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
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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.
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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.
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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
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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.).
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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.
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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).
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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.
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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.
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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.
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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.
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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.
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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.
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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.).
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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
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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.
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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.
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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).
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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
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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).
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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.;
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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.
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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.).
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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