtedly reflects thedifficulty of obtaining certain identification of microspecies in this genus and a determined recordingeffort is required in order to provide a more accurate picture.
In the past, E. micrantha was often confused with E. scottica (Scottish Eyebright) and itoften forms hybrids, especially with E. nemorosa (Common Eyebright), greatly adding to theidentification difficulty. The BSBI Euphrasia Handbook details a total of 13 species with whichE. micrantha forms hybrids (Metherell & Rumsey 2018). There is one Irish record for thehybrid with E. scottica from south Co Down (H38); two records of the hybrid with E.salisburgensis (Irish Eyebright) from W Galway (H16); one record of the hybrid with E.nemorosa from Co Down (H38) and one record of the hybrid with E. tetraquetra (WesternEyebright) from Co Sligo (H28) (Stace et al. 2015).
Elsewhere in Ireland, E. micrantha has been recorded from 30 of the 40 Irish VCs, with a greaterfrequency in the northern half of the island (Cen Cat Fl Ir 2; An Irish Flora 1996). Thisis still apparent in the hectad map published in the New Atlas. At the same time, and purely forcomparison, the FNEI 3 lists only two post-1960 records for Cos Down (H38) and Londonderry (H40),and none at all for Co Antrim (H39).
Native, very rare, but probably under-recorded. European boreal-montane.
July 1946; MCM & D; northern slopes of Belmore Mountain.
This annual tetraploid species, which is associated with upland wet moors, flushes and bogs, has onlybeen recorded in Fermanagh on three occasions, all before 1976 and never by the current author(RSF) or RHN, the Joint BSBI County Recorders. All three records have been checked by Dr P. Yeo and avoucher for Miss Scannell's 1975 record exists in DBN. In common with other members of the genus,it is undoubtedly under-recorded.
Elsewhere in Ireland, E. scottica has been listed from a total of 22 Irish VCs (Cen Cat Fl Ir2) and its distribution was described in An Irish Flora (1996) as, "Occasional inthe West and Centre, rare elsewhere". The New Atlas hectad map shows it as rather thinly andwidely scattered, mainly in the northern half of Ireland.
In Britain, the New Atlas map indicates a definite N & W upland distribution of E.scottica although, as with all eyebrights, identification confusion with other microspecies andhybridisation means it is under-recorded (F.J. Rumsey, in: Preston et al. 2002).
E. scottica is a slender, erect, sparingly branched plant, with small white flowers. The upperleaves are small, narrow and bluntly toothed; they and the bracts are green (not bronze) and they areonly about twice as long as broad.
E. scottica is very similar to E. micrantha (Slender Eyebright) and the two may not bespecifically distinct. E. scottica flowers earlier than E. micrantha and has lessred-brown pigment on the flower corolla and vegetative parts, with the exception of the under-leaf,which is often distinctly purple (New Flora of the BI 1997). Hybrids between E. scotticaand E. micrantha are readily formed and are thought to be very common, particularly in Scotland.In addition, E. scottica is known to form hybrids with a total of eight other Euphrasiamicrospecies apart from E. micrantha (Metherell & Rumsey 2018). There are three Irish recordsof the hybrid between E. scottica and E. arctica (Arctic Eyebright), one from Co Dublin(H21) and two from W Galway (H16), and one record of the hybrid with E. micrantha from south CoDown (H38) (A.J. Silverside, in: Stace et al. 2015).
The details of the other two Fermanagh records of E. scottica are: Lough Jenkin, MullaghfadForest, 1948, MCM & D; and Meenagleragh Lough, NW of Little Dog, 29 July 1975, M.J.P. Scannell &D. McClintock, DBN.
Native, occasional and very local. European boreo-arctic montane.
1904; Praeger, R.Ll.; western end of the Cliffs of Magho.
May to September.
A distinctive annual species with small white flowers, the leaves and bracts are usually copper, bronzeor a deep chocolate brown in colour, narrower than the other Euphrasia microspecies and with verysharply or jaggedly toothed leaf margins. The plant is generally very dwarf (up to only 14 cm tall),especially when grazed, but it can sometimes become quite a branched, bushy little entity. If in anydoubt, examination of the capsule assists identification; in all other Irish eyebrights the upper marginof the capsule is fringed with hairs, but in E. salisburgensis it is glabrous (Flora ofConnemara and the Burren; Metherell & Rumsey 2018). E. salisburgensis is onlyrecorded in Ireland, not in Britain. Records of it from Yorkshire are errors (Metherell & Rumsey2018).
E. salisburgensis forms hybrids with E. tetraquetra, E. nemorosa, E.micrantha and E. arctica. It may also hybridise with E. scottica and E.frigida, but there is uncertainty with respect to these two pairings at present (Metherell &Rumsey 2018).
Irish Eyebright is found on shallow, dry, limestone soils and it is closely associated with Thymuspolytrichus (Wild Thyme) upon whose roots it is semi-parasitic. In Fermanagh, it has beenrecorded in 16 tetrads, all but one with post-1975 records. It grows on limestone or base-rich montanescarps, screes and shallow limestone pastures and, as the tetrad distribution map indicates, it isalmost confined to the limestone uplands around Knockmore Hill and Monawilkin on the Western Plateau.The only other two more remote sites are at the Monastair Gorge above the Marble Arch (or CladdaghRiver) Glen and at Banagher, 3 or 4 km NW of Enniskillen.
As the specific epithet might suggest, E. salisburgensis is common and ranges widely in the Alps,Pyrenees and other mountains of C & E Europe, but it also occurs at lowland sites further north inScandinavia (Grey-Wilson & Blamey 1995).
In the British Isles, the species is completely confined to suitable limestone soils in the westerncounties of Ireland, pre-eminently in the famous karst limestone districts of the Burren and Connemarafurther south. In NI, E. salisburgensis is confined to Fermanagh, where many of its sites are ofsufficient scientific interest to be designated as ASSIs for their conservation interest. E.salisburgensis is also listed by the NI Environment Agency as one of 68 vascular plant speciesof priority conservation concern. The Irish plants are recognised as a named endemic variety, var.hibernica Pugsley.
Native, locally frequent to abundant. Eurasian temperate, naturalised in eastern N America.
2 August 1970; Farrell, Ms L.; between Monawilkin Lough and Dromore Hill.
Throughout the year.
Growth form and preferred habitats
O. vernus is a stiff, erect, freely branched, annual, 30-50 cm tall, with wiry, 4-angled stemsbearing opposite leaves that are stalkless, narrow, linear-lanceolate, margins rather remotely toothedand surfaces heavily hispid with short, bulbous-based hairs. The flowers are a dull or palepurplish-red, but albinos also commonly occur.
Red Bartsia is a hemi-parasite, attaching its fibrous roots to the roots of grasses and purloining aproportion of their assimilates to assist its own growth and reproduction. In general, semi-parasitesare comparable to unspecialised parasites and since most attach themselves to the roots of grasses theremust be comparatively little risk of the seeds not being near a suitable host plant (Salisbury 1942).
In common with other ruderals, O. vernus is often associated with disturbed sites such assandpits, quarries and neglected or derelict waste ground, eg locally in Fermanagh around the disusedrailway station at Maguiresbridge. The established strategy of O. vernus is categorised as R/CR,ie intermediate between a straight Ruderal species and a Competitive Ruderal (Grime et al. 1988,2007).
O. vernus colonises a wide range of soils from the decidedly damp to the very dry, although somelakeshore sites may be variable in this respect, becoming dry in summer, yet winter wet. It appears toprefer somewhat heavier clay or loam soils that are moderately acid to calcareous and sunny (Salisbury1964; Sinker et al. 1985). Being a relatively poor competitor when among perennial weeds in opencommunities, although more competitive than some ruderal annuals, it is not surprising that Red Bartsiais absent from more fertile soils, except where they are trampled, heavily overgrazed (eg by cattle or,more likely, horses) or otherwise regularly disturbed. Thus O. vernus is quite often foundoccupying bare patches in moderately grazed, short-turf, permanent grassland, or on the margins ofarable fields (although these are rare nowadays in Fermanagh). Other sites are open situations aroundfield gateways and in vegetation gaps on trampled or otherwise regularly disturbed ground, such as alongwayside grass verges. Presumably these environments allow it to avoid competition from vigorousestablished perennials in more closed, taller, denser vegetation (Sinker et al. 1985).
Flowering reproduction
O. vernus flowers from June to August (Garrard & Streeter 1983). The flowers, 8-10 mm long,are held in long, leafy one-sided racemes terminal on the stem and side branches. The individual flowersare tubular, two-lipped, about 1.3 mm long and softly hairy. The four hairy anthers are held closetogether, slightly exserted from the mouth of the corolla tube; the anthers are aptly described as,"peeping out from under the hood of the upper corolla lip" (Hutchinson 1972). The style islong, slender and hairy. Pollination is carried out by bees and occasional butterflies (Proctor &Yeo 1973). The small brown seeds are roundly ovoid, 1.5 mm long, with longitudinal whitish ribs and finetransverse ridges connecting them. The fruit capsule is oblong and pubescent and is surrounded by thepersistent calyx. Each capsule contains about 15 seeds which are shed in autumn but do not germinateuntil early in the following spring. Some seed is also long-term persistent in the soil seed bank (iesurviving more than five years) (Butcher 1961; Salisbury 1964; Clapham et al. 1987; Thompsonet al. 1997).
The range in size of individual plants is considerable, the largest specimens examined by Salisbury(1942) bore 445 capsules and the smallest just nine. The mean capsule number on 124 specimens was 112per plant and the mean seed number was 14.8 per capsule, with a range from 1-36. The mean seed outputwas therefore approximately 1,660 ± 105 seeds per plant (Salisbury 1942).
Variation
O. vernus (= Bartsia odontites L.) Hudson, O. rubra Gilib.) is a very variablespecies which is difficult to separate adequately into infraspecific entities. Sometimes segregates aretreated as subspecies and at others as species. Three subspecies of O. vernus are now recognised(Stace 1991, 1997), but they have not been distinguished by most of the recorders working in Fermanagh.
Subsp. vernus has stems to 25 cm, with 1-4 pairs of branches, but occasionally has no branches instarved plants. It also has its lowest flower at node 6-9 and leaves lanceolate and distinctly toothed.It is recorded from most of B & I, but there are many recording errors involved in parts of the S ofEngland and Wales; it is scattered in Ireland, Wales and N Britain. Subsp vernus replaces subsp.serotinus in most of C & N Scotland and it is mainly aestival (summer flowering).
Subsp. serotinus (Syme) Corb. has taller stems, 2-50 cm, with 2-8 pairs of branches spreading atright angle and with their tips often upcurved. Leaves are linear-lanceolate, somewhat narrowed at thebase, margins obscurely toothed. It is considered frequent over most of B & I, except in C & NScotland. It is the only subspecies in most of C & S Britain, and its flowering is mainly autumnal(Stace 2019).
In the Fermanagh Flora Database, there are a total of just 19 records of subsp. serotinus, and asthe tetrad map for this taxon shows, it is widely scattered across 16 tetrads. This form of the plant isconsidered to be autumnal (Stace 2019). As the map for subsp. serotinus illustrates, a solitaryrecord (made by the late Nora Dawson at Aghiver, just N of Castle Archdale in April 1976) has apost-1975 date.
It is claimed that the majority of Irish O. vernus plants belong to subsp. serotinus(Hackney et al. 1992), but the current author (RSF) and RHN have not examined the local situationin Fermanagh and therefore cannot make any pronouncement.
The third subspecies, subsp. litoralis (Fr.) Nyman (= O. vernus subsp. pumilus auct. non (Nordst.) A. Pedersen), is a dwarfcoastal form with stems to 20 cm, 0-3 pairs of branches and the lowest flower at node 4-8. It occurs ongravelly and rocky sea-shores and in saltmarshes on the coasts of N & W Scotland from Kintyre (VC101) to Shetland (VC 112), Westmorland (a salt-marsh at Askam-in-Furness) (VC 69) and Merioneth on the WWales coast (VC 48). It is described as aestival (ie summer flowering) (Snogerup 1982; Halliday 1997;Stace 2019).
Fermanagh occurrence
The species has been recorded in 103 Fermanagh tetrads, 19.5% of those in the VC. Apart from disturbedpermanent pasture situations, locally it is mainly recorded on roadsides and along upper lakeshoregrasslands. As the species tetrad distribution map shows, it is thinly scattered throughout Fermanagh,but apparently more frequently around Upper Lough Erne. The excessively detailed sampling of the latterregion produced by the EHS Habitat Survey in 1986-7 has significantly skewed the overall speciesrepresentation towards this area of the county and, therefore, this concentration of O. vernusrecords should not be taken too seriously.
British and Irish occurrence
Widely distributed throughout B & I. Of the three subspecies, subsp. vernus is commonest innorthern Britain, but only local and infrequent in the south. Subsp. serotinus is common insouthern England, but is rare or absent from much of N Scotland. Subsp. litoralis is a dwarfplant confined to coastal grassland in a few places in N Scotland and the Outer Hebrides (Garrard &Streeter 1983; Clapham et al. 1987).
European and world occurrence
O. vernus is well distributed across temperate Europe and Asia, although while present in most ofthe W Mediterranean, it is absent from the eastern portion of the basin. The distribution thins markedlynorthwards of S Scandinavia, although it is not entirely absent. It is present in eastern N America, butnot native there (Hultén & Fries 1986, Map 1684).
Names
The genus name 'Odontites' was a name used by Pliny for a plant regarded as 'good for toothache' derivedfrom the Greek 'odonto', or 'odons', 'a tooth' and the Greek meaning 'connected with' (Gilbert-Carter1964). The Latin specific epithet 'vernus' means 'spring flowering' (Gledhill 1985).
The English common names include 'Eyebright', 'Eyebright Cow-wheat', 'Red Eyebright', 'Red Odontites','co*ck's comb', 'Hen-gorse', 'Poor Robin', 'Sanctuary' and 'Twiny-legs' (Grigson 1955, 1987; Melderis& Bangerter 1955). Grigson is particularly dismissive of the plant, regarding it as having no strongcharacteritics, "No peculiarities, no beauties, no virtues. A red, dullish, disregarded annual ofthe cornfields, a weed which has not even incurred the hatred of farmers." (Grigson 1955, 1987).
Threats
None.
Native, very rare. Mediterranean-Atlantic.
25 August 1996; Northridge, R.H.; E of Rossmore Point, Lower Lough Erne, Castle Archdale estate.
Growth form and preferred habitats
An attractive, erect, rigid, usually unbranched, grey-green, annual root hemi-parasite with fibrous rootsthat parasitise grasses and other associated herbs. Up to 50 cm tall, P. viscosa, as its speciesname indicates, is clothed overall with downy, sticky (viscid), glandular hairs. It has oblong, toothed,stalkless leaves that can be opposite or alternate.
Scarce or rare as a native plant species in both B & I, Yellow Bartsia usually grows in lowland,damp, sunny, rough- or pasture grasslands, often on nutrient-poor, stony, sandy soils close to the sea.Sites are often along tracksides or in pasture grassland that is patchily grazed, ie subject to moderatelevels of disturbance that help keep the short turf open, providing suitable vegetation gaps forgermination and establishment of P. viscosa on host plants (Garrard & Streeter 1983; Perring& Walters 1989; French 2020). Fixed dune grassland in dryish, dune slacks is one of itscharacteristic habitats and poor, infertile pasture reclaimed from dampish heathland is another (M.C.F.Proctor, in: Stewart et al. 1994).
Flowering reproduction
Seeds germinate in the spring and, being an annual species, late-flowering, from June or July to October,P. viscosa reproduces entirely by seed. The inflorescence is a terminal spike-like raceme offlowers in the axils of leaf-like bracts. The large, yellow two-lipped flowers, 16-25 mm in length,11-13 mm in diameter, have a long corolla tube (a little longer than the tubular calyx), the lower lipmuch longer than the upper hooded one (Perring & Walters 1989). The four stamens are included in theupper corolla lip. Pollination is carried out by Hymenoptera and Diptera (ie bees, butterflies andflies) (Fitter 1987).
The fruit is a hairy, narrow capsule, 7-9 mm, slightly longer than the calyx tube, and contains manysmall seeds, 0.4 × 0.3 mm, finely striate and brown in colour (Butcher 1961). There is no oil body onthe seed, so ant-assisted dispersal does not occur in this species (Sell & Murrell 2007). Thecurrent author (RSF) has not been able to find basic information on such topics as seed number (apartfrom 'many small seeds') per capsule, mean number of flowers per plant, mode of seed release anddispersal, mean level of seed production, duration of survival in the soil seed bank and percentagegermination. Clearly work is required to remedy this situation and any observations would be verywelcome.
Fermanagh occurrence
P. viscosa is usually associated with damp, open sites on sandy, grassland that typically becomeflooded in winter, such as occurs for example in fixed coastal dunes or in other low-lying ground nearthe sea, so its discovery for the first time at Castle Archdale on the shore of Lower Lough Erne made byRHN in 1996 was quite remarkable. A second Fermanagh site emerged in 2001 when RHN and F. Carroll cameacross three individual plants growing in damp ground at Cross Townland, on the northern outskirts ofEnniskillen town. In October 2004, RHN found a third site, again at Castle Archdale, where at least oneplant occurred in an old reseeded lawn at the front of the remains of the major house of the estate.
British and Irish occurrence
In Britain, there has been some awareness in recent times that this species is in a phase of expansionfrom its previous known distribution, the plant being now observed colonising in both northern andeastern directions. Very probably, this is the result of recent introductions from imported wild flowerseed mixtures (M.C. F. Proctor, in: Stewart et al. 1994; Webb et al. 1996).
Throughout B & I, the native distribution of P. viscosa is frequent to very local andis largely confined to the S & W of both islands, but with scattered occurrences further north in NWEngland, S Scotland and N Ireland. In England, it is mainly found from Kent to Cornwall and the ChannelIslands, but it extends north to Dunbartonshire in W Scotland. The species is limited to lowland sitesand is most often coastal, occasionally occurring in abundance (Garrard & Streeter 1983).
Taken together, this pattern strongly suggests the species is temperature limited, requiring both a mildwinter and a following mild spring growing season. The native distribution is roughly bounded by the 5°Cmean January isotherm, which fits with this notion of a temperature limit to distribution (M.C.F.Proctor, in: Stewart et al. 1994).
In Ireland, it occurs in SW, NW and Western regions. P. viscosa has its most northerly occurrencein Ireland on the shores of Lough Swilly in E Donegal (H34), but it is much more prevalent and frequentin the SW of Ireland and almost entirely absent from the C & E (BSBI Atlas 2). In the NE ofIreland, Harron (1986) found it around Lough Neagh in five sites over four VCs, and it also occurred atthe coast near Ballykelly, Co. Londonderry (H40) (Hackney et al. 1992).
The New Atlas map displays a major species decline in SW Ireland, probably the result ofre-seeding of older pastures. Elsewhere in Ireland, the distribution appears to be stable. In Britainand the Isle of Man, the New Atlas hectad map shows introduced plants are increasingly frequentand widely scattered from the S coast of England to near Inverness in Scotland (F.J. Rumsey, in: Prestonet al. 2002).
European and world occurrence
P. viscosa belongs to the Mediterranean-Atlantic phytogeographical element and is widespread in S& W Europe, the Mediterranean basin, the Iberian Peninsula and W France. It is also found inMacaronesia, including N Africa and the Azores, and has its northern limit in W Europe near Lough Lomondin Scotland (M.C.F. Proctor, in: Stewart et al. 1994; Sell & Murrell 2007).
Names
The genus name 'Parentucellia' was given in memory of T. Parentucelli who was an important figure in therevival of learning and became Pope Nicholas V (1447-55). He founded the Vatican Library and the BotanicGardens at Rome (Melderis & Bangerter 1955; Gilbert-Carter 1964). The previous genus name was'Bartsia' and one of the English common names is 'Viscid Bartsia' (Butcher 1961). The genus name'Bartsia' was given in memory of a Dutch physician, J. Bartsch (1709-38) (Melderis & Bangerter 1955;Gilbert-Carter 1964). The Latin specific epithet 'viscosa' means 'sticky' or 'viscid', literally 'fullof bird-lime' ('viscum') (Gilbert-Carter 1964).
Threats
None.
Native, common. European boreo-temperate, widely naturalised, including in N America.
1866-70; Stewart, S.A.; Enniskillen Town.
May to November.
Growth form and recognition
A conspicuous, obligate hemi-parasitic, almost glabrous annual up to 1-30(-50) cm tall, with a shallowfibrous root and erect, simple or branched, usually black-spotted, hard, angular stems and opposite,narrowly oblong or linear toothed leaves and yellow, two-lipped flowers arranged in loose, leafy,spike-like racemes (Perring & Walters 1989). Yellow-rattle is unmistakable thanks to its veryprominent, pale-yellow inflated but flattened calyx. Thanks to this distinctive, persistent calyx, eventhe still standing dead 'ghost' of the plant can be readily identified long after the seed has beenshed. The only plant that might conceivable be confused with it, is the sticky and, in Fermanagh,extremely rare, Parentucellia viscosa (Yellow Bartsia).
Obligate partial-parasite ecology
Species of Rhinanthus are mainly found in low-sward, open grassland communities, and thechicken-and-egg question is asked, are they present due to their habitat preference, or is thevegetation a reflection of their presence? R. minor is shade intolerant and an open sward islikely to be important to allow its initial seedling establishment in spring. Being a hemi- orpartial-parasite, the roots of seedling Yellow-rattle attach themselves by haustoria to the alreadyestablished roots of neighbouring plants, mostly grasses, but also legumes, eg Trifolium repens(White Clover), creating xylem to xylem connectivity (Hodgson 1973). When this root connection is made,it enables the hemi-parasite to steal a proportion of its nutrient and solute requirements directly fromthe host plant. Having said this, there is no evidence of the parasite benefitting from acquisition ofphloem-borne photosynthates from its host. The benefits are restricted to xylem-borne water, dissolvedminerals and some organic compounds such as reduced nitrogen in the form of amino-acids, plus someorganic carbon (Jiang et al. 2004). In order to withdraw sap from the host xylem, the parasitemust compete with the host's transpiration stream by maintaining an osmotic potential gradient belowthat of the host plant. High concentrations of sugar alcohols, such as mannitol, help R. minor tomaintain this osmotic balance (Hodgson 1973).
Subsequent survival of R. minor, especially in a productive grass-clover sward, is based on theparasite's ability to make use of its host and, at the same time, it modifies the competitiverelationship between the parasite and its host in the former's favour. Consequently, the presence ofR. minor in open grassland communities is strongly related to management and the associated soilfertility/productivity level of any particular site (Westbury 2004). Yellow-rattle can infect a widerange of host plants, enabling it to colonise and establish itself in different grassland types,although it cannot invade all grassland communities equally well. Nevertheless, hemi-parasites likeR. minor can invade closed grassland habitats where other annual species are either rare orcompletely absent (Westbury 2004). R. minor plants have been observed to infect several hostssimultaneously. In sand dune vegetation, of a population of 65 R. minor plants studied, five wereattached to just one host plant, and two to at least seven different hosts (Gibson & Watkinson1989). The list of potential hosts for R. minor determined from four studies in Britain (Gibson& Wilkinson 1989) and one in Europe (Weber 1976), included 50 species from 18 plant families. Ofthese the Fabaceae (Legumes) accounted for eleven species and the Poaceae (Grasses) 16. Nine of thespecies were annuals or short-lived perennials, 36 were herbaceous perennials and five were woodyspecies (Gibson & Watkinson 1989). Legume species are considered good hosts for hemi-parasiticspecies, the additional nitrogen supply enabling an increased biomass and fecundity (Westbury 2004).
Some degree of autotrophic growth without host support can occur in R. minor, but such individualsare frequently stunted, growing only 5-7.5 cm tall; they do not form branches and may fail to flower. Inany event, they do not produce seed at levels comparable with heterotrophic hemi-parasitic individuals(Westbury 2004).
Apart from its predominant association with meadows on soils of moderate to low fertility, R.minor also features, to a more limited extent, in pastures and from ungrazed grassland, mire androadside verges, as well as displaying atypical occurrences on rock outcrops (Grime et al. 1988,2007).
Soil and habitat preferences
R. minor grows in permanent, less disturbed, infertile, damp grassland in a wide variety of soiland habitat situations. It can occur on substrates ranging in character from clay to sand, chalk,limestone and occasionally peat, but it is absent from sites with a soil pH below 5.0. It can becomelocally abundant in neutral or base-rich grassland. One of the soil types it rather commonly frequentsis alluvial or glacial till laid down in moraines or in drumlins. This type of soil can have either poorto impeded drainage or be free-draining. In the latter case, the traditional farming practice ofapplying lime and farmyard manure will buffer the loss of minerals caused by leaching, thus maintainingthe soil pH and favouring the survival of R. minor (Westbury 2004).
Traditional agricultural management of meadow/pasture grassland by first cutting hay in late-July,followed by autumn grazing cattle on the same ground, leads to an opening up of the sward for manyspecies, including R. minor, by reducing standing biomass and creating bare ground throughtrampling. The effect of this management is greatest in soils of low to moderate fertility (Westbury2004).
In terms of soil moisture regimes, Yellow-rattle cannot tolerate drought, but it can cope with meadowsthat become temporarily waterlogged in winter and the species extends occasionally into the margins ofsoligenous mire (ie lake-shore fen conditions) (Grime et al. 1988, 2007; Westbury 2004).
However, Yellow-rattle is most typically a plant of hay meadows and is especially associated with thoseof high floristic diversity (Grime et al. 1988, 2007). Being an annual, its survival in anylocation depends on its ability to release seed before the hay crop is cut. The very widespreadagricultural move in B & I from hay to silage fodder production, the latter of which is cut muchearlier, together with the reseeding of meadows and the spread of artificial inorganic fertilizers andslurry, have all resulted in a decline in the field prevalence of R. minor populationsnationwide. Yellow-rattle persists, however, in sites that are more marginal and in areas whereintensive agriculture is either uneconomic or impossible, for instance due to being inaccessible tolarge, modern machinery.
Phenology and flowering reproduction
Seed germinates in spring and plants flower from May to August. Each terminal raceme inflorescencecontains less than 30 bisexual flowers, each one with a flattened and inflated calyx with four teeth andthe upper lip of the bright or pale yellow corolla flattened from the side, enclosing the four stamens.Nectar is secreted at the base of the ovary and the flowers attract insects, mainly bees and bumblebees,that carry out pollination. If this fails to occur, the flowers can self-pollinate. As commonly happensin numerous other tubular flowers (eg Gentiana spp.), some Bombus bee species bite a holeat the base of the calyx and corolla and steal nectar, thus bypassing the legitimate flower entrance andavoiding enacting the pollination mechanism of the species.
Seed is set from June through to September. The capsule, 12-17 mm, is brownish, ovate and compressed(Sell & Murrell 2007). Seed production obviously varies enormously with vegetation, environment andseason, but as an indication of typical reproductive success, 171 R. minor plants in pasture hada mean capsule number of 17.30 ± 0.96, each containing an average of 9.7 ± 0.1 seeds, resulting in acalculated seed output of 168 ± 11 per plant. "It does not therefore appear that the production ofa large seed output is in any sense a necessary concomitant of the semi-parasitic habit."(Salisbury1942).
Whenever the capsule is mature, the large pale yellow ovate seeds, 3 × 2 mm, each of which is laterallycompressed and possesses a broad semi-circular wing (Butcher 1961), become loose inside it, and thusolder fruiting racemes make a distinctive rattling noise when shaken, a feature reflected in the largerange of English common names given to the species (see below). The ripe capsules split open at theirapex and the stiff stems of the plant are shaken by wind or passing animals, thus scattering the wingedseeds on the breeze.
The distance travelled by seed in this manner is slight or poor, unless aided by man. Seedlings tend tobe located at no more than 1.45 m from the edge of an existing population. Mowing machinery has beenshown to assist seed dispersal both in terms of distance and seed number dispersed and hay cutting andtransportation have also been shown to be an effective means of seed dispersal (at least 4 m), bettereven than sheep grazing (a maximum of 0.9 m) (Westbury 2004).
Dormant seed is reported to be ± transient, ie not persistent for more than a year in the soil seed bank,germination taking place in the spring following production (Roberts 1986).
Variation in population density
Populations of R. minor are often observed to fluctuate greatly in density from year to year andfrom site to site. On a sand-dune grassland in Norfolk, natural population densities varied from 100-200flowering plants per m2 in dense patches, to 0-50 m2 in less dense areas (Gibson1986), while in a SSI meadow in W Yorkshire, populations ranged from 596 to 1004 individuals perm2. These figures are considered rather low in comparison to populations in a twelve year oldmeadow in Canada, where mean high density was measured at 4,382 ± 1,236 per m², and low density at 405 ±183 individuals per m². However, these Canadian figures were for densities at the beginning of thegrowing season, and losses of plants occur throughout the year, and obviously more noticeable at thehigher density at the start of the season (Westbury 2004).
Populations of R. minor are strongly regulated by sward composition and structure, the number ofindividuals being relatively low in more productive swards, indicating the limited competitive abilityof Yellow-rattle when faced with vigorous growth in its potential host plants (Westbury 2004). Theestablished strategy of R. minor was categorised as R/SR, meaning intermediate between a straightRuderal and a Stress-tolerant Ruderal (Grime et al. 1988, 2007).
Toxicity and grazing
Being totally dependent on seed production each year for its local survival, Yellow-rattle in pastureland is also extremely sensitive and vulnerable to heavy grazing pressure. However, as the plant issomewhat poisonous, containing glycosides that presumably render it unpalatable, for this reason animalstend to avoid grazing it. Stock will only browse on R. minor if they are really very hungry orare actually starving (Cooper & Johnson 1998).
On account of its semi-parasitic nature, Grigson (1955, 1987) referred to Yellow-rattle as, "anotherof the pretty infirmities of the land". He also quotes Gerard's Herbal, "What temperature orvirtue this herbe is of, men have not as yet been carefull to knowe, seeing it is accountedunprofitable." (Gerard 1597).
Fermanagh occurrence
Yellow-rattle is a common and widespread species in Fermanagh, having been recorded in 150 tetrads, 28.4%of those in the VC. It is found chiefly in and around damp or even flushed patches of ground inagricultural grasslands, but also on drier parts of fen-girdled lakeshores, by rivers and on cliffledges and in quarries. It is more prevalent in the SW half of the county.
Variation
The genus Rhinanthus is very variable and contains about 50 species in the north temperate regionsof the world (Sell & Murrell 2007). R. minor is a very variable species and a number ofdifferent schemes of subdivision have been proposed. While Praeger was referring plants of wetterhabitats to R. stenophyllus in 1936 (see the account below), now sometimes considered one of sixsubspecies in the flora of B & I (New Flora of the BI 1991, 1997), others feel we do not yethave sufficient information to put names on Irish variant forms (An Irish Flora 1996; Parnell& Curtis 2012).
Six intergrading and inter-fertile subspecies, based mostly on geographic and ecological evidence (F.J.Rumsey, in: Preston et al. 2002; Stace 2019), are now recognised in the flora of the BI, howeverthe variation is complex and continental plants are even more complex. Some forms in Britain simply donot fit comfortably into any of the proposed subspecies and Stace (2019) has suggested that it mightindeed be better to abandon the subspecies altogether.
Variation in R. minor is described by Sell & Murrell (2007) as, "a combination ofecotypes, geographical races and seasonal variants".
British and Irish occurrence
As the New Atlas hectad map indicates, R. minor is very widespread throughout the whole ofB & I with the exception of intensively cultivated arable land around Cambridgeshire (VC29), wetground in the English Wash, plus in parts of N Essex (VC19). In Ireland, it is least common or absent inparts of Co Wicklow (H20) and further west in N Kerry (H2) and Co Limerick (H8). Populations declined inboth B & I during the 20th century, but have recovered somewhat due to the use of R. minor asa component of colourful 'Wild-flower seed mixtures' widely and commonly sown in gardens and councilamenity areas (F.J. Rumsey, in: Preston et al. 2002).
European and world occurrence
R. minor is a mainly European species, being widespread throughout most of the continent fromIceland southwards, although becoming rare around the Mediterranean basin. Having said this, it doespenetrate the whole length of Italy from N to S. In N Sweden, it is regarded as an alien introduction(Seel & Press 1993). It is widely naturalised beyond its native range and consequently is probablynot native in Asia and N America, having been transported into the latter in imported hay (Fernald 1907;Hultén & Fries 1986, Map 1702).
Names
The genus name 'Rhinanthus' is a combination of the Greek 'rhinos', 'a nose' and 'anthos' 'aflower', ie 'nose flower', referring to the trunk-like upper corolla lip of the Mediterranean speciesR. elephas L., which is now placed in a completely different genus, Rhynchocorys(Gilbert-Carter 1964). The Latin specific epithet 'minor' means 'lesser' or 'smaller' (Gilbert-Carter1964).
Grigson (1955, 1987) lists a total of 48 English common names for Rhinanthus minor of which atleast twelve include 'rattle' as a word element, all clearly referring to the characteristic noise madewhen the ripe capsules are shaken. Numerous other names such as 'Money in the purse', or 'in the box' or'the basket' also allude to the same property of the fruit. Vickery (2019) notches up a list of 50English common names, indicating that 'Shackle- 'bags', or 'basket' or 'box' or 'caps' also refers to'rattle' in various parts of the country.
Threats
Changes in agricultural practice associated with intensification have reduced the incidence of thespecies, but it persists in more marginal sites and in areas where intensive agriculture is uneconomicor impossible.
Native, very rare, or rather very under-recorded. European boreo-temperate.
1934; Praeger, R.Ll.; Co Fermanagh.
June.
As mentioned under Rhinanthus minor on this website, this species is very variable and for over acentury or so several attempts have been made to subdivide it. Six subspecies are described and keyed inthe New Flora of the BI (Stace 1991, 1997 & 2019), two widespread and four others that aremuch more local and montane in character. The two widespread subspecies are subsp. minor andsubsp. stenophyllus and, of these, the latter is a plant of wetter soils in fens and dampgrasslands and is more common in the north of Britain, replacing subsp. minor which predominatesin southern regions of Britain (F.J. Rumsey, in: Preston et al. 2002). In Ireland, the NewAtlas hectad map shows subsp. stenophyllus widely but thinly scattered, mainly in NIreland and in the far SW tip of the island.
The variation in R. minor is such that even seasonal forms occur. These differ in their floweringperiods (ie some of the taxa are aestival (spring flowering) and others, like subsp.stenophyllus, are late-flowering or autumnal) –a situation that also occurs in the related generaEuphrasia and Odontites. These seasonal forms also differ considerably from each other intheir morphology. The differences include branching patterns, position (ie node number) of the lowermostflower and the number of leaves between the topmost branch and the lowermost bract. Taken individually,none of these characters allows fully confident identification of the group or subspecies and someplants may remain impossible to assign even when using the characters in combination (Parnell &Curtis 2012). Variation in R. minor is described by Sell & Murrell (2007) as, "acombination of ecotypes, geographical races and seasonal variants".
The situation in Ireland, with respect to this variation, is not yet properly surveyed and studied, soalthough the famous Edwardian botanist Robert Lloyd Praeger was keen to distinguish R.stenophyllus, and in the Census List of the Irish flora incorporated into his book TheBotanist in Ireland, he suggested that it might be, "?general" (Praeger 1934i).
Tall plants with 1-2 leaves between the topmost branch and lowermost bract have been labelled subsp.stenophyllus, but further examination of Irish plants in a European context is required. It isprobably wisest for Irish botanists to follow An Irish Flora (1996; Parnell & Curtis 2012)and await a detailed study of the variation on this island before assigning names to our particularrange of variants. For the moment, there are just two records of this taxon in the Fermanagh FloraDatabase, the site unspecific record listed above, plus a second Praeger listing from Belcoo, LoughMacnean dated 1934-8 (Praeger 1939).
Stace (2019) commented that some British populations of R. minor do not fit into any of the sixproposed subspecies and the pattern of variation on the Continent is even more complex than in Britain.He concluded that it might be better to just abandon the subspecies altogether.
Native, frequent. European boreo-temperate, very probably an introduction in eastern N America.
1881; Stewart, S.A.; Co Fermanagh.
March to December.
Growth form and preferred habitats
This species is a stiff, erect leafy annual or biennial hemi-parasite of somewhat pyramidal form thatattaches itself, xylem to xylem, to the roots of grasses, sedges and other herbs and extracts water andmineral solutes from its host. As is the case with the related Rhinanthus minor (Yellow-rattle),there is no evidence of the parasite benefitting from acquisition of phloem-borne photosynthates fromits host (Westbury 2004). P. palustris has stems that grow up to 60 or 70 cm, but usually lessthan this; it branches freely and produces deeply cut, bipinnate leaves 30-50 mm long, all of whichcharacters distinguish it from the closely related P. sylvatica (Lousewort), which is a shorterperennial with fewer, prostrate branches and leaves that are only 15-20 mm long (Webb et al.1996). The stem leaves can be partly opposite and partly alternate. P. palustris also has a calyxthat is pubescent and the upper lip of the dull red corolla has four teeth. In comparison, the calyx ofP. sylvatica is glabrous and the upper lip of the corolla has just two teeth (Garrard &Streeter 1983).
These two red-flowered Lousewort species are partial- or hemi-parasites, possessing chlorophyll anddisplaying both autotrophic and heterotrophic nutrition. The current author's (RSF's) account ofRhinanthus minor on this website provides more details of the benefits and consequences ofpartial parasitism on species and vegetation. The poverty of the pasture caused by the partialparasitism of this and other related species has led to the charge against these plants of producinglice in sheep grazing them and hence the English common name 'Lousewort' (Melderis & Bangerter1955).
Marsh Lousewort is also known as Red Rattle since the leaves and stems of the plant are very often abright red or dark purple colour, indicating a phosphate or a nitrogen mineral deficiency. It is a goodindicator of situations where there is mobile ground water (ie lateral and/or vertical water movement)and a moderate to poor supply of mineral nutrients, making the name 'Marsh Lousewort very appropriate(Sinker et al. 1985). P. palustris tolerates a wide range of wet soil conditions, fromacid to neutral, or less commonly calcareous base-rich, over clay or peat, but it most usually frequentsrather acid soils and can do so at altitudes up to 860 m. It is a plant of valley bogs, wet heaths, wetmeadows, fens, stream sides and hillside flushes, in fact almost anywhere acidic with a variable watertable (Garrard & Streeter 1983; Sinker et al. 1985). The thing it most definitely does nottolerate, is shade.
Flowering reproduction
P. palustris produces large, tubular, bisexual flowers 25-30 mm long, inshort, terminal leafy racemes from May to September. The flowers are shortly stalked (nearly sessile) inthe axils of the upper leaves, but they eventually form, especially when in fruit, a ± dense, if rathershort, truncated, spike-like appearance. The calyx lobes are much shorter than the inflated andreticulate tube. The calyx is large with two broad, irregularly toothed lobes that are hairy outside.The 2-lipped, laterally flattened corolla is tubular and longer than the calyx, deep purple-red orreddish-pink in colour, the upper lip, narrowly hood-like and with two very small teeth on each side ofthe hood. Following Proctor & Yeo (1973) description, but not word for word, "The corolla hoodencloses the four stamens with the stigma just protruding between them from the underside near the tip.The lower lip of the corolla has two large side lobes and a smaller middle lobe, forming a flat, butobliquely angled landing platform for insect visitors. The two pairs of stamens face one another in theconfines of the corolla hood and the pressure of the sides of the hood keeps them pressed together,preventing the released pollen from escaping." (Proctor & Yeo 1973; Garrard & Streeter1983).
Pollination mechanism
Based on Proctor & Yeo's description of the mechanism, the "visiting bee or bumble bee graspsthe base of the obliquely slanting lower lip of the corolla with its forelegs and the corolla tube justbelow the lip with its middle legs. Holding onto the flower, it then inserts its head into the widestpart of the entrance to the hood, touching the club-like stigma that protrudes obliquely downwards frombetween the anthers; in the process, pollen is transferred from a previous flower visit."
As the insect probes for the deeply hidden nectar secreted at the fleshy base of the ovary within thetubular corolla and calyx, "it prises apart the sides of the corolla hood, at the same time drawingforward the upper part of the hood and releasing the pressure on the stamens, allowing released pollento fall from between them and rain down onto its head". (Hutchinson 1972; Proctor & Yeo 1973).
Fruit and seed
The fruit capsule is 15-17 mm, ovoid, compressed and curved, longer than the calyx. The seeds are few,large, 2.5 × 1.6 mm, oblong, coarsely reticulate and brown in colour (Butcher 1961). They lie in thebottom part of the capsule, which when dry rattles with them when it is shaken (Sell & Murrell2007). Salisbury (1942) examined only a few plants but found the number of capsules per plant rangedfrom 27 to 400, with a mean of 165 capsules. The contents of 30 capsules ranged from 6 to 20 seeds witha mean of 12. He calculated from this that the mean seed output would be around 1,980 per plant.
As might be expected, seed dispersal involves wind and water: wind to shake the rather tall stems,sufficiently strong, not just to rattle the capsules, but to eject the large seed, which weigh anaverage of 1.48 gm (Salisbury 1942). Since P. palustris plants either stand in water in marshyground, or are growing near water, a proportion of the seeds may often land in water, where they canfloat for from one to six months (H.B. Guppy, quoted in Ridley 1930). [No date given for Guppy, butRidley lists a total of six references by this author. The six references are rather briefly, andprobably inadequately, listed for our purposes.] The seeds possess an air-cell structure in the seedcoat that enables floatation (F.K. Ravn 1894, quoted in Ridley 1930).
Seeds germinate in the spring and early summer (Fitter 1987).
o the European
Godwin (1975) lists the fossil finds of P. palustris seeds recorded from two English sites datingfrom the Middle Weichselian ice age, and additional records, "from zones I & III of the LateWeichselian in Kent and the Isle of Man and from zone IV of the Flandrian [the current warm interglacialperiod] in Northumberland. Finally, there is a Roman record from Newstead." He goes on to comment,"The fossil record strongly indicates periglacial survival in Britain", ie the species waspresent during the glacial stage, living near the ice ('peri' means 'around') (Godwin 1975).
Fermanagh occurrence
In Fermanagh, P. palustris has been recorded in 98 tetrads (18.6% of the VC total) and 86 of themhave post-1975 records. Typical habitats are marshy ground ranging from wet meadows to fens, to flusheson bogs and stream-sides. As the tetrad distribution map shows, Marsh Lousewort is widespread throughoutFermanagh, on lakeshores, especially around both sections of Lough Erne, on the Western Plateau and inthe extreme SE of the county.
P. palustris occurs frequently in the Lough Erne water meadows where it has to be able to tolerateboth light grazing pressure and trampling by cattle. It grows vigorously forming distinct bushy plantsin short, more or less flooded grazed grassland, but it also manages to persist and grow tall andspindly (up to 100 cm), even amid relatively tall tussock grass and sedge vegetation in wet ground.
British and Irish occurrence
Although P. palustris remains widely distributed throughout B & I, it has become very rare inC & SE England, but common in the N & W. There have been major population losses on both islandsstretching over since the 1960s and 1970s, largely due to drainage for agriculture and for urban oragricultural development (F.J. Rumsey, in: Preston et al. 2002). Some losses are probably alsothe consequence of neglected, wet lowland alluvial areas becoming overgrown by scrub (Brewis etal. 1996). In parts of S England, eg along the Thames valley, P. palustris has become ararity of ancient wet meadows on nutrient-rich soils and is now on the verge of local extinction, or hasactually become extinct (Crawley 2005).
European and world occurrence
Belongs to the European boreo-temperate phytogeographical element and is mainly restricted to Europe. Itextends throughout temperate middle latitude Europe southwards to the Pyrenees, N Italy, S Bulgaria, SUrals and the Caucasus. The plant in B & I is subsp. palustris, and a northern subspecies,subsp. borealis (Zett.) Hyl. replaces it in N Europe. A related species extends the distributioneastwards into C Asia and Siberia (P. karoi Freyn). P. palustris has been recorded ineastern N America, but it is very probably an alien introduction there (Hultén & Fries 1986, Map1694; Sell & Murrell 2007).
Uses
Previously, Pedicularis sylvatica (Lousewort), P. palustris and Rhinanthus minor(Yellow-rattle), being closely allied to the Euphrasia spp., Eyebrights, were used in herbalmedicine, but as remedies they have now fallen into disuse. In the late 18th century, the Rev. Dr JohnWalker visited the island of Jura in the Hebrides and discovered a large number of inhabitants werecrippled for life by a disease allegedly caused by a reddish worm, about an inch (2.5 cm) in length,which lodged under the skin of the knees or the ankles, causing intense pain. The affliction was knownby the Gaelic name of 'Fillan', and cases were recorded from elsewhere in the Hebrides and WesternHighlands by other early authors. The only known local cure was to take the root of what Walker (anexperienced botanist and scientist) later identified was P. palustris, pound it and mix it withthe marrow of beef bones or goat tallow. This was applied on a hot poultice to the part of the bodyaffected (Allen & Hatfield 2004). The root was used externally against lice and internally to treatinfestations of worms (Walker 1764, 1771).
It is now known that P. palustris contains a glycoside called 'aucubin' which is an effectiveinsecticide (Darwin 1996), supporting the above story.
Names
The genus name, 'Pedicularis' is derived from the Latin 'pediculus' , 'a louse' (Johnson & Smith1946) and the species get their 'Lousewort' common name from the old belief (or myth) that, "itfilleth sheep and other cattle, that feed in meadows where this groweth, full of lice" (Gerard1597). 'Red Rattle' refers to the ripe seed rattling inside the base of the capsule and, also, thecolour of the flowers (Prior 1879). The Latin specific epithet 'palustris' means 'of swampy places'.
In addition to 'Marsh Lousewort' and 'Red Rattle' Grigson (1966, 1987) lists ten more English commonnames several of which are shared with Rhinanthus minor, eg 'co*ck's Comb' (from a supersededspecific epithet of the latter, 'Crista galli'). Other interesting names for P. palustris include'Cow's wort' and 'Deadmen's bellows', to the latter Grigson adds "(ie pillies, male members)",a reference the current author (RSF) does not comprehend. Three names make mention of nectar as'Honey-cap', 'Honeysuckle' and 'Wild Honeysuckle'. There is also 'Suckies', from sucking the honey. Apair of the names listed refer to the wet ground habitat, 'Moss-crop' (ie 'moor') and 'Moss-flower'.There is, of course, the notion of seeds rattling in the dry capsule, 'Rattleweed', 'Rattle-baskets' and'Rattle-pods' as well as 'Red Rattle' (Grigson 1955, 1987; Vickery 2019).
Grigson (1955, 1987) explains that in Germany, P. palustris and P. sylvatica were bothreferred to as 'Läusekraut', our 'Lousewort' being a direct translation, "bycause the cattell thatpasture where plenty of this grasse groweth become full of lice" (Lyte 1578). The plants weresupposed to give the cows the fluke-worms of liver-rot. Other plants that were reputed to give theanimals the rot included Drosera rotundifolia (Round-leaved Sundew), Pinguicula vulgaris(Common Butterwort) and Hydrocotyle vulgaris (Marsh Pennywort), all plants of similar constantlydamp or wet ground habitats. "The herbe is not onely unprofitable, but also hurtfull, and aninfirmitie of the medowes.", according to Gerard (1597, 1633).
Threats
None in Fermanagh, at any rate, but subject to habitat loss through drainage and management neglect ofwet ground in many areas elsewhere.
Native, common. European temperate, adventive in N Europe and very probably introduced in eastern NAmerica.
1881; Stewart, S.A.; Co Fermanagh.
January to November.
Growth form and preferred habitats
This is a low-growing (almost dwarf), monocarpic perennial with a thick, fleshy taproot and stemsreaching a height of only 25 cm or less. It is a very variable plant and dwarf forms, only 6-7 mm tall,have been reported (Blamey & Grey-Wilson (1989). As the technical term indicates, monocarpicperennials fruit only once and they may take two or more years to develop sufficient energy reserves toallow them to trigger flowering behaviour (Grime et al. 1988, 2007). Lousewort produces several ±prostrate, decumbent stems 7-20 cm long and the erect or ascending flowering stem bears mostly alternateleaves 15-20 mm in length, fern-like, deeply divided pinnately and with small, toothed or pinnatelylobed segments (Parnell & Curtis 2012). Like other members of the family, it is a hemi-parasite onthe roots of its neighbours, principally grasses, but doubtless occasionally on the commonnitrogen-fixing legume Lotus corniculatus (Common Bird's-foot-trefoil), with which it frequentlyassociates.
Flowers of P. sylvatica can be easily separated from those of the more scarce or rare P.palustris (Marsh Lousewort) by having only one tooth on each margin of the upper lip of thecorolla near the tip of the hood, as compared to the two teeth of the latter.
P. palustris is an abundant and virtually constant species of heathland, moorland and the betterdrained parts of acidic soils and peat (including that developed directly over limestone rocks). Assuch, it very regularly frequents blanket bog and heathy upland pastures, marshy lakeshores and thedrier parts of raised or valley bogs. It is found at all altitudes, although predominantly in theuplands where it reaches 915 m. Essentially a species of fairly wet acid soils, including flushes, itcan, however, tolerate some degree of summer drought (Sinker et al. 1985). In Fermanagh, with aclose proximity to the Atlantic Ocean from where most weather arrives, Lousewort is rarely much testedwith respect to drought conditions!
The established strategy of P. sylvatica is categorised as SR, meaning it is a Stress-tolerantRuderal species (Grime et al. 1988, 2007).
Flowering reproduction
P. sylvatica flowers from April to July or August and is mainly pollinated by bumble-bees (Garrard& Streeter 1983). The pollination mechanism is the same as that for P. palustris described indetail in the current author's (RSF's) species account on this website. The flowers, 15-30 mm long, arebright pink or red (rarely white) and 3-10 of them are borne in a short, loose (or lax), terminalspike-like cluster. The calyx is bright green, tubular, angular, inflated when the flower is in fruit,hairless, dividing above into five fimbriate, leafy lobes (ie it is not 2-lipped) (Butcher 1961; Claphamet al. 1987). The corolla tube exceeds the calyx in length and it is 2-lipped: the upper hoodedlip and lower lip are of about equal length. As with the taller and more floriferous P.palustris, the flowers produce copious nectar for their size and children, at least in pasttimes, made a habit of sucking the flowers to drink their honey, giving rise to numerous related Englishcommon names for the two 'Louseworts' or 'Red Rattles'.
The fruit is a two-valved capsule, 12-15 mm, ovoid, shortly acuminate at apex, shorter than or equal tothe calyx tube (Sell & Murrell 2007). The capsule contains a few angular seeds, each 3.0 × 1.6 mm,ovoid, netted, finely striate, brown (Butcher 1961; Melderis & Bangerter 1955).
The seeds are said to possess an elaiosome food body that is designed to attract ants which should helpin seed dispersal. The oil-containing food body is unusual in this instance because in P.palustris it is derived from a protruding endosperm-haustorium (Berg 1954). The haustorium isthe organ with which the roots of the hemi-parasite species attach to the roots of their host plant(s),and this is a very unusual modification in this species. The seeds are also reported to be usuallypartially winged (Stace 2019) and, unlike those of P. palustris, seeds of P. sylvatica aretransient in the soil seed bank, persisting for less than one year (Thompson et al. 1997). Thecurrent author (RSF) has not been able to gather information on the level of seed production for thisspecies from the literature available to him, nor any study of its seed dispersal.
The seeds rattle inside the capsule when the fruit is ripe, giving rise to one of the English commonnames.
Fermanagh occurrence
P. sylvatica has been recorded in 196 Fermanagh tetrads, 37.1% of those in the VC. It is commonand widespread, especially in the more upland western half of the county.
Variation
Two subspecies are recognised in B & I, subsp. sylvatica with a hairless calyx, which is themost common form in Fermanagh, and an endemic Irish variant with a hairy calyx referred to as subsp.hibernica, which so far has only been recorded at four sites in Fermanagh and is doubtlessregularly overlooked (Revised Typescript Flora; An Irish Flora 1996). A third subspecies,subsp. lusitanica (Hoffmanns. & Link) Coutinho, is described in Flora Europaea3 and is endemic to Portugal and SW Spain (E. Mayer, in: Tutin et al. 1972).
British and Irish occurrence
Frequent throughout the whole of B & I in damp heathy places, except for parts of the EnglishMidlands where it is scarce and local. There has been a widespread decline in S & E England due tothe loss of heathland and unimproved grassland habitats. Analysis of BSBI databases indicates that mostof these losses have taken place since 1950 (F.J. Rumsey, in: Preston et al. 2002).
European and world occurrence
P. sylvatica belongs to the European temperate phytogeographical element and is widespreadin W & C Europe, having its greatest frequency in W Europe. It is, however, completely absent from SEurope and the entire Mediterranean region. It extends northwards to C Sweden and eastwards to Lithuaniaand W Russia (Clapham et al. 1987). Subsp. sylvatica is distributed through the range ofthe species, except Portugal. Subsp. hibernica occurs across NW Europe, from Ireland to Norway.P. sylvatica has been recorded in SE Newfoundland, but it is not considered native in N America(Hultén & Fries 1986, Map 1697).
Uses
Previously P. sylvatica and Rhinanthus minor (Yellow-rattle), being closely allied to theEuphrasia spp., Eyebrights, were used in herbal medicine, but as remedies they have now falleninto disuse. Old herbalists such as Culpepper considered P. sylvatica, which they referred to as'Dwarf Red Rattle', a wound herb and styptic (Grieve 1931). Culpepper (1653) says of 'Red Rattle',"accounted profitable to heal up fistulas and hollow ulcers, and to stay the flux of humours inthem, as also the abundance of women's courses, or any other fluxes of blood, being boiled in red wineand drunk".
Names
The genus name, 'Pedicularis' is derived from the Latin 'pediculus', 'a louse' (Johnson & Smith 1946)and the species get their 'Lousewort' common name from the old belief (or myth) that, "it fillethsheep and other cattle, that feed in meadows where this groweth, full of lice" (Lyte 1578; Gerard1597, 1633). Two of the other English common names are 'Dwarf-' or 'Lesser-Red Rattle' which refer tothe ripe seed rattling inside the base of the dry capsule and, also, the colour of the flowers (Prior1879; Grieve 1931). The Latin specific epithet 'sylvatica' is from 'silva' meaning 'woodland' andtranslates as 'growing in woods', which is completely incorrect, making it a misnomer (Gilbert-Carter1964).
A total of eight additional English common names are listed by Grigson (1955, 1987), several of whichrefer to the practice of children sucking the nectar from the flowers, eg 'Bee-sookies', 'Honeysookies', 'Honeysuckle' and 'Sookies'. The other names refer to the rattle of seeds in the ripe capsule,eg 'Rattle baskets', 'Rattle pods' and 'Shackle-boxes'.
Threats
Common and widespread and not under threat.
Native, rare, but certainly under-recorded. European temperate.
Pre-1957; MCM & D; Lough Achork.
The post-1975 Fermanagh flora survey did not examine P. sylvatica for this mainly Irish subspeciesand there are just four records in the Flora database made by Meikle and co-workers in the pre-1957period. The reason for this omission is that for most of the period of the survey the field Flora usedhas been An Irish Flora (1977), which makes no reference to these two subspecies, despite thefact that it was Webb himself who described and named them (Webb 1956).
Recognition of the variant forms is a simple matter and should be addressed in future fieldwork. Subsp.hibernica differs from subsp. sylvatica in having the outside of the calyx and flowerstalks clothed with long, white, curled hairs (subsp. sylvatica is hairless in these areas,though hairy around the throat, and rarely sometimes also on the angles of the calyx) (BSBI PlantCrib 1998). The hairiness of subsp. hibernica is so pronounced that the hairs catch mistand the plants then appear greyer in colour. The two subspecies can occur together in mixed populationsin damp upland grassland, heath and bog. In some areas, where this overlap in distribution occurs (eg ineastern Ireland), a few intermediate plants may occur, with short or sparse hairs on the calyx (AnIrish Flora 1996).
The sites of the other three MCM & D records, again undated but pre-1957, are as follows: summitarea, Tappaghan Mountain; Lough Corry, Kimran Td; and Derrynacarbit Lough.
Despite its name, this subsp. is not confined to Ireland, having been recorded in W Scotland, SWales and N England (BSBI Plant Crib 1998).
