THE FERN GAZETTE Eomifi in Chief: M. PsoiHJCTtON Editor: A. Eeoaanl llEviEW Editors: A. F. l^er M B, THE FERN GAZETTE is a journal of the British Pteridological Society and contains peer-reviewed papers on all aspects of pteridology. Manuscripts may be submitted, and books etc. sent for review, to: Prof. M. Gibby, Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 SLR, UK, FemGazette@eBPS.org.uk The Fern Gazette Editorial Board 2013-2016 The purpose of the Editorial Board is to widen the range of disciplines within pteridology that are covered in the Fern Gazette, and to widen its geographic focus. The role of Editorial Board members is two-fold, first to help in reviewing manuscripts in their particular area of expertise and, second to solicit papers for publication among their local contacts. Members of the Editorial Board are asked to serve for three years initially. The Review Editors commission review articles for the Fern Gazette. The editorial team of the Fern Gazette comprises the following people: Editor in Chief Mary Gibby, Royal Botanic Garden Edinburgh, Edinburgh, UK Editorial Board Wen-Liang Chiou, Taiwan Forestry Research Institute, Taipei, Taiwan Stephen Jury, Lei^ton Buzzard, UK Klaus Mebltreter, Institute de Ecologia, A.C., Xalapa, Mexico Barbara Parris, Fern Research Foundation. Kerikeri, New Zealand Jefferson Prado, Institute de Botanica, Sao Paulo, Brasil Fred Rumsey, Angela Marmont Centre, NHM. London, UK Harald Schneider, Life Sciences IJepartment, NHM, London, UK Proctaction Editor Andrew Leonard, Portsmouth, UK Review Editors Adrian Dyer, afdyer499@googlemail.com Bridget I Instructions for authors are on page 240 of this volume and also available at htip://www.eBPS.org.uk Copyright © 2013 British Pteritte logical Society. All rights reserved. No part of this publication may be reproduced in any material form (including photocopying or storing it in any medium by electronic means) without the permission of the British Pteridological Society. THE FERN GAZETTE Volume 19 Part 5 was publidied on 1 8th October 2013 Published by THE BRITISH PTERIDOLOGICAL SOCIETY o'o Department of Life Sciences, The Natural Histtny Museum, Cromwell Road, London SW7 5BD, UK http:/7www.bishops.co. uk FERN GAZ. 19(6): 193-202. 2013 193 CAUSES AND CONSEQUENCES OF THE VARIABILITY OF LEAF LIFESPAN OF FERNS K. mehltreter’ & j. M. SHARPS MISSOURI BOTANICAL ' Institute de Ecologia, A.C., Xalapa, Mexico Email: kmehltreter@gmail.com Sharplex Services, Edgecomb, Maine, USA Email: joannesharpe@email.com JAN 2 ? 20U GARDEN LIBRARY Key words: deciduousness, indeterminate growth, leaf dimorphism, leaf traits, phenology, seasonality. ABSTRACT Leaf lifespan (LLS) is a fundamental ecological trait of special importance to growth and survival of ferns. The LLS of ferns varies between two and 49 months depending on species and site conditions. Whereas temperate ferns are mainly summergreen, tropical ferns are mostly evergreen. However, some tropical ferns have seasonal or even deciduous leaf phenologies or dimorphism with shorter- lived fertile leaves that are produced only during a specific time of the year. The median LLS of tropical ferns of 19.2 months is nearly twice as long as in seed plants ( 1 0.0 months). Possible correlates of leaf lifespan such as leaf mass per area, nutrient content and herbivore damage, and future research questions are INTRODUCTION Leaves are the most conspicuous parts of ferns, with the exception of tree ferns that also have tmnks of up to 15 m height. Ferns have developed leaves of all sizes, from large 4- 5 m long leaves (e.g. Angiopteris evecta (G Forst.) Hoffm.) to the smallest leaves of 1-3 mm length of Azolla filiculoides Lam. and some species of filmy ferns such as Didymoglossum ovale E. Foum. Because of the importance of leaves as “energy transformers”, their number, size and functional characteristics such as lifespan are of special importance to growth and survival of plants. Whereas the number of leaves, their size and other morphological traits can be easily measured at any time, the direct measurement of leaf lifespan (LLS) requires their entire ontogeny to be followed from the first appearance of a developing crozier through expansion, maturation and death (though not necessarily detachment). Some authors measure the LLS as the time from the moment the leaf is fully expanded to leaf death (Kikuzawa & Lechowicz, 201 1), a definition which would result in estimates of LLS which are from two to eight weeks shorter because of the time needed for the crozier to complete leaf expansion. Leaf lifespan can also be estimated by dividing the mean leaf number by the annual leaf production (see Mehltreter, 2008). Both direct measurement and estimation require several observations throughout at least one year and consequently few data are available on fern LLS. For example, Wright et al. (2004) included 13 fern species in a dataset of 2548 vascular plant species, but only two of those fem species contained data on LLS. For the current review, we compiled a dataset of 71 fern species, which is still less than one percent of the currently known 9500 fem species (excl. lycophytes. Smith, 2010). 194 FERN GAZ. 19(6): 193-202. 2013 The LLS of ferns varies in temperate climates as well as in tropical climates, and species of different LLS may coexist at the same sites (Shiodera et al 2010). Consequently, we asked; How much does LLS vary in ferns? Does mean LLS differ between ferns and angiosperms? How does LLS vary between species, sites and within the geographic range of a single species? Which leaf traits are correlated with LLS? VARIATION IN FERN LEAF LIFESPANS Leaves of the shortest lifespan may belong to some species of grape ferns (Botrychium), which may persist for 1.8 months (B. gallicomontanum Farrar & Johnson-Groh) to 2.8 months (B. mormo W.H. Wagner) in Minnesota, U.S.A. (Johnson-Groh & Lee, 2002), whereas the longest mean LLS of 4.9 years was reported for fertile leaves of Terpsichore asplenifolia (L.) A.R. Sm. (Monge Gonzalez, 2007), a Mexican epiphyte with overhanging leaves of indeterminate growth. Leaves of indeterminate growth have a very long potential LLS, supposedly unlimited by intrinsic factors (e.g. plant hormones), but die mainly as a consequence of abiotic factors (e.g., lack of water, mechanical damage) or biotic interactions (e.g., herbivores, fungal infections). Leaves of smaller, younger plants of T. asplenifolia are still of determinate growth, remain sterile and are shorter lived than the fertile leaves of larger plants. As a consequence of their indeterminate growth, leaves of T. asplenifolia are capable of continuous spore production because they possess all stages of sporangia, old sporangia which already have released their spores near the leaf base, fully developed sporangia with mature spores in the mid-seetion of the leaf and immature new sporangia near the leaf tip. The LLS can differ considerably for species that occur over a wide geographical range. Botrychium virginianum (L.) Sw. occurs from Canada to Pern under distinct climatic conditions and growth seasons of different duration. In North America, leaves of B. virginianum appear in early spring and die in late summer, a period of about four months (Wagner & Wagner, 1993). However, in Mexico the LLS of B. virginianum is 1 7. 1 months, and thus a new leaf may coexist for several months together with the leaf of the year before. On the other hand, Acrostichum danaeifolium Langsd. & Fisch., one of the few species that has been studied at different sites, maintained a similar LLS in Mexico (7.7. months; Mehltreter & Palacios-Rios, 2003) and Puerto Rico (9.6 months; Sharpe, 2010). The median LLS for our dataset of 71 fem species was 15.6 months (range 1.8-48.9 months), much longer than the median LLS of 8.5 months for seed plants (range 1.1- 288 months, Wright et al, 2004). However, it is difficult to assess the validity of such a comparison because it could be biased by factors such as species selection, disturbance regimes, phylogenetic constraints, or geographic location. In our dataset, 91.5% of fem species came from tropical areas, where we expect LLS to be longer. Restricting the analysis to tropical species, the median LLS for ferns, 19.2 months (first quartile = 11 months, third quartile = 24.6 months, n = 66), is still significantly longer than that for angiosperms, 10.0 months (first quartile = 7.0 months, third quartile =19.1 months, n = 163, Mann-Whimey U = 3633, P < 0.001). No significant differences of LLS among four of the five tropical fem floras could be detected, but ferns of Puerto Rico had a significantly longer LLS than ferns of Hawaii and Mexico (F = 3.71, P = 0.009, Table 1). Further studies are needed to determine whether these differences are consequence of the selection of sampled species we used or if they represent a real contrast among fem MEHLTRETER & SHARPE : VARIABILITY OF LEAF LIFESPAN OF FERNS 1 95 : Sterile LLS of fer (in months) at different 1 sites, S.E. = Standard Site Mean LLS ± 1 S.E. 1 Minimum LLS Maximum LLS * Hawaii (n = 14) 15.0 ±1.75 5.0 ^ 24.6 Jamaica (n = 3) 17.3 ±3.76 11.0 24.0 Puerto Rico (n = 14) | 26.5 ± 3.44 8.4 48.9 Mexico (n= 17) 16.4 ±1.89 , 5.5 32.0 1 Taiwan (n = 18) | 17.5 ±1.82 5.1 30.3 TEMPERATE FERNS There are three general LLS categories for temperate ferns: summergreen, wintergreen and evergreen. Summergreen ferns produce a flush of new leaves in spring (e.g. Adiantum pedatum L., Matteuccia stnithiopteris (L.) Tod.) that die at the end of the summer growing season. In contrast, wintergreen species retain some green leaves through the winter until the following spring or summer, just before or after a new set of leaves begins to develop. The leaves of evergreen ferns are not produced in a seasonal flush, but are replaced as older leaves die and consequently the plants are never leafless. Although leaves of most evergreen species live for more than a year, LLS may still differ considerably among species. Phenograms for much of the temperate fern flora of Britain and Ireland have been published (Page 1997) and from them we estimated LLS by counting the months from leaf maturity (ignoring expansion phase) to full senescence. Using terminology defined by Lellinger (2002) we then classified the taxa in this flora as 27 summergreen, 9 wintergreen and 1 1 evergreen species with mean LLS of approximately five, 1 1 and 16 months, respectively. In the British flora, LLS varies between four months {Botry chium lunaria (L.) Sw.) and 25 months {Polystichum lonchitis (L.) Roth). Combining the 13 phenological types identified by Sato & Sakai (1980) for mature leaves into the same three categories of Lellinger (2002) results in a temperate fem flora for Hokkaido, Japan that has 42 summergreen, 1 1 wintergreen and 15 evergreen species, with the high ratio of summergreen to evergreen species at high latitudes contrasting with a very low ratio at lower latitudes in Japan. Sato (1982) further documents the correspondence between frost tolerance and LLS category, with few frost tolerant leaves found among the summergreen species. In the flora of one area of Japan (Yoshida & Takasu, 1 993) mean LLS of 21 species ranges from five months {Deparia pycnosora var. alvosquamata M. Kato) to 37 months (Asplenium wrightii D.C. Eaton ex Hook.). Whereas summergreen ferns store all their nutrients and energy in the rhizome or roots during the winter and then reinvest in new leaves during the following season, wintergreen and evergreen species expose their leaves to cold temperatures and even 196 FERN GAZ. 19(6): 193-202. 2013 frost, but still benefit from photosynthetic gains during mild winter days (e.g., Dryopteris filix-mas (L.) Schott; Bauer etal., 1991). Because longer living leaves of wintergreen and evergreen ferns have to survive a wider variation of environmental conditions than shorter-lived leaves of summergreen ferns, their longer LLS are often correlated with robust leaf structure and greater leaf mass per area (LMA, see below). TROPICAL FERNS Ferns of tropical habitats are often evergreen due to the lack of seasonality in temperature and rainfall. Flowever ferns in tropical as well as subtropical areas that undergo dry seasons of several months may be deciduous because drought may cause stress to the leaves resulting in leaf drop patterns similar to those caused by cold temperatures in temperate areas. Some epiphytic fern species are also deciduous and sprout new leaves at the beginning of the rainy season (e.g., Phlebodium areolatum (Flumb. & Bonpl. ex Wind.) J. Sm., Polypodium rhodopleuron Kunze). Such epiphytes may be even more exposed to water stress because they are rooted in shallower substrate than terrestrial ferns. Other species, such as the Mexican tree femAlsophila firma (Baker) D.S. Conant which grows in the understory of montane forests, are not exposed to severe drought stress at their natural sites (Mehltreter & Garcia-Franco 2008). However shorter LLS (Chabot and Hicks, 1982) and deciduousness followed by synchronous leaf flush have been reported to reduce herbivore damage (Lieberman & Lieberman, 1984; Karban, 2007). Consequently, A. firma might be deciduous during the wet season when herbivorous insects thrive and develop all new leaves synchronously two to three months after leaf fall. Water is never scarce in mangrove forests but plants have to deal with increased salinity of soil water. The mangrove fern Acrostichum danaeifolium Langsd. & Fisch does not possess any specific mechanism for eliminating sodium such as the salt glands of other mangrove species. Consequently, sodium is accumulated in the leaves and is eliminated only when leaves die, perhaps accounting for the LLS of 7-10 months, relatively short for a tropical fern (Mehltreter & Palacios-Rios 2003, Sharpe 2010). Indeterminate leaves of climbing ferns such as Lygodium venustum Sw. suffer from disturbances such as branch falls and their measured LLS of 5.6 months might be far below their potential LLS without external disturbance factors (Mehltreter 2006). POSSIBLE CORRELATES OF LEAF LIFESPAN In the following, we discuss several general hypotheses that have been suggested as correlations between LLS and other plant characteristics. These hypotheses are often based on knowledge of angiosperms, and we relate it to our dataset of ferns. A longer LLS increases the time availa ble for photosvnthetic gains, but requires more drought. According to this hypothesis, evergreen leaves have a higher leaf mass per area (LMA) than deciduous leaves (see Kikuzawa and Lechowicz, 2011). For eight fern species of our data set for which LLS and LMA are available, this correlation is significant (Pearson correlation coefficient r = 0.75, P = 0.03, Figure, la). In addition, LMA is negatively correlated with nitrogen (N) content (r = -0.81, P = 0.01). Greater environmental tolerance of longer-lived leaves has a tradeoff resulting in lower photosvnthetic rates and lower N concentrations . Available data for ferns are not significant (both P = 0.11, Figure lb and Ic). Most ferns have generally lower photosynthetic rates than angiosperms (see Mehltreter, 2010), supposedly because of their lower water transport efficiency and lesser control of water transpiration (Orians &. MEHLTRETER & SHARPE; VARIABILITY OF LEAF LIFESPAN OF FERNS 1 97 Figure 1. Correlation between sterile LLS and a) LMA b) Net C02-assimilation rate and c) N content in tropical ferns. Data from Baruch & Goldstein 1999^^, Chiu et al. 2013^, Durand & Goldstein 200 1^^^, Funk & Amatangelo 201 3^, Tanner 1983®^, Winkler et al. 2005‘^. MEHLTRETER & SHARPE: VARIABILITY OF LEAF LIFESPAN OF FERNS 1 99 dead leaves continue to ftinction as humus accumulator. In other fern species, dead leaves possess such a rigid petiole and rachis that they still work as a protective shield for adjacent living leaves against wind, high irradiance or frost such as in the xerophytic species of Cheilanthes or the paramo fern genus Jamesonia. In the climbing fern genus Lygodium, the rachises of dead leaves are the main climbing structure for young leaves to reach the canopy (Mehltreter, 2006). Fern species with faster growth rates have a shorter LLS . The faster growing fern species develop in sunny, open habitats and can produce more leaves in a year than shade tolerant species. Siman & Sheffield (2002) have shown in greenhouse experiments with Polypodium vulgare that leaf production rates increase with increased light and temperature. Leaves of sun species have shorter LLS than shade species, and sun leaves have a shorter LLS than shade leaves of the same individual (Reich et al., 2004; Vincent, 2006). In Hawaii the introduced Australian tree fern Sphaeropteris cooperi (F. Muell.) R.M. Tryon has shorter LLS than the native, shade tolerant species of Cibotium (Durand & Goldstein, 2001). Because of its fast growth, S. cooperi invades and outcompetes the native Hawaiian tree ferns (Medeiros et al, 1992; Chau et al, 2013). months) but relatively short-lived fertile leaves (4.6 months). Lines indicate linear regressions for monomorphic and dimorphic species. 200 FERN GAZ. 19(6): 193-202. 2013 CHALLENGES FOR THE FUTURE We need to establish a larger and more consistent database of ecological life histories of ferns that includes, at a minimum, leaf characteristics such as LLS, LMA, nutrient content and photosynthetic rates and then to detect functional consequences of these traits to allow for more profound analyses of the role of ferns in an ecosystem. Some data on each of these characteristics exist, yet are scattered throughout the literature. For example, Watkins et al. (2010) took photosynthetic measurements on 21 species of tropical ferns but summarized the results by habitat and did not include LLS in their analyses. Karst & Lechowicz (2007) compare fem foliar traits with those of seed plants, and include LMA and photosynthetic rates but not LLS, and base their fem conclusions on the very limited data from Wright etal. (2004). Measurements of a single species over a wide geographic range could reveal the limits on adaptability of various leaf traits in changing environments. Experimental greenhouse studies could be used to investigate which factors such as nutrient concentrations may be correlated with longer LLS. Physiological experiments are also required to better understand which factors trigger the ontogenetic development into sterile or fertile leaves, especially in dimorphic fem species. Long- term studies over decades may allow the identification of responses to global climate change and should be established in protected sites such as those within the International Long Term Ecological Research network. ACKNOWLEDGEMENTS We thank Tom Ranker, Adrian Dyer and Paulo Windisch for helpful comments on the REFERENCES BAUER, H.C., GALLMETZER, C. & SATO, T. 1991. Phenology and photosynthetic activity in sterile and fertile sporophytes of Dn>opteris filix-mas (L.) Schott. Oecologia 86: 159-162. BARUCH, Z. & GOLDSTEIN. G 1999. Leaf constmction cost, nutrient concentration, and net CO2 assimilation of native and invasive species in HawaiT. Oecologia 121: 183-192. BRODRIBB, T.J., HOLBROOK, N.M., ZWIENICKII, M.A. & PALMA, B. 2005. Leaf hydraulic capacity in ferns, conifers and angiosperms: impacts on photosynthetic maxima. New Phytologist 165: 839-846. CHABOT. B.F. & HICKS, D.J. 1982. The ecology of leaf life spans. Annual Review of Ecology and Systematics 13: 229-259. CHAU, M., WALKER. L.R. & MEHLTRETER, K. 2013. An invasive tree fem alters soil and plant nutrient dynamics in Hawaii. Biological Invasions 15- 355-370 CHIOU, W.L.. LIN, J.C. & WANG J-Y. 2001. Phenology of Cibotium taiwanense (Dicksoniaceae). Taiwan Journal of Forestry Science 16: 209-215. CHIU, T.Y., WANG H.H., KUO, YL. & KUME, T. 2013. Effect of habitat preference on frond life span in three Cyathea tree ferns. Geophysical Research Abstracts, 1 5. COLEY P.D. 1980. Effects of leaf age and plant life history patterns on herbivory. Nature 284: 545-546. COOKE, J., & LEISHMAN, M.R. 2011. Silicon concentration and leaf longevity: is silicon a player in the leaf dry mass spectrum? Functional Ecology 25: 1181-11 88. DURAND, L.Z. & GOLDSTEIN, G 2001. Growth, leaf characteristics, and spore production in native and invasive tree ferns in Hawaii. American Fem Journal 91 : 25-35. MEHLTRETER & SHARPE: VARIABILITY OF LEAF LIFESPAN OF FERNS 2C FUNK, J.L., & AMATANGELO, K.L. 2013. 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Herbivory in epiphytic bromeliads, orchids and ferns in a Mexican montane forest. Journal of Tropical Ecology 21: 147-154. WRIGHT. I.J., REICH, P.B., WESTOBY, M., ACKERLY, D.D., BARUCH, Z., BONGERS, F., CAVENDER-BARES, J., CHAPfN, F.S., CORNELISSEN, J.H.C., DIEMER, M., FLEXAS, J., GARNIER, E., GROOM, P.K., GULIAS, J., HIKOSAKA, K., LAMONT, B.B., LEE, T, LEE, W., LUSK, C., MIDGLEY, J.J., NAVAS, M.-L., NIINEMETS, U., OLEKSYN, J., OSADA, N., POORTER, H., POOT P PRIOR L , PYANKOV V.I., ROUMET, C., THOMAS, S.C., TJOELKER, M.G, VENEKLAAS, E. & VILLAR, R. 2004. The world-wide leaf economics spectrum. Nature 428: 821-827. YOSHIDA, C. & H. TAKASU. 1993. Leaf life-span in some ferns of the Kii Peninsula 1. Acta Phytotaxonomica et Geobotanica 44: 59-66. 203 FERNGAZ. 19(6). 2013 BIOGRAPHY - JOANNE M. SHARPE FERN GAZ. 19(6). 2013 204 205 FERN GAZ. 19(6). 2013 BOOK REVIEW Who found our ferns? John Edgington. 2013. British Pteridological Society Special Publication No. 12 ISBN 978-0-9926120-1-6 216pp. £15. 245 x ITOmm. This meticulously compiled book has several different strands to document the history of discovery of each species of fern and Lycophyte recorded from Britain and Ireland. The chapters are ordered using modem nomenclature and the species are shown subdivided by Classes, Orders and Families. A short account is given at the beginning of the chapter for each Family that includes any fossil evidence and a summary of the present knowledge of fem classification. Early identifications were of necessity based largely on gross morphology. By using molecular and chromosome information we now have access to many more resources than were available to the pioneer taxonomists, and the increase in species numbers within the flora in the twentieth century has resulted from recognition of species complexes in genera like Polypodium or Dryopteris. The author shows that relating current taxonomic concepts to early records can be quite a challenge. The main part of the text chronicles a complicated search through many references and early herbarium records to unravel exactly which species is being described, sometimes from rather limited or inaccurate descriptions. An added complication in deciding which taxon was intended is that species which were abundant in the past might have a more restricted distribution now. The first record from each of the countries within Britain is given. The whole combines to provide a history of botanical recording and the developing understanding of plant classification for ferns and Lycophytes. Before Linnaeus formalised the names of plants people used a descriptive Latin phrase which was considerably more cumbersome than the binomial names that evolved Two varieties of Asplenium scolopendrium, treated as separate species (Matthieu de L’Obet, 1581) FERNGAZ. 19(6). 2013 206 into the names that are now use. Each chapter describes not simply a possible first record, but the first record that is unambiguously the taxon which we currently recognise by the stated name. In the midst of the confusion over early classification and multiple applications of the same plant names, one can sympathise with the difficulties and complications people encountered as they struggled to make sense of different species or varieties. In the search for early records of British species it was necessary to span across Europe to compare descriptions from the botanical community that were being applied within Britain. In interpreting the taxa it is sometimes necessary to go back to the names given to the plants in classical times that come to us as Greek or Latin words, some of which are still used as part of the scientific name. At the end of the book is a very useful summary of early names, synonyms and Latin phrase names showing how similar descriptive words have been used across so many species and even families that we now know were un-related. Individual species accounts show the difficulties botanist encountered in attempting to describe structures that were not understood. Isoetes, for example, like many other species were first thought to be flowering plants strangely lacking both in flowers and seed. Further confusion was shown in the idea that the fertile cones of Equisetum arvense were a different species from the vegetative shoots. The situation improved with the understanding the of the fern life cycle in the late 18^h century, but it was hardly surprising that Pilularia globulifera was not initially recognised as a fern. It was interesting to find that the first recorded specimen of Asplenium septentrionale was from Holyrood Park in Edinburgh in 1669, where the ferns still thrive, despite Victonan predation. In a discussion of the crested variety of Asplenium scolopendrium, first mentioned in the 17^^ century, reference was omitted to a forked variety carved in the 15 century on the Apprentice Pillar in Rosslyn Chapel, but perhaps illustration alone does not qualify as publication. Some species first recognised in this county during the Victorian fern collecting craze seemed to disappear almost as quickly as they were found, Trichomanes speciosum being a good example. There were others now known from very few herbarium specimens, such as Cystopteris alpina, possibly becoming extinct before they had been fully recognised. At least the species first found within Britain in the 19^^ century were better described and supported by herbarium specimens, but it is of note that more remote montane species such at Athyrium distentifolium and Cystopteris montana were only difterentiated comparatively late. A recurring problem was the difficulty of determining w et er morphologically distinctive taxa merited species or varietal rank. A major a vance IS acknowledged in Irene Manton’s work on chromosome counts, published in h bridT"^ particularly useful in separating Dryopteris and Polypodium species and This IS not merely an account of long-resolved taxonomic problems as controversies continue and new discoveries continue to appear. There is the Trichomanes speciosum gametophyte stoiy that started in 1989 with the unexpected finding of gametophytes where the sporophytes had long since been lost. The gametophytes were subsequently found to be extraordinanly widespread, apparently wery long-loved and remarkably resilient. Cystopteris diaphana was added to the British flora in 2000 and although our tern flora is comparatively small there is still scope for more discoveries. This account is a detailed detective work through old records and specimens, and illuminates the excitement of discovery of our fern flora. H.S. McHafFie FERN GAZ. 19(6):207-211. 2013 207 NEW COMBINATIONS AND LECTOTYPIFICATIONS FOR SOME SOUTH-EAST ASIAN, MALESIAN AND PACIFIC GRAMMITID FERNS (POLYPODIACEAE) B S PARRIS Fern Research Foundation, 21 James Kemp Place, Kerikeri, Bay of Islands. New Zealand 0230. Email; barbara2parris@gmaiLcom Key Words: New combinations, lectotypifications. South East Asia, Malesia, Pacific, Grammitidaceae, Polypodiaceae New combinations are made for Ctenopterella nhatrangensis (C.Chr. & Tardieu) Parris, Oreogrammitis pubinervis (Blume) Parris, Oreogrammitis subevenosa (Baker) Parris, Prosaptia brassii (Copel.) Parris, Prosaptia javanica (Copel.) Parris, Prosaptia samoensis (C.Chr.) Parris, Stenogrammitis minutissima (J.W.Moore) Parris, Tomophyllum duriusculum (Christ) Parris, T. foersteri (Rosenst.) Parris, T. lividum (Mett.) Parris, T. secundum (Ridl.) Parris and T sesquipinnatum (Copel.) Parris. Lectotypes are chosen ior Polypodium emersonii var. samoense C.Chr., P. minutissimum J.W.Moore, P. secundum Ridl., P. subcoriaceum Copel. & P. subsecundodissectum var. novoguineense Rosenst. INTRODUCTION isions of grammitid ferns for Flora Malesiana and regional accounts in and the Pacific necessitates the publication of the following new NEW COMBINATIONS 1. Ctenopterella nhatrangensis {C.Chr. & Tardieu) Parris comb, nov. Based on Ctenopteris nhatrangensis C.Chr. & Tardieu, Not. Syst. (Paris) 8(4): 181 (1939). Type: Vietnam, Nhatrang, Song Ko, 700 m alt., 18 May 1922, Poilane 3338 (BM 001039878!). RANGE. Vietnam; endemic. 2. Oreogrammitis pubinervis (Blume) Parris comb. nov. Based on Grammitis pubinervis Blume, Enum. PI. Javae: 1 16 (1828). Type; Java [Blume s. n.] (lectotype LI, chosen by Parris, 2007). Synonym: Polypodium pubinerve (Blume) Christ, Ann. Jard. Buitenzorg 15: 146 (1897). Oreogrammitis pubinervis has previous been regarded as a synonym of O. setosa (Parris, 2007), but differs from the latter in having the lamina glabrous on both surfaces. 3. Oreogrammitis subevenosa (Baker) Parris comb. nov. Based on Polypodium subevenosum Baker in Hook. & Baker, Syn. Fil.: 320 (1867). Type: Penang, Mactier s. n. (holotype K!; isotypes E 00195211!, E 00195212!). Synonyms: Grammitis subevenosa (Baker) C.Chr. & Tardieu, Notul. Syst. (Paris) 8: 179 (1939). P. sessilifolium Hook., Sp. Filic. 4; 168 (1863) non Liebm. (1849). G. sessilifolia J.Sm., Hist. Filic.; 181 (1875) nomen novum pro P. sessilifolium Hook. (1863) non Liebm. (1849). P. malaicum Alderw., Malayan Ferns: 577 (1909), nomen novum pro P. 208 FERN GAZ. 19(6);207-211. 2013 sessilifolium Hook. (1863) non Liebm. (1849). G. malaica (Alderw.) Tagawa, Acta Phytotax. Geobot. 8: 173 (1939), nomen superfl. illegit. pro G sessilifolia J.Sm. P. maxM>ellii Baker, Bull. Misc. Inform., Kew 1893; 211 (1893). G. max^^’ellii (Baker) Parris, FemGaz. 12(2): 118 (1980). RANGE. Thailand?, Vietnam, Peninsular Malaysia (Perak, Penang, Kelantan, Negeri Sembilan, Terengganu, Pahang, Melaka, Johor), Borneo (Kalimantan, Sarawak), Philippines (Palawan, Sibuyan), Moluccas (Seram), New Guinea (West Papua). Oreogrammitis adspersa (Blume) Parris has only been collected once in Thailand according to Tagawa & Iwatsuki (1989, as Grammitis adspersa); perhaps the specimen belongs to O. subevenosa, as does all Peninsular Malaysian material examined. Oreogrammitis subevenosa was previously regarded as a synonym of O. adspersa (Copeland 1952, Parris 1983, 2007, 2010), but differs from it in having glabrous rather than setose sporangia. Oreogrammitis nuda (Tagawa) Parris of Taiwan also has glabrous sporangia, but differs from O. subevenosa in having elongate, rather than ± circular, sorus receptacles. The receptacles are prominent on the adaxial surface of the lamina in both species. Oreogrammitis nuda and O. subevenosa always have a white deposit (sometimes hard to see) on the hydathodes of the veins on the adaxial surface of the lamina; some material of O. adspersa also has a white deposit on the hydathodes, but it is not charactenstic of the species. 4. Prosaptia brassii (Copel.) Parris comb. nov. Based on Ctenopteris brassii Copel., Univ. Calif. Publ. Bot. 18: 225 (1942). Type: New Guinea, Brass 9302 (holotype MICH 1 1 15939!; isotypes A, BRI, K!, L 005 1 1 96!, PE!, ^NGE. New Guinea (West Papua, Papua New Guinea); endemic. /-05apna has been treated as a synonym of P circumvallata (Rosenst.) Copel. (Pams 1990), but the former has denser stipe and lamina hairs than the latter, and the abaxial surface of the rachis concolorous with the lamina rather than darker than it. 5. Prosaptia javanica (Copel.) Parris comb. nov. Based on Polypodium javamcum Copel., Ph.lipp. J. Sci.. C. Bo., 8: 142 (1913). Type: 1350-1850 m alt., CHv-cn Bry>ant Expedition 298 (US!) RANGE. Java; endemic. Polypodium javanicum was regarded as a synonym of Polypodium cryplosorum C.Chr. fekeTurp"*™'!." “ synonym of Prosaptia barathrophyUa 1 has shorter darker frond hairs than Prosap, 6. Prosaptia samoensis (C.Chr.) Parris comb Based on Polypodia l. 4c (1943). Type: Tau, ii var. samoense C.Chr., B. P. Bishop Mus. Bull. 1 77: 1 1 1 , lAcccc P^^k back of Tau, 430 m, 28 Jau. \922. D W Garber 7 19 BISH^lMsfs f mcu ' ™ here; isolectotypes pm ! ”1'’- 'f Residual syntypes: Samoa: Olosega, top of m.^Carber mi (BISH 105561-); Savaii, above S.l. 400 m, Garber 726 (BISH 105559!), Garber 731 Christophersen 3178 (BISH 1055581 (BISH 105554 105554 p. p.!), Garber 747 (BISH 105617!); Tutuila Alava Ridge 4( “P Of ,z,o 105556') r* ’■ 700 m, Christophersen 27 (BISH 1055561), Chrtstophersen 35 (BISH 1055651); Vaisingano Canyon, 500 m. PARRIS: NEW COMBINATIONS IN GRAMMITIDS 209 Christophersen 78 (BISH 105557!, BM!, MICH 1190851!). RANGE. Samoa (Olosega, Savaii, Ta’u, Tutuila, Upolu) and Tonga (Tafahi; Buelow 1178, 1230 (both CHR!)). Christophersen 78 is the only syntype represented in Christensen’s herbarium at BM and there is no indication of its type status on the specimen. The material chosen as lectotype is that illustrated when the variety was described. Prosaptia samoensis differs from P. alata (Blume) Christ, of which P. emersonii (Hook. & Grev.) C.Presl is a synonym, in having denser short simple eglandular hairs on the sorus pouch, longer pinnae and a narrower wing along the rachis. 7. Stenogrammitis minutissima (J. W.Moore) Parris comb. nov. Based on Polypodium minutissimum J. W.Moore, Bull. B. P. Bishop Mus. 102: 1 1 (1933). Type: Society Is., Raiatea, Mount Temehani, E facing slope, 400 m alt., 7 Oct. 1926, Moore 187 (lectotype BISH 498922!, chosen here; isolectotype BISH 498923!). RANGE. Society Islands (Raiatea); endemic to Mount Temehani and known only from the type collection. Smith et al. ( 1 99 1 ) did not include this species as a member of Lellingeria, nor did Labiak (2011) mention it when he separated Stenogrammitis from Lellingeria. Stenogrammitis minutissima differs from S. subcoriacea (Copel.) Labiak, also from the Society Islands (Tahiti) and in the Marquesas Islands, in the presence of forked hairs on the lamina margin, particularly at the apex of the pinnae, and on the abaxial surface of the rachis; no other types of hair are present. In S. subcoriacea, forked hairs, if present, are only at on the abaxial surface of the lamina near its base, and simple glandular and simple catenate hairs, if present, are only on the abaxial surface of the rachis. 8. Stenogrammitis subcoriacea (Copel.) Labiak Syn. Polypodium subcoriaceum Copel., Bernice P. Bishop Mus. Bull. 93; 12 (1932). Type: Society Islands, Tahiti, S. of Orohena, 1500 m, 16 May 1927, L. H. MacDaniels 1478. Both Labiak (2011) and the TROPICOS on-line data base cite US material as the holotype and UC material as the isotype. The US collection is not the holotype, however. Copeland (1932) states in the introduction to his account of the pteridophytes of the Society Islands (where Polypodium subcoriaceum is described) that he identified material from the Bernice P. Bishop Museum (BISH) for this publication, so BISH is likely to have the holotype, not US. In BISH three sheets of MacDaniels 1478 were seen during a visit in 1992: BISH 500016, BISH 105458 and BISH 105459. In 2012 BISH 500016 and BISH 105458, but not BISH 105459, were seen again in BISH. As there is more than one sheet in BISH a lectotype needs to be chosen from the available material there. BISH 500016 barcode 1000428 bears the words “Polypodium subcoriaceum Copel. n. sp. TYPE” in Copeland’s handwriting and is selected here as lectotype. BISH 105458 barcode 1000429 has a typed label, with “DUPL. TYPE” typed in the top right hand comer; this and BISH 105459, together with material in MICH!, UC! and US, are isolectotypes. 9. Tomophyllum duriusculum (Christ) Parris comb. nov. Based on Polypodium duriusculum Christ, Verb. Natur. Gesell. Basle 11: 213 (1895). Type: Celebes, G Matinang N, c. 1000 m alt., 27 Aug. 1894, Sarasin 733 (P 00632869!). RANGE. Sulawesi and Philippines (Luzon, Barcelona 541 et al. (PNH 190564!)). 210 FERN GAZ. 19(6):207-211. 2013 10. Tomophyllum foersteri (Rosenst.) Parris comb. nov. Based on Polypodium bipinnatifidum wax. foersteri Rosenst., Repert. Spec. Nov. Regni Veg. 12: 179(1913). Type: Nova Guinea germanica,BolanMts, 2400-3000 malt., 1912, Keysser B 84 (S P7206!). RANGE. New Guinea (West Papua, Papua New Guinea): endemic. Tomophyllum foersteri was previously regarded as a variety of T bipinnatifidum (Baker) Parris (Copeland 1953) or as a synonym of it (Parris 1990). The former differs from the latter, however, in having the rachis prominent and darker than the lamina on both surfaces and in having occasional to dense simple eglandular hairs on the abaxial surface of the lamina, including as receptacular paraphyses; the latter has the rachis prominent and concolorous on the abaxial surface and sunken and darker on the adaxial surface, with the abaxial surface of the lamina glabrous or rarely with occasional simple eglandular hairs that do not occur in the sorus as receptacular paraphyses. 11. Tomophyllum lividum (Mett.) Parris comb. nov. Based on Polypodium lividum Mett., Ann. Mus. Lugd. Bat. 2: 221 (1866). Type: Java, Korthals s. n. (L!). RANGE. Java; endemic. 12. Tomophyllum secundum (Ridl.) Parris comb. nov. Based on Polypodium secundum Ridl., Trans. Linn. Soc. London, Bot. 2, 9: 262 (1916). Type: Dutch New Guinea, Mt Carstensz, Camps X to XI, 6700-8300 ft alt., Boden Kloss s. n. (lectotype BM 000800777!, chosen here). Synonyms: P. subfalcatum var. semiintegrum Copel., Philipp. J. Sci., C. Bot. 2: 138 (1907). k subsecundodissectum var. novoguineense Rosenst., Nova Guinea 8: 725 (1912). Type: Nova Guinea neerlandica, summit of Hellwig Mts, 2583 m alt., Oct. 1909, von Romer 716 (lectotype L 913.103... 129!, chosen here). RANGE. Borneo (Sabah), Philippines (Mindoro), Moluccas (Seram), New Guinea (West Papua, Papua New Guinea). Tomophyllum secundum was previously treated as a synonym of T subsecundodissectum (Zoll.) Parris (Copeland 1953, Parris 2007), but the former differs in having all pinnae completely dimidiately lobed, or occasional pinnae in middle part of frond with a few basiscopic lobes in the apical part, while the latter has pinnae regularly with basiscopic lobes in the apical part. 13. Tomophyllum sesquipinnatum (Copel.) Parris comb. nov. Based on Ctenopteris sesquipinnata Copel., Univ. Calif Publ. Bot. 1 8: 225 ( 1 942). Type: mT Uct ^ ^ (holotype MICH 1115918!; isotypes A, BRI, K 00049 1 452 ! , L ^NGE. New Guinea (West Papua); endemic to Lake Habbema and known only from the type collection. Tomophyllum sesquipinnatum was previously treated as a synonym of T. bipinnatifidum ams ) ut the former differs from the latter in having a membranous, rather than a conaceous lamina, and pinnae lobed less deeply, to a wing 0.3-0.5 mm wide along the costa rather than 0.I-0.3 mm wide PARRIS: NEW COMBINATIONS IN GRAMMITIDS 211 ACKNOWLEDGEMENTS I am most grateful to Alison Paul (BM) for helpful information on the type of Ctenopteris nhatrangensis and on the material of Polypodium emersonii var. samoense in Christensen’s herbarium, the Keepers of BISH^ BM, CHR, K, L, MICH, PE and UC for access to and/or loan of specimens and Ewen Cameron at AK for expediting the loans. REFERENCES BACKER, C.A. & POSTHUMUS, O. 1931. De op Java voorkomende soorten van Polvpodium L. sensu latiore. Natuurkundig Tijdschrift voor Nederlandische Indie 9 1 : 223- 307. COPELAND, E.B. 1932. Pteridophytes of the Society Islands. Bernice P. Bishop Mus. Bull. 93:86 pp. COPELAND, E.B. 1952. Grammitis. Philipp. J. Sci. 80(2): 93-269. COPELAND, E.B. 1953. Grammitidaceae of New Guinea. Philipp. J. Sci. 81(2): 81-1 19. LABIAK, P. 2011. Stenogrammitis, a new genus of grammitid ferns segregated from Lellingeria (Polypodiaceae). Brittonia 63(1): 139-149. PARRIS, B.S. 1983. A taxonomic revision of the genus Grammitis Swartz (Grammitidaceae: Filicales) in New Guinea. Blumea 29(1): 13-222. PARRIS, B.S. 1990. Noteworthy species of Grammitidaceae from South-east Asia. Hooker’s leones Plantarum XL(IV): 129 pp. PARRIS, B.S. 2007. Five new genera and three new species of Grammitidaceae (Filicales) and the re-establishment of Oreogrammitis. Gardens’ Bull. Singapore 58(2): 233-274. PARRIS, B.S. 2010. Grammitidaceae, in Flora of Peninsular Malaysia I, 1: 131-206, ed. PARRIS B.S. et al.. Forest Research Institute Malaysia. SMITH, A.R., MORAN, R.C. & BISHOP, L.E.1991. Lellingeria, a new genus of Grammitidaceae. Amer. Fern J. 81(3): 76-88. TAGAWA, M. & IWATSUKI, K. 1989. Grammitidaceae, in Flora of Thailand 4: 581-599, ed. SMITINAND, T. & LARSEN, K. 212 FERN GAZ. 19(6). 2013 FERN GAZ. 19(6):2I3-239. 2013 213 THE NOMENCLATURE, TYPIFICATION AND TAXONOMY OF ASPLENIUM FALCATUM, A. POLYODONAISD CONFUSED SPECIES A.E. SALGADO' & C.R. FRASER- JENKINS' Christian Brothers University, Dept, of Biology, 650 East Parkway South, Memphis, Tennessee, TN 38104, U.S.A.; email: esalgado@cbu.edu. C.R. Fraser-Jenkins, Student Guest House, Thamel, P.O. Box 5555, Kathmandu, Nepal; e-mail: chrisophilus@yahoo.co.uk Key words: Pteridophytes, Asplenium falcatum, Asplenium polyodon, Asplenium aethiopicum, Trichomanes adiantoides, lectotype. ABSTRACT Asplenium falcatum Lam., described from Sri Lanka was a part of Linnaeus' concept of Trichomanes adiantoides L. The taxonomic history of T. adiantoides is outlined, showing that Linnaeus had a mixed concept and that the name had subsequently been taken in two different senses by later authors. It is lectotypified here in the sense of the African species known as Asplenium aethiopicum (Burm.f.) Bech. and a proposal to reject the name T adiantoides and all combinations based on it is being prepared. This is intended to enable the Sri Lankan element within T. adiantoides to continue to be known by its well-known name, A. falcatum Lam., and the African as A. aethiopicum, which species are outlined further and their ranges in Africa and Asia are given. Although the name A. polyodon GForst. had recently been applied to A. falcatum, it is a distinct species confined to Australasia. Other related S.E. Asian and Polynesian species are discussed briefly and contrasting diagnostic descriptions are provided to distinguish between the often confused species, A. polyodon, A. falcatum and A. macrophyllum Sw. INTRODUCTION A widespread and distinctive Asplenium species which occurs in Sri Lanka, South and N.E. India, S. China, S.E. Asia and Madagascar and the Mascarenes, had long been mentioned in Indian literature and elsewhere under the name A. falcatum Lam. But Morton (1967) concluded that the name A. falcatum was illegitimate due to the misapplication of the Linnaean name Trichomanes adiantoides L. by Lamarck, and took up the name^. polyodon GEorst. to replace it. Asplenium polyodon is a name that has been applied to a well-known species in Australasia and Oceania and was described from a plant that was most probably from New Zealand. It is superficially similar to the above A. falcatum, but after some years of doubt about the conspecificity of the Sri Lankan etc. plant and A. polyodon, we now- report here that A. polyodon is a distinct species confined to Australasia and the Asian. etc. plant must therefore be known by another name. However, due to nomenclatural problems, it has not been clear by which name it should be known now that it is clear that A. polyodon is applicable to a different species. Linnaeus (1753) created a considerable nomenclatural and taxonomic problem when he described Trichomanes adiantoides L., referring to previously published descriptions and illustrations by himself (Linnaeus, 1747), Burman (1737) and Plukenet (1691, 1696). 214 FERNGAZ. 19(6):2 13-239. 2013 Figure 1. Hermann’s fem. Herb. Hermann No. 385 (BM). SALGADO & FRASER- JENKINS: ASPLENIUM FALCATUM & A. POLYODON 2 1 5 Burman’s illustration was of a fern from Ceylon collected by Paulus Hermann, but Plukenet’s plant was of a different species from Africa. The relevant plates show two very distinct ferns neither of which belongs in Trichomanes in its modem circumscription. Hermann collected the plants in Sri Lanka and these are presently found in Leiden, in the Academic des Sciences (Institut de France, Paris) and the British Museum of Natural History. He gave a one-line description of a fern in his Musaeum Zeylanicum (1717) volume 3, fol. 47: ""Filix Zeylanica,foliis adianthi in modum incisis" (Figure. 1). Johannes Burman (1737) published an illustration (Figure 2) of Hermann’s fern, which is found at BM, and wrote a detailed description of the plant. He also added a reference to the African fern of Leonard Plukenet (1696), ""Adianthum africanum rutae murariae aemulum, segmentis longioribus acutis”, illustrated by Plukenet (1691) (Figure 3, 4). It was thus Burman who first made the incorrect connection between the African and Sri Lankan plants. Linnaeus maintained the confusion by including reference to both Burman and Plukenet under his Trichomanes adiantoides in Flora Zeylanica (1747: 35) and later in Species Plantarum ( 1 753) and thus validated a name for a mixed concept based on two different species. Both voucher-specimens can be found in the historical collections at BM. Hermann’s plant is in Herb. Hermann 3: 47, no. 385, and Plukenet’s specimen is in Herb. Sloane 95,: 16, at BM-SL (Figure 4). Burman’s herbarium is in Geneva (G), but is not relevant to the case. In his EncyclopMie Methodique, Lamarck (1786) noticed the problem created by Burman and perpetuated by Linnaeus, and distinguished both species transferring both separately to the genus Asplenium, listing them under the French vernacular name for that genus, ‘Doradille’. He remarked under "Doradille adiantoide' that neither of the cited illustrations was of a Trichomanes but both were of true ‘doradilles’, and that the two species were quite different from each other. He treated the Sri Lankan species as Asplenium falcatum Lam., and the African fern asAsplenium adiantoides (L.) Lam. and noted that Linnaeus had included within T adiantoides both the African plant illustrated by Plukenet and the Sri Lankan plant described by Burman. Hermann’s words, "'Filix Zeylanica, foliis adianthF etc., might appear to be related to the origin of the name T adiantoides, but Linnaeus did not cite Hermann. He cited Burman (1737) and his own Flora Zeylanica (1747) both of which mention the African and the Sri Lankan plants. It could be argued that Lamarck should have applied the name Asplenium adiantoides to the Sri Lankan plant. On the other hand, it can be concluded that, as both the Burman and the Plukenet plants were mentioned by Linnaeus, either can have equal claim to his epithet adiantoides. Lamarck’s choice was to apply the name A. adiantoides to the African element rather than the Sri Lankan. Tardieu-Blot & Ching (1936) listed A. adiantoides Lam. as a synonym of A. praemorsum Sw., the name then being utilised instead of^. aethiopicum, and stated that Plukenet illustrated it for the first time but that Linnaeus had reported it in error under his Trichomanes adiantoides, which is a completely different plant. They also said that the type of T. adiantoides is a synonym of Asplenium falcatum Lam. and it can be found at Hermann’s herbarium at Kew [ 51 c] and it was well illustrated by Burman (1737). Following Christensen (1905) where he renamed the Sri Lankan plant Asplenium adiantoides - a later homonym of Lamarck’s A. adiantoides - they concluded that the name adiantoides must be retained for the Sri Lankan plant - the Asplenium falcatum of Lamarck - which they called “la plante de Linne”, the Linnaean plant, while A. adiantoides Lam. must be called A. praemorsum, the African plant. While this is one 216 FERN GAZ. 19(6):2 13-239. 2013 possible interpretation, it is also entirely arbitrary and went against Lamarck’s conclusion. As they stated that the type is the specimen in Hermann’s herbarium (actually at BM, not Kew), this cannot be taken as valid lectotypification as the type has to be the illustration Linnaeus cited. h. gore 2. Type plate Barman t. 43 cited by Linnaeus under Trichomanes adian, aides in Flora Zeylamca (1748) and Species Plantarum (1753). This plant is from Ceylon and was renamed Asplenium falcatum by Lamarck. SALGADO & FRASER-JENKINS: ASPLENIUM FALCATUM & A. POLYODON 217 Morton (1967) similarly was of the clear, if arbitrary opinion that the Sri Lankan plant should have been the element identified as Trichomanes adiantoides . According to ramosa Zeylanica..." and then mentioned its leaves as being serrate in the way of an Adiantum, he reasoned that Linnaeus’ main intention must therefore have been to use the Ceylon plant as the basis for T adiantoides. He went so far as to say that the type of A. adiantoides “must be the Ceylon planf though without formally selecting a type. He therefore concluded that de Lamarck should not have applied the name A. adiantoides to the African plant, though in reality he was free to do so. Two pieces of evidence linking Linnaeus’ account to Hermann’s specimen (Figure 1) are shown by the citation in Flora Zeylanica of what was actually Linnaeus’ number 385, and by the name Trichomanes, which was written in Linnaeus’ own handwriting on Hermann’s original specimen. Linnaeus cited his Flora Zeylanica account of Trichomanes adiantoides in Species Plantarum, and this account was based on the Hermann specimen that he unambiguously annotated, though he did not make direct reference to the specimen in Flora Zeylanica or Species Plantarum. The Hermann material at BM is original and should be considered for lectotypification purposes. Therefore, the type specimen of Trichomanes adiantoides must be chosen from either the Hermann specimen (Herb. Hermann 3: 47, no. 385 at BM), Plukenet specimen (Herb. Sloane 95: 16, at BM-SL) or either of the illustrations mentioned by Linnaeus in his Species Plantarum. According to Morton’s interpretation of T. adiantoides, when Lamarck named the and not a new species, since the Sri Lankan plant was in his view the basis and type of T. adiantoides. He therefore treated A. falcatum as an illegitimate nomen superfluum. Furthermore he concluded that Lamarck’s Asplenium adiantoides from Africa, a synonym of A. aethiopicum (Burm.f.) Bech., was not a new combination, as would appear to be the case, but was independent of Trichomanes adiantoides L. (from Sri Lanka), despite Lamarck’s citation of it. Morton concluded, ''Lamarck was therefore unjustified in typifying the Linnaean species on the basis of the African plant illustrated by Plukenet. One subsequent statement on typification was made by Brownsey (1998), who under T. adiantoides L. in the synonymy of A. polyodon, cited: “T: Ceylon [Sri Lanka]; holo: BM (Herb. Burman 385).” At first sight this appears to constitute lectotypification despite the statement in the introduction that "these type statements are not to be interpreted as lectot\pifiications'\ because it also says that the type citations under taxa "reflect the authors' belief in their current status" and under the Code (McNeill, 2012: Art 9.9), the use of a wrong term for a type is merely treated as an error to be corrected (in this case holotype instead of lectotype). The process of lectotypification under the ICBN does not allow for disavowal of intent. However, Herb. Burman is in G, not BM as cited (it should have been Herb. Hermann) and there is no such specimen as Burman no. 385, which was the reference given in Linnaeus’ Flora Zeylanica to Hermann’s specimen no. 385. Thus this statement cannot be taken as lectotypification as no actual existing element was cited, nor the correct herbarium. Lectotypification of Trichomanes adiantoides L. Lamarck was correct in separating the two species, but the present difficulty concerning the correct application of names is due to subsequent disagreement about his application of the epithet adiantoides and his choice to name the Sri Lankan plant independently. 218 FERN GAZ. I9(6):213-239. 2013 Figure 3. Plukenet’s illustration cited by Burman (1737) and Linnaeus (1753). SALGADO & FRASER-JENKINS: ASPLENIUM FALCATUM & A. POLYODON 219 Linnaeus’ simple diagnosis fits either the Sri Lankan or the African fern almost equally, either of which can therefore be chosen as lectotype. Thus, in contrast to Morton’s argument, it can be concluded alternatively that the name A. falcatum Lam. is legitimate as a new Sri Lankan species, when the name A. adiantoides (L.) Lam., understood as being a new combination based on Linnaeus’ name, is applied to the African species, as Lamarck did. There also remains the question of the typification of T. adiantoides because neither Linnaeus, Lamarck, Christensen, Tardieu-Blot & Ching, nor Morton selected a single element that can nowadays be taken as being a lectotype. As pointed out to C.R. Fraser- Jenkins by C.E. Jarvis (in litt. 26 Oct. 2005), though they made reference to various descriptions and illustrations, they either chose a specimen incorrectly or did not unequivocally select a single element as type, this was therefore insufficient to be taken as modem lectotypification. It has now become essential to select between either the Sri Lankan element or the African element as a permanent lectotype of P. adiantoides in order to elucidate the nomenclature definitively. As discussed above, there are two options for the choice, both being consistent with Linnaeus’ protologue and both being tenable. The nomenclatural consequences of the two possible options for lectotypification are as follows: 1. Lectotypification of T. adiantoides based on the Sri Lankan element (following Morton): a. Trichomanes adiantoides L., non A. adiantoides Lam., becomes a synonym of Aspleniiim sp. from Sri Lanka and S. India, etc. The epithet adiantoides cannot be used for the Sri Lankan species, as the combination is preoccupied in Asplenium by A. adiantoides Lam, the African species. b. Asplenium adiantoides Lam., the African species, becomes a synonym of A. aethiopicum (Burm.f) Bech. The African plant remains A. aethiopicum, which name was first applied to it by Becherer (1936) in lieu of^. praemorsum. c. Asplenium falcatum Lam. becomes an illegitimate name as Lamarck cited T. adiantoides in its synonymy so it is a nom. nov. for T adiantoides as he should have chosen the epithet adiantoides instead, which was available. d. The Sri Lankan and South Indian plant would need to be known by a different name from either A. falcatum or A. polyodon, but it is unclear whether another name exists or whether it would have to be described as a new species due to the existence in literature of several related but obscure names, no longer in use, from different parts of Africa and e. Providing a name for Linnaeus’ Sri Lankan plant is therefore very difficult and would require years of further investigate research in two continents and several different regions. 2. Lectotypification of T adiantoides L. based on the African plant (following Lamarck): for A. aethiopicum and the a. Asplenium adiantoides (L.) Lam. becomes the correct i 220 SALGADO & FRASER-JENKINS: ASPLENIUM FALCATUM & A. POLYODON 221 et ai, in prep.) for the rejection of the name T. adiantoides and all combinations based upon it. This will enable the African/S. Asian plant to continue to be known as A. aethiopicum and the Sri Lankan/Indian plant to continue to be called A. falcatum. Accordingly we now select as lectotype, here designated, of Trichomanes adiantoides L, Sp. PL. 2:1098 (1753), the following illustration, Plukenet, Phytographia 3: t. 123. f 6 (1691; Figure 3). We also select an epitype, here designated, to assist in the precise and critical identification of the above lectotype, “South Africa, Cape Province, Kalk Bay Mountain, in rock crevices, c. 1000 ft. A.F. Braithwaite 75, 25 Aug. 1960 (K)”, octaploid sexual, det. A.F. Braithwaite (1986). Figure 5. Asplenium aethiopicum (Burm.f ) Bech., Sonnerat s.n. (Herb. Lamarck, ) 222 FERN GAZ. 19(6):2 13-239. 2013 Lectotypiflcation of Asplenium falcatum Lam. The Sri Lankan etc. plant Linnaeus was referring to is now once more to be called A. falcatum Lam., as it used to be widely known previously. Lamarck, however, did not Figure 6. Asplenium falcatum Lam., Sledge 873 (epitype, BM). SALGADO & FRASER-JENKINS: ASPLENIUM FALCATUM & A. POLYODON 223 specify the type of Asplenium falcatum, though he applied the name to Linnaeus’ Sri Lankan element. We therefore select as lectotype, here designated, of A. falcatum Lam., Encycl. Meth. Bot. 2(1); 306 (1786), the illustration in Burman’s Thesaurus Zeylanicus, t. 43 (1737). Burman wrote a detailed description of this plant published together with the plate (Figure 2). This illustration is a very good and accurate representation of the Hermann specimen in Herb. Hermann 3: 47 (Figure 1, BM) and most usefully shows the long son on the underside, though the specimen is now pasted down to show only the top surface. Linnaeus cited this reference, among others, under Trichomanes adiantoides in Flora Zeylanica (1747) and Species Plantarum (1753). The plant is from Sri Lanka and was renamed Asplenium falcatum by Lamarck. In order to help with identification, we here designate as epitype of the above lectotype: “Ceylon, in forest in the upper part of Ritigala, N. Cent. Province, alt. 2250 ft., W. A. Sledge 873, 14 Jan. 1 95 1 (BM)’’ (cited by Sledge 1965; Fig 6). Similar material from Sri Lanka was found by Manton & Sledge (1954) to be octaploid sexual, but we are as yet unable to find their Sri Lankan voucher- specimen, which should be among their extensive unincorporated material at BM. which Morton confused it with (though, as he stated, taken in a broad sense). Pending rejection of T. adiantoides L., nomina ubique rejic. prop., the African (and S. Asian) plant referred to by Linnaeus should continue to be called A. aethiopicum (Burm.f.) Bech. subsp. aethiopicum (see below re subspecies), with T. adiantoides L. and A. adiantoides (L.) Lam. in its synonymy. Asplenium aethiopicum (Burm.f) Bech. Lamarck also mentioned two further specimens of A. adiantoides, one collected in Afiica by Sonnerat (Figure 5) but without specific locality, and the other in Mauritius by Commerson (Figure 7). Both specimens are preserved in Herb. Lamarck at P. The Sonnerat specimen bears Lamarck’s annotation ""vide Pluk. Fig. 123, t. 6” and belongs to A. aethiopicum, while the Commerson specimen was a misidentification by Lamarck as it belongs to A. falcatum. However neither plant cited by Lamarck has type status as they were not included by Linnaeus as part of his concept. Morton (1967), followed by Schelpe & Anthony (1986), designated the Commerson specimen as lectotype of A. adiantoides Lam. (Figure 7), but taking the name as if independent from Linnaeus’ name. This lectotypification therefore does not stand as it was not one of the elements cited by Linnaeus. Christensen (1905) again made the combination A. adiantoides (L.) C.Chr., as if independent of Lamarck’s name, which latter he listed separately and identified as being A. praemorsum Sw. taken in a wide sense, meaning A. aethiopicum (as at that time no- one knew the identity of Trichomanes aethiopicum Burm.f). But his new combination was mistaken as it was an illegitimate later homonym of Lamarck’s and had also been utilised independently in the meantime by Raddi (1819) and Raoul (1844). He took the Linnaean basionym and his attempted combination in the sense of the Sri Lankan plant, placing A. falcatum Lam. in its synonymy. By the time Lamarck mentioned the African plant, it had already been described and named three times. The oldest description was by Nicolaas Burman (1768) under the name Trichomanes aethiopicum Burm.f Becherer (1936) transferred this to Asplenium and the species is now known as Asplenium aethiopicum (Burm.f) Bech. This is a widespread, phenotypically and cytologically variable subtropical species found 224 FERN GAZ. 19(6):2 13-239. 2013 SALGADO & FRASER-JENKINS: ASPLENIUM FALCATUM & A. POLYODON 225 throughout Africa, the Mascarenes, Macaronesia, S.E. and South Asia (including Myanmar, Thailand, China, Vietnam, Sri Lanka, S. India, and in one locality in N.E. India), with the closely related species or subspecies, A. praemorsum Sw., occurring in C. and S. America. In Africa it is a member of a highly critical allopolyploid pillar complex, whose members have frequently been confused with each other, but nevertheless usually have distinct frond-morphology. The subspecies of the complex were elucidated and delineated morphologically and cytologically by Braithwaite ( 1 984, 1986) and are now well established in African, European and Asian literature. Asplemim aethiopicum is typified by a specimen from the Cape of Good Hope, in Herb. Burman (G). It is known to be an octaploid sexual taxon, subsp. aethiopicum; exactly the same taxon also occurs commonly, and has been confirmed through chromosome counts, in South India and Sri Lanka, and the same subspecies has probably been discovered recently in Assam, N.E. India, as well (Odyuo et al. 2012). As stated above, T. adiantoides L. and A. adiantoides (L.) Lam. are being proposed as nomina ubique rejic. prop, and will thus be synonyms of A. aethiopicum if the proposal is accepted. The detailed range of .4. aethiopicum is Macaronesia (Madeira, cytologically distinct). West, South and throughout East Africa, the Comoros, Madagascar, Mauritius, La Reunion, Yemen, South and N.E. India (Andamans, Tamil Nadu, Kerala, Karnataka, Andhra Pradesh, Assam), Sri Lanka, Myanmar, Thailand, Vietnam, S. and S.W. China, Malaysia, Java, the Philippines, Australia, New Caledonia, Oceania. Records from Hawai’i and C. and S. America refer to the closely related A. praemorsum Sw., which is probably best treated as a further subspecies of A. aethiopicum, and which is generally rather less lobed in its pinnae. It should be noted that whereas in much recent Indian literature following Morton ( 1 973), the name A. gueinzianum Mett. ex Kuhn, described from Natal, is applied to what used to be known in North India and China as A. laciniatum sensu Hook., non D.Don, the latter species is not known from Africa and Morton misidentified A. ginenzianum. Although Morton thought the type of A. gueinzianum looked indistinguishable from the Indian plant, Schelpe & Anthony (1986) also saw the type and placed it within A. aethiopicum, but it has subsequently been precisely identified by Roux (2009) as being conspecific with the purely Afncan species, A.friesiorum C.Chr., whose name it therefore replaces. Recent unresearched misapplication of the name A. gueinzianum in India and China is entirely erroneous. Fraser- Jenkins et al (2011) renamed the Indian and Chinese plant as A. lacinioides Fraser- Jenk., which has been overlooked among a number of other taxonomic and nomenclatural errors in the genus in the Flora of China (2013). Asplenium falcatum Lam. Until the time of Morton’s (1967) publication, the nwtvQ A. falcatum was widely used for plants from Africa, India (partly in error for the species known there as ‘M. macrophyllum" auct., non Sw. and for A. finlaysonianum Wall, ex Hook, in N.E. India). Sri Lanka, S.E. Asia and China to Polynesia (the latter in error for related species). Several of these plants are not conspecific with A. falcatum, though of the same general habit. But following Morton (1967) in believing that A. falcatum was an illegitimate superfluous name, botanists did not hesitate to use the name A. polyodon, newly proposed as applying to the plants previously named as A. falcatum; see for example Sledge (1982), who had previously utilised the name A. falcatum (Sledge, 1965) from Sri Lanka. In South India and mainland China, however, Morton’s publications remained almost 226 FERN GAZ. 19(6):2 13-239. 2013 Figure 8. Asplenium polyodon GForst., “Insulae Oceani Pacifici”, G. Forster s.n (lectotype,UPS). SALGADO & FRASER-JENKINS: ASPLENIUM FALCATUM & A. POLYODON 227 unknown and use of the name A.falcatum persisted until recently. It was still used by Wu (1999) in the Flora Reipublicae Popiilaris Sinicae, with A. polyodon given in synonymy, but without being aware of the complications, while the Flora of China (2013) simply followed Morton without necessary research and misapplied the name A. polyodon to it. Otherwise the name A. falcatum has practically disappeared from new collections in many herbaria, which have instead adopted A. polyodon as if the correct name. But as mentioned above, the two are different species and A. falcatum is now restored as the correct name for the Sri Lankan and Indian etc. plant. A.falcatum has been found to be a sexual tetraploid species in Sri Lanka (Manton & Sledge, 1954) and N. India (Bir 1960), both voucher-specimens checked and confirmed by CRFJ, though other records from India are unreliable and without voucher-specimens. Various reports from other countries sub A. polyodon do not refer to A. falcatum. The detailed range of A.falcatum as construed here is Madagascar, the Mascarenes (Mauritius, La Reunion, Rodriguez), S., C. and N.E. India (Andamans, Nicobars, Tamil Nadu, Kerala, Karnataka, Orissa, W. Bengal (Teesta - Siliguri), Assam, Nagaland, Arunachal Pradesh), ?Bangladesh, Sri Lanka, across mainland S. China (Yunnan to Hainan), Taiwan, Myanmar, Thailand, Vietnam, Malaysia and Indonesia to the Philippines. Reports from N.E. India are mainly in error for ‘M. macrophyllum" sensu auct. Ind., non Sw. (see below), or sometimes for A. finlaysonianum Wall, ex Hook., or A. khasianum Sledge, but A. falcatum also occurs there as a rarity. A more deeply lobed or pinnatifidly pinnate plant from S. India, Sri Lanka and Malaya is^. mysorense Roth (syn. A.falcatum var. bipinnatum (Sledge) Sledge; A. bipinnatum (Sledge) Philcox, non C.Chr.). But the plants from Australasia are A. polyodon (see below), which does not occur in India or Africa, despite reports in error for A.falcatum. Asplenium falcatum can also be confused with the S. Indian and Sri Lankan species, A. decrescens Kunze, especially when the pinnae of A. falcatum are narrower than usual, as often occurs as part of its individual variation. But A. decrescens has a long-creeping rhizome and apart from in the basal acroscopic lobe, the sori are parallel to and almost confined to a line along the midrib of the pinna, and do not point directly outwards nearly to the margin. None of these species occurs in the C. or W. Indo-Himalaya, as sometimes reported in Polynesian plants previously reported as A. falcatum may represent one or more distinct species, and require further study. Asplenium polyodon GForst. {non Wall. nom. nud.). Georg Forster described .4. polyodon from a plant he probably collected in New Zealand during Captain James Cook’s second trip to the Pacific (Figure 8). The species was described in his Australian Prodromus (1786), the same year as A. falcatum Lam., together with many other new species collected during the voyage of H.M.S. Challenger. Unfortunately Forster did not specify anything that could now be taken as a holotype, nor gave the locality, and apparently the only surviving specimen of^. polyodon collected by him is a small, sterile, juvenile frond found in the Thunberg Herbarium in Uppsala, UPS (Figure 8). Nicolson & Fosberg (2004) designated this specimen as the lectotype of A. polyodon. Hooker (1860) considered A. polyodon identical to A.falcatum Lam. and commented on the variability of the latter species and that under his rather wider species concepts it intergrades imperceptibly into A. macrophyllum Sw., A. caudatum GForst. and others. Christensen (1905) listed A. polyodon as both a synonym and a variety of his sense of A. adiantoides. However we now report from study of and familiarity with the 228 S ALGADO & FRASER-JENKINS: ASPLENIUM FALCA TUM & A. POL YODON 229 Caledonia has other closely related plants that may represent at least two perhaps infraspecific taxa, and one new species with a creeping rhizome. Despite previous reports, true A. polyodon also does not occur in Polynesia, where different species appear to replace it. Wallich (1828: 63) mentioned Asplenium polyodon Wall. {nom. nud.) as the name for what he had previously called Asplenium porrectum Wall. (Num. List no. 224), from Mauritius, which was equated by Christensen (1905) with A. protensum Schrad. It is independent of A. polyodon GForst. Asplenium macrophyllum Sw. Aspelnium macrophyllum Sw. (1801) is much closer to A. falcatum and belongs to the Figure 10. Asplenium polyodon GForst. Pinna shape, serration ; R.Schodde3\(){BM). al arrangement. 230 FERN GAZ. 19(6):2 13-239. 2013 polyodon to replace the supposedly illegitimate name A. falcatum and some previous taxonomists had even considered it a large form of A. falcatum. But it is clearly a distinct species. It is based on a plant collected by Groendal in Mauritius (Figure 15), of which the holotype is a rather poor specimen, preserved in S. Asplenium macmphyllum differs with longer, less prominently serrate and straight apices, sometimes with long-acute subapical lobes and ending in a long-acute or short acuminate tip. The stipe is also less scaly. Asplenium macrophyllum does not agree with Hermann’s specimen and Burman’s illustration. It is certainly known only from the Mascarenes, though it has often been SALGADO & FRASER-JENKINS; ASPLENIUM FALCATUM & A. POLYODON 23 1 reported from N.E. India (W. Bengal, Darjeeling Distr.; south Sikkim; and Assam (from where it has been confirmed cytologically as a sexual tetraploid)), Bangladesh, Myanmar, Thailand, through South-East Asia and Borneo to the Philippines. At least the Indian subcontinental plant, also including Myanmar and Thailand, is not the same as true A. macrophyllum from the Mascarenes and appears to represent a distinct species, closer to, but distinct from A. falcatum. This currently unidentified species does not have the ovate pinna-bases of A. macrophyllum, and also has fewer pinnae, with fewer lobes and less teeth than in A. falcatum. It is currently under study by the present authors in comparison with S.E. Asian species. It is an uncommon fern, poorly represented in herbaria and in the past has sometimes been confused with A. fmlaysonianum Wall, ex Hook. {e.g. from Nepal). 232 FERN GAZ. 19(6):2 13-239. 2013 Figure 13. Asplenium falcatum Lam. Lowest pinnae not decreasing. , lie de France (Mauritius Commerson s.n., (Herb. Lamarck, not a type, sheet 2, P). SALGADO & FRASER-JENKINS: ASPLENIUM FALCATUM & A. POLYODON 233 Asplenium macrophyllum is absent from Polynesia and Australasia and reports from the former refer to apparently different taxa. Asplenium cultratum Gaudich. Gaudichaud-Beaupre (1828) described Asplenium cultratum Gaudich. from a plant collected in Guam, Marianas, during the voyage of Captain Freycinet around the world, the type being at P (Figure 16). Considered for a long time to be a synonym of A.falcatum Lam., this is the next oldest name without previous homonyms that would have been a possible replacement for it, if applied in a broad sense. But the type and other collections of A. cultratum do not quite match the Hermann specimen or other collections from Sri Lanka very well and might perhaps represent a distinct species. Particularly for this reason it is fortunate that there does not have to be a replacement for the name A. falcatum. Asplenium urophyllum Wall. nom. nud. Asplenium urophyllum Wall, is another name that has occasionally been used for plants of A. falcatum that are more deeply serrate. Otherwise, it is practically indistinguishable from A.falcatum. It was a nomen nudum mentioned by Wallich (1828) in his Numerical List or Catalogue as collected from Penang, Malaya, by George Porter. It has occasionally been listed as A. macrophyllum var. urophyllum Hooker (1860), who validated the name at varietal rank. Christensen (1905) listed it as a synonym of either A. adiantoides (L.) C.Chr. or A. macrophyllum Swartz. It seems likely that it is only a more serrate-lobed stage or form of A. falcatum. Polynesian species related to A. falcatum and .4. polyodon Polynesian plants have traditionally been identified as A. polyodon, A. macrophyllum or A. falcatum, depending on the period when the specimens’ identity was determined by botanists. True A. polyodon, A. macrophyllum and A. falcatum do not occur in the Marianas, Fiji, Tonga, Samoa or other Polynesian islands further east. The Polynesian plants differ in details of the frond apical segment, pinna margin and shape, and sori. The paucity of samples from the major groups of Polynesian islands in European herbaria makes it difficult to determine the ranges of variation that may exist. This species-complex clearly requires further study in Polynesia as the species have not been fully elucidated there. Diagnostic comparisons of A.falcatum, A. macrophyllum and A. polyodon 1 . Rachis scales narrow and ribbon-like or folded forming a concave cavity at the base of the scale, with 2-4 narrow branches along the margin. Lowest pinna pairs reducing; frond apical segment different from pinnae, narrow, pinnatifid and incised, base narrow and unevenly cuneate, formed by two adnate pinnae, margin incised and serrate, two rows of sori, one parallel to the rachis; pinnae narrowly trapeziform, base unevenly cuneate; pinna margin incised, forming serrate lobes; veins oblique to the costa, forking two or three times; sori very prominent, forming two rows, first row parallel then diverging from the costa, second row of sori after the second fork, never reaching near to the margin; sori 16 - 20 cm long polyodon (Figs. 10, 12, 14) 2. Rachis scales wide and flat, not forming a concave cavity at the base of the scale, with 234 FERN GAZ. 19(6):2 13-239. 2013 3-6 short branches around the base. Lowest pinna pairs of the same length as middle pinnae; frond apical segment wide to very wide, ovate-lanceolate, base broadly and evenly cuneate, margin finely serrate throughout, often with two long acuminate basal lobes resembling attached pinna-like lobes, with all sori in a single row at an oblique angle to the costa; pinnae wide, ovate-trapeziform, base unevenly cuneate; veins oblique Figure 14. Asplenium polyodon GForst. Lowest pinna pair reducing. R Schodde 316 (BM). SALGADO & FRASER-JENKINS: ASPLENIUM FALCA TUM & A. POL YODON 235 Figure 15. Aspleniu ophyllum Sw., Groendal s.n., Mauritius (type, S). 236 FERN GAZ. 19(6):213-239. 2013 Figure 16 . Asplenium cultratum Gaudich., Syntype: Gaudichaud s.n., Guam, Marianas Is. (with permission, courtesy Herb. P.) SALGADO & FRASER-JENKINS: ASPLENIUM FALCATUM & A. POLYODON 237 3b. Pinnae few, trapeziform (2.6-)3-6(-7) cm wide, 10-18(-20) cm long, 2-3 times longer than wide, straight, pinna apex acute to shortly acuminate, pinna margin notched to unevenly serrate throughout the length of the pinna A. macrophyllum (Figure 15) CONCLUSIONS 1 . Trichomanes adiantoides L. is lectotypified by Linnaeus’ African plant, now called A. aethiopicum (Burm.f ) Bech. Trichomanes adiantoides and all combinations based upon it are soon to be proposed as nom. ubique rejic. prop. 2. Asplenium falcatum Lam. is a legitimate name described from Sri Lanka and should 3. Asplenium polyodon GForst. is specifically distinct ^rom A. falcatum Lam. This name should only be applied to plants from Australia, New Zealand and adjacent islands. 4. Asplenium macrophyllum Sw. is specifically distinct from A. falcatum Lam. (and A. polyodon GForst.) and occurs in the Mascarenes. The name should not be applied to A. falcatum, nor to the urmamed species reported as ""A. macrophyllum” from India etc. 5. Plukenet’s African fern (also in S. Asia, etc.) should continue to be known as Asplenium aethiopicum (Burm.f) Bech. subsp. aethiopicum in anticipation of the rejection of T. adiantoides L. and vl. adiantoides (L.) Lam. 6. Polynesian plants are different from the typical A. polyodon GForst. dead A. falcatum Lam. and require further study. ACKNOWLEDGEMENTS We are most grateful to Dr. C.E. Jarvis, of the Linnaean Typification Project, Botany Dept., The Natural History Museum, London, for kindly advising CRFJ of the lack of typification status of Lamarck’s, Morton’s and Brownsey’s comments concerning T. adiantoides and for helpful discussion. We are also grateful to Prof J. McNeill, of the Royal Botanic Garden, Edinburgh, for his helpful comments and suggested revision prior to finalisation of the paper. REFERENCES BECHERER, A. 1936. Note sur deux especes d’ Asplenium, Candollea 6; 22-24. BIR, S.S. 1960. 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A numerical list of dried specimens of plants in the East India Company’s Museum, collected under the superintendence of Dr. Wallich of the Company’s Botanic Garden at Calcutta, pp. 1-253. London and Hereford. WU, S.-H. 1999. Aspleniaceae, in Flora Reipublicae Popularis Sinicae 4(2): 16-163. Beijing: Science Press. THE BRITISH PTERIDOLOGICAL i Registered Charity No. 1092399 Patron: HRH The Prince of Wales Officers and Comniittee from April 2013 President: Proll J.A. EdgB^ton, 19 Nfedclenbutgh Square. London WCIN 2AD 3 1753 00403 8573 THE FERN GAZETTE VOLUME 19 PART 6 i variability of leaf lifespan of ferns K. Mehltreter & JM. t i short biography of Joanne M. Sharpe k sfe»rt bio^^hy of lOans Mehltreter MAIN ARTICLES New combinations and lectotypifications for some South-East Asian, Maksian and Pacific grammitid ferns (Polypodiaceae) AX Pmr^ The nomendature, typification and taxonomy of A^mum falcatum, A. poiyvdtfH and confused species A£. Salgado & C.R. 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