Abswacta Botanica 2\(1): 15-19, 1997 ‘© Department of Pant Taxonomy and Ecology, ELTE, Budapest The taxonomic diversity of epiphyllous bryophytes S. Rob Gradstein Systematisch-Geobotanisches Institut, Universitat Gottingen, Untere Karspiile 2, D-37073 Géitingen, Germany Keywords: Epiphyllous, Bryophyte, Taxonomy, Diversity, Ecology, Liverworts “Abstract: The first report of an epiphyllous bryophyte probably dates back to 1788 when Swartz described Jungermannia flava Sw. (= Lejeunea flava (Sw.) Nees) from Jamaica, growing on old fer fronds, Since then, hundreds of epiphyllous bryophyte species have been described. Compared with about 5-6,000 species of bryophytes occurring in the topics, the “epiphyllae" are, however, a relatively modest group. Moreover, many of them are facultatively epiphyllous and are also common on bark or other substrates. "Typically" epiphyllous species (ie. bryophytes growing exclusively or almost exclusively on leaves) usually occur as shade epiphytes in the understorey of the rain forest. They are small, pale-coloured, apressed-growing plants with thizoids in bundles that differentiate into large, adhesive discs, serving as an effective hold-fast. Short-distance dispersal by means of ‘gemmae is very commen in typical epiphylls and some species produce neotenic gametophytes as an adaption to survival in the ephemerous habitat, The distribution within the taxonomic system of typically epiphyllous ‘bryophytes is very restricted. Most of them are hepatics of the Lejeuneaceae subfamily Lejeuneoideae; other typical cpiphylls include species of Radulaceae (Radula sect. Epiphyllae), Hookeriacese sensu lato (e.g. Crossomitrium sect. Crossomitrium) and Nemataceae (Ephemeropsis rjibodensis). The restricted occurrence of typical epiphylls in ‘bryophytes is tentatively explained by the special morphological and ecological; requirements posed upon these ‘organisms for their survival in the phyllosphere. Introduction, Since the end of the 18th century, hundreds of epiphyllous (= foliicolous) bryophytes have been described from localities throughout the tropics. Most of them grow only occasionally in the phyllosphere, however, and are common inhabitants of bark or other substrates. "Exclusive" or “obligate” epiphylls seem to very rare in bryophytes and the British hepaticologist E.W. Jones (in Richards 1984, p. 1256) claimed that obligate epiphylis possibly do not exist in bryophytes. To prove that a species is an obligate epiphyll is obviously very difficult and would require a very large number of habitat observations from the whole range of the species, since taxa may be obligately epiphyllous in one region, non-obligate in another. As such proof is normally lacking, usage of the term "obligately epiphyllous" does not seem practical, This paper focuses on those bryophytes thal have thus far been found exclusively or almost exclusively on leaves. They may loosely be called “typically epiphyllous” (compare "typically epiphytic", Oliver 1930), being more or less equivalent to "eu-epiphyllous" (Sérusiaux 1977), “truely epiphyllous," or ‘"haufig epiphyl* (Licking 1995). The rationale for restricting this paper to the typical epiphylls is that these are more likely to possess special adaptations to the phyllosphere and thus may offer the best clues as to how "epiphyllae” ‘may differ from other bryophytes. Early history The oldest report of an epiphyllous bryophyte was probably by Swartz (1788), who described the hepatic J. flava Sw. (= Lejeunea flava (Sw.) Ness) from Jamaica, growing on old fem fronds adnascitur frondibus senescentibus Filicum', see Swartz 1806). Though frequently occurring on16 leaves, Lejeunea flava is by no means a typical epiphyll and is also common on bark or other substrates. The species is widespread throughout the topics and occurs also in warm temperate regions, 2g. in western Europe. ‘Table 1. Families and genera of bryophytes having typically epiphyllous species. Genera which are predominantly epiphyllous are in bold, Groups containing only facultative epiphylls are omitted, a Muse HOOKERIACEAE sens lato CCrassomitrum section Crossomirium.- tropical America (Allen 1990) (Chaetomitrium sp.» pantropical Daitonia spp. - pantropical Dimorphocladon sp. - Borneo (ide Richards 1984) Distichoplyllidium spp, - topical Afvican species “Lesteodon spp. Aftian and Asiatic species (fide Pécs, in it.) NewaTaceas Ephemeropsis(E ibodensts!)- SE Asia, Australasia Note: cpipiyious members of Sematophyllaceae subfim, Clastobryoideae, some of which have been considered obligate epiphylls (Richards 1984), all seem tobe facultative epiphyis. Hepaticae Leseuneaceat Prrauxmoness Caudalejeunea spp. Aftican species Loseincoment Acentholejeunea spp. - Asia ‘Aphanolejeunea spp. - pantropical Ceratolejeunea spp. (only facultaively epiphyllous?) Coloicjeunea spp. - pantropical Cyclolejeunea - topical America CCotura spp. - pantropical Diplasiotejeunea spp. - pantropical Drepanolejewrea spp, ~ paniropical Lepiotejeunea - pantropical ‘Metcgeriopss- Malesia Microlejeunea spp. - pantropical Odontolejeunea - tropical America and E. Afvica Ototejeunea - Asia and E, Africa Prionolejeunea spp. -pantvopical ‘Rhaphidolejeunea - Asia and wopical America Stenolejeunea spp. Asia Sphaerolejeunea - Colombia Tuyamaella sp. - Asia RADULACEAE Rasdula sect. Epiphyliae = subg. Metaradula, Schuster 1991) Note: epipiyilous members of Frullaniaceae (e.g. Frullania Steph. enAbstracta Botanica 21 (1997) Fulford, M. H. 1956, Young stages of the leafy Hepaticee: résumé, Phytomorphology 6: 199-235. Goebel, K. von 1888, Morphologische und biologische Studien. ‘Annales du Jardin Botunique de Buitenzorg 7: 1-73. Grolle, R. 1985. Zur Kenntnis der Oiolejeunea, Hanssknechtia 2: 45-56. Grditein, S. R. 1992, The vanishing rain forest as an environment for bryophytes and lichens. In: Bates, J. W. & A.W. Farmer (eds), Bryophytes and lichens in a changing ‘environment, pp. 232-256, Clarendon Press, Oxford Gndstein, S, R 1994, Lejeuneacese: _Prychantheae, Brachiolejuneae, Flora Neotropica Monograph 62: 1-216, Gnadstein, $.R. Pécs, T. 1989. Bryophytes. In: Lieth, H. & M. J.A, Werger (eds.), Tropical rain forest ecosystems, pp. 311.325, Elsevier, Amsterdam. Grole, R. 1984, Zur Kenntnis der Lejeuneoideae in Cuba (1): Cyelolejeunea, Wissenschafliche Zeitschrift der Friedrich Schille- Universitit Jena, Naturwiss. R., 33: 759-764, Lebermoosgattung Jover-Ast, §. 195% Le genre Colura. Revue Bryologique et Lichénologique 22: 206-312, Licking, A. 1995, Diversitit und Microhabitapriferenzen epinhyller Moote in einem tropischen Regenwald in Costa Rica. Disseration, Ulm Montfoor, D. & R. Ek. 1990. Verical distribution and ecology of epiphytic bryophytes in a topical lowland rain forest in French Guiana. Institue of Systematic Botany, Unrecht, Oliver, W. RB. 1930, New Zealand epiphytes. J. Eco. 8: 1-50, Pécs, T. 1978. Epipyllous communities and their disibution in East Aftca. Bryophyt Biblioth. 13: 681-713, 19 P6cs, T. 1995. Epiphyllous liverwort diversity at world-wide level 8nd its threat and conservation. In: Delgadillo, C. (ed), Abstracts, Intemational Bryclogical Conference, August 7 12, 1995, Mexico City, p. 40-41. Instiuo de Biologia, UNAM, Mexico. Richards, P.W, 1984, The ecology of topical rain forest bryophytes. In: Schuster, RM. (ed.), New Manual of Bryology, pp. 1233-1270. Hattori Botanical Laboratory, Nichinan, Schuster, RM, 1991. On neotenic species of Radula, Journal of the Hattori Botanical Laboratory. 70: $1-62, ‘Sérusiaux, E, 1977. Les lichens folicoles: concept, classification écologique et postion systématique, Les Naturalistes Belges| 58: HIB, ‘Swartz, 0. 1788, Nova Genera & Species Plantarum, Stockholm, ‘Swartz, O. 1806, Flora Indize Occidentalis, Vo. 3. Erlangen ‘Teeuwen, M, 1989, A revision of the genus Odontolejeunea. Nova Hedwigia 48: 1-32, Thiers, B. M. 1988, Morphological adaptations of the Jungermanniales (Hepaticae) to the tropical rainforest habitat, Joumal of the Hattori Botanical Laboratory. 64: 5- 14 Tinxier, P. 1973. Le genre Tuumaella Hat (Lajeunéacées) Monographie. Revue Bryologique et Lichénologique 38: 221-244, Weber, F. von, 1815, Historiae Muscorum Hepaticarum Prodromus. Kiel, ‘Winkler, S. 1967. Die epiphyllen Moose der Nebelwalder von EI Salvadar, C. A. Revue Bryologique et Lichénologique 35: 303-369,