BLUMEA 51: 221– 280 Published on 27 July 2006 http://dx.doi.org/10.3767/000651906X622210 A TAXONOMIC REVISION OF HYMENOPHYLLACEAE ATSUSHI EBIHARA 1, 2, JEAN-YVES DUBUISSON 3, KUNIO IWATSUKI 4, SABINE HENNEQUIN 3 & MOTOMI ITO 1 SUMMARY A new classification of Hymenophyllaceae, consisting of nine genera (Hymenophyllum, Didymoglossum, Crepidomanes, Polyphlebium, Vandenboschia, Abrodictyum, Trichomanes, Cephalomanes and Callistopteris) is proposed. Every genus, subgenus and section chiely corresponds to the monophyletic group elucidated in molecular phylogenetic analyses based on chloroplast sequences. Brief descriptions and keys to the higher taxa are given, and their representative members are enumerated, including some new combinations. Key words: filmy ferns, Hymenophyllaceae, Hymenophyllum, Trichomanes. INTRODUCTION The Hymenophyllaceae, or ‘filmy ferns’, is the largest basal family of leptosporangiate ferns and comprises around 600 species (Iwatsuki, 1990). Members are easily distinguished by their usually single-cell-thick laminae, and the monophyly of the family has not been questioned. The intrafamilial classification of the family, on the other hand, is highly controversial – several fundamentally different classifications are used by individual researchers and/or areas. Traditionally, only two genera – Hymenophyllum with bivalved involucres and Trichomanes with tubular involucres – have been recognized in this family. This scheme was expanded by Morton (1968) who hierarchically placed many subgenera, sections and subsections under two larger genera Hymenophyllum and Trichomanes and recognized four monotypic genera, Cardiomanes, Hymenoglossum, Rosenstockia and Serpyllopsis, and all of which have specialized fronds. Since his system carefully covered New World taxa, most of the recent floristic works of Central and South America adopted his scheme (Stolze, 1976; De la Sota, 1977; Kramer, 1978; Smith, 1981; Tryon & Tryon, 1982; Proctor, 1985, 1989; Mickel & Beitel, 1988; Lellinger, 1989; Tryon & Stolze, 1989; Marticorena & Rodriguez, 1995; Pacheco, 1995; Baksh-Comeau, 2000; Sánchez, 2000). Conversely, Copeland (1938b, 1947) split the 1) Department of System Sciences, Graduate School of Arts and Sciences, The University of Tokyo, 3-8-1 Komaba, Meguro-ku, Tokyo 153-8902, Japan. Present address: Department of Botany, The National Science Museum, 4-1-1 Amakubo, Tsukuba 305-0005, Japan; ebihara@kahaku.go.jp. 2) Research fellow of the Japan Society for the Promotion of Science. 3) Équipe ‘Paléobiodiversité, Systématique et Évolution des Embryophytes’ UMR 5143, Laboratoire de Paléobotanique et Paléoécologie, IFR 101 CNRS ‘Institut d’Ecologie Fondamentale et Appliquée’, Université Pierre et Marie Curie, 12 rue Cuvier, F-75005 Paris, France. 4) The Museum of Nature and Human Activities, Hyogo, Yayoigaoka 6-chome, Sanda 669-1546, Japan. © 2006 Nationaal Herbarium Nederland, Leiden University branch 222 BLUMEA — Vol. 51, No. 2, 2006 family into 34 genera, basing his work mainly on in situ and herbarium observation of Asian material. His scheme has been adopted in floristic works of Asia (Copeland, 1958; Ching, 1959; De Vol, 1975; Nakaike, 1975; Tagawa & Iwatsuki, 1979; Parris, 1992), but has not yet been applied to New World taxa. Another system consisting of 47 genera was proposed by Pichi Sermolli (1977b), but it has been adopted only in a limited number of publications on African plants (e.g., Pichi Sermolli, 1983). Finally, Iwatsuki integrated a series of his studies of morphological characters (Iwatsuki, 1975, 1977a, b, 1978, 1981, 1982) into a new hierarchical system consisting of eight genera (Iwatsuki, 1984, 1985). Although his system has been adopted in a few recently published floristic works (Iwatsuki, 1995; Du Puy & Orchard, 1993; Green, 1994; Bostock & Spokes, 1998), generic combinations for taxa in regions other than Asia and Australia are still wanting. These taxonomic inconsistencies were compounded with the typification problem of the Linnaean genus Trichomanes (see Holttum, 1976 and Pichi Sermolli, 1981) – Trichomanes crispum was considered the type in the systems devised by Copeland (1933, 1938b), Pichi Sermolli (1977b) and Iwatsuki (1984), while T. scandens was considered the type by Morton (1968) and Nakaike (1975). Recent molecular phylogenetic studies (Dubuisson, 1997; Pryer et al., 2001; Dubuisson et al., 2003a; Hennequin et al., 2003; Ebihara et al., 2004; Ebihara et al., submitted) have provided new insights into the systematic relationships within the Hymenophyllaceae. The presence of two large monophyletic groups (Pryer et al., 2001) can be interpreted to support the traditional bigeneric system; however some traditional Trichomanes taxa (Cardiomanes, Microtrichomanes, Pleuromanes) belong in fact to the Hymenophyllum lineage (Pryer et al., 2001; Hennequin et al., 2003; Ebihara et al., 2004; Ebihara et al., submitted). Moreover, the trichomanoid and hymenophylloid lineages, both consisting of nearly the same number of taxa, may possess different historical backgrounds – the Hymenophyllum lineage has likely diversified more recently than the Trichomanes lineage (Ebihara et al., 2004; Schuettpelz & Pryer, in press). This may explain why the Trichomanes lineage displays a greater variety of morphological features and ecological habits than the Hymenophyllum lineage. Earlier studies on Trichomanes (Dubuisson, 1997; Dubuisson et al., 2003a), as well as our latest study (Ebihara et al., submitted), also based on rbcL sequences and representing nearly all taxonomic lineages, did not support any of the above classifications. These inconsistencies are probably caused by strong homoplasy of taxonomic characters due to adaptive evolution (Dubuisson et al. 2003b; Ebihara et al., submitted). We therefore believe that a fundamental taxonomic revision of the Hymenophyllaceae is in order. Even today, the principles of generic delimitation maintained by Copeland (1938b) seem quite reasonable: naturalness and convenience, and the latter is “subject always to the preceding principle (naturalness)”. It is often argued that a classification completely based on the results of molecular phylogeny is difficult to put into practice, but recognition of unnatural (i.e., polyphyletic/paraphyletic) groups is a disadvantage for biological studies in general. Hence we prefer to reorganize the species into natural (i.e., monophyletic) and also morphologically recognizable groups. We propose to place all species of the hymenophylloid lineage under a single genus Hymenophyllum and to subdivide the trichomanoid lineage into eight genera corresponding to eight robust clades in our global analysis (Ebihara et al., submitted). We also treat many of the subclades retrieved (Ebihara et al., submitted) as subgenera if relationships within the A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 223 Fig. 1. An explanatory figure of the correspondence between the present system and the results of molecular phylogeny of Hymenophyllaceae. The phylogram is primarily based on the MP tree using chloroplast rbcL gene (Ebihara et al., submitted); the topology within the ‘H’ clade follows that of the MP tree using rbcL, rbcL-accD and rps4-trnS regions (Hennequin et al., in press). Bootstrap values (> 50%) supporting the clades are shown on their branches (bootstrap supports > 80% are shown by bold lines), but the values are replaced by asterisks in the taxa comprising a single sample. All the groups adopted as taxa in the present system are supported by high posterior probabilities (> 0.99) of Bayesian inference (Ebihara et al., submitted; Hennequin et al., in press). The clade names indicated in the tree correspond to those used in Ebihara et al. (submitted). — Ab: Abrodictyum; Ac: Achomanes; Ca: Callistopteris; Ce: Cephalomanes; Cr: Crepidomanes; Da: Davalliopsis; Di: Didymoglossum; Fe: Feea; H: Hymenophyllum s.l.; Le: Lecanium; Mg: Microgonium; Ne: Nesopteris; NT: Neotropical; Pa: Pachychaetum; PT: Paleotropical; Se: Selenodesmium; T: Trichomanes s.l.; TP: Trichomanes pixidiferum; Va: Vandenboschia. clade are well resolved in our analysis (Fig. 1); in the absence of support, infrageneric classification within some genera is deferred. Although Ebihara et al. (submitted) illustrated that anatomical and cytological characters are more taxonomically informative than macroscopic characters, we selected macroscopic characters for the keys in this paper even though this makes the keys more complex. 224 BLUMEA — Vol. 51, No. 2, 2006 To aid in characterization of taxa recognized by us, representative species of each genus, subgenus or section are enumerated here. Please note that the list includes the minimum number of taxa commonly accepted in recent floristic works, and therefore does not necessarily represent all the members. A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 225 TAXONOMIC TREATMENT KEY TO THE GENERA 1a. Rhizomes long-creeping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 b. Rhizomes short-creeping or erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 2a. Rhizomes nearly glabrous or sparsely covered with light-coloured hairs. Involucres usually bivalvate (Fig. 2a) . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Hymenophyllum b. Rhizomes covered with reddish to dark-coloured hairs. Involucres usually not bivalvate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Roots absent or replaced by root-like shoots (Fig. 2c) . . . . . . . . . . . . . . . . . . . 4 b. Roots present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4a. Continuous false veinlets parallel to true veins (Fig. 2d) present. Blade venation catadromous (Fig. 2e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Didymoglossum b. Continuous false veinlets parallel to true veins absent. Blade venation anadromous (Fig. 2f). . . . . . . . . . . . . . . . . . . 3. Crepidomanes (subg. Crepidomanes) 5a. Roots few and fine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Polyphlebium b. Roots numerous and robust . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6a. Blades once-pinnate to bipinnatifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 b. Blades bipinnate to highly divided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7a. Pinnae asymmetric . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Vandenboschia b. Pinnae symmetric . . . . . . . . . . . . . . . . . . . . . 7. Trichomanes (subg. Lacostea) 8a. Marginal laminar trichomes present . . . . 7. Trichomanes (subg. Trichomanes) b. Marginal laminar trichomes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9a. Abortive fronds (Fig. 2g) present. . . . . . . 3. Crepidomanes (subg. Nesopteris) b. Abortive fronds absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a. Minute clavate hairs present on stipes and rachises . . . . . . . 5. Vandenboschia b. Minute clavate hairs on stipes and rachises absent . . . . . . . . . . 6. Abrodictyum 11a. Involucres bivalvate (Fig. 2a) . . . . . . . 1. Hymenophyllum (subg. Fuciformia) b. Involucres tubular (Fig. 2b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12a. Blade venation catadromous (Fig. 2e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 b. Blade venation anadromous (Fig. 2f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13a. Lamina tough, more than one cell thick . . . . . . . . 7. Trichomanes (subg. Feea) b. Lamina membranous, one cell thick . . . . 7. Trichomanes (subg. Trichomanes) ← Fig. 2. Examples of morphological characters used in the keys. a. Bivalvate involucres (Hymenophyllum deplanchei; Ebihara 001228-05, TI); b. tubular involucres with an exserted receptacle (Rc) (Crepidomanes humile; Iwatsuki 99H06, TNS); c. root-like shoots (RS) grown out from rhizomes (Rz) (Crepidomanes bipunctatum; Ebihara 000225-020, TNS); d. continuous false veinlets (FV) parallel to true veins (TV) (Didymoglossum bipunctatum; S. Matsumoto 01-820, TNS); e. catadromous venation – FAV: the first acroscopic vein; FBV: the first basiscopic vein in each pinna (Trichomanes polypodioides; T.A. Ohsawa 178-15, TNS); f. anadromous venation (Vandenboschia birmanica; A. Narita 1070b, TNS); g. abortive fronds (pointed by an arrow) (Crepidomanes aphlebioides; M. Kato et al. C-13883, TI); h. long bristle-like light reddish hairs on a stipe (Callistopteris apiifolia; Ebihara 040922-03, TNS); i. reduced lamina cells (Abrodictyum pluma; R. Jaman s.n., TNS). — Scale bars: a, b, d, h = 1 mm; c, e, f, i = 5 mm; g = 5 cm. 226 BLUMEA — Vol. 51, No. 2, 2006 14a. Blades once-pinnate to bipinnatifid . . . . . . . . . . . . . . . . . . . . . 8. Cephalomanes b. Blades bipinnate or more divided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15a. Long bristle-like light reddish hairs (Fig. 2h) present on stipes and rachises . 16 b. Long bristle-like light reddish hairs absent on stipes and rachises . . . . . . . . . 17 16a. Laminar cells reduced (Fig. 2i). Less than 3 rows of cells between midribs and margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. Abrodictyum b. Laminar cells not reduced. More than 3 rows of cells between midribs and margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. Callistopteris 17a. Laminae more than one cell thick . . . . . . 7. Trichomanes (subg. Davalliopsis) b. Laminae one cell thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18a. Laminar cell-walls thin and straight . . . . 2. Crepidomanes (subg. Nesopteris) b. Laminar cell-walls slightly or quite thick, wavy or pitted . . . . . 6. Abrodictyum 1. HYMENOPHYLLUM Hymenophyllum Sm. (1793) 418. — Lectotype: Hymenophyllum tunbrigense (L.) Sm. (selected by Presl (1843) 31). Rhizomes long-creeping, frequently branching, usually filiform or wiry, up to 2 mm diam., nearly glabrous or sparsely covered with multicellular hairs (exceptionally densely covered with hairs in subg. Fuciformia), protostele subcollateral, dorsiventral, or ‘reduced’ (with more or less central xylem-like parenchyma), cortices heterogeneous, with few and fine roots. Stipes various in length, at a distance from the adjacent ones. Blades usually pinnate to quadripinnate or occasionally simple, 165 by 17 cm, venation anadromous, laminae usually one cell thick, but sometimes two or more cells thick. Sori paratactic, lips usually bivalvate, receptacles usually included in involucres. Distribution — Throughout the tropics to temperate regions; c. 250 species. Habitat — Usually low- to middle-epiphytes on tree trunks, sometimes canopy sunepiphytes, occasionally epilithic or terrestrial. Chromosome base number — Various, x = 11 to 36. Note — Hymenophyllum in our sense stands as the largest genus of Hymenophyllaceae, and almost corresponds to Hymenophyllum in the traditional and broad sense but also includes some taxa formerly regarded as Trichomanes s.l. Iwatsuki (1984) segregated Sphaerocionium, which is characterized by distinctive stellate hairs, from the remaining Hymenophyllum (s.l.) taxa. Recent studies (Hennequin et al., 2003, in press) suggest that Sphaerocionium is nested within Mecodium, another easily recognizable group of Hymenophyllum s.l., as currently defined. Under our strategy of classification – making each taxon a natural group – Sphaerocionium is to be treated as a genus, species belonging to Mecodium should be split into more than two genera. To avoid such inconvenience, all species of Hymenophyllum s.l. are placed in our Hymenophyllum. Because the systematic relationships within this genus are not fully clarified, the current revision treats only subgenera, with each subgenus monophyletic based on our latest study using molecular data (Hennequin et al., in press). A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 227 KEY TO THE SUBGENERA 1a. Rhizomes short-creeping or erect . . . . . . . . . . . . . . . . . . . 8. Subg. Fuciformia b. Rhizomes long-creeping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Stellate hairs present on blades . . . . . . . . . . . . . . . . . 2. Subg. Sphaerocionium b. Stellate hairs absent on blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Blades dichotomously divided . . . . . . . . . . . . . . . . . 2. Subg. Sphaerocionium b. Blades not dichotomously divided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4a. Blades simple to pinnatifid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 b. Blades pinnate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 5a. Blades kidney-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . 10. Subg. Cardiomanes b. Blades not kidney-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6a. Veins of blades simple . . . . . . . . . . . . . . . . . . . . . . . . 1. Subg. Hymenophyllum b. Veins of blades pinnately divided . . . . . . . . . . . . . . . 7. Subg. Hymenoglossum 7a. Blades glaucous or covered with whitish multicellular hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Subg. Pleuromanes b. Blades neither glaucous nor covered with whitish multicellular hairs . . . . . . . 8 8a. Laminae more than two cells thick throughout . . . . . . 9. Subg. Diploöphyllum b. Laminae one cell thick, or only partly multi-layered. . . . . . . . . . . . . . . . . . . . . 9 9a. Stipes with multicellular hairs present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 b. Stipes glabrous or with only caducous hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10a. Sori on both acroscopic and basiscopic sides of pinnae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. Subg. Myrmecostylum b. Sori confined to acroscopic sides of pinnae . . . . . . . 1. Subg. Hymenophyllum 11a. Rhizomes thick, more than 0.4 mm diam., nearly glabrous or with scattered pale hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Subg. Globosa b. Rhizomes filiform, less than 0.4 mm diam., with scattered short brown hairs 12 12a. Margins of segments toothed . . . . . . . . . . . . . . . . . . . 1. Subg. Hymenophyllum b. Margins of segments entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13a. Lamina cell walls thin and straight, sori lacking a thickened base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Subg. Mecodium b. Lamina cell walls more or less thick, sori each with a thickened base, on short acroscopic segments near the rachis . . . . . . . . . . . . . 1. Subg. Hymenophyllum 1. Subgenus Hymenophyllum Didymoglossum Desv. subg. Chilodium C. Presl (1843) 23. — Hymenophyllum Sm. subg. Chilodium (C. Presl) Croxall (1975) 515. — Hymenophyllum Sm. sect. Chilodium (C. Presl) C. Chr. (1906) XVI. — Lectotype: Didymoglossum neesii (Blume) C. Presl (= Hymenophyllum denticulatum Sw.) (selected by Christensen (1906) XVI, as H. denticulatum Sw.). Meringium C. Presl (1843) 24, t. VIII, f. B. — Hymenophyllum Sm. subg. Meringium (C. Presl) Copel. (1937) 14. — Hymenophyllum Sm. sect. Meringium (C. Presl) Diem & J.S. Licht. (1959) 666. — Type: Meringium meyenianum C. Presl (= Hymenophyllum serrulatum (C. Presl) C. Chr.). Leptocionium C. Presl (1843) 26, t. XI, f. D. — Hymenophyllum Sm. sect. Leptocionium (C. Presl) Prantl (1875) 54. — Hymenophyllum Sm. subg. Leptocionium (C. Presl) H. Christ (1897) 20. — Hymenophyllum Sm. subsect. Leptocionium (C. Presl) C.V. Morton (1968) 170. — Type: Leptocionium dicranotrichum C. Presl (= Hymenophyllum dicranotrichum (C. Presl) Sadeb.). 228 BLUMEA — Vol. 51, No. 2, 2006 Hymenophyllum Sm. subg. Sphaerodium C. Presl (1843) 31. — Hymenophyllum Sm. sect. Sphaerodium (C. Presl) C. Chr. (1906) XVI. — Lectotype: Hymenophyllum wilsonii Hook. (selected by Copeland (1938b) 14). Hymenophyllum Sm. subsect. Dimidiata C. Presl (1843) 32. — Lectotype: Hymenophyllum unilaterale Bory ex Willd. (= Hymenophyllum peltatum (Poir.) Desv.) (selected by Morton (1968) 164). Hymenophyllum Sm. subg. Cycloglossum C. Presl (1843) 32. — Hymenophyllum Sm. sect. Cycloglossum (C. Presl) C. Chr. (1906) XV. — Lectotype: Hymenophyllum caespitosum Gaudich. (selected by Copeland (1938b) 14). Hymenophyllum Sm. subg. Craspedophyllum C. Presl (1843) 33. — Hymenophyllum Sm. sect. Craspedophyllum (C. Presl) C. Chr. (1906) XV. — Craspedophyllum (C. Presl) Copel. (1938b) 27. — Type: Hymenophyllum marginatum Hook. & Grev. Serpyllopsis Bosch (1861a) 318. — Trichomanes L. subg. Serpyllopsis (Bosch) H. Christ (1897) 23. — Type: Serpyllopsis caespitosa (Gaudich.) C. Chr. (= Hymenophyllum caespitosum Gaudich.). Hymenophyllum Sm. sect. Pachyloma H. Christ (1897) 15, based on Pachyloma Bosch (1861a) 318, nom. illeg., non DC. (1828) 122, nec Spach (1839) 194. — Type: Hymenophyllum marginatum Hook. & Grev. Hymenophyllum Sm. subg. Hemicyatheon Domin (1913) 20. — Hemicyatheon (Domin) Copel. (1938b) 27. — Hymenophyllum Sm. sect. Hemicyatheon (Domin) K. Iwats. (1984) 172. — Type: Hymenophyllum baileyanum Domin. Hymenophyllum Sm. sect. Acanthotheca Nakai (1926) 262. — Type: Hymenophyllum acanthoides (Bosch) Rosenst. Hymenophyllum Sm. subg. Buesia C.V. Morton (1932) 336. — Buesia (C.V. Morton) Copel. (1938b) 47. — Hymenophyllum Sm. sect. Buesia (C.V. Morton) C.V. Morton (1968) 164. — Type: Hymenophyllum miriicum C.V. Morton. Hymenophyllum Sm. subg. Amphipterum C. Presl ex Copel. (1937) 68, based on Amphipterum C. Presl (1849) 258, nom. nud. — Amphipterum (C. Presl ex Copel.) Copel. (1938b) 46. — Hymenophyllum Sm. sect. Amphipterum (C. Presl ex Copel.) C. Chr. ex K. Iwats. (1984) 171. — Type: Hymenophyllum fuscum (Blume) Bosch. Hymenophyllum Sm. subg. Myriodon Copel. (1937) 73. — Myriodon (Copel.) Copel. (1938b) 47. — Hymenophyllum Sm. sect. Myriodon (Copel.) C.V. Morton (1968) 168. — Type: Hymenophyllum odontophyllum Copel. (= Hymenophyllum brassii C. Chr.). Rosenstockia Copel. (1947) 36. — Hymenophyllum Sm. subg. Rosenstockia (Copel.) R.M. Tryon & A.F. Tryon (1981) 134. — Type: Rosenstockia rolandi-principis (Rosenst.) Copel. (= Hymenophyllum rolandi-principis Rosenst.). Hymenophyllum Sm. sect. Eupectinum Diem & J.S. Licht. (1959) 635. — Type: Hymenophyllum pectinatum Cav. Hymenophyllum Sm. sect. Pseudomecodium K. Iwats. (1984) 172. — Pseudomecodium (K. Iwats.) Satou (1997) 269. — Type: Hymenophyllum exsertum (Wall. ex Hook.) Copel. Hymenophyllum Sm. subg. Euhymenophyllum C. Presl (1843) 31, nom. illeg.; originally includes the lectotype of the genus, Hymenophyllum tunbrigense (L.) Sm. Hymenophyllum Sm. sect. Serrulata C. Presl (1843) 32, nom. illeg.; originally includes the lectotype of the genus, Hymenophyllum tunbrigense (L.) Sm. Hymenophyllum Sm. subsect. Evoluta C. Presl (1843) 32, nom. illeg.; originally includes the lectotype of the genus, Hymenophyllum tunbrigense (L.) Sm. Hymenophyllum Sm. sect. Plumosa K. Iwats. (1984) 172, nom. illeg., non Hymenophyllum Sm. subsect. Plumosa Prantl (1875) 55. — Type: Hymenophyllum levingei C.B. Clarke. Rhizomes long-creeping, filiform, nearly glabrous or sparsely covered with thick-walled hairs. Stipes to 15 cm long. Blades simple to quadripinnatifid, narrowly elliptic to subdeltate, 30 by 15 cm, margins of segments entire. Sori usually at the tips of acroscopic segments of pinnae, lips bivalvate, entire or serrate, receptacles included or slightly extruded from involucres. Distribution — Cosmopolitan; c. 100 species. Habitat — Usually epiphytic, occasionally epilithic. A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 229 Chromosome base number — Various, x = 11 to 28. Note — This is the largest subgenus of Hymenophyllum and shows great diversity both in morphology and cytology. Since recent molecular results (Ebihara et al., 2002; Hennequin et al., 2003; Hennequin et al., in press) suggest most of the subgroups included in this subgenus are probably polyphyletic, further study is necessary for the classification within this subgenus. REPRESENTATIVE SPECIES: New combinations: Hymenophyllum trichophorum (Alderw.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes trichophorum Alderw. (1924) 53, the earlier name for the plant formerly called Amphipterum humatoides Copel. (1941) 465]. Other species: Hymenophyllum acanthoides (Bosch) Rosenst.; H. armstrongii (Baker) Kirk; H. baileyanum Domin; H. barbatum (Bosch) Baker; H. bivalve (G. Forst.) Sw.; H. blandum Racib.; H. brassii C. Chr.; H. caespitosum Gaudich.; H. cupressiforme Labill.; H. dentatum Cav.; H. denticulatum Sw.; H. deplanchei Mett. ex Kuhn; H. dicranotrichum (C. Presl) Sadeb.; H. dimidiatum Mett. ex Kuhn; H. exsertum Wall. ex Hook.; H. feejeense Brack.; H. foersteri Rosenst.; H. fucoides (Sw.) Sw.; H. fuscum (Blume) Bosch; H. geluense Rosenst.; H. gorgoneum Copel.; H. holochilum (Bosch) C. Chr.; H. johorense Holttum; H. ledermanni Brause; H. levingei C.B. Clarke; H. lobbii T. Moore ex Bosch; H. macrosorum Alderw.; H. marginatum Hook. & Grev.; H. melanosorum (Copel.) C.V. Morton; H. minimum A. Rich.; H. miriicum C.V. Morton; H. multiidum (G. Forst.) Sw.; H. oligosorum Makino; H. pachydermicum Ces.; H. pectinatum Cav.; H. peltatum (Poir.) Desv.; H. penangianum C.G. Matthew & H. Christ; H. pumilum C. Moore; H. revolutum Colenso; H. rolandi-principis Rosenst.; H. rubellum Rosenst.; H. rugosum C. Chr. & Skottsb.; H. secundum Hook. & Grev.; H. serrulatum (C. Presl) C. Chr.; H. sibthorpioides (Bory ex Willd.) Mett.; H. simonsianum Hook.; H. tenellum (Jacq.) Kuhn; H. triangulare Baker; H. tunbrigense (L.) Sm.; H. walleri Maiden & Betche; H. wilsonii Hook. 2. Subgenus Sphaerocionium (C. Presl) C. Chr. Hymenophyllum Sm. subg. Sphaerocionium (C. Presl) C. Chr. (1934) 5. — Sphaerocionium C. Presl (1843) 33, t. IV, f. B, t. X, f. B, C. — Hymenophyllum Sm. sect. Sphaerocionium (C. Presl) C. Chr. (1906) XV. — Lectotype: Hymenophyllum hirsutum (L.) Sw. (selected by Copeland (1937) 10). Trichomanes L. sect. Flabellata C. Presl (1843) 16. — Lectotype: Trichomanes digitatum Sw. (= Hymenophyllum digitatum (Sw.) Fosberg) (selected by Morton (1968) 199). Hymenophyllum Sm. sect. Pilosa C. Presl (1843) 32. — Lectotype: Hymenophyllum valvatum Hook. & Grev. (selected by Morton (1968) 168). Sphaerocionium C. Presl sect. Pilosa C. Presl (1843) 34. — Lectotype: Sphaerocionium schiedeanum C. Presl (= Hymenophyllum crispum Kunth) (selected by Morton (1968) 168). Sphaerocionium C. Presl sect. Stellata C. Presl (1843) 34. — Lectotype: Sphaerocionium interruptum (Kunze) C. Presl (= Hymenophyllum plumieri Hook. & Grev.) (selected by Morton (1968) 169). Gonocormus Bosch sect. Microtrichomanes Mett. ex Prantl (1875) 51, based on Gruppe Microtrichomanes Mett. (1864) 413, nom. nud. — Trichomanes L. sect. Microtrichomanes (Mett. ex Prantl) C. Chr. (1906) XIV. — Microtrichomanes (Mett. ex Prantl) Copel. (1938b) 35. — Crepidomanes (C. Presl) C. Presl subg. Microtrichomanes (Mett. ex Prantl) K. Iwats. (1984) 175. — Lectotype: Gonocormus digitatus (Sw.) Prantl (= Hymenophyllum digitatum (Sw.) Fosberg) (selected by Christensen (1906) XIV). Hymenophyllum Sm. subsect. Ciliata Prantl (1875) 55. — Lectotype: Hymenophyllum lineare (Sw.) Sw. (selected by Morton (1968) 168). Hymenophyllum Sm. subsect. Hirsuta Prantl (1875) 55. — Lectotype: Hymenophyllum raddianum Müll.Berol. (= Hymenophyllum fragile (Hedw.) C.V. Morton) (selected by Morton (1968) 170). 230 BLUMEA — Vol. 51, No. 2, 2006 Hymenophyllum Sm. subsect. Plumosa Prantl (1875) 55. — Lectotype: Hymenophyllum plumosum Kaulf. (selected by Morton (1968) 169). Hymenophyllum Sm. subg. Apteropteris Copel. (1937) 176. — Apteropteris (Copel.) Copel. (1938b) 34. — Hymenophyllum Sm. sect. Apteropteris (Copel.) C.V. Morton (1968) 170. — Sphaerocionium C. Presl subg. Apteropteris (Copel.) K. Iwats. (1984) 173. — Type: Hymenophyllum malingii (Hook.f.) Mett. Rhizomes long-creeping, filiform, usually covered with stellate hairs. Stipes to 18 cm long. Blades pinnate to tripinnatifid, elliptic to subdeltate, 165 by 17 cm, covered with stellate hairs, or with simple hairs. Sori at the tips of ultimate segments, bivalvate or tubular, lips entire, receptacles included or slightly extruded from involucres. Distribution — Cosmopolitan, highly diversified in the Neotropics; c. 70 species. Habitat — Epiphytic on tree trunks. Chromosome base number — x = 36. Note — Though the Sphaerocionium species characterized by their stellate hairs have sometimes been treated as an independent genus (e.g., Copeland, 1938b; Iwatsuki, 1984), we reduced them to a subgenus of Hymenophyllum for the reason noted in the description of the genus. Our present definition of Sphaerocionium also includes most of the Microtrichomanes species (Ebihara et al., 2004) whose hairs have been transformed from stellate into simple. Although Morton (1947) subdivided this group chiefly by the nature and placement of the hairs, recent studies (Ebihara et. al., 2004, Hennequin et al., in press) suggest their phylogenetic relationships are more closely related to their geographical distributions. REPRESENTATIVE SPECIES: Hymenophyllum adiantoides Bosch; H. aeruginosum (Poir.) Carmich.; H. amabile C.V. Morton; H. angustum Bosch; H. antillense (Jenman) Jenman; H. braithwaitei Ebihara & K. Iwats.; H. capillare Desv.; H. consanguineum C.V. Morton; H. crispum Kunth; H. dependens C.V. Morton; H. digitatum (Sw.) Fosberg; H. elegans A. Spreng.; H. elegantulum Bosch; H. ferrugineum Colla; H. fragile (Hedw.) C.V. Morton; H. frankliniae Colenso; H. fusugasugense H. Karst. ex Sturm; H. glaziovii Baker; H. hemipteron Rosenst.; H. hirsutum (L.) Sw.; H. hirtellum Sw.; H. horizontale C.V. Morton; H. karstenianum Sturm; H. lanatum Fée; H. lanceolatum Hook. & Arn.; H. lindenii Hook.; H. lineare (Sw.) Sw.; H. lobato-alatum Klotzsch; H. lyallii Hook.f.; H. malingii (Hook.f.) Mett.; H. microcarpum Desv.; H. molle C.V. Morton; H. multialatum C.V. Morton; H. nitidulum (Bosch) Ebihara & K. Iwats.; H. obtusum Hook. & Arn.; H. palmatiidum (Müll. Berol.) Ebihara & K. Iwats.; H. pilosissimum C. Chr.; H. plumieri Hook. & Grev.; H. plumosum Kaulf.; H. pulchellum Schltdl. & Cham.; H. pyramidatum Desv.; H. roraimense C.V. Morton; H. ruizianum (Klotzsch) Kunze; H. semiglabrum Rosenst.; H. sericeum (Sw.) Sw.; H. sieberi (C. Presl) Bosch; H. simplex C.V. Morton; H. speciosum Bosch; H. splendidum Bosch; H. subobtusum Rosenst.; H. subrigidum H. Christ; H. tomaniiviense (Brownlie) Ebihara & K. Iwats.; H. tomentosum Kunze; H. trapezoidale Liebm.; H. trichophyllum Kunth; H. urbanii Brause; H. valvatum Hook. & Grev.; H. verecundum C.V. Morton. 3. Subgenus Mecodium C. Presl ex Copel. Hymenophyllum Sm. subg. Mecodium C. Presl ex Copel. (1937) 93, based on Mecodium C. Presl (1849) 258, nom. nud. — Mecodium (C. Presl ex Copel.) Copel. (1938b) 17. — Lectotype: Hymenophyllum polyanthos (Sw.) Sw. (selected by Copeland (1937) 10). Hymenophyllum Sm. sect. Glabra Prantl (1875) 54. — Lectotype: Hymenophyllum polyanthos (Sw.) Sw. (selected by Morton (1968) 172). Hymenophyllum Sm. sect. Corrugatae K. Iwats. (1984) 173. — Type: Hymenophyllum ooides F. Muell. & Baker. A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 231 Rhizomes long-creeping, filiform, nearly glabrous. Stipes up to 10 cm long. Blades pinnate to tripinnatifid, elliptic to subdeltate, 45 by 6 cm, margins of segments entire. Sori at the tips of ultimate segments, lips bivalvate, entire, receptacles included in involucres. Distribution — Cosmopolitan; more than 35 species. Habitat — Epiphytic on tree trunks. Chromosome base number — x = 28. Note — This subgenus corresponds to the cosmopolitan species H. polyanthos and its local derivatives; it also corresponds to the ‘H. polyanthos clade’ in Hennequin et al. (in press). The chromosome base number of x = 28 is a synapomorphy of this subgenus. Although Presl (1849) originally used the generic name Mecodium for H. sanguinolentum, which is now referred to subg. Myrmecostylum, Copeland (1937) selected H. polyanthos as the type when he legitimized the name – the type designation may not be correctable even it was later corrected to H. sanguinolentum by Copeland himself (1938b). REPRESENTATIVE SPECIES: New combinations: Hymenophyllum novoguineense (Rosenst.) K. Iwats., comb. nov. [based on Hymenophyllum blumeanum A. Spreng. var. novoguineense Rosenst. (1908b) 371]. Other species: Hymenophyllum abruptum Hook.; H. apiculatum Mett. ex Kuhn; H. axillare Sw.; H. brevifrons Kunze; H. copelandii C.V. Morton; H. corrugatum H. Christ; H. cuneatum Kunze; H. darwinii Hook.; H. fendlerianum Sturm; H. fumarioides Bory ex Willd.; H. inaequale (Poir.) Desv.; H. kuhnii C. Chr.; H. mnioides Baker; H. myriocarpum Hook.; H. ooides F. Muell. & Baker; H. paniculilorum C. Presl; H. polyanthos (Sw.) Sw.; H. rarum R. Br.; H. recurvum Gaudich.; H. siliquosum H. Christ; H. undulatum Sw.; H. whitei Goy; H. wrightii Bosch. 4. Subgenus Globosa (Prantl) Ebihara & K. Iwats., comb. nov. Based on Hymenophyllum Sm. sect. Globosa Prantl (1875) 55. — Lectotype: Hymenophyllum junghuhnii Bosch (selected by Morton (1968) 172). Sphaerocionium C. Presl sect. Glabra C. Presl (1843) 34, non Hymenophyllum Sm. sect. Glabra Prantl (1875) 54. — Lectotype: Hymenophyllum caudiculatum Mart. (selected by Morton (1968) 172). Rhizomes long-creeping, filiform to wiry, nearly glabrous. Stipes to 15 cm long. Blades tri- to quadripinnate, elliptic to subdeltate, 60 by 15 cm, margin of segments usually entire, minutely serrate in some species. Sori at the tips of ultimate segments, lips bivalvate, entire or serrate, receptacles included in involucres, often capitate. Distribution — Tropics to temperate regions of Asia and Pacific areas; at least one species is distributed in the southern part of South America; c. 25 species. Habitat — Epiphytic, or occasionally epilithic. Chromosome base number — x = 36. Note — This subgenus corresponds to the ‘H. australe clade’ in Hennequin et al. (in press). It has been confused with the H. polyanthos group (subg. Mecodium in the present system), but features of the rhizome (e.g., dorsiventral protostele) are clearly different (Hennequin et al., in press). The earliest name given to this group, ‘Sphaerocionium sect. Glabra’ (Presl, 1843) becomes a later homonym of Hymenophyllum sect. 232 BLUMEA — Vol. 51, No. 2, 2006 Glabra (Prantl, 1875) when it is placed under Hymenophyllum according to the latest Code (Greuter et al., 2000, Art. 21.3, Note 1). Therefore the name of this subgenus should be Globosa. REPRESENTATIVE SPECIES: Hymenophyllum australe Willd.; H. badium Hook. & Grev.; H. caudiculatum Mart.; H. demissum (G. Forst.) Sw.; H. emarginatum Sw.; H. imbriatum J. Sm. ex Hook.; H. lexuosum A. Cunn.; H. imbricatum Blume; H. javanicum A. Spreng.; H. junghuhnii Bosch; H. longifolium Alderw.; H. productum Kunze; H. reinwardtii Bosch; H. riukiuense H. Christ; H. salakense Racib.; H. thuidium Harr.; H. treubii Racib. 5. Subgenus Pleuromanes (C. Presl) Ebihara & K. Iwats., comb. nov. Based on Trichomanes L. subg. Pleuromanes C. Presl (1849) 17. — Pleuromanes (C. Presl) C. Presl (1849) 258. — Crepidomanes (C. Presl) C. Presl subg. Pleuromanes (C. Presl) K. Iwats. (1984) 174. — Type: Trichomanes acutum C. Presl (1843) 42 (= Hymenophyllum acutum (C. Presl) Ebihara & K. Iwats., comb. nov., non Hymenophyllum acutum Mett., nom. nud.). Craspedoneuron Bosch (1861a) 322. — Trichomanes L. sect. Craspedoneuron (Bosch) Prantl (1875) 52. — Type: Trichomanes album Blume (= Hymenophyllum pallidum (Blume) Ebihara & K. Iwats., comb. nov. [based on Trichomanes pallidum Blume (1828) 225]). Rhizomes long-creeping, filiform, sparsely covered with whitish/yellowish hairs. Stipes to 12 cm long. Blades bi- to tripinnate, elliptic to subdeltate, 35 by 6 cm, covered with whitish/yellowish multicellular hairs or almost glabrous, margins of segments entire. Sori at the tips of ultimate segments, lips slightly to distinctly bivalvate, entire, receptacles included or slightly exserted from involucres. Distribution — Old World tropics, especially in the Pacific region; 5 species. Habitat — Epiphytic on tree trunks (including tree ferns). Chromosome base number — x = 36. Note — This subgenus contains H. flabellatum plus its local forms and Pleuromanes, formerly considered a member of Trichomanes s.l.; it also corresponds to ‘H. flabellatum clade’ in Hennequin et al. (in press). REPRESENTATIVE SPECIES: Hymenophyllum acutum (C. Presl) Ebihara & K. Iwats.; H. labellatum Labill.; H. leratii Rosenst.; H. pallidum (Blume) Ebihara & K. Iwats.; H. rufescens Kirk. 6. Subgenus Myrmecostylum (C. Presl) Ebihara & K. Iwats., comb. nov. Based on Myrmecostylum C. Presl (1843) 27, t. X, f. A. — Hymenophyllum Sm. sect. Myrmecostylum (C. Presl) C. Chr. (1906) XVI. — Lectotype: Myrmecostylum tortuosum (Hook. & Grev.) C. Presl (= Hymenophyllum tortuosum Hook. & Grev.) (selected by Christensen (1906) XVI). Ptychophyllum C. Presl (1843) 28, t. XI, f. E. — Hymenophyllum Sm. sect. Ptychophyllum (C. Presl) C. Chr. (1906) XVI. — Type: Ptychophyllum plicatum (Kaulf.) C. Presl (= Hymenophyllum plicatum Kaulf.). Tetralasma Phil. (1860) 208. — Type: Hymenophyllum quadriidum Phil. (= Hymenophyllum plicatum Kaulf.), no generic combination has been made for this genus. Rhizomes long-creeping, filiform to wiry, sparsely covered with thick-walled hairs. Stipes to 15 cm long. Blades pinnatifid to quadripinnate, elliptic to subdeltate, 55 by 14 cm, margins of segments entire or serrate. Sori at the tips of ultimate segments, lips bivalvate, entire or serrate, receptacles included in involucres. A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 233 Distribution — Southern Chile, New Zealand and New Caledonia; at least 8 species are known. Habitat — Epiphytic on tree trunks, occasionally terrestrial. Chromosome base number — x = 36. Note — This subgenus corresponds to the ‘H. sanguinolentum clade’ in Hennequin et al. (in press), and is characterized by brown multicellular hairs on the stipes or blade margins, or alternatively by dentate blade margins. In exceptional cases, H. sanguinolentum has nearly glabrous fronds, almost indistinguishable from H. polyanthos – but containing of oils that make an impression of the blades on the mounting paper; this feature is a characteristic of this group. REPRESENTATIVE SPECIES: Hymenophyllum krauseanum Phil.; H. magellanicum (Desv.) Willd.; H. paninese Ebihara & K. Iwats.; H. plicatum Kaulf.; H. sanguinolentum (G. Forst.) Sw.; H. scabrum A. Rich.; H. tortuosum Hook. & Grev.; H. villosum Colenso. 7. Subgenus Hymenoglossum (C. Presl) R.M. Tryon & A.F. Tryon Hymenophyllum Sm. subg. Hymenoglossum (C. Presl) R.M. Tryon & A.F. Tryon (1981) 134. — Hymenoglossum C. Presl (1843) 35. — Hymenophyllum Sm. sect. Hymenoglossum (C. Presl) T. Moore (1857) cxii. — Type: Hymenoglossum cruentum (Cav.) C. Presl (= Hymenophyllum cruentum Cav.). Hymenophyllum Sm. sect. Integra C. Presl (1843) 32. — Lectotype: Hymenophyllum asplenioides Sw. (selected by Morton (1968) 172). Rhizomes long-creeping, filiform, nearly glabrous. Stipes 7–15 cm long. Blades simple to pinnatifid, ovate to narrowly ovate, 15 by 4 cm, margins of segments entire. Sori at the frond margins, slightly immersed in the laminae, lips bivalvate, entire, receptacles included in involucres. Distribution — Central to South America and Madagascar; 3 species known. Habitat — Epiphytic on tree trunks. Chromosome base number — x = 36? Note — Hennequin et al. (in press) revealed that H. heimii of Madagascar probably has a sister relationship to H. cruentum of southern Chile (formerly treated as a monotypic genus, Hymenoglossum). A Central-South American species, H. asplenioides, is also placed here because of its morphological resemblance to the two species above. REPRESENTATIVE SPECIES: Hymenophyllum asplenioides Sw.; H. cruentum Cav.; H. heimii Tardieu. 8. Subgenus Fuciformia Ebihara & K. Iwats., subg. nov. A subgenere Globosa rhizomatibus brevibus et erectis differt. Based on Hymenophyllum Sm. sect. Fuciformia H. Christ (1897) 20, nom. nud. — Type: Hymenophyllum fuciforme Sw. Rhizomes erect or short-creeping, thick, densely covered with reddish hairs. Stipes 3–15 cm long. Blades tri- to quadripinnate, ovate to narrowly ovate, 65 by 15 cm, margins of segments entire. Sori at the tips of ultimate segments, lips bivalvate, entire or wavy, receptacles included or slightly extruded from involucres. 234 BLUMEA — Vol. 51, No. 2, 2006 Distribution — Southern temperate regions (Chile and New Zealand); 2 species. Chromosome base number — x = 36. Habitat — Epiphytic, epilithic or terrestrial. Note — This subgenus comprises only two species having tufted rhizomes, an exceptional feature in Hymenophyllum. Evidence suggests that these two species, which are disjunctly distributed in southern Chile and New Zealand, form a monophyletic group (Hennequin et al., in press). REPRESENTATIVE SPECIES: Hymenophyllum fuciforme Sw.; H. pulcherrimum Colenso. 9. Subgenus Diploöphyllum (Bosch) Ebihara & K. Iwats., comb. nov. Based on Diploöphyllum Bosch (1861a) 322, non Diplophyllum (Dumort.) Dumort. (1835) 15, nom. cons., ICBN 2000. — Hymenophyllum Sm. sect. Diploöphyllum C. Chr. (1906) XV. — Hymenophyllum Sm. subsect. Diploöphyllum (C. Chr.) C.V. Morton (1968) 174, as ‘Diplophyllum’. — Lectotype and sole species (in the present treatment): Hymenophyllum dilatatum (G. Forst.) Sw. (selected by Christensen (1906) XV). Rhizomes long-creeping, wiry, sparsely covered with dark hairs. Stipes 2–15 cm long. Blades tri- to quadripinnate, ovate or narrowly ovate, 55 by 15 cm, laminae more than two cells thick, margins of segments entire. Sori at the tips of ultimate segments, lips bivalvate, entire, receptacles included in involucres. Distribution — New Zealand, monotypic. Habitat — Epiphytic on tree trunks. Chromosome base number — x = 36. Note — This monotypic subgenus is characterized by multilayered laminar cells throughout its blades and long-creeping rhizomes. Molecular data (Hennequin et al., in press) suggest this is closely related to the subgenera Fuciformia and Cardiomanes. Van den Bosch’s (1861a) original spelling of this taxon was Diploöphyllum, neither ‘Diploophyllum’ as used by Copeland (1938b) nor ‘Diplophyllum’ by Morton (1968), and its inclusion of a diacritical sign as a diaeresis is legitimate in the latest Code (Greuter et al., 2000, Art. 60.6). 10. Subgenus Cardiomanes (C. Presl) Ebihara & K. Iwats., comb. nov. Based on Cardiomanes C. Presl (1843) 12. — Trichomanes L. subg. Cardiomanes (C. Presl) H. Christ (1897) 33. — Type and sole species: Trichomanes reniforme G. Forst. (1786) 84 (= Hymenophyllum nephrophyllum Ebihara & K. Iwats., nom. nov. = Cardiomanes reniforme (G. Forst.) C. Presl, non Hymenophyllum reniforme Hook.). Rhizomes long-creeping, wiry, sparsely covered with dark hairs. Stipes 5–25 cm long. Blades simple, kidney-shaped, 35 by 13 cm, laminae more than two cells thick, margins of segments entire. Sori at the blade margins, tubular, lips entire, receptacles exserted from involucres. Distribution — New Zealand, monotypic. Habitat — Epiphytic. Chromosome base number — x = 36. Note — One of the most distinct subgenera of Hymenophyllum (formerly included in Trichomanes s.l. due to its cup-shaped involucres). A replacement name is necessary when Cardiomanes reniforme is moved to Hymenophyllum. A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 235 2. DIDYMOGLOSSUM Didymoglossum Desv. (1827) 330. — Trichomanes L. sect. Didymoglossum (Desv.) T. Moore (1857) cx. — Trichomanes L. subg. Didymoglossum (Desv.) C. Chr. (1906) XIV. — Lectotype: Didymoglossum muscoides (Sw.) Desv. (= Didymoglossum hymenoides (Hedw.) Copel.) (selected by Christensen (1906) XIV, as ‘Trichomanes hymenoides Hedw.’). Distribution — Throughout the tropics; more than 30 species. Chromosome base number — x = 34. Note — Corresponding to the Di clade in Ebihara et al. (submitted), and also to Morton’s (1968) Trichomanes subgenus Didymoglossum. This genus contains three genera sensu Copeland (1938b): Didymoglossum, Microgonium and Lecanium (Lecanolepis). All species in Didymoglossum are dwarf epiphytes, mainly in tropical regions, and false veinlets are present. Though both Copeland (1938b) and Morton (1968) used the presence of marginal setae as the primary character to distinguish Microgonium from Didymoglossum, there was evidently secondary loss of marginal hairs in some ‘Microgonium’ species (Ebihara et al., submitted). Therefore, submarginal false veinlets stand as the primary character to separate the two subgenera. KEY TO THE SUBGENERA 1a. Submarginal false veinlets absent . . . . . . . . . . . . . . . . 1. Subg. Didymoglossum b. Submarginal false veinlets present . . . . . . . . . . . . . . . . . 2. Subg. Microgonium 1. Subgenus Didymoglossum Hemiphlebium C. Presl sect. Lecanium Prantl (1875) 46, based on Lecanium C. Presl (1843) 11, t. I, nom. illeg., non Reinw. (1825) 48. — Trichomanes L. sect. Lecanium (Prantl) H. Christ (1897) 25. — Lecanolepis Pic.Serm. (1973) 449. — Type: Lecanium membranaceum (L.) C. Presl (= Didymoglossum membranaceum (L.) Vareschi). Didymoglossum Desv. sect. Flabellata C. Presl (1843) 23. — Lectotype: Didymoglossum sphenoides (Kunze) C. Presl (= Didymoglossum punctatum (Poir.) Desv.) (selected by Morton (1968) 191). Didymoglossum Desv. sect. Pinnata C. Presl (1843) 23. — Lectotype: Didymoglossum muscoides (Sw.) Desv. (= Didymoglossum hymenoides (Hedw.) Copel.) (selected by Morton (1968) 191). Hemiphlebium C. Presl (1843) 25, t. IX. — Trichomanes L. sect. Hemiphlebium (C. Presl) T. Moore (1857) cx. — Trichomanes L. subg. Hemiphlebium (C. Presl) H. Christ (1897) 23. — Type: Hemiphlebium pusillum (Sw.) C. Presl (= Didymoglossum pusilum (Sw.) Desv.). Rhizomes long-creeping, frequently branching, usually filiform, to 0.5 mm diam., densely covered with dark-coloured hairs, protostele collateral to regressed, cortex petiole-like, roots absent, root-like shoots present (sensu Schneider, 2000). Stipes often quite reduced, up to 1 cm, at a distance from the adjacent ones. Blades usually simple, or sometimes lobed to pinnatifid, elliptic to narrowly ovate, 6 by 6 cm, venation catadromous, often flabelliform, submarginal false veinlets absent, longitudinal false veinlets parallel to the true veins present, internal cell walls thin and straight, setae or scale-like projections sometimes. Sori epitactic or pantotactic, often immersed in the laminae, campanulate, lips bilabiate or occasionally truncate, often dark-margined, receptacles exserted. Distribution — Throughout the tropics, mainly in the New World; more than 20 species. 236 BLUMEA — Vol. 51, No. 2, 2006 Habitat — Epilithic or low-epiphytic. Note — Corresponding to the Le subclade in Ebihara et al. (submitted). REPRESENTATIVE SPECIES: New combinations: Didymoglossum hildebrandtii (Kuhn) Ebihara & Dubuisson, comb. nov. [based on Trichomanes hildebrandtii Kuhn (1879) 70]; D. lorencei (Tardieu) Ebihara & Dubuisson, comb. nov. [based on Trichomanes lorencei Tardieu (1977) 147]; D. motleyi (Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes motleyi Bosch (1861b) 145]; D. sublimbatum (Müll.Berol.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes sublimbatum Müll.Berol. (1854) 737]; D. tahitense (Nadeaud) Ebihara & K. Iwats., comb. nov. [based on Trichomanes tahitense Nadeaud (1873) 18]. Other species: Didymoglossum angustifrons Fée; D. curtii (Rosenst.) Pic.Serm.; D. exiguum (Bedd.) Copel.; D. gourlianum (Grev. ex J. Sm.) Pic.Serm.; D. hymenoides (Hedw.) Copel.; D. krausii (Hook. & Grev.) C. Presl; D. liberiense (Copel.) Copel.; D. lineolatum Bosch; D. melanopus (Baker) Copel.; D. membranaceum (L.) Vareschi; D. nummularium Bosch; D. ovale E. Fourn.; D. petersii (A. Gray) Copel.; D. pinnatinervium (Jenman) Pic.Serm.; D. punctatum (Poir.) Desv.; D. pusillum (Sw.) Desv.; D. reptans (Sw.) C. Presl; D. rhipidophyllum (Sloss.) Pic.Serm. 2. Subgenus Microgonium (C. Presl) Ebihara & K. Iwats., comb. nov. Based on Microgonium C. Presl (1843) 19, t. VI, f. A, B. — Hemiphlebium C. Presl sect. Microgonium (C. Presl) Prantl (1875) 46. — Trichomanes L. sect. Microgonium (C. Presl) H. Christ (1897) 24. — Lectotype: Microgonium cuspidatum (Willd.) C. Presl (= Didymoglossum cuspidatum (Willd.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes cuspidatum Willd. (1810) 499]) (selected by Christensen (1906) XIV). Rhizomes long-creeping, frequently branching, filiform, to 0.5 mm diam., densely covered with dark hairs, protostele collateral to regressed (see Ebihara et al., submitted), cortex petiole-like, roots absent, root-like shoots present. Stipes often quite reduced, to 1 cm, at a distance from the adjacent ones. Blades usually simple, or sometimes lobed to pinnatifid, elliptic to narrowly ovate, 3 by 2 cm, venation catadromous, continuous submarginal false veinlets present, longitudinal false veinlets parallel to the true veins present, internal cell walls thin and straight, margins entire and glabrous. Sori epitactic or pantotactic, often immersed in the laminae, tubular, lips often dilate, receptacles exserted. Distribution — Throughout the tropics, mainly in the Old World; more than 10 species. Habitat — Epilithic or low-epiphytic. Note — Corresponding to the Mg subclade in Ebihara et al. (submitted). REPRESENTATIVE SPECIES: New combinations: Didymoglossum bimarginatum (Bosch) Ebihara & K. Iwats., comb. nov. [based on Microgonium bimarginatum Bosch (1861b) 143]; D. ekmanii (Wess. Boer) Ebihara & Dubuisson, comb. nov. [based on Trichomanes ekmanii Wess.Boer (1962) 319, f. 33a, b]; D. godmanii (Hook.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes godmanii Hook. in Baker (1866) 337, t. 8, f. A]; D. kapplerianum (Sturm) Ebihara & Dubuisson, comb. nov. [based on Trichomanes kapplerianum Sturm in Martius (1859) 276]; D. kirkii (Hook.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes kirkii Hook. in Hook. & Baker (1867) 78]; D. lenormandii (Bosch) Ebihara & Dubuisson, comb. nov. [based on Trichomanes lenormandii Bosch (1861b) 144]; D. mindorense (H. Christ) K. Iwats., comb. nov. [based on Trichomanes mindorense H. Christ (1908a) 270]; D. pygmaeum (C. Chr.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes pygmaeum C. Chr. (1920) 10, f. 2]; D. sinuatum (Bonap.) A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 237 Ebihara & Dubuisson, comb. nov. [based on Trichomanes sinuatum Bonap. (1920) 25]; D. wesselsboeri Ebihara & Dubuisson, nom. nov. [based on Trichomanes hookeri C. Presl (1843) 16, non Didymoglossum hookeri C. Presl]. Other species: Didymoglossum cuspidatum (Willd.) Ebihara & Dubuisson. 3. CREPIDOMANES Crepidomanes (C. Presl) C. Presl (1849) 258. — Trichomanes L. subg. Crepidomanes C. Presl (1849) 17. — Trichomanes L. sect. Crepidomanes (C. Presl) Prantl (1875) 51. — Type: Trichomanes intramarginale Hook. & Grev. (= Crepidomanes intramarginale (Hook. & Grev.) C. Presl). Distribution — Throughout the Old World tropics to northern temperate regions; more than 30 species. Chromosome base number — x = 36. Note — This genus corresponds to the PT clade in Ebihara et al. (submitted), and also to Copeland’s (1938b) genera Crepidomanes, Gonocormus, Crepidopteris (Reediella) p.p., Vandenboschia p.p. and Microtrichomanes p.p. Copeland’s (1938b) genera Crepidomanes and Gonocormus are placed here, and a part of Crepidopteris, Vandenboschia and Microtrichomanes are also included. Most species of Crepidomanes are epiphytes, mainly in rain forests of the Paleotropics; however, Nesopteris, with a short, erect rhizomes, is also included here. KEY TO THE SUBGENERA AND SECTIONS 1a. Fronds smaller, less than 15 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 b. Fronds larger, more than 15 cm long . . . . . . . . . . . . . . . . . . 2. Subg. Nesopteris 2a. Elongate marginal cells present in laminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1c. Subg. Crepidomanes sect. Crepidium b. Elongate marginal cells absent in laminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Proliferation present on stipes . . . . 1b. Subg. Crepidomanes sect. Gonocormus b. Proliferations absent on stipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4a. Blades flabellate. . . . . . . . . . . . . . . 1b. Subg. Crepidomanes sect. Gonocormus b. Blades pinnate . . . . . . . . . . . . . . . 1a. Subg. Crepidomanes sect. Crepidomanes 1. Subgenus Crepidomanes Distribution — Throughout the Paleotropics. Note — Corresponding to the Cr subclade in Ebihara et al. (submitted). Three sections are recognized in this subgenus. 1a. Section Crepidomanes Trichomanes L. subg. Minora Prantl (1875) 51. — Lectotype: Trichomanes intramarginale Hook. & Grev. (= Crepidomanes intramarginale (Hook. & Grev.) Copel.) (selected by Morton (1968) 181). Trichomanes L. sect. Taschneria C. Presl ex C. Chr. (1906) XV, based on Taschneria C. Presl (1849) 258, nom. nud. — Crepidomanes (C. Presl) C. Presl sect. Taschneria (C. Presl ex C. Chr.) K. Iwats. (1984) 175. — Type: Trichomanes ilicula Bory ex Willd. (= Crepidomanes bipunctatum (Poir.) Copel.). 238 BLUMEA — Vol. 51, No. 2, 2006 Rhizomes long-creeping, frequently branching, filiform, 0.1– 0.5 mm diam., densely covered with dark hairs, protostele subcollateral, cortex with centrifuge sclerification, roots absent, root-like shoots present. Stipes sometimes reduced, or to 8 cm long, at a distance from the adjacent ones. Blades simple to quadripinnatifid, elliptic to subdeltate, 12 by 6 cm, venation anadromous, false veinlets often present, submarginally (continuous or interrupted) and/or parallel (but not connected) to true veins, laminae almost always one cell thick, exceptionally thicker (C. intramarginale), internal cell walls thin and straight. Sori paratactic, tubular, lips usually bilabiate, often deltate, sometimes dilate, receptacles exserted. Distribution — Throughout the Paleotropics; more than 20 species. Habitat — Usually epilithic on wet rocks along streams, sometimes epiphytic on tree trunks. Note — Crepidomanes in the strict sense contains only those species that have false veinlets, but recent study (Ebihara et al., submitted) has shown that some species lacking false veinlets (formerly placed under Vandenboschia by Copeland) are closely related to species that have them and/or are nested in a clade consisting of species with false veinlets. REPRESENTATIVE SPECIES: New combinations: Crepidomanes africanum (H. Christ) Ebihara & Dubuisson, comb. nov. [based on Trichomanes africanum H. Christ (1909a) 21]; C. chevalieri (H. Christ) Ebihara & Dubuisson, comb. nov. [based on Trichomanes chevalieri H. Christ (1908b) 106]; C. draytonianum (Brack.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes draytonianum Brack. (1854) 252, pl. 36, f. 3]; C. fallax (H. Christ) Ebihara & Dubuisson, comb. nov. [based on Trichomanes fallax H. Christ (1909b) 24]; C. mettenii (C. Chr.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes mettenii C. Chr. (1906) 644]. Other species: Crepidomanes barnardianum (F.M. Bailey) Tindale; C. bipunctatum (Poir.) Copel.; C. christii (Copel.) Copel.; C. clarenceanum (F. Ballard) Pic.Serm.; C. frappieri (Cordem.) J.P. Roux; C. inopinatum (Pic.Serm.) J.P. Roux; C. intramarginale (Hook. & Grev.) Copel.; C. kurzii (Bedd.) Tagawa & K. Iwats.; C. latealatum (Bosch) Copel.; C. latemarginale (D.C. Eaton) Copel.; C. melanotrichum (Schltdl.) J.P. Roux; C. rupicolum (Racib.) Copel.; C. schmidtianum (Zenker ex Taschner) K. Iwats.; C. vitiense (Baker) Bostock. 1b. Section Gonocormus (Bosch) K. Iwats. Crepidomanes (C. Presl) C. Presl sect. Gonocormus (Bosch) K. Iwats. (1984) 174. — Gonocormus Bosch (1861a) 321. — Trichomanes L. sect. Gonocormus (Bosch) H. Christ (1897) 27. — Trichomanes L. subg. Gonocormus (Bosch) C. Chr. (1906) XIV. — Lectotype: Gonocormus prolifer (Blume) Prantl (= Crepidomanes minutum (Blume) K. Iwats.) (selected by Christensen (1906) XIV). Rhizomes long-creeping, frequently branching, filiform, c. 0.15 mm diam., covered with dark brown hairs or glabrescent, protostele subcollateral, cortex with centrifugal sclerification, roots absent, root-like shoots present. Stipes very short, or to 3 cm long, at a distance from the adjacent ones, proliferations often observed. Blades simple to quadripinnatifid, flabellate to narrowly ovate, 12 by 3 cm, venation anadromous, false veinlets absent, internal cell walls thin and straight. Sori paratactic or pantotactic, campanulate, lips dilate, receptacles long-exserted. A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 239 Distribution — Throughout the Paleotropics; 1 polymorphic species, Crepidomanes minutum (Blume) K. Iwats., is recognized here. Habitat — Epilithic or epiphytic on tree trunks. Note — The single species ranges from Africa to the Pacific according to Yoroi & Iwatsuki (1977). There is much genetic variation in the rbcL sequences of this species from various localities, but it corresponds neither to morphological features nor to geographical distribution (A. Ebihara, unpublished data). Further studies are needed. 1c. Section Crepidium (C. Presl) Ebihara & K. Iwats., comb. nov. Based on Didymoglossum Desv. subg. Crepidium C. Presl (1843) 23. — Trichomanes L. sect. Crepidium (C. Presl) C. Chr. (1906) XV. — Crepidopteris Copel. (1938b) 57, nom. illeg., non Benth. (1862). — Crepidophyllum C.F. Reed (1948) 88, nom. illeg., non Herzog (1926) 356. — Reediella Pic.Serm. (1970) 719. — Crepidomanes (C. Presl) C. Presl subg. Crepidium (C. Presl) K. Iwats. (1984) 174. — Type: Didymoglossum humile (G. Forst.) C. Presl (= Crepidomanes humile (G. Forst.) Bosch). Rhizomes long-creeping, frequently branching, filiform, 0.3–0.4 mm diam., densely covered with dark brown or blackish hairs, protostele subcollateral, cortex with centrifugal sclerification, roots absent, root-like shoots present. Stipes 0.2–1.2 cm long, at a distance from the adjacent ones. Blades bipinnatifid to tripinnatifid, ovate to narrowly ovate, 1–8 by 2–3 cm wide, venation anadromous, double rows of elongate marginal cells present, false veinlets absent, internal cell walls thin and straight. Sori paratactic, tubular, lips dilate, receptacles long-exserted. Distribution — Old World tropics (Asia to Pacific); at least 1 species, Crepidomanes humile (G. Forst.) Bosch. Habitat — Epilithic or low-epiphytic. Note — Crepidium (Reediella), a group characterized by marginal elongated cell rows, has been shown to be an unnatural group (Ebihara et al., submitted) – species having single rows of elongated cells are attributed to another genus, Polyphlebium, in the present system. As a result, this section contains only C. humile. 2. Subgenus Nesopteris (Copel.) Ebihara & K. Iwats., comb. nov. Based on Nesopteris Copel. (1938b) 65. — Trichomanes L. sect. Nesopteris (Copel.) C.V. Morton (1968) 190. — Cephalomanes C. Presl subg. Nesopteris (Copel.) K. Iwats. (1984) 176. — Type: Nesopteris grandis (Copel.) Copel. (= Crepidomanes grande (Copel.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes grande Copel. (1911) 70]. Rhizomes suberect, ascending or long-creeping, quite thick, 2.8–3.5 mm diam., densely covered with dark hairs, protostele massive, cortex homogeneous, roots numerous and robust. Stipes to 25 cm long, very close to each other except in C. aphlebioides. Blades 4- or 5-pinnate, elliptic to narrowly ovate, 15–50(–70) by 10–20(–25) cm, venation anadromous, false veinlets absent, internal cell walls thin and straight. Sori paratactic, tubular, lips often dilate, receptacles long-exert. Distribution — Old World tropics (Asia to Pacific); at least 4 species. Habitat — Terrestrial or epilithic, occasionally climbing tree trunks. Note — Large fronds and stout rhizomes observed in members of this subgenus are apparently quite different from those of subgenus Crepidomanes. Our study (Ebihara 240 BLUMEA — Vol. 51, No. 2, 2006 et al., submitted) revealed that the erect rhizomes of the Nesopteris species are probably the result of an exceptional evolution from creeping rhizomes to erect ones, which is in the direction opposite to the evolutional trait of Hymenophyllaceae. We assume that C. aphlebioides, the only species of this subgenus having long-creeping rhizomes, is transitional between the two subgenera. REPRESENTATIVE SPECIES: New combinations: Crepidomanes intermedium (Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes intermedium Bosch (1861b) 179]; C. thysanostomum (Makino) Ebihara & K. Iwats., comb. nov. [based on Trichomanes thysanostomum Makino (1899) 46]. Other species: Crepidomanes aphlebioides (H. Christ) I.M. Turner; C. grande (Copel.) Ebihara & K. Iwats. 4. POLYPHLEBIUM Polyphlebium Copel. (1938b) 55. — Phlebiophyllum Bosch (1861a) 321, non Phlebophyllum Nees in Wallich (1832) 75, 102. — Trichomanes L. sect. Phlebiophyllum (Bosch) Prantl (1875) 52. — Trichomanes L. subg. Polyphlebium (Copel.) Allan (1961) 34. — Type: Polyphlebium venosum (R.Br.) Copel. Rhizomes long-creeping, frequently branching, filiform, 0.1–0.8 mm diam., densely covered with light brown hairs, protostele subcollateral, cortex with centrifuge sclerification, roots few and fine. Stipes 0.3–5.5 cm long, at a distance from the adjacent ones. Blades pinnate to quadripinnate, ovate to narrowly ovate, 30 by 6.5 cm, venation anadromous, sometimes a single row of elongate marginal cells present, false veinlets absent, internal cell walls thin and straight. Sori paratactic, tubular, lips usually dilate, receptacle long-exserted. Distribution — Temperate regions of the southern hemisphere, and mountain forests of low altitude regions; c. 15 species. Habitat — Epilithic on wet rocks or epiphytic on tree-fern trunks. Chromosome base number — x = 36. Note — Corresponding to the Tp clade in Ebihara et al. (submitted). Though constituents of this genus have so far been confused with those of Crepidomanes and Vandenboschia of the present system, their stem and root features well differentiate them from the others. REPRESENTATIVE SPECIES: New combinations: Polyphlebium angustatum (Carmich.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes angustatum Carmich. (1818) 513]; P. borbonicum (Bosch) Ebihara & Dubuisson, comb. nov. [based on Trichomanes borbonicum Bosch (1861b) 158]; P. capillaceum (L.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes capillaceum L. (1753) 1099]; P. colensoi (Hook.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes colensoi Hook. (1854) t. 979]; P. diaphanum (Kunth) Ebihara & Dubuisson, comb. nov. [based on Trichomanes diaphanum Kunth (1815) 25]; P. endlicherianum (C. Presl) Ebihara & K. Iwats., comb. nov. [based on Trichomanes endlicherianum C. Presl (1848) 333]; P. exsectum (Kunze) Ebihara & Dubuisson, comb. nov. [based on Trichomanes exsectum Kunze (1837) 47, t. 29, f. 2]; P. hymenophylloides (Bosch) Ebihara & Dubuisson, comb. nov. [based on Trichomanes hymenophylloides Bosch (1863) 209]; P. ingae (C. Chr.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes ingae C. Chr. in C. Chr. & Skottsb. (1920) 3, f. 2]; P. philippianum (Sturm) Ebihara & Dubuisson, comb. nov. [based on Trichomanes philippianum Sturm (1858) 188]; P. pyxidi- A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 241 ferum (L.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes pyxidiferum L. (1753) 1098]; P. vieillardii (Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes vieillardii Bosch (1861c) 90]; P. werneri (Rosenst.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes werneri Rosenst. (1908a) 35]. Other species: Polyphlebium venosum (R.Br.) Copel. 5. VANDENBOSCHIA Vandenboschia Copel. (1938b) 51. — Trichomanes L. subg. Vandenboschia (Copel.) Allan (1961) 34. — Type: Vandenboschia radicans (Sw.) Copel. Distribution — Throughout the tropics, extending to northern temperate regions; more than 15 species. Habitat — Hemi-epiphytic on tree trunks or epilithic, occasionally terrestrial. Chromosome base number — x = 36. Note — Corresponding to the Va clade in Ebihara et al. (submitted). Our molecular analysis shows Copeland’s genus Vandenboschia evidently of polyphyletic origin – he included species of at least four different lineages (the Va, Po, PT, NT clades in Ebihara et al., submitted). Vandenboschia, in the present system, refers to V. radicans, type of Vandenboschia, and its relatives. Nakaike (1975) advocated the application of the generic name Lacosteopsis instead of Vandenboschia on the grounds that Vandenboschia was an invalid generic name since Copeland (1938b) included the type of an existing genus (T. scandens) when he established Vandenboschia (T. Nakaike, pers. comm.). This confusion was attributed to the controversial typification of Trichomanes as noted above. Recent nomenclatural conservation of T. crispum against T. scandens (Greuter et al., 2000) denied the claim that Vandenboschia is an illegitimate name (Morton, 1968; Holttum, 1976). KEY TO THE SUBGENERA 1a. Blades bipinnate or more finely divided, hemi-epiphytic or epilithic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Subg. Vandenboschia b. Blades simply pinnate, hemi-epiphytic . . . . . . . . . . . . . . . 2. Subg. Lacosteopsis 1. Subgenus Vandenboschia Rhizomes short- or long-creeping, irregularly branching, rather thick, to 1.7 mm diam., densely covered with brown to bright brown multicellular hairs, protostele reduced, cortex heterogeneous, roots numerous and robust. Stipes 1–16 cm long, clustered or irregularly distanced. Blades bipinnate to 5-pinnatifid, ovate to linear-ovate, 40 by 20 cm, venation anadromous, false veinlets absent, laminae often reduced, regular arrangement of elongate cells observed in some species, internal cell walls various (thin to thick, straight to coarsely pitted). Sori paratactic, tubular to campanulate, lips sometimes dilate, receptacles long-exserted. Distribution — Throughout the tropics, extending to northern temperate regions; more than 15 species. 242 BLUMEA — Vol. 51, No. 2, 2006 Habitat — Terrestrial in humid places, or epiphytic on tree trunks including tree ferns. REPRESENTATIVE SPECIES: New combinations: Vandenboschia boschiana (Sturm ex Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes boschianum Sturm ex Bosch (1861b) 160]; V. collariata (Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes collariatum Bosch (1859) 368]; V. gigantea (Bory ex Willd.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes giganteum Bory ex Willd. (1810) 514]. Other species: Vandenboschia birmanica (Bedd.) Ching; V. cyrtotheca (Hillebr.) Copel.; V. davallioides (Gaudich.) Copel.; V. johnstonensis (F.M. Bailey) Copel.; V. liukiuensis (Y. Yabe) Copel.; V. maxima (Blume) Copel.; V. radicans (Sw.) Copel.; V. speciosa (Willd.) G. Kunkel; V. subclathrata K. Iwats. 2. Subgenus Lacosteopsis (Prantl) Ebihara & K. Iwats., comb. nov. Based on Trichomanes L. sect. Lacosteopsis Prantl (1875) 53. — Lacosteopsis (Prantl) Nakaike (1975) 21. — Lectotype: Trichomanes luschnatianum C. Presl (= Vandenboschia rupestris Ebihara & K. Iwats., comb. nov. [based on Hymenophyllum rupestre Raddi (1825) 67, t. 80]) (selected by Christensen (1906) XV, as Trichomanes rupestre (Raddi) Bosch). Rhizomes suberect, or short- to long-creeping, irregularly branching, wiry, densely covered with brown to bright brown multicellular hairs, or nearly glabrous in climbing parts, protostele reduced, cortex heterogeneous, roots numerous and robust at the terrestrial parts. Stipes 1–16 cm long, clustered or irregularly distanced. Blades usually pinnate, narrowly elliptic to linear-ovate, 50 by 6 cm, venation anadromous, false veinlets absent, laminae often reduced, regular arrangement of elongated cells observed in some species, internal cell walls various (thin to thick, straight to coarsely pitted). Sori paratactic, tubular to campanulate, lips sometimes dilate, receptacles long-exserted. Distribution — Both New World and Old World tropics (excl. Africa); at least 2 species. Habitat – Hemi-epiphytic, climbing on tree trunks. REPRESENTATIVE SPECIES: Vandenboschia auriculata (Blume) Copel.; V. rupestris (Raddi) Ebihara & K. Iwats. 6. ABRODICTYUM Abrodictyum C. Presl (1843) 20, t. VII. — Trichomanes L. sect. Abrodictyum (C. Presl) T. Moore (1857) cx. — Cephalomanes C. Presl subg. Abrodictyum (C. Presl) K. Iwats. (1984) 176. — Habrodictyon C. Presl ex Bosch (1861a) 321. — Type: Abrodictyum cumingii C. Presl. Distribution — Throughout the tropics; c. 25 species. Chromosome base number — x = 33 (a few doubtful records x = 36). Note — Corresponding to the Pa clade in Ebihara et al. (submitted). Circumscription of this genus is very similar to Morton’s subgenus Pachychaetum except for the inclusion of Abrodictyum and the exclusion of Davalliopsis. Dissected fronds consisting primarily of the axis (lacking laminar cells) is a unique character observed in some species of this genus, but according to molecular results such fronds evolved several times in parallel within this genus. A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 243 KEY TO THE SUBGENERA 1a. b. 2a. b. 3a. b. Distribution in Asia/Pacific . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Distribution in Africa/America . . . . . . . . . . . . . . . . . . . . 2. Subg. Pachychaetum Rhizomes long-creeping . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Subg. Abrodictyum Rhizomes erect or short-creeping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Lamina texture membranous . . . . . . . . . . . . . . . . . . . . . . . 1. Subg. Abrodictyum Lamina texture not membranous . . . . . . . . . . . . . . . . . . . 2. Subg. Pachychaetum 1. Subgenus Abrodictyum Trichomanes L. sect. Leptomanes Prantl (1875) 52, nom illeg.; one of the five original species, Trichomanes smithii Hook. (= Abrodictyum cumingii C. Presl), is also the type of Trichomanes L. sect. Abrodictyum (C. Presl) T. Moore (1857). Rhizomes suberect, or short- to long-creeping, irregularly branching, rather thick, to 1.7 mm diam., densely covered with brown to bright brown multicellular hairs, protostele massive, cortex homogeneous, roots numerous and robust. Stipes 1–16 cm long, clustered or irregularly distanced. Blades bipinnate to 5-pinnatifid, narrowly ovate to linear-ovate, 40 by 6 cm, venation anadromous, false veinlets absent, laminae often reduced, regular arrangement of elongate cells observed in some species, internal cell walls various (thin to thick, straight to coarsely pitted). Sori paratactic, tubular to campanulate, lips sometimes dilate, receptacles long-exserted. Distribution — Paleotropics (Asia to the Pacific); c. 15 species. Habitat — Terrestrial in humid places, or epiphytic on tree trunks including tree ferns. Note — Corresponding to the Ab clade in Ebihara et al. (submitted). REPRESENTATIVE SPECIES: New combinations: Abrodictyum asae-grayi (Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes asaegrayi Bosch (1861b) 180]; A. brassii (Croxall) Ebihara & K. Iwats., comb. nov. [based on Macroglena brassii Croxall (1975) 543]; A. caudatum (Brack.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes caudatum Brack. (1854) 256, pl. 36, f. 5]; A. clathratum (Tagawa) Ebihara & K. Iwats., comb. nov. [based on Trichomanes clathratum Tagawa (1939) 164]; A. lavofuscum (Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes lavofuscum Bosch (1861c) 88]; A. idoneum (C.V. Morton) Ebihara & K. Iwats., comb. nov. [based on Trichomanes idoneum C.V. Morton (1973) 272] (Old World plants formerly called ‘T. gemmatum’ are this species, because T. gemmatum J. Sm. ex Baker was originally applied to a Neotropical plant (see Morton, 1973)); A. pluma (Hook.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes pluma Hook. (1854) t. 997] (Asian plants formerly identified as ‘T. meifolium’ are this species); A. schlechteri (Brause) Ebihara & K. Iwats., comb. nov. [based on Trichomanes schlechteri Brause (1912) 10]; A. strictum (Menzies ex Hook. & Grev.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes strictum Menzies ex Hook. & Grev. (1831) t. 122]. Other species: Abrodictyum boninense Tagawa & K. Iwats.; A. cumingii C. Presl. 2. Subgenus Pachychaetum (C. Presl) Ebihara & K. Iwats., comb. nov. Based on Trichomanes L. subg. Pachychaetum C. Presl (1843) 16. — Cephalomanes C. Presl subg. Pachychaetum (C. Presl) K. Iwats. (1984) 177. — Lectotype: Trichomanes rigidum Sw. (= Abrodictyum rigidum (Sw.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes rigidum Sw. (1788) 137]) (selected by Christensen (1906) XV). 244 BLUMEA — Vol. 51, No. 2, 2006 Trichomanes L. subg. Macroglena C. Presl (1848) 333. — Macroglena (C. Presl) Copel. (1938a) 49. — Cephalomanes C. Presl subg. Macroglena (C. Presl) K. Iwats. (1984) 176. — Lectotype: Trichomanes meifolium Bory ex Willd. (1810) 509 (= Abrodictyum meifolium (Bory ex Willd.) Ebihara & K. Iwats., comb. nov.) (selected by Christensen (1906) XV). Trichomanes L. sect. Selenodesmium Prantl (1875) 53. — Selenodesmium (Prantl) Copel. (1938b) 80. — Lectotype: Trichomanes rigidum Sw. (= Abrodictyum rigidum (Sw.) Ebihara & Dubuisson) (selected by Christensen (1906) XV). Rhizomes erect or short-creeping, quite thick, about 2 mm diam., covered with brown to blackish hairs, protostele massive, cortex homogeneous, roots numerous and robust. Stipes 7–15 cm long, clustered or more or less distanced. Blades bipinnate to quadripinnatifid, linear-ovate to subdeltate, 30 by 18 cm, venation anadromous, false veinlets absent, laminae reduced in some species, internal cell walls thick and coarsely pitted. Sori paratactic, tubular, lips truncate, receptacles long-exserted. Distribution — Throughout the tropics; more than 10 species. Habitat — Usually terrestrial in humid places, occasionally low-epiphytic. Note — Corresponding to the Pa subclade in Ebihara et al. (submitted). REPRESENTATIVE SPECIES: New combinations: Abrodictyum cellulosum (Klotzsch) Ebihara & Dubuisson, comb. nov. [based on Trichomanes cellulosum Klotzsch (1844) 531]; A. cupressoides (Desv.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes cupressoides Desv. (1827) 330]; A. dentatum (Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes dentatum Bosch (1861b) 182]; A. elongatum (A. Cunn.) Ebihara & K. Iwats., comb. nov. [based on Trichomanes elongatum A. Cunn. (1836) 368]; A. kalimantanense (K. Iwats. & M. Kato) Ebihara & K. Iwats., comb. nov. [based on Macroglena kalimantanensis K. Iwats. & M. Kato (1980) 31, f. 1]; A. laetum (Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes laetum Bosch (1861c) 90]; A. obscurum (Blume) Ebihara & K. Iwats., comb. nov. [based on Trichomanes obscurum Blume (1828) 227] (there are two varieties under this species: var. obscurum and var. siamense (H. Christ) K. Iwats., comb. nov. [based on Trichomanes siamense H. Christ (1901) 103]); A. setaceum (Bosch) Ebihara & K. Iwats., comb. nov. [based on Trichomanes setaceum Bosch (1861b) 176]; A. sprucei (Baker) Ebihara & Dubuisson, comb. nov. [based on Trichomanes sprucei Baker in Hook. & Baker (1867) 87]; A. tamarisciforme (Jacq.) Ebihara & Dubuisson, comb. nov. [based on Trichomanes tamarisciforme Jacq. (1789) 285, t. 21, f. 2]. Other species: Abrodictyum meifolium (Bory ex Willd.) Ebihara & K. Iwats.; A. rigidum (Sw.) Ebihara & Dubuisson. 7. TRICHOMANES Trichomanes L. (1753) 1097. — Type: Trichomanes crispum L., type. cons., ICBN 2000 (vs Trichomanes scandens L.). Distribution — The Neotropics, and at least one species in continental Africa; more than 60 species. Unpublished molecular data suggest inclusion of a few other Paleotropical taxa (from Madagascar and the Indian Ocean), but further phylogenetic investigations are needed. Chromosome base number — x = 32 (a few doubtful records x = 36). Note — Corresponding to the NT clade in Ebihara et al. (submitted). Constituents of this genus include Morton’s subgenus Achomanes and sect. Davalliopsis (placed under subg. Pachychaetum) along with T. scandens (placed under subg. Trichomanes). In A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 245 contrast to its long-term acceptance in New World floras, Morton’s subgenus Achomanes shared few morphological characters other than geographical distribution and chromosome base number. Recent inclusion of Davalliopsis in this group (Dubuisson et al., 2003a; Ebihara et al, submitted) made it even more difficult to define morphologically. We therefore recognize four subgenera under this genus, which are easily definable. Monophyly of subgenus Davalliopsis and subg. Lacostea is clearly shown by our molecular analysis (as the Da subclade), but they are very different in most morphological features. KEY TO THE SUBGENERA 1a. b. 2a. b. 3a. b. 4a. b. 5a. b. Lamina texture membranous, one cell thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Lamina texture tough, more than one cell thick . . . . . . . . . . . . . . . . . . . . . . . . . 5 Rhizomes short-creeping or erect . . . . . . . . . . . . . . . . . . . 1. Subg. Trichomanes Rhizomes long-creeping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Blades tripinnate or more finely divided . . . . . . . . . . . . . . 1. Subg. Trichomanes Blades once-pinnate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Margins of blades hairy . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Subg. Trichomanes Margins of blades glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Subg. Lacostea Venation catadromous in sterile blades . . . . . . . . . . . . . . . . . . . . . . 2. Subg. Feea Venation anadromous in sterile blades . . . . . . . . . . . . . . . . 3. Subg. Davalliopsis 1. Subgenus Trichomanes Trichomanes L. subg. Eutrichomanes C. Presl (1843) 16. — Trichomanes L. sect. Eutrichomanes (C. Presl) T. Moore (1857) cx. — Lectotype: Trichomanes scandens L. (selected by Morton (1968) 180; his idea that this and several subsequent synonyms include the type species of Trichomanes led him to designate T. scandens as lectotypes for these taxa. Although T. crispum has recently been fixed to the type of the genus as noted above, Morton’s lectotypifications do not need to be changed since they were validly published). Trichomanes L. sect. Pinnata C. Presl (1843) 16. — Lectotype: Trichomanes scandens L. (selected by Morton (1968) 180). Ragatelus C. Presl (1843) 16. — Trichomanes L. sect. Ragatelus (C. Presl) C. Chr. (1906) XIV. — Type: Ragatelus crinitus (Sw.) C. Presl (= Trichomanes crinitum Sw.). Homoeotes C. Presl (1848) 331. — Ptilophyllum Bosch sect. Homoeotes (C. Presl) Prantl (1875) 48. — Trichomanes L. sect. Homoeotes (C. Presl) C. Chr. (1906) XIV. — Type: Homoeotes heterophylla (Willd.) C. Presl (= Trichomanes humboldtii (Bosch) Lellinger). Trichomanes L. subg. Pseudoachomanes C. Presl (1849) 16. — Trichomanes L. sect. Pseudoachomanes (C. Presl) C. Chr. (1906) XIV. — Lectotype: Trichomanes sinuosum Rich. ex Willd. (= Trichomanes polypodioides L.) (selected by Christensen (1906) XIV, as T. polypodioides L.). Odontomanes C. Presl (1849) 20. — Trichomanes L. sect. Odontomanes (C. Presl) C. Chr. (1906) XIV. — Type: Odontomanes hostmannianum (Klotzsch) C. Presl (= Trichomanes hostmannianum (Klotzsch) Kunze). Trichomanes L. sect. Neurophyllum T. Moore (1857) cx, based on Neurophyllum C. Presl (1843) 18, t. IV, f. C, nom. illeg., non Torr. & A. Gray. (1840) 612. — Neuromanes Trevis. ex Bosch (1859) 347. — Type: Neurophyllum pinnatum (Hedw.) C. Presl (= Trichomanes pinnatum Hedw.) (selected by Christensen (1906) XIV). Ptilophyllum Bosch subsect. Sinuosa Prantl (1875) 47. — Lectotype: Ptilophyllum sinuosum (Rich. ex Willd.) Prantl (= Trichomanes polypodioides L.) (selected by Morton (1968) 198). 246 BLUMEA — Vol. 51, No. 2, 2006 Ptilophyllum Bosch sect. Acarpacrium Prantl (1875) 48. — Trichomanes L. sect. Acarpacrium (Prantl) C. Chr. (1906) XIV. — Lectotype: Ptilophyllum ptilodes (Bosch) Prantl (= Trichomanes alatum Sw.) (selected by Christensen (1906) XIV, as T. alatum Sw.). Ptilophyllum Bosch sect. Trigonophyllum Prantl (1875) 48. — Trichomanes L. sect. Trigonophyllum (Prantl) C. Chr. (1906) XIV. — Lectotype: Ptilophyllum bancroftii (Hook. & Grev.) Prantl (= Trichomanes arbuscula Desv.) (selected by Christensen (1906) XIV, as T. arbuscula Desv.). Ptilophyllum Bosch subsect. Lamellata Prantl (1875) 48. — Trichomanes L. subsect. Lamellata (Prantl) C.V. Morton (1968) 194. — Type: Ptilophyllum martiusii (C. Presl) Prantl (= Trichomanes martiusii C. Presl). Trichomanes L. sect. Maiora Prantl (1875) 52. – Crepidomanes (C. Presl) C. Presl subg. Maiora K. Iwats. (1984) 174. — Lectotype: Trichomanes scandens L. (selected by Morton (1968) 181). Trichomanes L. subg. Holophlebium H. Christ (1897) 27. — Lectotype: Trichomanes scandens L. (selected by Morton (1968) 181). Mortoniopteris Pic.Serm. (1977a) 243. — Trichomanes L. sect. Mortoniopteris (Pic.Serm.) C. Sánchez (2000) 39. — Type: Mortoniopteris scandens (L.) Pic.Serm. (= Trichomanes scandens L.). Pteromanes Pic.Serm. (1977a) 244. — Type: Pteromanes martiusii (C. Presl) Pic.Serm. (= Trichomanes martiusii C. Presl). Trichomanes L. subg. Achomanes C. Presl (1843) 15, nom. illeg.; originally including Trichomanes crispum L., the type of the genus. Ptilophyllum Bosch (1861a) 321, nom. illeg., non Morris in C.W. Grant (1840) 327, nec (Nutt.) Rchb. (1841) 169; originally including Trichomanes crinitum Sw., the type of Ragatelus C. Presl (1843). Ptilophyllum Bosch subsect. Crispa Prantl (1875) 47, nom. illeg.; originally including Trichomanes crispum L., type of the genus. Rhizomes erect, short- or long-creeping, usually fine to quite thick, 1–5 mm diam., occasionally filiform, sparsely or sometimes densely covered with brown to blackish hairs, protostele massive, cortex homogeneous, or occasionally heterogeneous (in T. scandens), roots numerous and robust. Stipes to 30 cm long, usually close to each other, sometimes more or less distant. Blades usually pinnate with symmetric pinnae, occasionally simple or more finely pinnate, ovate to linear-ovate, 70 by 25 cm, venation usually catadromous, or anadromous (in T. scandens and T. anadromum), sometimes pubescent with unicellular or multicellular hairs, false veinlets present in some species (perpendicular to true veins), internal cell walls slightly thick and wavy, margins sometimes hairy with simple or stellate hairs. Sori paratactic, or epitactic (in T. scandens), often immersed in the laminae, rarely pedicellate (in T. pinnatum), tubular, lips sometimes dilate, receptacles usually long-exserted. Distribution — Same as that of the genus; more than 30 species. Habitat — Usually terrestrial in humid places, sometimes epiphytic on tree trunks, exceptionally as canopy epiphytes (T. pilosum). Some taxa such as T. crispum appear ecologically opportunistic. Note — Corresponding to the Ac subclade in Ebihara et al. (submitted). REPRESENTATIVE SPECIES: Trichomanes accedens C. Presl; T. alatum Sw.; T. anadromum Rosenst.; T. anomalum Maxon & C.V. Morton; T. arbuscula Desv.; T. bicorne Hook.; T. crenatum Bosch; T. crinitum Sw.; T. crispum L.; T. cristatum Kaulf.; T. dactylites Sodiro; T. delicatum Bosch; T. egleri P.G. Windisch; T. imbriatum Backh. ex T. Moore; T. galeottii E. Fourn.; T. guidoi P.G. Windisch; T. holopterum Kunze; T. hostmannianum (Klotzsch) Kunze; T. humboldtii (Bosch) Lellinger; T. kalbreyeri Baker; T. lucens Sw.; T. ludovicinum Rosenst.; T. macilentum Bosch; T. martiusii C. Presl; T. micayense Hieron.; T. pellucens Kunze; T. pilosum Raddi; T. pinnatum Hedw.; T. plumosum Kunze; T. polypodioides L.; A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 247 T. robustum E. Fourn.; T. roraimense Jenman; T. scandens L.; T. spruceanum Hook.; T. steyermarkii P.G. Windisch & A.R. Sm.; T. trigonum Desv.; T. vandenboschii P.G. Windisch; T. vaupesense Lellinger; T. vittaria DC. ex Poir. 2. Subgenus Feea (Bory) Hook. Trichomanes L. subg. Feea (Bory) Hook. (1844) 114. — Feea Bory (1824) 446. — Ptilophyllum Bosch sect. Feea (Bory) Prantl (1875) 48. — Trichomanes L. sect. Feea (Bory) H. Christ (1897) 29. — Lectotype: Feea polypodina Bory (= Trichomanes osmundoides Poir.) (selected by Smith (1875) 349, as Trichomanes spicatum Hedw. ex Hook.). Hymenostachis Bory (1824) 588. — Trichomanes L. subg. Hymenostachis (Bory) Hook. (1844) 114. — Trichomanes L. subsect. Hymenostachis (Bory) C.V. Morton (1968) 197. — Type: Trichomanes elegans Rudge (= Trichomanes diversifrons (Bory) Mett.). Maschalosorus Bosch (1861a) 320. — Trichomanes L. sect. Maschalosorus (Bosch) C. Chr. (1906) XIV. — Type: Maschalosorus mougeotii (Bosch) Bosch (= Trichomanes mougeotii Bosch). Rhizomes suberect or ascending (leading to a dwarf tree-fern habit), quite thick, 1.5–8 mm diam., apex covered with brown to blackish hairs, basal parts covered by roots, protostele massive, cortex homogeneous, root numerous and robust. Stipes to 8 cm long, usually close to each other. Fronds dimorphic (except in T. mougeotii), simple or without lamina in the fertile fronds, once-pinnate in the sterile fronds, 30 by 8 cm, venation catadromous (in sterile frond), false veinlets absent, laminae usually more than one cell thick in fertile fronds, internal cell walls thick and coarsely pitted. Sori paratactic or pantotactic, immersed in the lamina, tubular, lips truncate or slightly dilate, receptacles long-exserted. Distribution — The Neotropics; more than 5 species. Habitat — Terrestrial or epilithic. Note — Corresponding to the Fe subclade in Ebihara et al. (submitted), and also to Copeland’s (1938b) genus, Feea. REPRESENTATIVE SPECIES: Trichomanes botryoides Kaulf.; T. diversifrons (Bory) Mett.; T. mougeotii Bosch; T. osmundoides Poir.; T. trollii Bergdolt. 3. Subgenus Davalliopsis (Bosch) Ebihara & K. Iwats., comb. nov. Based on Davalliopsis Bosch (1861a) 323. — Trichomanes L. sect. Davalliopsis (Bosch) Prantl (1875) 53. — Cephalomanes C. Presl subg. Davalliopsis (Bosch) K. Iwats. (1984) 177. — Lectotype: Trichomanes prieurii Kunze (= Trichomanes elegans Rich.) (selected by Prantl (1875) 53). Rhizomes erect or ascending, quite thick, 4–15 mm diam., densely covered with dark hairs, protostele massive, cortex homogeneous, roots numerous and robust. Stipes to 50 cm long, close to each other. Blades tri- to quadripinnatifid, deltate, 60 by 36 cm, venation anadromous, false veinlets absent, laminae usually more than one cell thick, internal cell walls thick and coarsely pitted. Sori paratactic, tubular, lips truncate or slightly dilate, receptacles long-exserted. Distribution — The Neotropics; at least 1 species. Habitat — Terrestrial. Note — Corresponding to a part of the Da subclade in Ebihara et al. (submitted). REPRESENTATIVE SPECIES: Trichomanes elegans Rich. 248 BLUMEA — Vol. 51, No. 2, 2006 4. Subgenus Lacostea (Bosch) C. Chr. Trichomanes L. subg. Lacostea (Bosch) C. Chr. (1906) 634. — Lacostea Bosch (1861a) 320. — Trichomanes L. sect. Lacostea (Bosch) H. Christ (1897) 29. — Lectotype: Lacostea brachypus (Kunze) Prantl (= Trichomanes pedicellatum Desv.) (selected by Christensen (1906) XIV, as ‘T. pedicellatum Desv.’). Rhizomes long-creeping, frequently branching, rather thick, 0.5–1.2 mm diam., sparsely to densely covered with brown hairs, protostele massive, cortex heterogeneous, roots numerous and robust terrestrially, rootless at the climbing part. Stipes very short and unclear, subdistant to remote. Blades once-pinnate with symmetric pinnae, occasionally finer (up to tripinnatifid), elliptic, 30 by 9 cm, venation anadromous, false veinlets absent, internal cell walls slightly thick and wavy. Sori paratactic, tubular, lips truncate or sometimes dilate, receptacles long-exserted. Distribution — The Neotropics; more than 4 species. Habitat — Lianas on tree trunks. Note — Corresponding to a part of the Da subclade in Ebihara et al. (submitted). REPRESENTATIVE SPECIES: Trichomanes ankersii C. Parker ex Hook. & Grev.; T. pedicellatum Desv.; T. tanaicum Hook.; T. tuerckheimii H. Christ. 8. CEPHALOMANES Cephalomanes C. Presl (1843) 17, t. V. — Lacostea Bosch sect. Cephalomanes (C. Presl) Prantl (1875) 50. — Trichomanes L. subg. Cephalomanes (C. Presl) C. Chr. (1906) XIV. — Trichomanes L. sect. Cephalomanes (C. Presl) C.V. Morton (1968) 189. — Type: Cephalomanes atrovirens C. Presl. Rhizomes erect to short-creeping, thick, usually densely covered with old stipes, protostele massive, cortex homogeneous, roots numerous and robust. Stipes 2–15 cm long, close to each other. Blades once-pinnate with asymmetric pinnae, narrowly elliptic, 10–20 by 3–4(–5) cm, venation anadromous, false veinlets absent, internal cell walls thick and coarsely pitted. Sori paratactic, campanulate, lips sometimes dilate, receptacles long-exserted. Distribution — The Paleotropics (Asia to the Pacific); c. 4 species. Habitat — Terrestrial or epilithic along streams. Chromosome base number — x = 32. Note — Corresponding to the Ce clade in Ebihara et al. (submitted). REPRESENTATIVE SPECIES: Cephalomanes atrovirens C. Presl; C. crassum (Copel.) M.G. Price; C. javanicum (Blume) C. Presl. The Malagasian Trichomanes madagascariense (Bosch) Moore has traditionally been placed in this group, but unpublished molecular data suggest its exclusion. Further investigations are needed. 9. CALLISTOPTERIS Callistopteris Copel. (1938b) 64. — Trichomanes L. sect. Callistopteris (Copel.) C.V. Morton (1968) 190. — Cephalomanes C. Presl subg. Callistopteris (Copel.) K. Iwats. (1984) 176. — Type: Callistopteris apiifolia (C. Presl) Copel. Rhizomes erect to short-creeping, quite thick, hairy at apices, protostele massive, cortex homogeneous, roots numerous and robust. Stipes 5–25 cm long, close to each other, A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 249 densely hairy with long bristle-like light reddish hairs. Blades tripinnate to quadripinnate, elliptic to narrowly ovate, 40 by 16 cm, venation anadromous, false veinlets absent, internal cell walls thin and straight. Sori paratactic, campanulate, lips truncate, or sometimes bilabiate, receptacles long-exserted. Distribution — The Paleotropics (Asia to the Pacific); c. 5 species. Habitat — Usually epilithic on moist rocks, occasionally epiphytic on tree trunks. Chromosome base number — x = 36. Note — Corresponding to the Ca clade in Ebihara et al. (submitted). REPRESENTATIVE SPECIES: New combination: Callistopteris superba (Backh. ex T. Moore) Ebihara & K. Iwats., comb. nov. [based on Trichomanes superbum Backh. ex T. Moore (1862) 44]. Other species: Callistopteris apiifolia (C. Presl) Copel.; C. baldwinii (D.C. Eaton) Copel.; C. baueriana (Endl.) Copel.; C. polyantha (Hook.) Copel. ACKNOWLEDGEMENTS We are grateful to Ms. E. Wood (Harvard University Herbaria) for checking and correcting the English manuscript; to Dr. P.D. Bostock (Queensland Herbarium) for checking the Latin diagnosis; to the directors and curators of herbaria (BM, K, L and P) for supporting our investigation of their specimens; to Dr. N. Murakami and Dr. H. Nishida for providing some literature; to Dr. Leon Perrie (Museum of New Zealand Te Papa Tongarewa) for giving advices about the replacement for Cardiomanes reniforme and to the anonymous reviewer for valuable comments on the manuscript. This study was partly supported by the JSPS Grant for Oversea Research No. 12575012 (for K.I.) and by JSPS Fellows (for A.E.). REFERENCES Alderw. — See Van Alderwerelt van Rosenburgh, C.R.W.K. Allan, H.H.B. 1961. Flora of New Zealand, vol. 1. Government Printer, Wellington. Baker, J.G. 1866. Some new species of Hymenophyllaceae. J. Linn. Soc., Bot. 9: 335 – 341. Baksh-Comeau, Y.S. 2000. Checklist of the pteridophytes of Trinidad & Tobago. Fern Gaz. 16: 11–122. Bentham, G. 1862. Leguminosae. Dalbergiae. In: C.F.P. Martius & A.G. Eichler (eds.), Flora Brasiliensis. Vol. 15, 1: 1– 349. Fleischer, Leipzig. Blume, C.L. 1828. Enumeratio Plantarum Javae. Van Leeuwen, Lugduni Batavorum. Bonaparte, R. 1920. Les ptéridophytes de Madagascar, première partie. 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Acta Phytotax. Geobot. 28: 152–159. A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 255 Appendix. Taxonomic placement (genus, subgenus and section) of each basionym of Hymenophyllaceae (excl. later homonyms) in the present classification. Basionym Abrodictyum boninense Tagawa & K. Iwats. cumingii C. Presl Adiantum decurrens Jacq. tenellum Jacq. trifoliatum L. Amphipterum humatoides Copel. Apteropteris applanata A.M. Gray & R.G. Williams Buesia megistocarpa Copel. Callistopteris calyculata Copel. mulensis K. Iwats. Cephalomanes atrovirens C. Presl australicum Bosch curvatum Bosch madagascariense Bosch oblongifolium C. Presl rhomboideum Bosch singaporianum Bosch wilkesii Bosch Crepidomanes agasthianum Madhus. & C.A. Hameed chuii Ching & P.C. Chiu dilatatum Ching & Chu H. Wang hainanense Ching indicum C.A. Hameed & Madhus. liboense P.S. Wang lunulatum Madhus. & C.A. Hameed malabaricum C.A. Hameed & Madhus. nanophyllum Tagawa omeiense Ching & P.C. Chiu paucinervium Ching pinnatiidum Ching & P.C. Chiu pseudonymanii Hosok. sarawakense K. Iwats. smithiae Ching tagawanum K. Iwats. tiendongense Ching & C.F. Zhang yunnanense Ching & P.C. Chiu zayüense Ching & S.K. Wu Davallia tegularis Desv. Didymoglossum acanthoides Bosch aculeatum Bosch afine Bosch angustifrons Fée Genus Subgenus / Section Abrodictyum Abrodictyum Abrodictyum Abrodictyum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Hymenophyllum ? Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum Hymenophyllum Callistopteris Abrodictyum – Abrodictyum Cephalomanes Cephalomanes Cephalomanes Trichomanes Cephalomanes Cephalomanes Cephalomanes Cephalomanes – – – ? – – – – Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidium ? Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Hymenophyllum ? Hymenophyllum Hymenophyllum Hymenophyllum Didymoglossum Hymenophyllum Hymenophyllum Hymenophyllum Didymoglossum 256 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Didymoglossum (cont.) anomalum Bosch braunii Bosch brevipes C. Presl cordifolium Fée decipiens Desv. dilatatum Bosch euphlebium Bosch ferox Hassk. fructuosum Fée grifithii Bosch hedwigii C. Presl holochilum Bosch hookeri C. Presl insigne Bosch laceratum Fée latealatum Bosch lineolatum Bosch longisetum C. Presl magellanicum Desv. nummularium Bosch ovale E. Fourn. palmarum Vareschi plicatum Bosch preslii Bosch racemulosum Bosch serrulatum C. Presl sociale Fée undulatum C. Presl wesselsboeri Ebihara & Dubuisson Feea boryi Bosch humboldti Bosch nana Bory polypodina Bory Gonocormus australis Ching samoensis Copel. siamensis Tagawa & K. Iwats. Hymenophyllum abietinum Hook. & Grev. abruptum Hook. acrosorum Bosch aculeolatum Bosch adiantoides Bosch aequabile Kunze afine Brack. alatum Sm. alfredii Rosenst. alishanense De Vol alpinum Colenso alternatum Fosberg alveolatum C. Chr. Genus Subgenus / Section Crepidomanes Hymenophyllum Crepidomanes Didymoglossum Polyphlebium Crepidomanes Crepidomanes Hymenophyllum Didymoglossum Crepidomanes Didymoglossum Hymenophyllum Didymoglossum Crepidomanes Didymoglossum Crepidomanes Didymoglossum Abrodictyum Hymenophyllum Didymoglossum Didymoglossum Didymoglossum Crepidomanes Hymenophyllum Crepidomanes Hymenophyllum Didymoglossum Crepidomanes Didymoglossum Crepidomanes /Crepidomanes Hymenophyllum Crepidomanes /Crepidomanes Didymoglossum – Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Hymenophyllum Didymoglossum Crepidomanes /Crepidomanes Didymoglossum Hymenophyllum Didymoglossum Crepidomanes /Crepidomanes Didymoglossum Crepidomanes /Crepidomanes Didymoglossum Pachychaetum Myrmecostylum Didymoglossum Didymoglossum Didymoglossum Crepidomanes /Crepidomanes Hymenophyllum Crepidomanes /Crepidomanes Hymenophyllum Didymoglossum Crepidomanes /Crepidomanes Microgonium Trichomanes Trichomanes Trichomanes Trichomanes Feea Trichomanes Feea Feea Crepidomanes Crepidomanes Crepidomanes Crepidomanes /Gonocormus Crepidomanes /Gonocormus Crepidomanes /Gonocormus Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Vandenboschia ? Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Mecodium Mecodium Hymenophyllum ? Sphaerocionium Sphaerocionium Hymenophyllum Vandenboschia Mecodium Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum ? A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) amabile C.V. Morton amoenum Sturm andinum Bosch angulosum H. Christ angustatum Kunze angustifrons H. Christ angustum Bosch anisopterum Peter antarcticum C. Presl apicale Bosch apiculatum Kuhn apterum Bosch arbuscula Desv. asperulum Kunze assamense Gand. asterothrix Kunze atrosanguineum Bosch atrovirens Colenso attenuatum Hook. aucklandicum Bosch australe Willd. austrosinicum Ching axillare Sw. babindae Watts badium Hook. & Grev. baileyanum Domin bakeri Copel. balansae E. Fourn. baldwinii D.C. Eaton balfourii Baker bamlerianum Rosenst. batuense Rosenst. berteroi Hook. beyrichianum Kunze bibraianum Sturm bicolanum Copel. bismarckianum H. Christ blandum Racib. blepharodes C. Presl blumeanum A. Spreng. bontocense Copel. borneense Hook. boryanum Bory ex Willd. boschii Rosenst. botryoides Bosch bougainvillense Jeff W. Grimes boutoni Baker brachyglossum A. Braun ex Kunze brachypus Sodiro braithwaitei Ebihara & K. Iwats. brassii C. Chr. breve Rosenst. Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Callistopteris Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Sphaerocionium Mecodium Globosa Sphaerocionium Sphaerocionium Sphaerocionium Sphaerocionium Hymenophyllum Mecodium Mecodium Sphaerocionium Sphaerocionium Hymenophyllum Globosa Sphaerocionium Mecodium Globosa Myrmecostylum Globosa Globosa Hymenophyllum Mecodium Hymenophyllum Globosa Hymenophyllum Hymenophyllum Globosa – Mecodium Globosa Hymenophyllum Sphaerocionium Sphaerocionium Myrmecostylum Hymenophyllum Globosa Hymenophyllum Hymenophyllum Mecodium Hymenophyllum Sphaerocionium Sphaerocionium Hymenophyllum Mecodium Hymenophyllum Sphaerocionium Hymenophyllum Sphaerocionium ? Sphaerocionium Hymenophyllum Mecodium ? 257 258 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) brevidens Alderw. brevifrons Kunze brevistipes Liebm. bridgesii Hook. bryophilum C. Chr. buchtienii Rosenst. caespitosum Gaudich. calodictyon Bosch campanulatum H. Christ caparaoense Brade capense Schrad. capillaceum Roxb. capillare Desv. capurroi de la Sota cardunculus C. Chr. carnosum H. Christ catherinae Hook. caudatellum H. Christ caudatum Bosch caudiculatum Mart. caulopteron Fée ceratophylloides H. Christ cernuum A. Gepp cheesemannii Baker chiloense Hook. chrysothrix Sturm cincinnatum A. Gepp clemensiae Copel. cocosense A. Rojas coloratum A. Braun ex Bosch compactum Bonap. consanguineum C.V. Morton constrictum H. Christ contextum Rosenst. contortum Bosch contractile Sodiro convolutum Bosch copelandianum Alderw. copelandii C.V. Morton coreanum Nakai corrugatum H. Christ costaricanum Bosch crassipetiolatum Stolze crispato-alatum Hayata crispatulum Bosch crispatum Wall. ex Hook. & Grev. crispum Kunth cristatum Hook. & Grev. cristulatum Rosenst. cruegeri Müll.Berol. cruentum Cav. cubense Sturm Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Mecodium Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Mecodium Mecodium ? Sphaerocionium Sphaerocionium Hymenophyllum Mecodium Sphaerocionium Sphaerocionium Globosa Globosa Sphaerocionium Sphaerocionium Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum Hymenophyllum Sphaerocionium Mecodium Mecodium Sphaerocionium Sphaerocionium Mecodium Mecodium Sphaerocionium Mecodium Globosa Mecodium Mecodium Mecodium Mecodium Sphaerocionium Globosa Sphaerocionium Globosa Sphaerocionium Hymenophyllum Myrmecostylum Sphaerocionium Hymenoglossum Sphaerocionium A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) cumingii C. Presl cuneatum Kunze cupressiforme Labill. daedaleum Blume darwinii Hook. dejectum Baker delavayi H. Christ delicatissimum Fée delicatulum Sehnem deltoideum C. Chr. dendritis Rosenst. densifolium Phil. dentatum Cav. denticulatum Sw. dependens C.V. Morton deplanchei Mett. ex Kuhn dichotomum Cav. dimidiatum Mett. ex Kuhn dimorphum H. Christ dipteroneuron A. Braun ex Kunze discosum H. Christ divaricatum Bosch dregeanum C. Presl durandi H. Christ dusenii H. Christ eboracense Croxall ectocarpon Fée elasticum Bory ex Willd. elatius H. Christ elberti Rosenst. elegans A. Spreng. elegantissimum Fée elegantulum Bosch ellipticosorum Alderw. elongatum Jeff W. Grimes emarginatum Sw. emersum Baker endiviaefolium Desv. epiphyticum J.W. Moore erecto-alatum Colenso eriophorum Bosch erosum Blume exiguum Bedd. eximium Kunze exsertum Wall. ex Hook. falklandicum Baker farallonense Hieron. fastigiosum H. Christ fecundum Bosch feejeense Brack. fendlerianum Sturm ferax Bosch Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllopsis Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Didymoglossum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Mecodium Hymenophyllum Globosa Mecodium – Hymenophyllum Sphaerocionium Sphaerocionium Hymenophyllum Mecodium Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum Mecodium Sphaerocionium Hymenophyllum Sphaerocionium Hymenophyllum Globosa Hymenophyllum Sphaerocionium Sphaerocionium Hymenophyllum Sphaerocionium Sphaerocionium Sphaerocionium Hymenophyllum Hymenophyllum Globosa Hymenophyllum Mecodium Mecodium Globosa Sphaerocionium Globosa Didymoglossum Globosa Hymenophyllum Hymenophyllum Mecodium Hymenophyllum Mecodium Hymenophyllum Mecodium Mecodium 259 260 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) ferrugineum Colla ilicula Bory ex Willd. imbriatum J. Sm. ex Hook. irmum Alderw. labellatum Labill. laccidum Bosch lavo-aureum Bory lexile Makino lexuosum A. Cunn. loribundum Kunth foersteri Rosenst. formosum Brack. foxworthyi Copel. francavillei Bosch frankliniae Colenso franklinianum Colenso fraseri Mett. fraternum C. Presl fuciforme Sw. fucoides Cav. fuertesii Brause fujisanense Nakai fulvum Bosch fumarioides Bory ex Willd. funckii Bosch fusugasugense H. Karst. ex Sturm gardneri Bosch gardnerianum Sturm geluense Rosenst. glaziovii Baker glebarium H. Christ gollmeri Bosch gorgoneum Copel. gracile Bory ex Willd. gracilescens Domin gracilius Copel. gratum Fée grevilleanum C. Presl gunnii Bosch ex Baker haematochroum Sodiro halconense Copel. hallieri Rosenst. hamuliferum Alderw. hayatai Masam. heimii Tardieu helicoideum Sodiro hemidimorphum R.C. Moran & B. Øllg. hemipteron Rosenst. henkelii Sim henryi Baker l’herminieri Mett. ex Kuhn herterianum Brause Hymenophyllum Crepidomanes Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Crepidomanes /Crepidomanes Globosa Hymenophyllum Pleuromanes Mecodium Sphaerocionium Globosa Globosa Mecodium ? Hymenophyllum Globosa Pleuromanes Sphaerocionium Sphaerocionium Sphaerocionium Sphaerocionium Mecodium Fuciformia Fuciformia Sphaerocionium Mecodium Sphaerocionium Mecodium Mecodium Sphaerocionium Hymenophyllum Sphaerocionium Hymenophyllum Sphaerocionium Hymenophyllum Mecodium Hymenophyllum Mecodium Hymenophyllum Mecodium Sphaerocionium Mecodium Mecodium Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Hymenoglossum Mecodium Sphaerocionium Sphaerocionium Mecodium Hymenophyllum Mecodium ? Hymenophyllum A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) herzogii Rosenst. himalaianum Bosch hirtellum Sw. holotrichum Peter hookeri Bory horizontale C.V. Morton hosei Copel. houstonii Jenman howense Brownlie humbertii C. Chr. humboldtianum E. Fourn. humile Nees & Blume imbricatum Blume inclinatum Bosch infortunatum Bory integrivalvatum C. Sánchez integrum Bosch intercalatum H. Christ interruptum Kunze intricatum Bosch involucratum Copel. ivohibense Tardieu jalapense Schltdl. & Cham. jamesoni Hook. japonicum Miq. javanicum A. Spreng. johorense Holttum junghuhnii Bosch kaieteurum Jenman karstenianum Sturm kerianum Watts khasianum Baker klabatense H. Christ kohautianum C. Presl krauseanum Phil. kuhnii C. Chr. kurzii Prantl laciniosum H. Christ lamellatum Stolze laminatum Copel. lanatum Fée lanceolatum Hook. & Arn. latifrons Bosch latilobum Bonap. ledermanni Brause lehmannii Hieron. leptocarpum Copel. leptodictyon Müll.Berol. leratii Rosenst. levingei C.B. Clarke limminghei Bosch lindenii Hook. Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Mecodium Sphaerocionium Sphaerocionium Pleuromanes Sphaerocionium Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Globosa Hymenophyllum Globosa Globosa Mecodium Hymenophyllum Mecodium Sphaerocionium Sphaerocionium Globosa Mecodium Sphaerocionium Mecodium Hymenophyllum Hymenophyllum Globosa Hymenophyllum Globosa Sphaerocionium Sphaerocionium Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Myrmecostylum Mecodium Hymenophyllum Mecodium Hymenophyllum Hymenophyllum Sphaerocionium Sphaerocionium Sphaerocionium Globosa Hymenophyllum Mecodium Hymenophyllum Globosa Pleuromanes Hymenophyllum Mecodium Sphaerocionium 261 262 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) lindigii Mett. lindsaeoides Baker lingganum Alderw. lobatoalatum Klotzsch lobatopapillosum Klotzsch ex Sadeb. lobbii T. Moore ex Bosch longifolium Alderw. lophocarpum Colenso lyallii Hook.f. macroglossum Bosch macrosorum Alderw. macrothecum Fée maderense Gibby & Lovis malaccense Gand. mannianum Mett. ex Kuhn marginatum Hook. & Grev. marlothii Brause martii Sturm martinicense Bosch matthewsii Bosch maxonii H. Christ ex C.V. Morton mazei E. Fourn. ex H. Christ megachilum C. Presl megalocarpum Colenso melanocheilos Colenso mentitum Gand. menziesii C. Presl merrillii H. Christ mettenii Bosch meyeri C. Presl micans H. Christ micranthum Bosch microcarpon Fée microcarpum Desv. microphyllum Mett. microsorum Bosch millefolium Schltdl. & Cham. minimum A. Rich. minutidenticulatum Ching & P.C. Chiu miriicum C.V. Morton mnioides Baker modestum Bosch molle C.V. Morton montanum Kirk moorei Baker moritziaum Sturm mortonianum Lellinger multialatum C.V. Morton multilorum Rosenst. myriocarpum Hook. nahuelhuapiense Diem & J.S. Licht. nanostellatum Lellinger Hymenophyllum Sphenomeris Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium – Hymenophyllum Sphaerocionium Sphaerocionium Hymenophyllum Globosa Myrmecostylum Sphaerocionium Hymenophyllum Hymenophyllum Mecodium Hymenophyllum Globosa Hymenophyllum Hymenophyllum Sphaerocionium Globosa Mecodium Mecodium Sphaerocionium Mecodium Hymenophyllum Globosa Hymenophyllum Globosa Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Globosa Sphaerocionium Sphaerocionium Mecodium Mecodium Mecodium Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Globosa Sphaerocionium Globosa Hymenophyllum Sphaerocionium Sphaerocionium Sphaerocionium Mecodium Mecodium Hymenophyllum Sphaerocionium A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 263 Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) nanum Sodiro natalense Bosch neozelandicum Gand. nephrophyllum Ebihara & K. Iwats. nigrescens Liebm. nigricans Colla nitens R.Br. nitiduloides Copel. notabile Fée nutantifolium Alderw. obtusum Hook. & Arn. odontophyllum Copel. okadai Masam. oligocarpum Colenso oligosorum Makino omeiense H. Christ ooides F. Muell. & Baker opacum Copel. orbignianum Bosch organense Hook. osmundoides Bosch ovatum Copel. oxyodon Baker pachydermicum Ces. palmatum Bosch palmense Rosenst. paniculilorum C. Presl paniense Ebihara & K. Iwats. pannosum H. Christ pantotactum Alderw. parallelocarpum Hayata parvifolium Baker parvulum C. Chr. pastoensis Hook. patagonicum Gand. paucicarpum Jenman pectinatum Cav. pedicellatum Kunze ex Klotzsch pedicularifolium Ces. penangianum C.G. Matthew & H. Christ pendulum Bory perissum Copel. perparvulum Alderw. perrieri Tardieu peruvianum Hook. & Grev. physocarpum H. Christ piliferum C. Chr. pilosissimum C. Chr. pilosum Alderw. platylobum Bosch pleiocarpum Alderw. plicatum Kaulf. Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Crepidomanes Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Globosa Cardiomanes Mecodium Myrmecostylum Pleuromanes Mecodium Sphaerocionium Hymenophyllum Sphaerocionium Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Globosa Sphaerocionium Sphaerocionium Mecodium Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Mecodium Myrmecostylum Sphaerocionium Globosa Mecodium Crepidomanes /Crepidomanes Mecodium Sphaerocionium Globosa Mecodium Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Globosa Hymenophyllum Sphaerocionium Hymenophyllum Sphaerocionium Globosa Myrmecostylum 264 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) plumieri Hook. & Grev. plumosum Kaulf. podocarpon Fée poeppigianum C. Presl poilanei Tardieu & C. Chr. pollenianum Rosenst. polyanthum Hook. polycarpum Mett. ex Kuhn polychilum Colenso polyodon Baker poolii Baker portoricense Kuhn praetervisum H. Christ prionema Kunze ex Sturm procerum Bosch proctoris C. Sánchez producens Fée productoides J.W. Moore productum Kunze protrusum Hook. pseudotunbridgense Watts pteropodum Bosch puellum Ces. pulchellum Schltdl. & Cham. pulcherrimum Colenso pulchrum Copel. pumilio Rosenst. pumilum C. Moore punctisorum Rosenst. pusillum Schott ex Sturm pycnocarpum Bosch pygmaeum Colenso pyramidatum Desv. pyriforme Bosch quadriidum Phil. quetrihuense Diem & J.S. Licht. raapii Gand. raddianum Müll.Berol. ramosii Copel. ramosissimum Ham. ex D. Don rarum R.Br. recurvum Gaudich. reductum Copel. refrondescens Sodiro reinwardtii Bosch remotipinna Bonap. remotum Bosch reniforme Hook. repens Dulac retusilobum Hayata revolutum Colenso ricciaefolium Bory ex Willd. Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Callistopteris Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Abrodictyum ? Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphenomeris Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Sphaerocionium Hymenophyllum Mecodium Hymenophyllum Mecodium – Mecodium Globosa Hymenophyllum Sphaerocionium Sphaerocionium Hymenophyllum Sphaerocionium Sphaerocionium Sphaerocionium Sphaerocionium Globosa Globosa Mecodium Hymenophyllum Sphaerocionium Abrodictyum ? Sphaerocionium Fuciformia Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Mecodium Mecodium Hymenophyllum Sphaerocionium Hymenophyllum Myrmecostylum Hymenophyllum Globosa Sphaerocionium Hymenophyllum – Mecodium Mecodium Hymenophyllum Sphaerocionium Globosa Mecodium Sphaerocionium Mecodium Hymenophyllum Globosa Hymenophyllum Hymenophyllum A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) rimbachii Sodiro ringens H. Christ riukiuense H. Christ rolandi-principis Rosenst. roraimense C.V. Morton rosenstockii Brause rubellum Rosenst. rufescens Kirk ruifolium Alderw. ruifrons Alderw. rufoibrillosum Ching & Z.Y. Liu rufum Fée rugosum C. Chr. & Skottsb. rupestre Raddi sabinifolium Baker saenzianum L.D. Gómez salakense Racib. samoense Baker sampaioanum Brade & Rosenst. scabrum A. Rich. schomburghii C. Presl secundum Hook. & Grev. semibivalve Hook. & Grev. semiissum Copel. semiglabrum Rosenst. semiteres Colla serra C. Presl seselifolium C. Presl shirleyanum Domin siliquosum H. Christ silvaticum C.V. Morton silveirae H. Christ simonsianum Hook. simplex C.V. Morton skottsbergii C. Chr. smithii Hook. sodiroi C. Chr. spathulatum Colenso speciosum Bosch spectabile Mett. ex Kuhn sphaerocarpum Bosch spicatum H. Christ spinosum Ching spinulosum Kunth splendidum Bosch sprucei Baker steerei C. Chr. streptophyllum E. Fourn. sturmii Bosch subdeltoideum H. Christ subdemissum H. Christ subdimidiatum Rosenst. Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Vandenboschia Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Hymenophyllum Globosa Hymenophyllum Sphaerocionium Hymenophyllum Hymenophyllum Pleuromanes Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum Lacosteopsis Hymenophyllum Sphaerocionium Globosa Globosa Sphaerocionium Myrmecostylum Mecodium Hymenophyllum Mecodium Hymenophyllum Sphaerocionium Fuciformia Hymenophyllum Myrmecostylum Globosa Mecodium Sphaerocionium Sphaerocionium Hymenophyllum Sphaerocionium Mecodium Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Sphaerocionium Mecodium Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Sphaerocionium ? Globosa Globosa Mecodium Mecodium Mecodium Hymenophyllum 265 266 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Hymenophyllum (cont.) subirmum Alderw. sublabellatum Ces. subobtusum Rosenst. subrigidum H. Christ subrotundum Alderw. subtilissimum Kunze superbum C.V. Morton suprapaleaceum Ching surinamense Bosch tablaziense H. Christ tabulare Bosch taliabense Alderw. tarapotense Stolze tasmannicum Bosch tayloriae Farrar & Raine tenellum D. Don tenerrimum Bosch tenerum Bosch terminale Bosch thomassetii C.H. Wright thuidium Harr. thunbergii Eckl. ex C. Presl todjambuense Kjellb. tomentosum Kunze torquescens Bosch torricellianum Alderw. tortuosum Hook. & Grev. trapezoidale Liebm. treubii Racib. trianae Hieron. triangulare Baker trichocaulon Phil. trichomanoides Bosch trichophyllum Kunth triidum Hook. & Grev. truncatum Colenso turquinense C. Sánchez ulei H. Christ & Giesenh. umbratile Diem & J.S. Licht. uncinatum Sim unilaterale Bory ex Willd. urbanii Brause urofrons Ching & C.F. Zhang vacillans H. Christ valvatum Hook. & Grev. venustum Desv. verecundum C.V. Morton veronicoides C. Chr. viguieri Tardieu villosum Colenso vincentinum Baker viride Rosenst. ex Copel. Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Sphaerocionium Hymenophyllum Sphaerocionium Sphaerocionium Hymenophyllum Sphaerocionium Mecodium Mecodium Hymenophyllum Sphaerocionium Globosa Sphaerocionium Mecodium Sphaerocionium Mecodium Sphaerocionium Hymenophyllum Globosa Mecodium Globosa Sphaerocionium Hymenophyllum Hymenophyllum Myrmecostylum Sphaerocionium Globosa Mecodium Hymenophyllum Hymenophyllum Mecodium Sphaerocionium Sphaerocionium Hymenophyllum Sphaerocionium Sphaerocionium Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum Hymenophyllum ? Sphaerocionium Sphaerocionium Sphaerocionium Mecodium Sphaerocionium Myrmecostylum Mecodium Hymenophyllum A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae Appendix. Continued. Basionym Hymenophyllum (cont.) viridissimum Fée vittatum Copel. walleri Maiden & Betche wercklei H. Christ whangshanense Ching & P.C. Chiu whitei Goy wilsonii Hook. wrightii Bosch zeelandicum Bosch zeyheri Bosch zollingerianum Kunze Hymenostachys diversifrons Bory elegans C. Presl Leptocionium barbatum Bosch dicranotrichum C. Presl edentulum Bosch laccidum Bosch sororium C. Presl violaceum Meyen ex Bosch Macroglena brassii Croxall kalimantanensis K. Iwats. & M. Kato obtusa Copel. toppingii O. Deg. & I. Deg. truncata Copel. Mecodium acrocarpum H. Christ ex Ching archboldii Copel. contiguum D.A. Sm. diversilabium Copel. edanoi Copel. gongboense Ching hainanense Ching jinfoshanense Ching & Z.Y. Liu kansuense Ching & Y.P. Hsu likiangense Ching & P.C. Chiu lineatum Ching & P.C. Chiu lofoushanense Ching & P.C. Chiu longissimum Ching & P.C. Chiu lushanense Ching & P.C. Chiu mexiae Copel. mikawanum Seriz. ovalifolium Ching & P.C. Chiu paramnioides H.G. Zhou & W.M. Chu propinquum Ching & P.C. Chiu stenochladum Ching & P.C. Chiu szechuanense Ching & P.C. Chiu taiwanense Tagawa tenellum Parris tenuifrons Ching Genus Subgenus / Section Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Hymenophyllum Hymenophyllum Sphaerocionium Hymenophyllum Mecodium Hymenophyllum Mecodium Hymenophyllum Mecodium Hymenophyllum Trichomanes Trichomanes Feea Feea Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Diploophyllum Hymenophyllum Abrodictyum Abrodictyum Vandenboschia Callistopteris Abrodictyum Abrodictyum Pachychaetum Vandenboschia – Abrodictyum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Mecodium Mecodium Mecodium Globosa Mecodium ? Mecodium Mecodium Mecodium Mecodium Globosa Mecodium ? Mecodium ? Mecodium Mecodium Mecodium Mecodium Mecodium Globosa Globosa Mecodium Globosa Mecodium Globosa 267 268 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Mecodium (cont.) wangii Ching & P.C. Chiu wenhsienense Ching & Y.P. Hsu Meringium archboldii Copel. bartlettii Copel. blumeanum C. Presl latifolium Copel. laxum Copel. melanosorum Copel. meyenianum C. Presl Microgonium benlii Pic.Serm. berteroanum C. Presl bimarginatum Bosch falsinervulosum M. Nishida minutifolium Tagawa & K. Iwats. omphalodes Vieill. ex E. Fourn. Microtrichomanes zamboanganum Copel. Neuromanes abruptum Bosch immersum Bosch kaulfussii Bosch Neurophyllum hedwigianum Fée hostmannianum Klotzsch thecaphyllum Fée Pleuromanes retusum Copel. Ragatelus eriophorus C. Presl Selenodesmium recurvum Ching & P.C. Chiu Sphaerocionium aureum C. Presl breutelli C. Presl commutatum C. Presl diversilobum C. Presl glanduliferum C. Presl grevilleanum C. Presl macrocarpum C. Presl nigricans C. Presl productum C. Presl ruizianum Klotzsch schiedeanum C. Presl sieberi C. Presl × tucuchense Jermy & T.G. Walker vestitum C. Presl Trichomanes abrotanifolium Bosch abruptum Fée ex H. Christ accedens C. Presl Genus Subgenus / Section Hymenophyllum Hymenophyllum Mecodium Mecodium Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Mecodium Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Didymoglossum Didymoglossum Didymoglossum Didymoglossum Crepidomanes Didymoglossum Microgonium Microgonium Microgonium Microgonium Crepidomanes /Crepidomanes Didymoglossum Hymenophyllum Sphaerocionium Trichomanes Trichomanes Trichomanes Trichomanes Trichomanes Trichomanes Trichomanes Trichomanes Trichomanes Trichomanes Trichomanes Trichomanes Hymenophyllum Pleuromanes Trichomanes Trichomanes Abrodictyum Pachychaetum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Hymenophyllum Sphaerocionium Mecodium Sphaerocionium Sphaerocionium Myrmecostylum Sphaerocionium Globosa Mecodium Globosa Sphaerocionium Sphaerocionium Sphaerocionium Sphaerocionium Sphaerocionium Vandenboschia ? Trichomanes Trichomanes ? Trichomanes Trichomanes A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 269 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) achilleaefolium Bory ex Willd. acranthum H. Itô acrocarpon Fée acrosorum Copel. acutilobum Ching acuto-obtusum Hayata acutum C. Presl adianthinum Bory adiantoides L. adscendens Kunze aerugineum Bosch aeruginosum Poir. aethiopicum Burm.f. africanum H. Christ alagense H. Christ alatum Sw. album Blume alternans Carruth. ex Seem. amabile Nakai amazonicum H. Christ ambiguum Mart. ambongoense Bonap. anadromum Rosenst. anceps Hook. andrewsii Newman angustatum Carmich. angustilaciniatum Bonap. angustimarginatum Bonap. angustissimum C. Presl ankersii C. Parker ex Hook. & Grev. anomalum Maxon & C.V. Morton antillarum Bosch aphlebioides H. Christ apicilare E. Fourn. apiifolium C. Presl apodum Hook. & Grev. arbuscula Desv. armstrongii Baker asae-grayi Bosch asnykii Racib. asplenioides Sw. assimile Mett. ex Kuhn astylum Kaulf. attenuatum Hook. auratum Fée aureovestitum Proctor aureum Bosch auriculatum Blume axillare Sodiro badium E. Fourn. baileyanum Watts ballardianum Alston Abrodictyum Cephalomanes Didymoglossum Cephalomanes Crepidomanes Crepidomanes Hymenophyllum Didymoglossum Asplenium Trichomanes ? Didymoglossum Hymenophyllum Asplenium Crepidomanes Crepidomanes Trichomanes Hymenophyllum Polyphlebium Vandenboschia Trichomanes Polyphlebium Crepidomanes Trichomanes Trichomanes Vandenboschia Polyphlebium Crepidomanes Abrodictyum Polyphlebium Trichomanes Trichomanes Vandenboschia Crepidomanes Crepidomanes Callistopteris Didymoglossum Trichomanes Hymenophyllum Abrodictyum Crepidomanes Hymenophyllum Crepidomanes Trichomanes Trichomanes Trichomanes Trichomanes Polyphlebium Vandenboschia Polyphlebium ? Trichomanes Didymoglossum Didymoglossum Pachychaetum – Didymoglossum – Crepidomanes/Crepidomanes Crepidomanes/Crepidomanes Pleuromanes Microgonium – ? Microgonium Sphaerocionium – Crepidomanes /Crepidomanes Crepidomanes /Gonocormus Trichomanes Pleuromanes – Vandenboschia Trichomanes – Crepidomanes /Gonocormus Trichomanes Davalliopsis Vandenboschia – Crepidomanes /Gonocormus Pachychaetum – Lacostea Trichomanes Vandenboschia Nesopteris Crepidomanes /Crepidium – Didymoglossum Trichomanes Hymenophyllum Abrodictyum Crepidomanes /Crepidomanes Hymenoglossum Crepidomanes /Gonocormus Trichomanes Trichomanes Trichomanes Trichomanes – Lacosteopsis – Trichomanes Microgonium Microgonium 270 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) bancroftii Hook. & Grev. barklianum Baker barnardianum F.M. Bailey batrachoglossum Copel. bauerianum Endl. beccarianum Ces. beckeri Krause ex Phil. belangeri Bory bicorne Hook. biidum Vent. ex Willd. bifolium Blume bilabiatum Nees & Blume bilobatum Alderw. bipunctatum Poir. birmanicum Bedd. bissei C. Sánchez bivalve G. Forst. blepharistomum Copel. blumei Hassk. bojeri Hook. & Grev. bonapartei C. Chr. bonincolum Nakai boninense Koidz. borbonicum Bosch borneense Alderw. boryanum Kunze boschianum Sturm ex Bosch botryoides Kaulf. bovini Bosch brachyblastos Mett. ex Kuhn brachypus Kunze bradei H. Christ brasiliense Desv. braunii Bosch brevisetum R.Br. brooksii Copel. bryoides Goldm. bucinatum Mickel & Beitel caespifrons C. Chr. caliginum Lellinger calufii C. Sánchez calvescens Bosch campanulatum Roxb. canariensis L. capillaceum L. capillatum Taschner carolianum Vareschi cartilagineum Vieill. & Pancher caruifolium Roxb. caudatum Brack. cavifolium Müll.Berol. cellulosum Klotzsch Trichomanes Didymoglossum Crepidomanes Abrodictyum ? Callistopteris Didymoglossum Hymenophyllum Vandenboschia Trichomanes Abrodictyum Crepidomanes Crepidomanes Crepidomanes Crepidomanes Vandenboschia Trichomanes Hymenophyllum Crepidomanes Hymenophyllum Didymoglossum Crepidomanes Crepidomanes Crepidomanes Polyphlebium Cephalomanes Cephalomanes Vandenboschia Trichomanes Trichomanes Vandenboschia Trichomanes Polyphlebium Polyphlebium Hymenophyllum Vandenboschia Crepidomanes ? Didymoglossum Abrodictyum Trichomanes Didymoglossum Hymenophyllum Crepidomanes ? Davallia Polyphlebium Crepidomanes Trichomanes Abrodictyum Vandenboschia ? Abrodictyum Polyphlebium Abrodictyum Trichomanes Didymoglossum Crepidomanes /Crepidomanes ? – Didymoglossum Myrmecostylum Lacosteopsis Trichomanes Pachychaetum Crepidomanes /Gonocormus Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Vandenboschia Trichomanes Hymenophyllum Nesopteris Sphaerocionium Microgonium Crepidomanes /Crepidomanes ? Crepidomanes /Gonocormus Crepidomanes /Crepidomanes – – – Vandenboschia Feea ? Vandenboschia Lacostea – – Pleuromanes Vandenboschia Crepidomanes /Gonocormus ? Didymoglossum Pachychaetum Trichomanes ? Sphaerocionium ? – – Crepidomanes /Crepidomanes Lacostea Pachychaetum ? Abrodictyum – Pachychaetum A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 271 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) chamaedrys Taton charophylloides Poir. chevalieri H. Christ chinensis L. christii Copel. ciliatum Sw. clarenceanum F. Ballard clathratum Tagawa clavatum Sw. cochinchinense Poir. cocos H. Christ cognatum C. Presl colensoi Hook. collariatum Bosch commutatum Sturm compactum Alderw. compressum Desv. concinnum Mett. ex Kuhn contiguum G. Forst. corcovadense Bosch coriaceum Kunze cormophylum Kaulf. cornutum C. Chr. corticola Hook. ex Bedd. craspedoneurum Copel. crassipilis Weath. crassum Copel. crenatum Bosch crinitum Sw. crispiforme Alston crispulum Bosch crispum L. cristatum Kaulf. cultratum Baker cuneatum H. Christ cuneiforme G. Forst. cupressifolium Hayata cupressoides Desv. curranii Weath. curtii Rosenst. cuspidatum Willd. cyrtotheca Hillebr. cystoseiroides H. Christ ex Tardieu & C. Chr. dactylites Sodiro daguense Weath. daucoides C. Presl davallioides Gaudich. debile Bosch delicatum Bosch demissum G. Forst. densinervium Copel. dentatum Bosch Didymoglossum Davallia Crepidomanes Sphenomeris Crepidomanes Hymenophyllum Crepidomanes Abrodictyum Hymenophyllum Lygodium Polyphlebium Trichomanes Polyphlebium Vandenboschia Trichomanes Abrodictyum Abrodictyum Crepidomanes Davallia Vandenboschia ? Trichomanes Cyathea Trichomanes ? Hymenophyllum Didymoglossum Trichomanes Cephalomanes Trichomanes Trichomanes Trichomanes Didymoglossum Trichomanes Trichomanes Didymoglossum Hymenophyllum Odontosoria Crepidomanes Abrodictyum Trichomanes Didymoglossum Didymoglossum Vandenboschia Vandenboschia Trichomanes Trichomanes Abrodictyum Vandenboschia Polyphlebium Trichomanes Hymenophyllum Cephalomanes Abrodictyum Microgonium – Crepidomanes /Crepidomanes – Crepidomanes /Crepidomanes Sphaerocionium Crepidomanes /Crepidomanes Abrodictyum Mecodium – – Trichomanes – Vandenboschia Lacostea Abrodictyum Pachychaetum Crepidomanes /Crepidium – ? Trichomanes – ? Sphaerocionium Microgonium Trichomanes – Trichomanes Trichomanes Trichomanes Microgonium Trichomanes Trichomanes Didymoglossum Hymenophyllum ? – Crepidomanes /Crepidomanes Pachychaetum Trichomanes Didymoglossum Microgonium Vandenboschia Vandenboschia Trichomanes Trichomanes Pachychaetum Vandenboschia – Trichomanes Globosa – Pachychaetum 272 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) denticulatum Houtt. depauperatum Bory diaphanum Kunth dichotomum Kunze diffusum Blume digitatum Sw. dilatatum G. Forst. dimidiatum C. Presl dimorphum Mett. ex Sadeb. dissectum J. Sm. ex Hook. divaricatum Poir. draytonianum Brack. dregei Bosch egleri P.G. Windisch ekmanii Wess.Boer elatum G. Forst. elegans Rich. elongatum A. Cunn. eminens C. Presl endlicherianum C. Presl englerianum Brause epiphyllum G. Forst. erectum Brack. ericoides Hedw. ex Hook. erosum Willd. europaeum Sm. exaltatum Brack. eximium Kunze ex Sturm exsectum Kunze extravagans Copel. fallax H. Christ fargesii H. Christ ferrugineum E. Fourn. iliculoides H. Christ iliforme Sturm imbriatum Backh. ex T. Moore irmulum C. Presl labellatum Bory labelliforme Hassk. labellula d’Urv. laccidum G. Forst. lavofuscum Bosch loribundum Hook. & Grev. foeniculaceum Bory ex Willd. foersteri Rosenst. fontanum Lindm. formosanum Y. Yabe fragile Hedw. francii H. Christ frappieri Cordem. fraseri Jenman frondosum Fée Davallia Crepidomanes Polyphlebium Hymenophyllum Crepidomanes Hymenophyllum Hymenophyllum Vandenboschia Trichomanes Vandenboschia Hymenophyllum Crepidomanes Abrodictyum Trichomanes Didymoglossum Davallia Trichomanes Abrodictyum Callistopteris Polyphlebium Abrodictyum Davallia Polyphlebium Abrodictyum Didymoglossum Vandenboschia Callistopteris Polyphlebium Polyphlebium Abrodictyum Crepidomanes Vandenboschia Abrodictyum Crepidomanes Abrodictyum Trichomanes Abrodictyum Hymenophyllum Hymenophyllum Hymenophyllum Dennstaedtia Abrodictyum Trichomanes Abrodictyum Cephalomanes Didymoglossum Crepidomanes Hymenophyllum Hymenophyllum Crepidomanes Didymoglossum Vandenboschia – Crepidomanes /Crepidium – Sphaerocionium Crepidomanes /Gonocormus Sphaerocionium Diploophyllum Lacosteopsis Feea Lacosteopsis ? Crepidomanes /Crepidomanes Pachychaetum Trichomanes Microgonium – Davalliopsis Pachychaetum – – Pachychaetum – – ? Microgonium Vandenboschia – – – Pachychaetum Crepidomanes /Crepidomanes Lacosteopsis ? Pachychaetum Crepidomanes /Crepidium Pachychaetum Trichomanes Pachychaetum Hymenophyllum Sphaerocionium Hymenophyllum – Abrodictyum Trichomanes Pachychaetum – Didymoglossum Crepidomanes /Crepidomanes Sphaerocionium Hymenophyllum ? Crepidomanes /Crepidomanes Didymoglossum Lacosteopsis A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 273 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) fruticulosum Jenman fucoides Sw. fulgens C. Chr. fulvum Klotzsch furcatum Kaulf. ex Spreng. fuscum Blume galeottii E. Fourn. gardneri Bosch gemmatum J. Sm. ex Baker gibberosum G. Forst. giesenhagenii C. Chr. giganteum Bory ex Willd. glaucofuscum Hook. godmanii Hook. goebelianum Giesenh. goetzii Hieron. gourlianum Grev. ex J. Sm. gracile Bosch gracillimum Copel. grande Copel. graucescens Bosch guianense Sturm guidoi P.G. Windisch guineense Afzel. ex Sw. haenkei C. Presl hartii Baker harveyi Carruth. ex Seem. haughtii C.V. Morton henzaianum Parish ex Hook. l’herminieri Fée herzogii Rosenst. hibernicum A. Spreng. hieronymi Brause hildebrandtii Kuhn hirsutum L. hispidulum Mett. ex Kuhn hispidum Poir. holopterum Kunze hookeri C. Presl hosei Baker huberi H. Christ humile G. Forst. hygrometricum Poir. hymenoides Hedw. hymenophylloides Bosch hypnoides H. Christ idoneum C.V. Morton ignobile Ces. imbricatum Sodiro inaequale Poir. incisum Thunb. inerme Bosch Didymoglossum Hymenophyllum Didymoglossum Polyphlebium Abrodictyum Hymenophyllum Trichomanes Trichomanes Abrodictyum Asplenium Didymoglossum Vandenboschia Hymenophyllum Didymoglossum Didymoglossum Polyphlebium Didymoglossum Crepidomanes Crepidomanes Crepidomanes Hymenophyllum Trichomanes Trichomanes Abrodictyum Trichomanes Abrodictyum ? Crepidomanes Polyphlebium Didymoglossum Trichomanes Polyphlebium Vandenboschia Hymenophyllum Didymoglossum Hymenophyllum Callistopteris Hymenophyllum Trichomanes Didymoglossum Hymenophyllum Trichomanes Crepidomanes Hymenophyllum Didymoglossum Polyphlebium Polyphlebium Abrodictyum Callistopteris ? Trichomanes Hymenophyllum Cyathea Hymenophyllum Didymoglossum Hymenophyllum Didymoglossum – Pachychaetum ? Hymenophyllum Trichomanes Trichomanes Pachychaetum – Didymoglossum Vandenboschia Pleuromanes Microgonium Didymoglossum – Didymoglossum Crepidomanes /Gonocormus Crepidomanes /Crepidium Nesopteris Pleuromanes Lacostea Trichomanes Pachychaetum Trichomanes ? Nesopteris – Didymoglossum Trichomanes – Vandenboschia Hymenophyllum Didymoglossum Sphaerocionium – Sphaerocionium Trichomanes Microgonium Hymenophyllum Trichomanes Crepidomanes /Crepidium Sphaerocionium Didymoglossum – – Abrodictyum ? Trichomanes Mecodium – Sphaerocionium 274 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) infundibulare Alderw. ingae C. Chr. intermedium Bosch intramarginale Hook. & Grev. intricatum Farrar japonicum Thunb. ex Murray javanicum Blume jenmanii Lellinger johnstonense F.M. Bailey junceum H. Christ jungermannioides E. Fourn. kalamocarpum Hayata kalbreyeri Baker kapplerianum Sturm kaulfussii Hook. & Grev. killipii Weath. kingii Copel. kirkii Hook. krausii Hook. & Grev. krugii H. Christ kunzeanum Hook. kurzii Bedd. labiatum Jenman laceratum Desv. laciniatum Roxb. laciniosum Alston laetevirens Fée laetum Bosch lambertianum Hook. lanceolatum Poir. lanceum Bory ex Willd. langsdorfii Bosch lasiophyllum Alderw. lastreoides C. Presl latemarginale D.C. Eaton latevirens Fée latifrons Bosch latilabiatum E.D. Br. latilobium Bonap. latipinnulata Bonap. latipinnum Copel. latisectum H. Christ lauterbachii H. Christ laxum Klotzsch lechleri Bosch ledermanni Brause lehmannii Hieron. lenormandi Bosch lentum Poir. lepervanchii Cordem. leprieurii Hook. leptophyllum A. Cunn. Cephalomanes Polyphlebium Crepidomanes Crepidomanes Vandenboschia ? Onychium Cephalomanes Trichomanes Vandenboschia Polyphlebium Polyphlebium Vandenboschia Trichomanes Didymoglossum Trichomanes Trichomanes Cephalomanes Didymoglossum Didymoglossum Abrodictyum Vandenboschia Crepidomanes Didymoglossum Trichomanes Cephalomanes Trichomanes Vandenboschia Abrodictyum Trichomanes Davallia Hymenophyllum Trichomanes Hymenophyllum Trichomanes Crepidomanes Vandenboschia Crepidomanes Crepidomanes Crepidomanes Trichomanes Abrodictyum Abrodictyum ? Crepidomanes Trichomanes Polyphlebium Cephalomanes Didymoglossum Didymoglossum Davallia Crepidomanes ? Trichomanes Abrodictyum – – Nesopteris Crepidomanes /Crepidomanes ? – – Trichomanes Vandenboschia – – Vandenboschia Trichomanes Microgonium Trichomanes Trichomanes – Microgonium Didymoglossum Pachychaetum Vandenboschia Crepidomanes /Crepidomanes Didymoglossum Lacostea – Trichomanes Vandenboschia Pachychaetum Trichomanes – Sphaerocionium Trichomanes Hymenophyllum Davalliopsis Crepidomanes /Crepidomanes Vandenboschia Crepidomanes /Crepidomanes Crepidomanes /Gonocormus Crepidomanes /Crepidomanes ? Pachychaetum ? Crepidomanes /Crepidium Trichomanes – – Didymoglossum Microgonium – Crepidomanes /Gonocormus ? Davalliopsis Abrodictyum A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 275 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) levissisum Fée liberiense Copel. lindenii C. Presl lindigii E. Fourn. lineare Sw. liukiuense Y. Yabe longicollum Bosch longifolium Desv. longifrons Nakai longilabiatum Bonap. longisetum Bory ex Willd. lorencei Tardieu loreum Bory & Bél. lozanii M.T. Murillo lucens Sw. lucidum Roxb. ludovicinum Rosenst. luerssenii F. Muell. luschnatianum C. Presl luzonicum C. Presl macgillivrayi Baker macilentum Bosch macroclados Kunze majorae Watts makinoi C. Chr. malayanum Roxb. malingii Hook.f. maluense Brause mandiocanum Raddi mannianum Mett. ex Kuhn mannii Hook. marchantioides Zippel ex Moritz marginatum Mett. martinezii Rovirosa martiusii C. Presl matthewii H. Christ maximum Blume megistostomum Copel. meifolium Bory ex Willd. melanopus Baker melanorhizon Hook. melanotrichum Schltdl. membranaceum L. merrillii Copel. mettenii C. Chr. mexicanum Bosch micayense Hieron. microchilum Baker microlirion Copel. microphyllum Mett. ex Kuhn micropubescens Proctor mildbraedii Brause Vandenboschia ? Didymoglossum Trichomanes Trichomanes Hymenophyllum Vandenboschia Abrodictyum Trichomanes Crepidomanes Crepidomanes Abrodictyum Didymoglossum Hymenophyllum Trichomanes Trichomanes Davallia Trichomanes Abrodictyum Vandenboschia Crepidomanes Hymenophyllum Trichomanes Trichomanes Crepidomanes Crepidomanes Sphenomeris ? Hymenophyllum Cephalomanes Abrodictyum Crepidomanes Crepidomanes Didymoglossum Abrodictyum Vandenboschia Trichomanes Crepidomanes Vandenboschia Crepidomanes Abrodictyum Didymoglossum Crepidomanes Crepidomanes Didymoglossum Abrodictyum Crepidomanes Vandenboschia Trichomanes Hymenophyllum Crepidomanes Crepidomanes Didymoglossum Hymenophyllum ? Didymoglossum Trichomanes Trichomanes Sphaerocionium Vandenboschia Pachychaetum Trichomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes ? Pachychaetum Didymoglossum Sphaerocionium Trichomanes Trichomanes – Trichomanes Pachychaetum Lacosteopsis Crepidomanes /Crepidium Hymenophyllum Trichomanes Trichomanes Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes – Sphaerocionium – Pachychaetum Crepidomanes /Gonocormus Crepidomanes /Gonocormus Didymoglossum Pachychaetum Vandenboschia Trichomanes Crepidomanes /Gonocormus Vandenboschia Crepidomanes /Crepidomanes Pachychaetum Didymoglossum Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Didymoglossum Pachychaetum Crepidomanes /Crepidomanes Vandenboschia Trichomanes Hymenophyllum Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Didymoglossum Hymenophyllum ? 276 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) millefolium Desv. milnei Bosch mindorense H. Christ minimum Alderw. minutissimum Alderw. minutulum Gaudich. minutum Blume miyakei Y. Yabe molle E. Fourn. montanum Hook. moritzii Bosch mosenii Lindm. motleyi Bosch mougeotii Bosch multiidum G. Forst. muscoides Sw. musolense Brause myrioneuron Lindm. myriophyllum Desv. myrioplasium Kunze naseanum H. Christ naumannii Kuhn & Luerss. ex H. Christ neesii Blume neilgherrense Bedd. nipponicum Nakai nitidulum Bosch niveum Burm.f. novoguineense Brause nudum Poir. nymanii H. Christ obscurum Blume olivaceum Kunze ex Klotzsch opacum Bosch orbiculare H. Christ orientale C. Chr. ornatulum Bosch osmundoides Poir. pabstianum Müll.Berol. pachyphlebium C. Chr. paciicum Hedw. padronii Proctor pallidum Blume palmatiidum Müll.Berol. palmatum C. Presl palmicola Bosch palmifolium Hayata paniculatum Alderw. pannosum Ces. papillatum Müll.Berol. pappei Kunze papuanum Brause paradoxum Domin Trichomanes Abrodictyum Didymoglossum Crepidomanes Didymoglossum Crepidomanes Crepidomanes Vandenboschia Trichomanes Didymoglossum Vandenboschia Didymoglossum Didymoglossum Trichomanes Hymenophyllum Didymoglossum Crepidomanes Didymoglossum ? Callistopteris Vandenboschia Polyphlebium Hymenophyllum Didymoglossum Vandenboschia Hymenophyllum ? Crepidomanes Hymenophyllum Crepidomanes Abrodictyum Polyphlebium Trichomanes Didymoglossum Vandenboschia Didymoglossum ? Trichomanes Didymoglossum Trichomanes ? Hymenophyllum Trichomanes Hymenophyllum Hymenophyllum Crepidomanes Didymoglossum Crepidomanes Crepidomanes Didymoglossum Abrodictyum Abrodictyum Didymoglossum Didymoglossum Davalliopsis Abrodictyum Microgonium Crepidomanes /Crepidomanes Didymoglossum Crepidomanes /Crepidomanes ? Crepidomanes /Gonocormus Vandenboschia Trichomanes Didymoglossum Lacosteopsis Didymoglossum Didymoglossum Feea Hymenophyllum Didymoglossum Crepidomanes /Gonocormus ? Didymoglossum ? – Vandenboschia – Hymenophyllum Microgonium Vandenboschia Sphaerocionium ? Crepidomanes /Gonocormus Sphaerocionium Crepidomanes /Crepidomanes Pachychaetum – Davalliopsis Didymoglossum Vandenboschia ? Feea Didymoglossum Davalliopsis ? Hymenophyllum Trichomanes Pleuromanes Sphaerocionium Crepidomanes /Gonocormus Microgonium Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes Didymoglossum Pachychaetum Pachychaetum Didymoglossum Didymoglossum A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 277 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) parvilorum Poir. parvulum Poir. parvum Copel. pedicellatum Desv. pellucens Kunze pellucidum Goldm. peltatum Poir. pennatum Kaulf. perpusillum Alderw. pervenulosum Alderw. petersii A. Gray philippianum Sturm piliferum Alderw. pilosum Raddi pinnatiidum Bosch pinnatinervium Jenman pinnatum Hedw. platyderon E. Fourn. platyrachis Domin pluma Hook. plumosum Kunze plumula C. Presl poeppigii C. Presl polyanthos Sw. polyodon Colenso polyphlebius V. Marcano polypodioides L. polysperma Poir. polystromaticum Bierh. powellii Baker preslianum Nakai preslii C.V. Morton prieurii Kunze procerum Fée proliferum Blume pseudo-arbuscula Alderw. pseudoblepharistomum Tagawa pseudocapillatum Alderw. ptilodes Bosch pulchellum Salisb. pulcherrimum Copel. pumilum Bosch punctatum Poir. pusillum Sw. pygmaeum C. Chr. pyxidiferum L. quelpaertense Nakai quercifolium Desv. racemulosum Bosch radicans Sw. ramitrichum Faden recedens Rosenst. Abrodictyum Hymenophyllum Vandenboschia Trichomanes Trichomanes Vandenboschia Hymenophyllum Trichomanes Crepidomanes Crepidomanes Didymoglossum Polyphlebium Hymenophyllum Trichomanes Trichomanes Didymoglossum Trichomanes Abrodictyum Trichomanes Abrodictyum Trichomanes Trichomanes Trichomanes Hymenophyllum Abrodictyum Trichomanes Trichomanes Sphenomeris Abrodictyum Crepidomanes Crepidomanes Cephalomanes Trichomanes Hymenophyllum Crepidomanes Abrodictyum Crepidomanes Crepidomanes Trichomanes Hymenophyllum Crepidomanes Abrodictyum Didymoglossum Didymoglossum Didymoglossum Polyphlebium Vandenboschia Trichomanes Abrodictyum Vandenboschia Crepidomanes Crepidomanes Pachychaetum Hymenophyllum Vandenboschia Lacostea Trichomanes Lacosteopsis Hymenophyllum Trichomanes Crepidomanes /Crepidium ? Crepidomanes /Crepidomanes Didymoglossum – Sphaerocionium Trichomanes Trichomanes Didymoglossum Trichomanes Pachychaetum Feea Abrodictyum Trichomanes Trichomanes Trichomanes Mecodium Pachychaetum Trichomanes Trichomanes – Pachychaetum Crepidomanes /Crepidomanes Nesopteris – Davalliopsis Sphaerocionium Crepidomanes /Gonocormus Pachychaetum Nesopteris Crepidomanes /Crepidium ? Trichomanes Hymenophyllum Nesopteris Pachychaetum Didymoglossum Didymoglossum Microgonium – Vandenboschia Trichomanes Pachychaetum Vandenboschia Crepidomanes /Crepidomanes Crepidomanes /Crepidomanes 278 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) reniforme G. Forst. repens Schott ex Sturm reptans Sw. resinosum R.C. Moran rhipidophyllum Sloss. rhizophyllum Cav. ridleyi Copel. rigidum Sw. robinsonii Hook. ex Baker robustum E. Fourn. roemerianum Rosenst. roraimense Jenman rosenstockii Alderw. rothertii Alderw. rotundifolium Bonap. rupicolum Racib. ruwenzoriense Taton samoense C. Chr. sandvicense Bosch sanguinolentum G. Forst. savaiense Lauterb. saxatile Brack. ex T. Moore saxifragoides C. Presl sayeri F. Muell. & Baker scandens L. schaffneri Schltdl. schiedeanum Müll.Berol. schlechteri Brause schmidianum Zenker ex Taschner schomburgkianum Sturm schomburgkii Bosch schultzei Brause seemannii Carruth. ex Seem. sellowianum C. Presl sericeum Sw. serratifolium Rosenst. serratulum Baker serricula Fée setaceum Bosch setiferum Baker ex Jenman setigerum Backh. ex T. Moore siamense H. Christ sibthorpioides Bory ex Willd. sinuatum Bonap. sinuosum Rich. ex Willd. smithii Hook. societense J.W. Moore solidum G. Forst. solitarium Jenman somai Nakai sonderi Bosch speciosum Willd. Hymenophyllum Vandenboschia Didymoglossum Trichomanes ? Didymoglossum Trichomanes Hymenophyllum Abrodictyum Didymoglossum Trichomanes Crepidomanes Trichomanes Cephalomanes Crepidomanes Didymoglossum Crepidomanes Crepidomanes Crepidomanes Vandenboschia Hymenophyllum Hymenophyllum Abrodictyum Crepidomanes Didymoglossum Trichomanes Didymoglossum Polyphlebium Abrodictyum Crepidomanes Trichomanes Trichomanes Abrodictyum Abrodictyum Trichomanes Hymenophyllum Polyphlebium Hymenophyllum Trichomanes Abrodictyum Didymoglossum Abrodictyum Abrodictyum Hymenophyllum Didymoglossum Trichomanes Abrodictyum Callistopteris Davallia Didymoglossum Vandenboschia Trichomanes Vandenboschia Cardiomanes Vandenboschia Didymoglossum Davalliopsis ? Didymoglossum Trichomanes Sphaerocionium Pachychaetum Didymoglossum Trichomanes Nesopteris Trichomanes – Crepidomanes /Crepidomanes Microgonium Crepidomanes /Crepidomanes Crepidomanes /Gonocormus Crepidomanes /Crepidomanes Vandenboschia Myrmecostylum Pleuromanes Pachychaetum Crepidomanes /Gonocormus Didymoglossum Trichomanes Didymoglossum – Abrodictyum Crepidomanes /Crepidomanes Trichomanes Trichomanes Abrodictyum Pachychaetum Trichomanes Sphaerocionium – Hymenophyllum Trichomanes Pachychaetum Didymoglossum Pachychaetum Pachychaetum Hymenophyllum Microgonium Trichomanes Abrodictyum – – Didymoglossum Vandenboschia Lacostea Vandenboschia A. Ebihara et al.: A taxonomic revision of Hymenophyllaceae 279 Appendix. Continued. Basionym Genus Subgenus / Section Trichomanes (cont.) spectabile Klotzsch sphenoides Kunze spicatum Hedw. ex Hook. spicisorum Desv. spinulosum Phil. splendidum Bosch spruceanum Hook. sprucei Baker squarrosum G. Forst. stenosiphon H. Christ steyermarkii P.G. Windisch & A.R. Sm. striatum D. Don strictum Menzies ex Hook. & Grev. strigosum Thunb. ex Murray stylosum Poir. subdeltoideum C. Chr. subexsertum Bosch sublabiatum Bosch sublimbatum Müll.Berol. subpinnatiidum Bosch subsessile Splitg. subtilissimum Brause subtriidum C.G. Matthew & H. Christ suffrutex Alderw. sumatranum Alderw. superbum Backh. ex T. Moore taeniatum Copel. tahitense Nadeaud tamarisciforme Jacq. tanaicum Hook. tenellum Hedw. tenerum A. Spreng. tenue Brack. tenuifolium Burm.f. tenuissimum Bosch tereticaulum Ching teysmanni Bosch thouarsianum C. Presl thuioides Desv. thysanostomum Makino tomaniiviense Brownlie torotumani Vareschi tosae H. Christ ex Matsum. tranninense Fée translucens Kunze trichodes Sw. trichoideum Sw. trichophorum Alderw. trichophyllum T. Moore trigonum Desv. trinerve Baker trollii Bergdolt Trichomanes Didymoglossum Trichomanes Trichomanes Hymenophyllum Trichomanes Trichomanes Abrodictyum Dicksonia Vandenboschia Trichomanes Vandenboschia ? Abrodictyum Microlepia Abrodictyum Abrodictyum Polyphlebium Trichomanes Didymoglossum Crepidomanes Trichomanes Crepidomanes Crepidomanes Cephalomanes Cephalomanes Callistopteris Hymenophyllum Didymoglossum Abrodictyum Trichomanes Polyphlebium Polyphlebium Polyphlebium Cheilanthes Polyphlebium Abrodictyum Crepidomanes Hymenophyllum Abrodictyum Crepidomanes Hymenophyllum Trichomanes Crepidomanes Polyphlebium Polyphlebium Polyphlebium Polyphlebium Hymenophyllum Abrodictyum Trichomanes Crepidomanes Trichomanes Trichomanes Didymoglossum Feea Feea Hymenophyllum Trichomanes Trichomanes Pachychaetum – Vandenboschia Trichomanes ? Abrodictyum – Pachychaetum Pachychaetum – Trichomanes Didymoglossum Crepidomanes /Gonocormus Lacostea Crepidomanes /Gonocormus Crepidomanes /Gonocormus – – – Sphaerocionium Didymoglossum Pachychaetum Lacostea – – – – – Pachychaetum Crepidomanes /Gonocormus Hymenophyllum Pachychaetum Nesopteris Sphaerocionium Lacostea Crepidomanes /Crepidomanes – – – – Hymenophyllum Pachychaetum Trichomanes Crepidomanes /Gonocormus Feea 280 BLUMEA — Vol. 51, No. 2, 2006 Appendix. Continued. Basionym Trichomanes (cont.) tuerckheimii H. Christ tunbrigense L. ujhelyii Kümmerle ulei H. Christ undulatum Sw. unilorum Cav. vandenboschii P.G. Windisch varians Alderw. vaupelii Brause vaupesensis Lellinger venosum R.Br. venustulum Colenso venustum Desv. vestitum Baker vieillardii Bosch virgatulum Bosch viridans Mett. ex Kuhn viride Bubani vitiense Baker vittaria DC. ex Poir. volubile Vell. walleri Watts wallii Thwaites ex Trimen warburgii H. Christ weddellii Bosch werneri Rosenst. wildii F.M. Bailey windischianum Lellinger yandinense F.M. Bailey zollingeri Bosch Vandenboschia assimilis Ching & P.C. Chiu inopinata Pic.Serm. lofoushanensis Ching subclathrata K. Iwats. titibuensis H. Itô tubilora F.S. Wagner Genus Subgenus / Section Trichomanes Hymenophyllum Trichomanes Polyphlebium ? Hymenophyllum Crepidomanes ? Trichomanes Didymoglossum Crepidomanes ? Trichomanes Polyphlebium Polyphlebium Vandenboschia Hymenophyllum Polyphlebium Vandenboschia Crepidomanes Asplenium Crepidomanes Trichomanes Trichomanes Crepidomanes Didymoglossum Abrodictyum Trichomanes Polyphlebium Crepidomanes Abrodictyum Didymoglossum Cephalomanes Lacostea Hymenophyllum Trichomanes – Mecodium Nesopteris ? Trichomanes Microgonium Crepidomanes /Crepidomanes ? Trichomanes – – Lacosteopsis Hymenophyllum – Vandenboschia ? Crepidomanes /Crepidomanes – Crepidomanes /Crepidomanes Trichomanes Lacostea Crepidomanes /Crepidomanes Didymoglossum Abrodictyum Davalliopsis – Crepidomanes /Crepidium Pachychaetum Microgonium – Vandenboschia Crepidomanes Vandenboschia Vandenboschia Crepidomanes Vandenboschia Vandenboschia Crepidomanes /Crepidomanes Vandenboschia ? Vandenboschia Crepidomanes /Crepidomanes Vandenboschia