Hymenophyllum Tunbrigense (L.) Sm. Author(s): P. W. Richards and G. B. Evans Source: Journal of Ecology, Vol. 60, No. 1 (Mar.

, 1972), pp. 245-268 Published by: British Ecological Society Stable URL: http://www.jstor.org/stable/2258054 Accessed: 06/07/2010 16:43
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245

BIOLOGICAL FLORA OF THE BRITISH ISLES

Hymenophyllum BY P. W. RICHARDS AND THE LATE G. B. EVANS Bangor Wales, University College ofNorth Biology, School ofPlant ListBr.Vasc.P1.(1958)No. 7.1 tunbrigense (L.) Sm. Hymenophyllum vertical or inclined flattened A mat-forming against growing perennial, bryophyte-like deeply pinnatisect pendulous, or tree numerous usually overlapping, rocks trunks, with rhizomes interlaced tufts bearing from much-branched leavesarising slender, wiry, very often roots at thenodes. when becoming Leavesbright young, palegreen ofadventitious 30-6-0 withmat surface, whenolder,translucent, bluishor brown(not blackish) at theveins), more orlessoblong onecellthick (except x 2 cm.Lamina 0-5-15-0) (rarely moreor less inoutline, thepinnae divided with winged rhachis, to ovate-oblong pinnate acroscopic segments, butirregularly, with5-11 oblong predominantly dichotomously in toothed. Cellsoflamina a single vein almost the finely apex:margin eachwith reaching thelateral lining c. 30-40rather chloroplasts large surface view60?1 x 39?1 ,um, with inexposed ofthe habitats the tending Petiole lamina, length walls. wiry, very slender, j-3at right to it. to become bent angles on the segment Sori usually5-10 per leaf and one per pinna,borneterminally toblack when purse-like and ripe, therhachis; indusia when brown green young, nearest to broadly triangular in theplane of thelamina, formed by two flattened flattened oftheir andirregularly arefree for length valves which aboutthree-quarters suborbicular indevelopment. ontheir distal annulus, gradate toothed with oblique margins. Sporangia 40-48gmin diameter. Spores in Britain, seems to be entirely as faras investigated, (Evans phenotypic Variation from appearing Europe. Populations orvarieties have been described No subspecies 1964). inthe North tothose ofEurope arefound Atlantic more orlesssimilar Islands, America, theexception ofthose ofthetropics with various andintheSouthern Hemisphere; parts and Jamaica, these of eastern North America are oftheAtlantic andperhaps Islands, critical and require re-examinawith H. tunbrigense notconspecific European probably tion. on trees and on peaty soilamong andlocally abundant on rocks, more Native rarely in moist, in thewestand north of GreatBritain, well-sheltered situations bryophytes, thanH. wilsonji and lesswidespread and abundant theSussex Wealdand in Ireland; a narrower with range. ecological from vice-counties in I. Geographical andaltitudinal thirty-seven Reported distribution. inIreland. inGreat Britain extends Britain andtwenty-five Thepresent distribution Great of an outlying group and Devonin thesouth to Skye in thenorth, with from Cornwall on thesandstone rocksof East Sussex:extinct sinceat least1875in its locus stations

246

Hymenophyllum tunbrigense (L.) Sm.

Wells in south, at theHighRocks, classicus west Tunbridge (Wolley-Dod 1937).Occurs Areasofhighest and north Ireland. are North frequency Scotland and Wales,western west andsouth-west Ireland. the this hassuffered from During pasthundred years species thedestruction of its habitats and from collectors: extinction unscrupulous in some hasincreased thediscontinuity localities ofitsdistribution. Thelarge oflocalities number in which it has notbeenseensince1930(Fig. 1) suggests thatit is still in diminishing much ofitsarea.Ithasoften been with H. wilsonii frequency over confused andsometimes
B
7/1

3
9

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2

TUN BRIGENSE (L) Sm. Fern Flora Filmy * 1930 onwards o refore 1930

PHYLLUM

HYMENO-

by p

~~~~~~70

94--* * 0

00`706 0

0~~~~~~~~~~~~ 0~~~~~~~~~~~~~~

0O~~~~~~~~~~~~~~~~~~~~~
9

00

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iforaiG.n1 Thce disrbuto ofeHmenohylum specbme gssr

(wig.)S.ithBrisIse.Ec

Atlstrofuthen BinotisheFloal byropermissino thegl BotanicalSociety of, thwBitih wislesyseandte

groups ofstations ('nidsatlantiques' ofChrist 1910);as in Britain itis decreasing andis nowextinct inmany former stations. Atpresent H. tunbrigense is certainly known only from France, Luxembourg, north Spain,Italyand theeastern coastof theBlackSea. In Luxembourg it is foundin numerous sites in the 'Suisse Luxembourgeoise',

AND G. B. EVANS P. W. RiCHARDS

247

1954).In West in thebasinoftheErnzNoire(Reichling rocks a district of sandstone close to the Luxembourg in Valley of the Sure, at Bollendorf, the Germany it occurred inthenorthlocalities there aremany & Hirzel 1967). In France frontier (Hess,Landolt the western (Basses Pyreenes): and Pyrenees west(Manche, Finistere) Cotes-du-Nord, in thenorth-east (Vosges) and has beenreported a few localities it is also known from In Spainitis recorded from Galiciaand Orneand Eure-et-Loire. Seine-et-Marne, from inseveral In Italy itoccurs intheSpanish 1885). onelocality Pyrenees (Lacoizqueta from 1936);alsoinMarche Toscania (Gams1935;Pichi-Sermolli localities inthe AlpeApuani,
6 30 24 70' 18 1218iE

12-

30

36

42

48

54

<

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72

60*

W~~~~~~~~~~~~~~~

X.

tunbrigense (L.) Sm. in Europe. o, Native FIG. 2. The distribution of Hymenophyllum status'; +, extinct; x, probablyextinct. occurrence; @, occurrence of 'uncertain on the easterncoast of theBlackSea are outside theFlora Europaea The occurrences area and are notshown. forMapping theFlora ofEurope. bypermission oftheCommittee Reproduced

on theeastern coastoftheBlackSea at (Pichi-Sermolli 1936). In theU.S.S.R. itis found 1962). three stations nearBatumi, Adzharya (Burchak-Abramovick 1910;Wilpert 1937) andin Germany intheBelgian Ardennes (Christ Formerly found sites at all recorded nearDresden, where it is now extinct in theSaichsische Schweiz Corsica(Jovet 1937).Formerly occurred in theislandof Elba, and probably (Wilpert years (Dostal1958). for many 1933).Reported from Czechoslovakia, butnotconfirmed Zagreb in 1900andhasbeendoubtfully near In Yugoslavia itis saidtohavebeenfound (Pevalek). recorded from three other localities

248

Hymenophyllum tunbrigense (L.) Sm.

H. tunbrigense is reliably recorded Outside from theAzores, Europe, Madeira andthe it havebeenfound Canaries. Populations with probably conspecific in SouthCarolina (twenty-one stations-Taylor & Bannister 1938;Anderson 1952)andin Jamaica (G. R. Proctor, personal communication). is from In Britain its altitudinal sea-level range (e.g. nearLlanfrothen, Merioneth; N. Ireland) Rhum, W. Inverness; to 1250ft(380 m) in an isolated Carlingford Lough, on Tryfan, at 1350ft(412 m) at Wistman's station Caernarvonshire, Wood,Devon, in Scully1916).Hart's and 1500ft(460 m) on Priestleap Mountain, Kerry (Carroll, is probably (1881)record from 2500ft(762m) in Tipperary due to confusion with H. inthelocality H. which still doesnot(D. A. Ratcliffe, wilsonii grows though tunbrigense a plantoflow altitudes and usually personal communication). Mainly replaced byH. above1000ft(305m).MostCaernarvonshire wilsonii localities arebetween 0 and 500ft limit decreases northwards: inthenorth (152m).Theupper ofEngland itis below500ft Foundat 1750m inthePyrenees (Turmel 1952)andto 700m nearBatumi, U.S.S.R. butmostofitspresent or former (Burchak-Abramovick 1962), localities in continental Europearebelow300m. in Britain, no doubt phyllum influenced tunbrigense though byhistorical factors suchas tobemainly offormer seems determined thedestruction of woodlands, bytheoccurrence microclimates and substrata. In themoist, sheltered in which suitable habitats itgrows, ofhumidity andtemperature arewidely different from therange theambient conditions ofBritain themacroclimate to be wellwithin andinmost thelimits appears oftolerance ofH. tunbrigense ofthespecies. Themicroclimatic differ from requirements ofH. those outunder thelatter. as pointed wilsonji ismost often found onoramong H. tunbrigense block rocky outcrops, e.g.cliffs, screes, areweathered so as to provide when crevices orfissures especially and they deepgullies, oroverhanging extent shaded rocks. aretosome often bytrees found Though near streams itis intolerant offlowing water andis absent andwaterfalls, from both thespray anddrip and thefloodzone of streams. The amount zones of waterfalls of shadeor shelter the ofthe seems tovary with macroclimate. demanded Forexample, inthe humidity very it sometimes moistwestof Scotland in NorthWales the growson open hillsides; oronrocks ofitshabitats areincrevices with some shelter from majority butitalso trees, inblock indeephollows screes arenotrees. where there In the grows level high oakwoods ofDartmoor in darkholes in the (Wistman's Wood,BlackTor Copse)itgrows mainly butin Higher PilesCopse,a woodat an intermediate 'clitter' (boulders), and elevation, inquite inthevalley woodsofthesamearea,itgrows on theboulders openpositions of thewoodfloor (Proctor 1962). In the SussexWeald H. tunbrigense colonies are foundon thevertical or slightly in outcrops of Tunbridge Wells(Neocomian; obliquewallsof deepfissures Wealden) tooneofthe the colonies areusually restricted twowalls. These Sussex sandstone; habitats former site near areinmany (andthe like Tunbridge Wells) respects those inLuxemvery thespecies has a similar of distribution where bourg pattern (Assa 1924;Klein 1916, in Luxembourg that andin Saxony the 1926;Heuertz 1933).Assastates colonies donot for faces thanone-third oftheir extend more in Britain up rock colonies height; arenot insomesites areliable tolong thus which restricted, except ofdesiccation. periods There Assa'sandKlein's is somediscrepancy between account oftheLuxembourg sites, butit ofthesites seems that most havea northerly andnone south. faces aspect Thestudy by
The distribution II. Habitat. (a) Climaticand topographical limitations. of Hymeno(152 m), in Scotlandbelow 300 ft(90 m).

P. W. RICHARDS

AND

G. B. EVANS

249

each withseveral localities, Caernarvonshire Evans (1964) of theaspectof twenty-three a NE or NW aspect; the N, that localities had seventeen sites,showed H. tunbrigense modified by in of the of aspect was all these effect faced SE, but six mostly remaining near waterfalls. in were deep gorges e.g. three local features,

and H. usuallygrowson the lowerpart of the treetrunks westIreland,H. tunbrigense of the two species on to the zones often the branches; extending at levels, wilsonii higher 261-2). meetbut scarcely overlap(see underH. wilsonii, pp. in both temperatures Isles mean and maximum the British Throughout Temperature. of thespecies,though thelimits of survival wellwithin and winter are probably summer by itis in somewayrestricted limits and upperaltitudinal itis possiblethatat itsnorthern above some average numberof day-degrees perhaps by an insufficient temperature, value. threshold of more than 170 C or a mean localitieshave a mean Julytemperature No British in continental Europe of morethan220 C. Localitiesin Italy and elsewhere maximum Doignon (1954) foundthatin 1947 summer temperatures. doubtlesshave muchhigher a respectively to Fontainebleaunear Paris survived fromFinistere plantstransplanted in the of40 and 44 successive daysin which30? C was exceeded.In cultivation maximum of c. 17?C, provided appearsto be normalat an averagetemperature growth laboratory, producedunderthesecondiis maintained, thoughsori are rarely thata highhumidity tions. above 3 90 C and the at all British localities is probably The meanJanuary temperature toleranceof the above about 10C. The lower limit of temperature mean minimum speciesis much lower than this. Johnson(Sowerby& Johnson1859), who cultivated to cold'. Verylow temperatures itwas 'little ifat all susceptible notedthat H. tunbrigense, at Fontainebleau in Saxonyand by Doignon's transplants in thelocalities weretolerated of -20? C in 1954; between1946 and 1954 theysurvived a minimum whichwithstood were of theseyearsthere and longperiodswithout thaw,whilein three 600 days offrost in a and in two of themtheplantswereenveloped weekswithdailyfrosts 5 consecutive Doignon's paperwhether ofice forperiodsof 15 days.It is notclearfrom thick covering In North records. meteorological weremeasuredin situor takenfrom his temperatures (see Vc). Wales snow coverforperiodsof up to a monthhas producedno ill effects of the Sussex localities, Withtheexception andpotential evapo-transpiration. Rainfall of about 30 in. (75 cm) and at least 175 rainydays,all whichhave an averagerainfall have over60 in. (150 cm) localities have 40 in. (100 cm) or moreand themajority British as in thedrier oflittle are however importance, and at least225 rainy days.Thesefigures are lacking. suitableforH. tunbrigense habitatsotherwise areas of Britain but in thecontinental localities, is availableabout therainfall No detailedinformation in all of themit is probably75 cm or more. in the British Isles is insufficiently Information about potentialevapo-transpiration of of H. tunbrigense but an examination withthe distribution detailedfora correlation the between waterdeficit (difference Green's(1964) map of theannualaveragepotential localities(but and rainfall) showsthatmostof theBritish evapo-transpiration potential of theSussexlocalities)lie in theareas withless than3 in. (7 6 again withtheexception cm) waterdeficit. areas subjectto highwinds,the it growsin coastal and mountainous Wind. Although and rockoutcrops are usuallysheltered bytrees, sitesactually occupiedbyH. tunbrigense features. othertopographical

as in southas epiphytes, andH. wilsonii occur H. tunbrigense both where In localities

250

Hymenophyllum tunbrigense (L.) Sm.

SeeH. wilsonii, 11(a). Assa(1924)gives dataonmicroclimatic conditions Microclimate. inLuxembourg. H. tunbrigense as a carpet over otherwise more orless (b) Subgtratum. commonly grows but humid invery situations itoccurs ontheground bare rock surfaces, among sphagnum, andother Leucobryum glaucum bryophytes covering peator uncompacted humus, e.g.in andin several woodsnearMaentwrog, or as an epiphyte west Scotland Merioneth, on with Igothecium associated Caernarvonshire often myosuroides, e.g.nearBetws-y-Coed, in Coed Cymerau and other in great woodsin Merioneth, (Griffith 1895), and,often in Derrycunihy in south-west Ireland and elsewhere abundance, Wood,Killarney (see as an epiphyte as wellas H. wilsonii, under Ila). NearBatumi, U.S.S.R.,it also grows ontrunks ofAlnus andPrunus andisfound barbata laurocerasus onrocks upto2 mabove theground (Burchak-Abramovick 1962). localities on sandstone, In itsBritish H. tunbrigense grows granite, quartzite andmany of rockof varied of the and chemical characteristics. The nature other types physical it is fairly rockseemsto be of no great importance, provided hardand its texture, moist for surface andposition allowittoremain Certain ofsandstone long periods. types seemto be particularly becauseof their favourable, probably porosity and watera store ofwater which in drought forthis holding properties provide periods. Perhaps rocksare theusual substratum reasonsandstone in areassuchas theSussexWeald, andSaxony which tobe only ofclimatic Luxembourg appear justwithin the plant's limits tolerance. mostoften on acidicrocks and Taylor H. tunbrigense grows (1960)states explicitly on limestone: thatit is notfound thisseems to be true, is no evidence though there this is because theplantis a calcifuge or becausethephysical as towhether properties of limestones are unsuitable. Near Derryclare Lodge,Connemara (westIreland)it on basicigneous or metamorphic grows rock(see III). or peat consisting of partly colonies on rocks, rawhumus Underold, established leaves andrhizomes ofup to 3 cm: thewhole to a depth mass decayed mayaccumulate turf actsas a ranker andbecomes when easily detachable, especially dry (seeVc). III. Communities. In theBritish IslesH. tunbrigense grows on rockfaces and trees in of deciduous various woodland types (mostly Quercetum petraeae, but occasionally or Fraxinetum. roboris Since itsimmediate associates Quercetum arechiefly bryophytes ferns andthedominant tree andother toaffect itmainly itsinfluence on appears through itcanperhaps themicroclimate, best be regarded to oneormore 'associules' as belonging or synusiae of bryophytes consisting mainly which form partof various woodland in certain associations In their account of thevegetation of topographical situations. Ireland, Braun-Blanquet & Tiuxen (1950)include bothH. tunbrigense and H. wilsonii as differential speciesof the Isothecium myosuroides sub-association of the Blechnoand oftheHymenophylletum. Quercetum In thevery humid areasin westIreland and westScotland where Hymenophyllum occurs on hillsides oftheFestuca-Agrostis tree it forms tunbrigense without cover part moorland associations. (andprobably other) Table 1 giveslistsof speciesfora fewselected standsincluding Hymenophyllum in North tunbrigense Walesand elsewhere. Higher plants other thantrees commonly itinclude associated with suchas Anthoxanthum mainly acidophilous species odoratum,
trees and shrubs (Quercus petraea,Ilex aquifolium, Callunavulgaris, Vaccinium myrtillus),

Blechnum Callunavulgaris, spicant, Digitalis purpurea, Dryopteris Deschampsiaflexuosa, pseudo-mas (D. borreri), Erica cinerea, Lonicerapericlymenum and Vaccinium myrtillus.

AND G. B. EVANS P. W. RICHARDS

251

Leucobryum S. quinquefarium), spp. (especially includeSphagnum Associatedbryophytes of hepaticae largenumber and a very tamariscinum Dicranum majus,Thuidium glaucum, and in distribution. Two of themostconstant and othermosses,manyof themAtlantic albicans and Isothecium abundant associates (very seldom absent) are Diplophyllum In Luxembourg Lefort& Lawalree(1951) and Gams (1938,p. 404) include myosuroides. Hymeno'associules'in which Listsforthebryophyte albicantis. itin theDiplophylletum Wood, Killarney,are given by Richards occurs in Derrycunihy phyllum tunbrigense on theassociatedbryophytes and their (1938). Paton (1954, 1956)givessomeinformation rocksof Sussex. on the sandstone succession occurson bouldersof basic whereH. tunbrigense At Derryclare Lodge, Connemara, avellana(withsome Fraxinusexcelsior petraeaand Corylus rock in a wood of Quercus mackaiiand Marchesinia molluscum, and Taxus baccata) it is associatedwithCtenidium as moreor less calcicole. usuallyregarded bryophytes Plagiochilaasplenioides, is rooting myrtillus itself whenVaccinium plantsassociatedwithit. It is able to maintain

with thehigher competition to be in direct rarely appears tunbrigense Hymenophyllum

glaucummore easily than withvarying success: it seemsto coexistwith Leucobryum in wilsonii (Evans 1964) but on thesteepslopes of wet,wooded ravines Hymenophyllum and dead or dying by Sphagnum quinquefarium NorthWales it is sometimes overgrown themossblanket. CompetiofHymenophyllum tunbrigense can be foundbeneath remains species(III, p. 263). tionwithH. wilsonii is dealtwithunderthelatter in theWestPyrenees. (1952) givesomeaccountofitsassociates Jovet (1933) and Turmel

seem tocompete with it itandvarious areovershadowing it.Bryophytes grasses through

lists forLuxembourg, and Gaume(1944)forFinistere. Reichling (1954)gives

on themicroclimate, and has been its effects Treefelling is usually harmful through
in the localitiesin Saxony of Hymenophyllum regarded as the chiefcause of extinction

are due to humanactivities. observed Most oftheeffects tobiotic factors. IV. Response

No doubt other in sites many localities (Rose 1957). 1937)andin someSussex (Wilpert in pastcenturies. havebeenlostfor thesamereason Britain hasbeenobserved when iflight, from rhizomes is notalways fatal:recovery Burning, havebeenkilled all theleaves byfire (Evans).

typical stands consistingof numerous Highly gregarious, V. (a) Gregariousness. a singleindividualor a clone colonies or small patches.Each of theseis presumably in shapeand very irregular fromone spore(but see Vlllc). The patchesare often arising form at thebase ofcliffs, e.g. 3 x 075 m nearLlanberis, strips longhorizontal frequently Caerns.,22 x 1 ft(7 x 0 3 m) in South Carolina (Taylor 1938): theymaybe as muchas Merioneth. Praeger (1934) Caerns.,Coed Crafnant, 150ml in area,e.g. at Betws-y-Coed, in Ireland. records patchesof24 ft2(2-2 m2) in Donegal and 50 ft2(4-6 m2) at Killarney The leavesoccasionally e.g. in NorthWales through grassand bryophytes, growscattered and in theWestern Isles, Scotland. varies with atmospheric humidity in varioushabitats.Performance (b) Performance the mass of foliage(deto desiccation)and light.Under averageconditions (liability scribedby Smith& Sowerby(1841) as like 'half-ruffled plumage')is about 4 cm deep. combining In exposed sitesthis may be reducedto about 1 cm, but in environments it may be 6 cm, some leaves reaching and good illumination constanthigh humidity soriare not moistshaded situations i.e. severeenoughto reduceleafexpansion.In very to desiccationthan the usuallydeveloped.Sporangiaand spores seem more sensitive and frequently leaves themselves show signs of damage, though the slow basipetal

ofmoderate under conditions ofsoriis found exposure, 15 cm.The maximum density

252

tunbrigense Hymenophyllum (L.) Sm. Table 1. Species lists and H. wilsonii tunbrigense for British Hymenophyllum localities
Site tunbrigense Hymenophyllum H. wilsonji Anthoxanthum odoratum Calluna vulgaris flexuosa Deschampsia Galium saxatile 1 la la If If o 0 r r va a o o r? la va If 2
-

3 la sd f f r r o o yr
-

4 30
-

5 ?
-

6 ?
-

7 ?
-

8 ?
-

9
-

Saxifraga spathularis Sedum anglicum Blechnum spicant dilatata Dryopteris selago Lycopodium vulgare agg. Polypodium Dicranum majus lucens Hookeria elegans Isopterygium Isothecium myosuroides undulatum Plagiothecium schreberi Pleurozium fasciculare RhJacomitrium R. lanuginosum loreus Rhytidiadelphus Sphagnum palustre S. plumulosum arbusculum Thamnium (T. alopecurum) Thuidium tamariscinum albicans Diplophyllum Peia epiphylla
D. scoparium

Oxalis acetosella Potentillaerecta

r r o

o If o y r r
-

20

? +

o r

? ? ? ? ?
-

? +

? ? ? + + o va ? la If if? ? ? ? ? ?? ? la + Plagiochila spinulosa + ia r Scapaniagracilis Detailsofsitesand ofspecies in onlyone list. occurring Site 1: Cerrig-y-rhiwdwr, Caernarvonshire. Groundand rocksunder100 m. Tremadoc, in lowerparts of streamvalleyin wood and H. tunbrigense Hymenophyllum in upperparts.(Quercus wilsonii petraeacd, Q. robur cd, Betulapubescens r, avellanaIsd, Fraxinus excelsior Corylus f,Ilex aquifolium o, Sorbusaucuparia a, Hederahelixf,Lonicera periclymenum f, Conopodium glomerata majusr, Dactylis montanum vr, Deschampsia cespitosa0, Digitalispurpurea vr, Epilobium vr, robertianum vr,Hieracium Geranium umbellatum agg.vr, vr, Hypericum pulchrum Luzula sylvatica nemorum la, Lysimachia vr, Melampyrum pratense la, Rubus agg. f, Silenedioica vr, Solidagovirgaurea, r, Umbilicus fruticosus rupestris o, Vaccinium myrtillus r, Athyrium filix-femina f, Dryopteris pseudo-mas va, + Pteridium bruntonii r, Thelypteris o, Cynodontium aquilinum limbosperma + foliosum +, Hedwigiaciliata +, Heterocladium Diphyscium heteropterum armoricum hornum la, Hypnum Hylocomium f,Mnium cupressiforme If,Plagiosucculentum thecium denticulata +, Polytrichum If, Rhabdoweisia formosum ?, recurvum delicatulum trichomanis Sphagnum o, Thuidium +, Calypogeia o, Plagiochila punctata +). Caernarvonshire. Boulder at c.320m. HymenoSite2: Cam Madryn, scree, north-facing wilsonil among boulders.(Andreaea o, Hypnum phyllum rupestris cupressiforme var. ericetorum heterostichum if, Polytrichum piliferum vr, Rhacomitrium f, r). Lophoziaventricosa Site3: Connor Pass, Dingle, Co. Kerry.Grazed grassland on north-facing slope at in scattered on rock and in c.410 m. Hymenophyllum wilsonii patches, mostly more heavilygrazed parts. (Achilleamillefolium tenuiscd, Carex f, Agrostis nerterioides vr, Epilobium caryophyllea If, Erica cinereao, Euphrasia frigida

va o o

y r a? d? r o
-

la o a o la

60 ? ?
-

? ?

? ?

??+ ??+

P. W. RICHARDSAND G. B. EVANS
Table 1 (cont.)

253

Site4: Site5:

Site6: Site7: Site8:

Site9:

montana (det. P. F. Yeo) o, Festucaovinao, F. vivipara cd, Jasione o, Juncus acrisvr, squarrosus la, Luzula campestris f, Polygalaserpyllifolia r, Ranunculus Rumexacetosa r, Sagina ciliatalf, Trifolium repensla, Veronica officinalis vr, commune Violariviniana r,Breutelia chrysocoma o, Polytrichum la). Co. Kerry.Slopingrockface. July1952 (C. D. Derrycunihy Wood, Killarney, Pigott). (Bazzaniatrilobata flava+, Lepidozia pinnata +). +, Lejeunea of Quercus Co. Kerry. Branches Derrycunihy Wood, Killarney, petraeanearto denudatum site 4. July 1952 (C. D. Pigott). (Dicranodontium +, Zygodon +, viridissimus tamarisci +, L. pulmonaria +, Lobaria laetevirens +, Frullania Stictina silvatica +). Co. Kerry.Bouldersbeside the waterfall. Lough Dreenaun,Waterville, May 1950(C. D. Pigott). Co. Kerry, wetter bases of boulders at site6. May LoughDreenaun, Waterville, 1950(C. D. Pigott). Co. Kerry. Boulders above Coomcallee, above LoughCurrane, at the Waterville, rothii summit of theridge.May 1950(C. D. Pigott). (Andreaea +, Hylocomium splendens J, Herbertahutchinsiae +, Pleuroziapurpurea+, Sphaerophorus +). globosus Wood of betweenKidstonesand Beech Hill, Long Sleddale,Westmorland. in cushionsof Diplophyllum wilsonii albicans Quercus petraea; Hymenophyllum withotherassociated species.April 1955 (C. D. Pigott).(Amphidium mougeotil +, Blepharostoma trichophylla +, Coenogonium +, Fissidensosmundoides ebeneum +). Lists4-9 weremadein areasof 1 m2or less.

ofthesporangia ineach ofripening often ensures that at leastsome sequence sporangia reach sorus maturity. exceeds butseveral ofsoriperleafin onegrowth flush Thenumber rarely ten, groups inconstantly arenotunusual moist Thenumber ofsporangia habitats. ofsori perleaf per arerelatively constant sorusand ofspores persporangium (Bower 1926). in viewof theobservations from is unlikely Franceand Germany quotedin11(a).In to subzero for several laboratory experiments plants exposed temperatures daysshowed no damage. with little snow cover extensive During long frosty periods damage may occur, ifthere is bright sunshine from cloudless butthisis probably due to especially skies, in theearly rather thanto freezing. In theexceptional frost in Snowdonia desiccation of 1963which lasted forsomeweeks one-third to halfofthecolonies months perhaps oftheir better leaves. survived thanthose which were lostmost Snow-covered colonies orlesswithout from to altermore where colonies were protection snow, but, subjected andthawing, wasincreased. In dry colonies natefreezing weather damage frosty maybe oflarge therock off from surface. s riously damaged bythepeeling portions of extensively The leavesare borneon a system VI. (a) Morphology ofsporophyte. muchbranched and extremely slender whichbear rather creeping, rhizomes, sparse oftherhizome is protostelic andcontains a roots. Thevascular adventitious system only small number ofwater-conducting elements. very of from oftheleaves an ecological The most features important morphological point ofthe for aretheir indeterminate view capacity apicalgrowth (seeVIc) andthestructure as in mostHymenophyllaceae is onlyone cellthick, lamina, which, (cf.Bower1926), instomata. at theveins, andentirely is present butitis very thin A cuticle except lacking There is no morphological sterile and and noteasyto demonstrate. difference between leaves. fertile
I
J.E.

etc.Damage bylow temperatures has notbeenreported (c) Effect offrost, drought, and

254

Hymenophyllum tunbrigense (L.) Sm.

means ofitsrhizome is probably system. Reproduction byspores (seeVllla) and rarely oftherhizome bythebreaking with attached awayofparts leaves.Clowes(quoted in Moore & Lindley thatin cultivation theleaveswereannualand many 1855)claimed floras haverepeated thisstatement, but our observations showthatin nature and in cultivation formorethan 1 yearand are capable of laboratory theyoften persist in moist continued growth. In many leavesthere areflushes ofgrowth in warm periods summer and winter. Not leaves can found both be with of infrequently spacedclusters these canbe dated backfor indusia and sometimes as much as five seasons. the Though ofunlimited fronds bepotentially the seldom may growth, great majority probably persist for more than 2 years to droughts which killthegrowing thewhole owing apexandoften leaf. is very rateobserved in Snowdonia is 6 cmin Vegetative spread slow;themaximum 3 years, with an average of2 5 cm(10 rhizomes). Onesetofmarked rhizomes waskilled 12months, 2 cmduring after butnotonehadextended more than bydrought this period. in Sussex Colonies marked all completely diedwhen byMrsJ.Patonhad nearly examinedbyEvans10 years ofcolonies later (1963).Longevity or clones is unknown, butis than a decadeandmayevenattain several centuries. undoubtedly longer LochLomond, The chromosomes, 1950)(from to Manton, Scotland). according showa ofsizethan ofH. wilsonii. wider those range data.Thefollowing (e) Physiological observations aresummarized from Evans(1964). In detached Water relations. leavesplacedin a series of controlled humidities (76, 43, 32, 15,10 and 0 % R.H.) at 200 + 10 C a steady in c. 2 h at about wasreached state fresh on H. wilsonii): 20-50% oftheinitial weight (cf.65-75 experiment % in a similar thefinal fellprogressively in which with therelative weight humidity (cf.H. wilsonii thesamefinal wasreached atallhumidities). When approximately theleaves weight were to a highhumidity returned a dryday)water conditions after (simulating night was from theatmosphere absorbed andtheoriginal fresh in 10h (cf.H. wasregained weight 10h andapparently which wasstill wilsonii 20-60 theoriginal after weight in a % below steady state).Mean ratesof loss variedbetween 23-36% fresh weight per hour(cf. 40-57 % in H. wilsonii) andrates ofuptake from airvaried saturated from 2 7-2 9 % (cf. 1 9-2 0% in H. wilsonii). The differences between thetwospecies bothwhen and gaining losing water suggest thatthegreater of H. wilsonii drought resistance notonlyon itsprotoplasm depends desiccation buton thecellsavoiding surviving better, severe diurnal mechanical stress H. tunbrigense during drought periods. fails to avoidthese stresses and appears to have even lesscontrol over water lossduring ambient conditions. changing The sensitivity ofH. tunbrigense to desiccation over periods up to 11 dayswasmanifested inprogressive tothecells ofexposure damage both with at a given length humidity and with saturation deficit at a given length ofexposure. After 5 daysat 50% R.H. at 200C abouthalf thecellswere killed. In H. wilsonji similar under conditions lessthan 11 days:evenafter killed after 5 daysat 0% R.H. damage 10% ofthecellswere was slight. ofH. wilsonii, Exceptional drought that in an isoresistance, approaching wasfound lated at 1000 inSnowdonia ft colony (305m)ina site (Caernarvonshire) unusually exposed to drought anddirect inthelaboraLeaveskept at continuously sunlight. high humidity
2n = 24 (Litardiere from (d) Chromosomes. 1921) (material France); n = 13 (Manton

(b) Mycorrhiza. Not reported. and vegetative Herbaceouschamaephyte, (c) Perennation reproduction. perennating by

P. W. RICHARDS

AND

G. B. EVANS

255

inH. tunbrigense resistance Drought to drought. sensitivity an increased developed tory with theplant's when combined which level, at cellular response to be a plastic appears in on colonies confer unusual drought resistance may plasticity morphological great habitats. exceptional to alternating are exposed thatwhenplants forms it was found growth In normal inthe amount there little difference was atmosphere of and saturated periods lowhumidity periods at thesamelow to continuous exposed ofplants with that compared ofdamage wasmoistened plant ifthe increased wasgreatly mortality hand cell other Onthe humidity. to theplant that damage Theseexperiments suggest liquidwater. contact with bydirect ofthe on theduration than oftime that it is desiccated lesson the totallength depends more being ofwater content a rapid change ofincreasing ordecreasing hydration, periods by plant ismoistened when the occurs which one.Thecelldamage a gradual than harmful the doesnotoccur near H. tunbrigense why may be a reason when desiccated water liquid ofrunning water. zoneofwaterfalls andin thecloseproximity spray with thatof H. is very low,contrasting of thegametophyte The drought resistance diedunder ofH. tunbrigense gametophytes In oneexperiment a large proportion wilsonii. ofH. wilsonii. gametophytes among lessthan15% mortality caused conditions which of method bythe investigated were points Compensation assimilation. Light andcarbon localities from Caernarvonshire ofH. tunbrigense Samples 1961). & Ashton (Lieth Lieth intensities than at lowerlight to reachcompensation exception without werefound In field samples thecompensation sites. adjoining of H. wilsonii from nearly samples inplants more situations from shady andlower variable ofH. tunbrigense wasquite point inwhich thecompensation in those places.In experiments from better-illuminated than inplants ofthe from 200to 500C wasmeasured temperatures at successively lower point it similar conditions under closely forsometime had beencultivated which twospecies constant whilein in H. tunbrigense thecompensation pointremained that was found ofH. tunbrigense. that equalled at 500 C italmost fell until H. wilsonji itgradually H. with as compared pointof H. tunbrigense lowercompensation The consistently illuminated for very poorly ofthe former tolerance toexplain thegreater wilsonii appears in point on thecompensation oftemperature effect (see IIa above).The differential sites at toreplace H. tunbrigense ofH. wilsonii onthe tendency the twospecies throw light may altitudes. higher ofHymenophyllaceae, especially thephotosynthetic pigments Athigh intensities light 1940). aredamaged (Gessner xanthophyll, in which leavesofthetwospecies & Jones, unpublished) In an experiment (Richards in cagesgiving to daylight and thenexposed werekeptin thedarkuntilstarch-free at lower in theleavesofH. tunbrigense ofshading, reappeared starch different degrees in those than ofH. wilsonii. values daylight weather conditions under favourable occur flushes which The growth VII. Phenology. a beenmentioned (VIc). Thesemayoccurat anytimeof year,though havealready Thisis orautumn. oneinlatesummer inspring anda minor main flush develops usually of manyEuropeanmosses(Hagerup1935).In Doignon's similar to the behaviour four times renewed during leaves were completely atFontainebleau (seeII) the transplants ofthe the soonafter toappear expansion ofobservation. Indusia begin the12-year period forseveral of theleafapex maycontinue As growth fewpinnaeof a newflush. first to one flush. each corresponding maybe found, ofindusia successive groups seasons, to oftheyear, owing partly can be found over a longperiod Plants with ripesporangia of individual to thesensitivity owing of thesorusand partly thegradate development

256

(L.) Sm. tunbrigense Hymenophyllum

in in their havebeenfound environment. Ripespores immediate colonies to conditions the colonies butin most inNorth oftheyear at onesiteoranother Wales, month every andwinter. is inautumn production peakofspore on thephenology ofthegametophyte. is available No information witha aboutthirty per sorus, Sporangia of thesporophyte. VIII. (a) Reproduction of Thepotential output 1926). (Bower number' of258-512 spores persporangium 'typical Though (cf.about3000in H. wilsonii). spores per sorusmaythusreach7500-15000 havenotbeen in abundance, sporophytes newly-established young, areproduced spores condiunder at least present ofcareful andeffective reproduction, search, in spite found and very seems to occur Germinating spores onlyvegetatively. tionsin GreatBritain, moist in are to inside the very indusia, especially often be found young gametophytes In the havebeenseenin nature. or wetseasons, gametophytes butno older conditions be found as an H. sometimes epiphyte where tunbrigense can Ireland, woodsofsouth-east totake place. Apospory establishment from must bepresumed spores onquite trees, young reported. has notbeen in droughts thesporangia: gapeopenexposing theindusial valves weather During dry incomplete inthis havean oblique way. Thesporangia be killed sporangia may immature nearcolonies with butattempts to catchairborne spores and dehisce laterally annulus numerous and contain have failed.Ripe sporesare usually3-celled ripe sporangia spore discharge As young inside theindusia, found areoften gametophytes chloroplasts. theclosed wall, perhaps inside indusium ofthesporangial bybreakdown takes place often be ofthedividing therefore may aboutbytheinternal spores. Dispersal pressure brought effected chiefly byrain-wash. andilluminahumidities under temperature laboratory conditions (moderate atvery high slow. In and subsequent of the prothallus is extremely growth tion). Germination stage in 12 months. notone gametophyte reached a 40-cell culture laboratory form which Goebel(1930)regards butnotinvariably, found to grow outinthe'tripolar' of thegenus. in Fig. 3. By 18 is shown as characteristic A 2-month-old gametophyte in size months nonehadproduced The cellsofthegametophyte aresimilar sexorgans. ofH. wilsonii. ofthegametophyte to those oftheleaflamina andaresmaller than those and only Mettenius a leaf-like it was sterile prothallus: after (1864)described 11 years 1 mmlong.Janckewski grown of gametophytes & Rostafinski (1875)givean account in France mature when from collected Thesewereribbon-like nearCherbourg. spores wereseen.The embryos sex organs:on two gametophytes attached and produced ofthegametophytes inother species ofHymenoorgans anditsreproductive morphology archegonia on the (1940,1948)whoobserved by Stokey havebeendescribed phyllum dorsalsurface. has notbeenreported. Apogamy to is frequent in theHymenophyllaceae, butdoesnotseem bygemmae Reproduction in this havebeenobserved species. Noneknown. (f) Hybrids. Wales, Nibbled leaveshavebeenseenin North orparasites. feeders IX. (a) Animal Slugs and woodlicehave been observed feeding by smallinvertebrates. suggesting and sporangia. indusia feeding bydayon theleaves, Noneobserved. In North epiphyllous hepaticae andinIreland Wales (b)Plantparasites. (1945-1962) are occasionally foundon theleaves.Duvigneaud Lejeuneaceae) (mostly
wereusually, In cultures thespores andreproduction ofgametophyte. (d), (e) Morphology and germination of spores.Spores remainviable forlong periodsif kept (c) Viability (b) Dischargeand dispersalof spores.The sporangiaripen in basipetal succession.

P. W. RICHARDSAND G. B. EVANS

257

ofH. tunbrigense on herbarium specimens found andlichens hepaticae epiphyllous lists andLuxembourg. Belgium from andH. wilsonii havebeenreported. diseases or other No fungal (c) Diseases. preplantby Ray (1686).Foundsomeyears as a British reported First X. History. by to shown Ray and later Sussex, Wells, Tunbridge viously byDaireat theHighRocks, 1962). Doody(seeEvans& Jermy
(b)

(a)

0000

00

0~~~~~~~~~

(L.) tunbrigense of Hymenophyllum sporesand younggametophyte FIG. 3. Germinating (b) Young Cain, Merioneth. Pistyll Sm. (a) Germinating sporesin sorus, x 400, from in sorus,x 180. gametophyte

Co. Galway, Ireland, at Gort, (1959)ina deposit found Andersen & Farrington byJessen, Interof theHoxnian In a deposit Interglacial. as dating theHoxnian from regarded numerous sporesof H. Watts(1959) discovered glacialat Kilbeg,Co. Waterford, at Kilbeg flora tree Theaccompanying inside. spores andtwosporangia with tunbrigense thepresent-day andresembles and Betula Alnus with associated trees includes evergreen is still found): tunbrigense Hymenophyllum ofthewestern flora Caucasus(where forest of probably Oceanicelement. Spores, theherbaceous speciesbelongto theWestern at Nechells, Birmingham, in Hoxnian deposits Interglacial arefrequent H. tunbrigense, are absentor rarein the they a stagewhen Picea and Abiesweredominant: during of HymenoRemains communication). personal Pinusphase(M. R. Kelly, succeeding in district in theBiarritz havebeenfound from thesameInterglacial, dating phyllaceae, occurs today(Oldfield, tunbrigense an areawhere Hymenophyllum thewestern Pyrenees, in West1961). is in Europe anditspresent area,which is undoubtedly H. tunbrigense a relict species of be theremains what must discontinuous, stilldecreasing and becoming increasingly Isles intheBritish Itsconcentration andmuch more continuous range. wasoncea larger further stations scattered outlying with widely andthe extreme Europe west ofcontinental As has beensuggested Oceanicbryophytes. eastis similar to thedistribution ofmany in which H. tunbrigense occupied period recent for thelatter themost 1957), (Richards

were to Hymenophyllum wilsonii, associated with largerspores attributed speciosum,

or to Trichomanes to H. tunbrigense either belonging Sporesof Hymenophyllaceae

258

(L.) Sm. tunbrigense Hymenophyllum

opticlimatic mayhavebeenthepost-glacial or lesscontinuous range a wideand more a still areain thewarm and itmay wellhaveoccupied Interglacials larger mum (though itsareahasbecome restricted this period optimum Tertiary). Since backas the even as far ofits centuries bythedestruction andinrecent deterioration, byclimatic andfragmented etc.(cf.Rose 1957). through forest clearance, habitats byman, in being from thatofH. wilsonii ofH. tunbrigense in Europe differs The distribution relict stations and in having Saxony, Luxembourg, (Sussex, moresouthern outlying in these to persist of H. tunbrigense themainarea. Theability etc.)farfrom Adzharia, its of on tolerance extremely poorly-illuminated stations depends probably outlying in deepcrevices in sandstone where etc., rocks, it to survive Thishas enabled habitats. it even in like are for Saxony suitable regions and regimes thehumidity temperature itslimits H. wilsonii oftolerance. mayalso have is faroutside themacroclimate where not survived in widely it but it has much area than does now, a larger onceoccupied it from its excludes the stations because light requirement relict relatively high dispersed for H. more which drought-resistant crevices Though tunbrigense. refuges provide shady in ofdesiccation to be ableto grow is notsufficiently tolerant H. wilsonii than thelatter, in demand situations which itslight except well-illuminated requirements therelatively Norway). France, west Britain, north-west (western ofhigh humidity atmospheric regions
wilsonii). see underHymenophyllum (For References

ListBr.Vasc.P1.(1958)No. 7.2 Hooker wilsonii Hymenophyllum auct.vix(Poir.)Desv.,H. unilaterale auct.vixBoryex Willd.) (H. peltatum ingeneral H. tunbrigense appearance A mat-forming resembling perennial, bryophyte-like and lessclosely to the as follows. Leavesmoreerect appressed and size,butdiffering in when dark olive never old,thicker bluish-green becoming blackish, substratum, green, curved when and morerigid, with thepinnae back(especially lesstranslucent texture, is notflat and appears to its relative so that thelamina narrower and overlapping dry) of 5-11),thevein Pinnasegments fewer (3-5 instead surface shiny. slightly length: regular Leafcells more larger (94+2 x 39+1 ,um), theapexonmature segments. reaching insurface more cf.30-40inH. tunbrigense). view, andwith chloroplasts (c.60-80 inshape indusia with convex strongly pear-shaped stalked; appearing Soriup to 10-15perleaf, more indiameter, Spores 62-78,um quite entire. overlapping, slightly valves, themargins than those ofH. tunbrigense. ornamented finely in size;no subspecies orvarieties havebeendescribed. Varies little, except and never withH. tunbrigense, the H. wilsonii is very distinct intergrades Though nowbe for after which would ofH. wilsonii wasdisputed plants many years status specific of saw specimens to it werefirst described by Ray (1686).Ray subsequently assigned to thesamespecies until from them as belonging H. tunbrigense Kent,but regarded 1962).Linnaeus himthatthey weredistinct (Evans& Jermy later persuaded Plukenet thebryolothem anditwasnotuntil 1830when afterwards and others againcombined inthe haddrawn Hooker's attention to both well field) species (whoknew gist W. Wilson united wasclearly established Hooker subsequently their distinctness (though them that oncemore). Taxa closely theEuropean H. wilsonji arefound resembling thetwospecies

P. W. RICHARDS ANDG. B. EVANS

259

and on mountains in thetropics: somehaveregarded H. in theSouthern Hemisphere ofthe (Poir.) Desv.orofH. unilaterale earlier described H. peltatum wilsonii as a synonym therelationship ofthe (Bourbon). As with H. tunbrigense, Bory exWilld. from Reunion investigation: meanwhile the populations requires further European andextra-European as a species usedforthisplantif it is regarded nameH. wilsonii can be legitimately andtheAtlantic islands. confined to north-west Europe
0 1 2 3 4

PHYLLUM WILSONII Hook.

HYMENO-

ll
9 2

Filmy Fern * 1930 onwards o Before1930 S t Ld

7

-___ 0 02 0

0~~~
0 5

006

*0O 00 I0 * 0 3 000 0 00

0~~~~~~~~
0
9

Soo

~~~~~~~~~~~~~~0
_

I~~~~~~~~~~~~~~~~

4~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

of Great Native.Widely distributed and locallyabundant in the westand north onrocks, onmountains andin Britain andin Ireland, growing inturf among bryophytes tolerant ofexposure anddesiccation than H. tunbrigense. somedistricts on trees, more I. Geographical and altitudinal distribution. ItsareaintheBritish Isles(Fig. 1) overlaps widely distributed (recorded inmore than twice as thatofH. tunbrigense, butitis more and from fifty-six vice-counties in GreatBritain and twentymany 10-km grid squares respectively). ninein Ireland compared with thirty-seven and twenty-five In southern inthesouth-east, allitsEnglish stations lying north ofa England itlacksoutlying stations linerunning from Start Point(Devon)to themouth oftheTees,butin Wales,north

260

wilsonii Hooker Hymenophyllum

andScotland itextends further east:italsoextends further north England (toShetland). Like H. tunbrigense it has sometimes in error to confusion beenrecorded owing with liverworts. from andnorth-west In Europe Britain itis known theFaeroes, outside only Norway itis found coastfrom Listato Stadwith France (Fig.2). In Norway alongthewest one outlying station further north at Stemshesten (630N) (seemapand comments in Faegri
6 30 24 70 18' 12 18'E 12 o 24: 30 36 42' 48'
54I.

_ 60-- -

72

5?

60'

630'w1

0r

of Hymenophyllum FIG. 2. The distribution wilsonii Hooker in Europe. 0, Native oftheCommittee for occurrence; x, probably extinct. Reproduced bypermission Mapping theFlora ofEurope.

ithasbeenrecorded from four in Finistere localities and Manche 1960).In France only it is less abundant and in these thanH. tunbrigense (Doignon1947).The onlyextrawhich are fairly thosein Europeare with European populations certainly conspecific those in theAtlantic islands (Azores, Madeira, Canaries). inBritain Thealtitudinal is from sealevel to3300ft(1000m)onMacgillicuddy's range Reeksin south-west Ireland (Hartin Scully to2000ft 1916).In England (610m) inwest and at least2850ft(864 m) on Yorks;in Walesto 2000ft(610 m) on Plynlimmon Snowdon. In Caernarvonshire, themaximum number of localities, in 500 ft grouped is at 500-1000 ft(152-305 in Evans1964).In Scotland intervals, to m) (see histograms 2500ft(762 m) in Breadalbane and 2012ft(613m) in theHebrides In the (Harris).

AND G. B. EVANS P. W. RICHARDS

261

300m (Hansen1966).In stations are under known out ofthefifteen Faeroesthirteen to be at in all appear French localities 1960). The to at least350m (Lid, Faegri Norway (Doignon1947). low altitudes

surfaces on shadedor exposed and amongscrees boulders on moss-covered outcrops, or on the trunks and branches vertical inclined), steeply butbyno means always, (often, that of thus overlaps Itshabitat range andbryophytes. andinturf among grasses oftrees conditions into more where it extends is more but open considerably H. tunbrigense LikeH. tunbrigense in dry weather. to desiccation andmore illuminated subject strongly itappears butin general less etc., boulders, in crevices, deepholesamong itoften grows shadeor in theopenprovided in moderate better the ofdeepshadeand grows tolerant it often in to H. in boulderThus contrast grows is not too tunbrigense dry. atmosphere in quite openwoodrocky ridges oron small streams ravines without permanent strewn inthesamelocality, occur ofHymenophyllum they arevery thetwospecies Where lands. there areseparate orzonesofeach, carpets cushions, usually intermingled; found rarely and well-illuminated themoreexposed in general situations and H. wilgonii occupying and darker ones.In north sheltered themore etc. Wales,westScotland, H. tunbrigense inwhich H. tunbrigense is confined to thelowermost to find stream valleys itis common andmore thezone far into theupper openparts: H. wilsonii extends while sheltered parts inthe there is a similar difference narrow. distribution of be quite On trees ofoverlap may thetwospecies (seebelow). butseems inthespray zoneofwaterfalls, absent itis usually rather LikeH. tunbrigense, on rocks liableto flooding, found and is occasionally of running water moretolerant Du, Merioneth. e.g.at Rhaeadr on factors partly clearly depends occupied by H. wilsonii The rangeof habitats oftherainfall, anddistribution theamount particularly humidity, affecting atmospheric wouldprobably be factors ofcloudandfog:these andthefrequency to wind exposure butadequate dataarelacking. in terms ofpotential evapo-transpiration, bestexpressed in the to sheltered it tends to be restricted etc.,while gullies, In areasoflow humidity to sunand on hillsides on rocks or in turf fully exposed humid areasitmaygrow most island on Slieve Ireland). (west More,Achill Merioneth, e.g.on theRhinogau, wind, thedistribution difference between of H. is an interesting there In Caernarvonshire it occurs of over60 in. (150 cm),where an annualrainfall in Snowdonia with wilsonii andinthe Peninsula where the ofhabitats andaltitudes, ina wide Lleyn range abundantly on to very small sites itis confined north-facing is 40-60in. (100-150 cm): here rainfall CarnMadryn, ofthehills(Yr Eifl, etc.) Moel-y-Gest, nearthesummits or screes rocks andsuitable much ofLleyn is cultivated because m.Thismay be partly at about200-550 ofclouds to persist on on thetendency also depends butitprobably arelacking, ravines thehilltops. to thatin similar and climate habitat there is a relation between range In Norway situait grows even in exposed archipelago Britain (Faegri1960, pp. 76-7).In theouter it becomes On themainland increasingly largeareas in purestands. covering tions, situaand sheltered restricted to moist from thecoast, distance with becoming exacting In factor. oftheairis thedecisive that thehumidity controlling concludes tions. Faegri common onsteep israther H. wilsonii ofsouth-west north-facing the Norway region fjord mountain slopes(Lye 1966). invery humid areas on trees occurs as an epiphyte IslesH. wilsonii only In theBritish of Detailsofthedistribution west Scotland). and south Merioneth, Ireland, (south-west

H. wilsonii is found on rock limitations. II. Habitat. (a) Climaticand topographical

262

wilsonii Hooker Hymenophyllum

byRichards in Derrycunihy aregiven Wood,Killarney, species thetwoHymenophyllum inlarge 30ft(9 m) abovetheground. mats on thetrees to over occurs (1938).H. wilsonii level at ground aretallest and theillumination thetrees In thepartofthewoodwhere aresmaller than9 m,butin parts where thetrees lower (Type1) it is notfound lowest descends basesand on thetree 2, 2a and 3), H. wilsonii andtheshadelessdeep(Types in all on tree basesand boulders mainly H. tunbrigense grows on boulders. also grows bothspecies farup thetrunks. Where ofHymenoofthewoodandnever ascends parts a higher zonethanH. tunbrigense H. wilsonii on thesametree, occupies occur phyllum is very little overlap. andthere can be seenat Coed Ganllwyd, Merioneth. ofthetwospecies distribution A similar as at than H. is in tunbrigense H. wilsonii an levels higher of epiphyte Theoccurrence has and that the former requirements higher light with the evidence 254-5) (pp. accordance to H. wilsonii in the level which the latter. The variation than tolerance drought greater parallel H. tunbrigense Woodis closely in Derrycunihy ascends) descends (andto which occur in of in which different tropical the levels at types variation epiphyte synusiae tothe in the variations similar (1911)observed pp. 121-2).Shreve rainforest 1952, (Richards in in ferns the Mountains Jamaica. of Blue distribution filmy height Isles it is absent south-east thepartoftheBritish England, from in Europegenerally; is noexperimental H. wilsonii that evidence summer There thehighest temperatures. with H. tunbrigense (X), ofhigh summer under and,as suggested temperatures is lesstolerant comin south-east maybe dueto a lackofhabitats ofH. wilsonii England theabsence is witha sufficiently humidmicroclimate. H. wilsonii adequateillumination bining that the normal temperature oflowwinter (seeVc) anditis unlikely tolerant temperatures in theBritish oftolerance Isles. itslimits anywhere exceed conditions with at least200 rainy 40 in. (1000mm)or more annualrainfall, arein areasreceiving days. forH. wilsonii lie within less than3 in. stations thearea with All British probably on Green's water deficit map(1964)(cf.H. tunbrigense, average potential (7 6 cm)annual II). Evans (1964)collected relative humidity Microclimate. manydata on temperature, inNorth for Waleswhere andH. wilsonil sites H. tunbrigense deficit andvapour pressure is in showing thatthough H. wilsonii The results are consistent growneartogether. in siteswarmer the and lesshumid thananyoccupied by H. tunbrigense found often ofthetwospecies overlap. frequently microclimate ranges and in Norway (Faegri1960)it is and chemical characteristics. In Britain age,texture suchas Carboniferous because oftheir limestone, whether on typical limestones, absent tuffs, e.g.at Cwm on basicvolcanic or their chemical butit grows properties, physical In the andCaderIdris, Merioneth. Caerns. IdwalandbyAfon Valley), (Conway Dulyn inonelocality, itoccurs Co. Clare(Ireland) limestone areaoftheBurren, Carboniferous rocks on Corylus butnotonlimestone (Webb and on sandstone as an epiphyte avellana of3 9wilsonii is found overa pH range Norway Hymenophyllum 1963).In south-west 4*3(Lye 1966). deadmossto peat on deadplant material from ranging H. wilsonji frequently grows petraea: on living it usually occurs on Quercus and rawhumus. As an epiphyte plants and the it has beenobserved also on Ilex aquifolium in Derrycunihy Wood,Killarney,
on rocksvery variedin geological Like H. tunbrigense, H. wilsonil grows (b) Substratum. All knownBritish stationsfor H. wilsonii evapo-transpiration. Rainfalland potential in Britain and has a morenortherly H. wilsonii rangethanH. tunbrigense Temperature.

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G. B. EVANS

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H. tunbrigense). in Table 1 (seeunder aregiven H. wilsonji desiccation, and temporary of exposure tolerance its greater Becauseof range a wider in and occurs tunbrigense H. than associates and varied numerous more has and in thesamewoodland growing Islesbesides In theBritish of plantcommunities. communities, in various other it is notuncommon as thelatter, moorland associations and chasmo(chomophyte on mountains rockledges and crevices ofbase-rich e.g.that species arctic-alpine include theassociates 1949)where of Tansley communities phyte In St Kilda plants. are non-vascular all itsassociates nearly where rockcommunities its greater of Because in 1927). (Turrill Callunetum occurs wilsonli HIymenophyllum associated often is more H. wilsonii situations, andwell-illuminated for exposed tolerance successto compete and seems mixed with bryophytes intimately grows It commonly and albicans are Diplophyllum associates withthem;two of its mostconstant fully altitude at a high scree on a boulder Onecasewas observed Rhacomitrium lanuginosum. andlaboraquinquefarium bySphagnum ithadbeenovergrown innorth Walesinwhich and Leucospecies thetwoHymenophyllum on competition between experiments tory
is less wilsonji moistconditions Hymenophyllum indicatethatundervery bryum glaucum withlichensthanis H. tunbrigense. it also occursin bare alpinum: and Thalictrum S. stellaris suchas Saxifragaoppositifolia,

On Foula (Shetland)it is commonon myrtillus. and Vaccinium stems of Callunavulgaris 1969). tussocks(Hawksworth squarrosus Juncus and sylvatica thebases ofLuzula wilsonji communities for including Hymenophyllum Lists of species III. Communities.

thanH. tunbrigense (Evans1964). successful overlaps, species theecological rangeof theHymenophyllum As has beenstressed, Theyusually and humidity are different. to light with respect their tolerances though in on trees as sometimes Where, growin mixture. areas and rarely discrete occupy in in north grow in sheltered Wales,they ravines and on rockfaces Ireland south-west must be determined by thetwospecies balancebetween thecompetitive closecontact, resistance anddrought (seeVId). different points their compensation itsassociates In Norway in Finistere. ofassociated details species Gaume(1944)gives chrysoas Breutelia such species Atlantic as in Britain, including, bryophytes, arechiefly
undulatum and Plagiothecium Isothecium myosuroides atrovirens, coma, Campylopus

withvarious together wilsonii, NorwayHymenophyllum (Lye 1966). In south-west It mayactas a pioneer onrock surfaces. ina cyclical succession a part plays bryophytes, peel often andbryophytes ofH. wilsonii carpets Later theheavy barerock. incolonizing to be re-colonized (Lye 1967). a baresurface leaving off,
than is less affected factors.In wooded habitatsH. wilsonii IV. Responseto biotic

forsometime condition in a depauperate andmaysurvive H. tunbrigense felling bytree from andregenerate burning It can survive light has beenremoved. cover thetree after therhizomes. in north Walesand Ireland or patches which Formscolonies V. (a) Gregariousness. than aresmaller the patches ft2 10-20 (1-2m2).In general mayreach 1950) 1934, (Praeger tend to be more diffusely 2 ft2 anddo notexceed (02 m2). Theleaves in H. tunbrigense and is growing theplant andwhen inH. tunbrigense bryophytes among than distributed are always thepatches whether It is notknown scattered. maybe widely they grasses more thanonegametophyte. or can arisefrom clones in therock faceat IdwalSlabs,CwmIdwal, In exposed e.g.cracks ofthehabitat. sites, sheltered placesthe butin moist than 05 cmlong, theleaves maybe no more Caerns.,
varies greatlywith the humidity in varioushabitats.Performance (b) Performance

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Hymenophyllum wilsonii Hooker

plant may form loosehanging masses upto 10cmdeepwith individual leaves upto20cm long. Sori are mostabundant on smallleavesin conditions thantheoptimum drier for vegetative growth; they are usually absent on etiolated or very luxuriant leaves.The number ofsoriperleafrarely exceeds fifteen, butseveral groups maybe found on one leaf, to flushes ofgrowth. corresponding

areharmful inSnowdonia selves andtheextensive to theleaves observed damage (Evans) andin Scotland in (Ratcliffe, personal ofearly communication)theunusually longfrost wasprobably 1963 duetothe combination ofsevere frost InNorandprolonged drought. was no traceoffrost waythere thesevere oftheearly1940's, damage during winters atitshighest even stations near at c.250m(Faegri do Bergen 1960). Though droughts may little about the off of colonies from rock physiological harm, they maybring peeling and thus surfaces leadto their destruction. Observations showthatH. wilsonjiis less liableto damageby drought thanH. intherateat which Thedifferences between thetwospecies water is lostfrom andtaken up bytheleaves from a saturated atmosphere suggests that there might be differences in thefine ofthecuticle structure ortheouter cellwalls, butno suchdifferences havebeen with detected thelight microscope.
(b) Mycorrhiza. No information. and vegetative (c) Perennation Herbaceouschamaephyte, reproduction. perennating by

(c) Effect offrost, drought, etc. Thereis no evidencethatlow temperatures by them-

tunbrigense. VI. (a) Morphology in mostrespects of sporophyte. Similar to thatof H. tunbrigense.

meansofitsrhizome is probably system. Reproduction byspores(see VIIIa), and by oftherhizome or wholecarpets. ofparts fragmentation Clowes(in Moore& Lindley that theleaves for 1855)stated were several innorth persistent years. Observations Wales on marked leavesin thefield confirm thatthey are normally long-lived withgrowth onweather flushes onceortwice a year, depending conditions; some leaves for persist at least5 years. is slow;ina setofthirty-seven Vegetative spread over rock surfaces pioneer rhizomes in Merioneth marked notone grew morethan2 cmin 12 months (all were ofclones orcolonies killed subsequently bydroughts). unknown. Longevity n = 18;the chromosomes inthe (c) Chromosomes. are'the recorded Pteridolargest yet phyta' (Manton1950, material from Borrowdale, Cumberland).

basedon Evans1964)is discussed under thelatter (mostly most (VId). Theecologically facts significant which havebeenestablished are: (i) theleafcellsofH. experimentally wilsonii areconsiderably more than those ofH. tunbrigense; of drought-resistant (ii)leaves H. wilsonii to a dry both losewater andgainwater from a saturated atmosatmosphere pheremoreslowly thanthoseof H. tunbrigense; (iii) thegametophyte of H. wilsonji is moredrought resistant thanthatofH. tunbrigense; between 10 (iv) at temperatures and 200 C, theleavesof H. wilsonli havea higher compensation pointthanthoseof H. tunbrigense, butat 5?C thecompensation is aboutthesamein bothspecies. point in thephenology VII. Phenology. No differences of H. wilsonji have beenobserved andH. tunbrigense. Theleaves oftheformer to be longer-lived than those ofthe appear butthis seems tobe duetothe ofH. tunbrigense todamage latter, greater susceptibility by on thephenology There is no information ofH. wilsonii. ofthegametophyte drought. from five the number ofsporangia wastwenty-five. The'typical' leaves) average persorus
VIII. (a) Reproduction of thesporophyte. In a sample fromnorthWales (thirty sori

(d) Physiological data. The comparative and H. tunbrigense physiology of H. wilsonii

AND G. B. EVANS P. W. RICHARDS

265

per sporeoutput is 128(Bower1926).The potential persporangium ofspores number with compared (about3000-4000, sorusis thusless thanhalfthatof H. tunbrigense fertile frequent than inH. tunbrigense: more is,however, ofsori Production 7500-15000). plant)and patches (or perindividual percolony morenumerous leavesare relatively in abundance, areproduced spores Though arerare. leaves without at leastsomefertile vegetative: ismainly asinH. tunbrigense, reproduction, effective that suggests the evidence havearisen might which smallsporophytes norvery gametophytes free-living neither on of H. wilsonji theoccurrence havebeenseen,though gametophytes from recently evidence is circumstantial Ireland etc.in south-west vulgaris stems, Calluna trees, young has notbeenobserved. takes place.Apospory from spores that reproduction
(a) (b)

Hooker.(a) x 70; (b) x 160. wilsonii ofHymenophyllum FIG. 3. Gametophytes

and dispersal tunbrigense. ofspores.As in Hymenophyllum (b) Discharge The sporesshowa highpercentage andgermination germination ofspores. (c) Viability oftenbegins withinthe sporangiumand under laboratoryconditions.Germination but theoldergametotheindusium, be foundwithin sporescan sometimes germinating is notas slowas ofgametophytes In culture, havenotbeenseenin thefield. growth phytes in H. tunbrigense. and reproduction Germinating sporesmayreacha ofgametophyte. (d), (e) Morphology are shownin in culture three-celled Gametophytes leavingthe sporangium. stagebefore have numerousrhizoids and Fig. 3. The largestprothalli,which are strap-shaped, ofarchegonia One prothallus of4 mmin 12 months. reacheda length produceda number died. Apogamyhas not been reportedand gemmaehave not been but subsequently observed. None reported. (f) Hybrids. IX. (a) Animal by smallanimalshave been obfeedersorparasites.Tracesof feeding Wales and woodliceand slugshave been seeneatingtheleaves, servedon plantsin north but the amountof damage is verysmall. lichensand hepaticaeare sometimes (b) Plantparasites.None observed.Epiphyllous foundon theleaves (see underH. tunbrigense IXb).

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Hymenophyllum wilsonii Hooker

X. History. First record: described byRay(1686)as 'Adiantum petraeum perpusillum anglicum foliis bifoliis veltrifidis from sent to himbyN. Newton from Newtoni' plants Buzzard RoughCragnearWrenose in theLake District. [Wrynose] Interglacial finds ofHymenophyllum spores, someofwhich may belong to H. wilsonii, are mentioned underH. tunbrigense (X). The onlyfossils definitely identified as H. wilsonji aretwospores andthree infine detritus mud sporangia containing SalixandPinus woodfrom Pollen ZoneVIc at Wybunbury inlitt.). Moss,Cheshire (H. J.B. Birks, The conjectural history ofHymenophyllum has already wilsonii beendiscussed above under H. tunbrigense andnoout(pp.257-8).It hasa lessfragmented areathan thelatter lying relict inEurope stations faroutside itsrange intheBritish France Isles,north-west andwest Norway. Though there is no direct evidence that itformerly occupied a much larger areathan itdoesnow, itnevertheless seems plausible that in somemoist climatic period(thepost-glacial or earlier) climatic optimum it extended further eastthanat present. If thiswereso, its absencein the SussexWeald and in continental Europe (except inFrance andNorway) would be difficult to account for that except byassuming to itshighcompensation owing point(and relative intolerance of deep shade), which it from excludes thekindofhabitats in which H. tunbrigense has beenable to survive, suitable for itdo notexist. totest itsability refuges In 1961oneofus (G.B.E.) attempted to grow on apparently suitable at twolocalities in Sussex and Kentby sandstone rocks transplanting plants from north Wales.The plants failed to establish and afterwards diedin a severe drought. Though H. wilsonii has disappeared from someof itslocalities, is no evidence there that itsareais diminishing seriously onan atthe present time. Because itislessdependent extremely moist and sheltered environment than H. tunbrigense, lessfrom ithassuffered deforestation and destruction in of itshabitats; also becauseit is relatively abundant ofitsarea and perhaps many parts it has rather lessconspicuous thanH. tunbrigense, beena lesseasyvictim to collectors. Faegri (1960, p. 77) supposes that itreached Norway from thewest bydirect immigrationacrosstheNorthSea, butif,as suggested above,its continental rangeformerly extended further eastthannow,thisis notnecessarily true.
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(c) Diseases. None known.

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EVANS

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