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The British Pteridological SocietyTHE FERN GAZETTEVOLUME 15 PART 61998CONTENTSPageMAIN ARTlCLESTrichomanes speciosum Willd. (HymenophyUaceae: Pleridophyta)in southern SpainF.J. Rumsey & J.C. Vogel197Population analysis of the type localities of some recently recognised taxaof British Pteridium (Dennstaedtiaceae: Pleridophyta)K.M.Bridge's. C.!. r\shcroft 1: Sh 'ffi 'ld205The addition of Acrosorus streptophyUus (Baker) Copel to theflora of ThailandB.S. Parris215BOOK REVIEWS204,214,216,218INSTRUCTIONS FOR AUTIIORS220Tl'lli H::.RN GAZE'ITE Volum 15 Part5wuspu ilihcdtlnI 'ilh Dcccmb1Jr. 1997PUblbhed by Tl m BRTTISJI Pl'f:RJDOl.OGICAL SOCIETY, c/u Ocl'llrllllcnt of BOtnny.The Natural Hiswry Museum, London SW7 5BDISSN 0308-0838Printed by Metloc Printers L CaxtonHouse, Old Station Road, Loughton, Essex IG 10 4PE

197FERN GAZ. 15(6) 1998TRJCHOMANES SPECIOSUM WILLD. (HYMENOPHYLLACEAE:PTERIDOPHYTA) IN SOUTHERN SPAINF.J. RUMSEY & J.C. VOGELConservation Biology Laboratory, Department of Botany,Natural History Museum, Cromwell Road, London, SW7 5BD, U.K.Key words: Macaronesian enclave, refugium, Tertiary relict flora, conservation.ABSTRACTThe history of the discovery, present distribution and status of Trichomanes speciosum insouthern Spain is given. A new site in the Sierra de Montecoche, supporting the gametophytegeneration only, is reported. The sporophyte generation has apparently declined in outlyingareas; the extent to which this is a natural cyclical process in drier periods is uncertain. Habitatdegradation through human activity has undoubtedly exacerbated climatic effects. Thespecies anay present supports the hypothesis that this area has acted as a European mainlandrefugium for Tertiary relicts otherwise largely restricted to the Atlantic Islands. Protection ofthe sites supporting Macaronesian floral elements is urgently required.INTRODUCTIONTrichomanes speciosum W illd. (syn. Vandenboschia speciosa (Willd.) Kunkel) is the sole nativeEuropean species of a widespread genus (Tutin et al., 1993). A Macaronesian-Europeanendemic, it has been considered one of the continent's most threatened pteridophytes. Excessivepast collection for horticulture (Allen, 1969), and the herbarium, in combination with therestriction of small sporophyte populations to few, scattered and vulnerable sites hasnecessitated legal protection throughout the species range under the Bern Convention (Anon,1979) and the Habitats Directive (Anon, 1992). Trichomanes speciosum has also been listed asglobally rare in the IUCN Red List (Walter & Gillet, 1998).Recent studies, summarised in Rumsey et al. ( 1998), have revealed that the gametophyte(sexual phase) of the life cycle, first described from cultivated material (Goebel, 1888), but notreported in the wild until over a century later (Rumsey et al., 1990), is scattered across Britainand Ireland and parts of continental Europe. Similar 'independent' Trichomanes gametophytepopulations have been reported in northern temperate areas of the U.S.A. (Farrar, 196 7 , 1985 ,1992) butT. speciosum would appear unique in that both populations with gametophytes onlyand those with both gametophyte and sporophyte generations are widespread.PAST RECORDS OF TRICHOMANES SPECIOSUM IN SOUTHERN SPAINKnowledge of the presence of Macaronesian floristic elements in the hills in the Algeciras areadates from the report of the discovery of Culcita macrocarpa C. Presl in the Miel Valley, afew kilometers W.S.W. of Algeciras (Allorge, 1934). However, both C. macrocarpa and thenow lost Dryopteris guanchica Gibby & Jermy had been collected in this area by Englishbotanists in the nineteenth century, who either failed to appreciate their significance ormisidentified the material (Fraser-Jenkins, 1982). Curiously Allorge ( 1934) did not recordTrichomanes speciosum, or Diplazium caudatum (Cav.) Jermy, both of which later authorsreport as growing close to the Culcita in this area. This omission led Nieschalk and Nieschalk(1965) to suspect that the Culcita site they subsequently discovered in the Miel Valley mightnot be the same as Allorge's, as they foundT. speciosum to be a frequent associate and indeedto be scattered elsewhere by the streambed lower in the valley. Contrary to the statement inSalvo Tierra ( 1982), this was not the first discovery of T. speciosum in this region, thatdistinction falling to the distinguished tropical botanist and bryologist P.W. Richards(Richards, 1934). He had previously drawn attention to the Atlantic bryophyte interest of the

FERN GAZE'ITE: VOLUME 15 PART 6 (1998)198"wateifall valley"[ Miel Valley] which he first visited in the spring of 1931 (Richards, 1932).Trichomanes was discovered on a return visit in April 1934 , growing "very sparingly on wetsandstone rocks in a small cave near a wateifall. He makes no mention of Culcita in his shortaccount. It seems most unlikely that this area should have been visited independently byRichards and Allorge. The date of Allorge's visit is not given in his account (Allorge, 1934) butboth he and Richards collected specimens of the Macaronesian moss Lepidopilum virens Card.(as Plagiothecium argyrophyllum Geh.) here in April of that year. These were the only knownfinds of this species on the European mainland until its recent rediscovery in this area byHakelier (Sergio et al., 1995) and it seems probable, although not stated anywhere, that the twobotanised together.Richards' (1934) report and the oblique reference in Nieschalk and Nieschalk (1965) wereoverlooked by many, including Molesworth-Allen ( 197 1). It was this paper, in which shereported T speciosum from two unspecified localities, that brought to wider attention thepresence not just of Trichomanes but also other pteridophytes with Macaronesian distributions,namely Diplazium caudatum, Pteris serrulata Forsk!\1 ( P. incompleta Cav.) and Cyclosorusdentatus (Forsk!\ 1) Ching ( Christella dentata (Forskal) Brownsey & Jermy), in Cadizprovince. Her careful researches in this region over the next few years revealed further sites forT speciosum in the Algeciras, Los Barrios and Tarifa districts, i.e. on the Sierra de Luna, Sierradel Nifio and Sierra de Ojen ranges (Molesworth-Allen, 1977). Her investigations theneffectively defined the species' known range, with no subsequent published repotts of additionallocalities, although the fern may additionally have occurred a little further west, on the Sierra deSaladavieja, the lower, westward continuation of the Ojen (Salvo, pers. comm.). It was searchedfor in this area during the 1995 Group of European Pteridologists (G.E.P.) excursion, butwithout success.The distribution of Trichomanes speciosum was mapped by Salvo Tierra ( 1982) in hiscomprehensive study of the pteridophyte flora of Andalucia, in two hectads, 60TFOO and70TFOO . The total number of sites and their location are not given in the published accounts(Molesworth-Allen, 197 1, 1977), which understandably sought to protect this species fromover-collection, a fate which had befallen another rare disjunct pteridophyte discovered by thesame author in this area, Psilotum nudum (L.) Beauv. (Molesworth-Allen, 1966). While sheregarded T speciosum as "locally abundant in a number ofarroyos" (Moles worth-Alien, 1977),the three areas from which she then knew the plant supported only "40-50 plants or patches",of which few had been fertile."AIMSTwo decades have passed since the above records from this phytogeographically important area,during which time patterns of land use have changed, levels of damaging human activity haveincreased, and higher local pollution loads and increasing aridity may have all affected thesevulnerable, unique, refugial areas. The distinct species assemblage which characterises theAtlantic fringe of Europe and Macaronesia is particularly rich in cryptogamic species, many ofwhich are rare and endangered. Since 1994 research has been carried out at the Natural HistoryMuseum into the distribution, ecology and historical biogeography of this community and itsconstituents. As part of this wider study these isolated sites in southern Spain were visited todetermine the current extent of both generations of Trichomanes speciosum, a model organismwhich had been well studied elsewhere (see Rumsey et al., 1998 , in press).MATERIALS AND METHODSWhere possible all previously reported sites of T speciosum were visited. In addition, sitesknown to suppor t, or to have supported, other disjunct Macaronesian elements were visited toascertain whether the sporophyte, or the less conspicuous gametophyte generation was present.

TRICHOMANES SPECIOSUM IN SOUTHERN SPAIN199We were fortunate to have access to the field notes made by Mrs Molesworth-Allen and her helpand encouragement in the field.RESULTSTrichomanes speciosum was refound in all of the areas from which it had previously beenreported. However, the species is now present only as the gametophy te generation in the Sierradel Nifio, and on the eastern slope of the Sierra de Luna. One new site, well to the north of theprevious discoveries , was detected while investigating the distribution of other Macaronesiantaxa. Thus, we were fortunate to be guided to a recently discovered site for Culcita macrocarpa,an arroyo draining into the Embalse del Guadan·anque on the N.E. side of the Sierra deMontecoche. The finder, Luis Federico Sanchez Tundidor, accompanied by Mrs .MolesworthAlien, a friend, and the authors were delighted to discover that the damp rock ledges by animpressive waterfall that supported the population of c. 15-20 Culcita plants also had extensivemats ofT. speciosum gametophytes (Fig. 1). This extends the known range of both these speciesby c. 13km to the north-east (the nearest known sites for each are on the Siena del Nifio). In thearea of the Siena del Nifoi , whereT. speciosum had previously been found, above La Granja,only a single site with a very small quantity of gametophy te was discovered in 1997 . The habitatappeared generally unsuitable and had been further disturbed by a serious land-slip the previouswinter. Only a single sporophyte colony had ever been known to Mrs Molesworth-Allen in thisarea (Moles worth-Allen, pers . comm.). No gametophytes were found in the extant site forC. macrocarpa elsewhere in the Siena del Nifio. Very small quantities of gametophy te were alsofound in a small streambed above the minor road between Tajo del Espino and Puerto de laZarza, on the east side of the Siena de Luna, high above the head of the Miel Valley at c. 550 maltitude. A single large plant of C. macrocarpa grew by a small waterfall nearby.Figure 1. Site of Trichomanes speciosum gametophyte population in the Siena de Montecoche,May 1997. (Photo F.J. Rumsey)

200FERN GAZETTE: VOLUME 15 PART 6 (1998)Figure 2. Trichomanes speciosum flourishing by small streams on the Dehesa de Ojen, May1997 .Also presentAthyriumfilix-femina (L.) Roth and, middle right, Pteris incomp!eta Cav.(Photo J.C. Vogel)

TRICHOMANES SPECIOSUM IN SOUTHERN SPAIN201Figure 3a. Distribution of Trichomanes speciosum Willd. in southern Spain as revealed by fieldsurvey (1995- 1997). Both generations recorded , Gametophyte generation only o. The 500 mcontour line is included.3b. The distribution within the SietTa de Ojen, Sierra de Luna and Sierra del Nifio mapped on al km grid basis. Contour lines at lOOm intervals.

202FERN GAZETTE: VOLUME 1 5 PART 6 ( 1998)In the Miel Valley only a single sporophytic plant of Trichomanes speciosum was seen in1995 and refound in 1997. This was growing adjacent to the only population of Culcitamacrocarpa seen on these visits. No Diplazium caudatum or Pteris incompleta were seen in thevalley in either year. The only area within which T. speciosum had maintained sporophyticnumbers was the northern flank of the Sierra de Ojen, the Dehesa de Ojen up to the ColladoPuerto del Viento on the S . E . flank of the Sierra de Luna. This population was apparentlythriving; many fertile plants were seen and juvenile sporophytes were common on gametophytemats (Fig. 2). The extent and condition of the sporophytes in this area would appear to dependon the condition of the mixed evergreen-deciduous woodland in which they occur, itselfdetermined by activities such as cork-cutting and, more harmfully, clearance or burning. Indeed,Molesworth-Allen (pers. comm.) had noted that in a previous visit (post her optimistic 1977account) few sporophytes could then be found in this area. Diplazium caudatum, Pterisincompleta and Polystichum setiferum (Forskal) Woynar were all seen in reasonable quantityand health, the last, in number and location almost exactly as reported by Molesworth-Allen( 1977). Of concern was the apparent absence of C. macrocarpa, the loss of which could onlyrealistically be explained by its selective removal. The terrain is however, difficult to search andthis species and other interesting finds might still be made in the more remote areas.The distribution of Trichomanes speciosum in southern Spain as revealed by our survey isgiven in Figure 3 .DISCUSSIONThe detection of the gametophyte generation of T. speciosum was to be expected given thefrequence of its occurrence in more northerly and continental areas of Europe (Vogel et al.,1993; Rumsey et al., 1998). What is, perhaps, surprising is that more sites were not locatedfringing those that currently maintain both generations. Evidence from elsewhere and the findsreported here from high on the Sierra de Luna and in the Sierra del Niiio, demonstrate the abilityof the gametophyte to persist in areas which once supported the sporophyte. The lack ofadditional gametophyte sites must reflect the general unsuitability of the habitat for bothgenerations; harsh environmental and climatic changes occur over such short distances that littlehabitat exists that could be considered suitable for gametophytes outside of the very restrictedareas suitable for sporophyte growth .W hen compared with earlier accounts (e.g. Nieschalk & Nieschalk, 1965; Molesworth Allen, 197 1, 1977) the population present in the best known and most accessible site, the MielValley, has suffered a severe and perhaps catastrophic decline. Trichomanes speciosum is not theonly taxon to have suffered in this way. Diplazium caudatum, first reported on the Europeanmainland from this site in 19 69 (Molesworth-Allen, 197 1 ) and suggested to have suffered fromcollection (Molesworth-Allen, 1977), may now be absent from the valley. Culcita macrocarpahas vanished from the more accessible of its past sites lower in the valley, but persists higher upwhere most casual visitors do not reach . Increased rambling and other outdoor pursuits, openingup of the undergrowth, felling in the wider area, higher incidence of fires, when coupled with arun of drier than average years is steadily reducing the conservation interest of this area. Onlythe inaccessibility of the still comparatively unspoilt and now unique vegetation on the Dehesade Ojen has acted to protect it. These very restricted woodland areas with their Tertiary relictMacaronesian cryptogamic flora, the sole Iberian sites for a range of disjunct flowering plants,e.g. Arisarum proboscideum (L.) Sa vi, are a national treasure and of international significance.Their active conservation must be addressed.ACKNOWLEDGEMENTSWe would like to thank the NERC who funded this research under grant GR3/0945 1 . We alsomust record our deep gratitude to Mrs Betty Molesworth-Allen for information, arrangingaccess with landowners, spritely company in the field and her optimistic navigation on the

TRICHOMANES SPECIOSUM IN SOUTHERN SPAIN203"road to Hell"! We also thank Fede Tundidor for sharing with us his exciting discovery of thenew Culcita locality.REFERENCESALLEN, D.E. 1969 . The Victorian fern craze. Hutchinson, London.ALLORGE, P. 1934 . Le Culcita macrocarpa Presl [Balantium Culcita (L' Herit.) Kauffm.] dansles montagnes d' Algesiras. Bull. Soc. Bot. France 81: 59 2-593 .ANON. 1979 . The Convention on the Conservation ofEuropean Wildlife and Natural Habitats.Council of Europe, Strasbourg.ANON. 1992. Council directive 9 2/43/EEC of 21 May 1992 on the conservation of naturalhabitats and of wild faunas and floras. Official journal of the European Communities 206:7-49.FARRAR, D.R. 1967. Gametophytes of four tropical fern genera reproducing independently oftheir gametophytes in the southern Appalachians. Science 155: 1266-1267.FARRAR, D.R. 1985 . Independent fern gametophytes in the wild. Proc. R. Soc. Edinb. 86B:3 61-369 .FARRAR, D.R. 1992. Trichomanes intricatum: the independent Trichomanes gametophyte inthe Eastern United States. Amer. Fern J. 82: 68-74 .FRASER- JENKINS, C.R. 1982. Dryopteris in Spain, Portugal and Macaronesia. Bol. Soc.Brot., Ser. 2, 55: 175-336.GOEBEL, K. 1888 . Morphologische und biologische Studien: 11. Zur Keimungsgeschichteeiniger Farne. Ann. lard. Buitenz. 7: 74- 1 19 .MOLESW ORTH-ALLEN, B . 1966. Psilotum nudum in Europe. Brit. Fern Gaz. 9: 249-25 1 .MOLESW ORTH-ALLEN, B . 197 1 . Observations on Spanish ferns. Brit. Fern Gaz. 10: 20020 2.MOLESW ORTH-ALLEN, B . 1977. Observations on some rare Spanish ferns in Cadizprovince, Spain. Fern Gaz. 11: 271-275 .NIESCHALK, A. & NIESCHALK, C. 1965 . Ein Stammfarn (Culcita macrocarpa) aufeuropaischem Boden. Natur und Museum 95: 495-498.RICHARDS, P.W. 193 2. Note on the bryophytes of the "Waterfall Valley" near Algeciras: anoutpost of the Atlantic flora. Rev. Bryol. et Lichen. 5: 1-9.RICHARDS, P.W. 1934 . Trichomanes radicans Sw. in the south of Spain. J. Bot. (London) 72:35 1 .RUMSEY, F.J., SHEFFIELD, E . & FARRAR, D . R . 1990 . British filmy-fern gametophytes.Pteridol. 2: 40-4 2.RUMSEY, F.J . , JERMY, A.C. & SHEFFIELD, E. 1998 . The independent gametophytic stage ofTrichomanes speciosum W illd. (Hymenophyllaceae), the Killarney Fern and its distributionin the British Isles. Watsonia 22: 1- 19 .RUMSEY, F.J., VOGEL, J.C., RUSSELL, S.J., BARRETT, J.A. & GIBBY, M . in press.Climate, colonisation and celibacy: population structure in Central European Trichomanesspeciosum (Pteridophyta). Bot. Acta.SALVO TIERRA, A.E. 1982. Flora pteridofitica de Andalucia. Unpublished PhD. Thesis,Universidad de Malaga, Spain.SERGIO, C., BRUGUE S, M. & CROS, R.M. 1995 . Threatened bryophytes in the IberianPeninsula - Some results and comments. Crypt. Helv. 18: 5 1-56.TUTIN, T.G., BURGES, N.A., CRATER, A.O., EDMONDSON, J.R., HEYWOOD, V.H.,MOORE, D.M., VALENTINE, D.H., WALTERS, S.M. & WEBB, D.A. 1993 . FloraEuropaea Vol. 1, 2nd ed. Cambridge University Press, Cambridge.VOGEL, J.C., JESSEN, S., GIBBY, M., JERMY, A.C. & ELLIS, L. 1993 . Gametophytes ofTrichomanes speciosum (Hymenophyllaceae: Pteridophyta) in Central Europe. Fern Gaz.14: 227-232.WALTER, K.S. & GILLETT, H.J. 1998 . IUCN Red List of threatened plants of the world. NewYork Botanical Garden, New York.

204FERN GAZETTE: VOLUME 1 5 PART 6 ( 1 998)BOOK REVIEWTHEFERNS OF BRITAIN AND IRELAND. C.N. Page. 1997 (2nd edition).Cambridge University Press, Cambridge. Hardback 95 , ISBN 0 52 1 58380 2; paperback 40 ,ISBN 0 52 1 58658 5 .The British Isles are not only blessed with a rich fern flora, they are also blessed with a wealthof books about pteridophytes. One of the most outstanding and innovative contributions hasbeen the first edition of The ferns of Britain and Ireland by Dr C.N. Page, published in 198 2 .This book has now appeared i n its second editi on, but at a considerably increased price. At sucha price, Page has to stand the test against the standard in European fern Floras, the IllustrierteFlora fiir Mitteleuropa, in short 'Hegi', published in its third edition by Prof. K. Kramer in1984 .Page starts h i s book with a general introduction t o ferns/fem allies and the factors,geography, soils and climate, which influence the distribution and occurrence of pteridophytesin the British Isles. Fern

Tl'lli H::.GAZE'ITE Volum RN 15 Part5 wus pu ilihcd tln I 'ilh Dcccmb1Jr. 1997 PUblbhed by Tl m BRTTISJI Pl'f:RJDOl.OGICAL SOCIETY, c/u Ocl'llrllllcnt of BOtnny. The Natural Hiswry Museum, London SW7 5BD ISSN 0308-0838 Printed by Metloc Printers L Caxton House, Old Station Road, Loughton, Essex IG 10 4PE

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