Bull. Natl. Mus. Nat. Sci., Ser. B, 33(2), pp. 55–68, June 22, 2007

The Hymenophyllaceae of the Pacific Area. 1.
Hymenophyllum subgenus Hymenophyllum
Atsushi Ebihara1 and Kunio Iwatsuki2
1

Department of Botany, National Museum of Nature and Science, Amakubo 4–1–1, Tsukuba 305–0005, Japan
E-mail: ebihara@kahaku.go.jp
2
The Museum of Nature and Human Activities, Hyogo, Yayoigaoka 6-chome, Sanda 669–1546, Japan
E-mail: iwatsuki@hitohaku.jp

Abstract Pacific species of Hymenophyllum subgen. Hymenophyllum redefined by Ebihara et al.
(2006) are enumerated. In total, 26 species are recorded in the studied area, and synonymies, information of the type material, distribution and cytological records of each species are provided.
Key words : Australasia, filmy fern, Hymenophyllaceae, Hymenophyllum, Oceania, Pteridophyta.

Hymenophyllaceae or filmy ferns are one of
the basal lineages of leptosporangiate ferns(Hasebe et al., 1995; Pryer et al., 2004) comprising about 600 species (Iwatsuki, 1990), mostly
distributed in the tropics and temperate regions.
In the Pacific area, this family has a great diversity, including many isolated lineages or monotypic ‘genera’, and occupies a high ratio in species
number of the pteridophyte flora (e. g., 15% in
New Zealand, Brownsey and Smith-Dodsworth,1989). Local floristic works have already been
published in most of the countries or regions of
this area. However, none of them overviews thefamily’s distribution throughout the Pacific. This
study covers the Mariana Islands, Caroline Islands, Bougainville Island (offshore Papua New
Guinea), Solomon Islands, Vanuatu (former New
Hebrides), New Caledonia, Fiji, Samoa, Tonga,
Cook Islands, Austral Islands, Society Islands,
Marquesas Islands, Australia, New Zealand andHawaiian Islands (Fig. 1). The authors are also
preparing a treatment of the family for Flora
Malesiana. Upon completion of the two publications, most of the Old World species can easily
be compared and identified. Since filmy ferns
usually prefer moist and shady habitats, desert
areas (e. g., Australian continent except for theGreat Dividing Range) and coral islands (e. g.,
Marshall Islands, Naulu, Tuvalu and Kiribati)

completely lack their distribution. After a longcontroversy on the classification of Hymenophyllaceae, Ebihara et al. (2006) advocated a new
system principally maintaining natural groups
recognized by molecular phylogeny, which isadopted here. See Ebihara et al. (2006) for synonymy of each higher taxon.
Genus 1. Hymenophyllum Sm., Mém. Acad.
Sci. Turin 5: 418 (1793).
The largest genus of the Hymenophyllaceae,
comprising about half (ca. 250 spp.) of thespecies of the family. Nine of the ten subgenera
are present in the Pacific area, usually epiphyticon tree trunks in foggy mountain forest in thetropics, or growing in lowland mossy forest at
southern high latitudes.
Subgenus 1. Hymenophyllum
The largest subgenus of the genus. Frond margins are serrate in most of the species.
1. Hymenophyllum peltatum (Poir.) Desv.,
Mém. Soc. Linn. Paris 6: 333 (1827).
[Fig. 2]— Trichomanes peltatum Poir. in Lamarck andPoir., Encycl. 8: 76 (1808).
Type: Bory de St.-Vincent s. n. (“Ile-deFrance”Mauritius) [P*] (*n. v., examined byTindale, 1963).

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Atsushi Ebihara and Kunio Iwatsuki

Fig. 1. The areas covered in this study (surrounded by the bold line) and the Malesia region (surrounded by thebroken line).

Hymenophyllum falklandicum auct. non Baker:
Puy et Orchard, Fl. Australia 50: 539 (1993).
Hymenophyllum unilaterale auct. non Bory ex
Willd.: Dobbie, N. Z. Ferns ed. 2, 58 (1921).
Distribution: Australia (QLD, NSW, VIC and
TAS), New Zealand, Macquarie Isl.; Malesia(Borneo and Papua New Guinea), Mascarene Isl.,
South Africa, Southern Chile.
Chromosome number: n11 (Brownlie, 1958,
New Zealand; Laws in Tindale and Roy, 2002,
Australia); 2n22 (Tindale, 1963, Australia).
Note: This is the only filmy fern species sharedby New Zealand and the Malesia region, and ourmolecular data show that materials from Tasmania and Malaysia are actually close (Ebihara, unpublished). In Malesia, this species is apparently
rare and known only from Mt. Kinabalu,
Malaysia (Parris, 1992) and Mt. Victoria, Papua
New Guinea (Iwatsuki, unpublished), both locat-

ed at high altitudes. It should be noted that its
chromosome number n11 is the smallest of all
pteridophytes.
2. Hymenophyllum multifidum (G. Forst.)
Sw., J. Bot. (Schrader) 1800(2): 102, 1801.
[Fig. 3]— Trichomanes multifidum G. Forst., Fl. Ins.
Austr. 85 (1786) — Davallia multifidum (G.
Forst.) A. Spreng., J. Bot. (Schrader) 1799(2):271 (1800) — Meringium multifidum (G.
Forst.) — Copel., Philipp. J. Sci. 67: 44 (1938).
Type: Forster Herb. 304 (Dusky Bay?, New
Zealand) [BM*] (see Nicolson and Fosberg,
2004).
Hymenophyllum truncatum Colenso, Trans.
and Proc. New Zealand Inst. 23: 390 (1891) —Hymenophyllum multifidum (G. Forst.) Sw. var.
truncatum (Colenso) Domin, Biblioth. Bot.

Pacific Hymenophyllaceae I
Table 1.

57

Distribution of the species of Hymenophyllum subgen. Hymenophyllum in the Pacific area.
Caroline Solomon
New
Vanuatu
Isls.
Isls.
Caledonia

Hymenophyllum
subgen. Hymenophyllum
1 H. peltatum
2 H. multifidum
3 H. minimum
4 H. marginatum
5 H. armstrongii
6 H. pumilum
7 H. gracilescens
8 H. bivalve
9 H. revolutum
10 H. cupressiforme
11 H. baileyanum
12 H. deplanchei
13 H. feejeense
14 H. serrulatum
15 H. edentulum
16 H. holochilum
17 H. praetervisum
18 H. dimidiatum
19 H. rolandi-principis
20 H. macgillivrayi
21 H. gorgoneum
22 H. denticulatum
23 H. kerianum
24 H. brassii
25 H. walleri
26 H. francii



Fiji

Samoa

Marquesas
New
Australia
Isls.
Zealand































































: present; : endemic.

20(85): 25 (1913).
Type: Colenso s. n., 1887–1890 (south of Dannevirke, Waipawa, New Zealand) [K].
Hymenophyllum alpinum Colenso, Trans.
Proc. New Zealand Inst. 31: 263 (1899) — Hymenophyllum multifidum (G. Forst.) Sw. var
alpinum (Colenso) Domin, Biblioth. Bot. 20(85):
25 (1913).
Syntypes: Hill s. n., 1845–1852; Colenso s. n.,
1898 (Ruahine Moutain-range, New Zealand) (n.
v.).
Hymenophyllum oligocarpum Colenso, Trans.
Proc. New Zealand Inst. 31: 264 (1899) — Hymenophyllum multifidum (G. Forst.) Sw. var
oligocarpum (Colenso) Domin, Biblioth. Bot.
20(85): 25 (1913).
Type: Hill s. n., 1898 (Waikaremoana, Hawke’s
Bay, New Zealand) (n. v.).
Hymenophyllum howense Brownlie, PacificSci. 14: 244 (1960).
Type: Moore 4, Nov. 1877 (Mt. Gower, Lord
Howe Isl., Australia) [K].
Distribution: Vanuatu, Fiji, Lord Howe Isl.,
New Zealand.
Chromosome number: n26 (Brownlie, 1954,
New Zealand; Braithwaite, 1975, Vanuatu andFiji).
Note: Recent studies have shown that the distribution of this species is wider northward thanrecognized before. “Mountain form” which has
curled fronds was reported from New Zealand(Holloway, 1923). The material we examined as“Mountain form” showed no obvious difference
from a usual one in rbcL sequence (Ebihara, unpublished).
3. Hymenophyllum minimum A. Rich., Fl. N.
Zel. 91, t. 14, fig. 2 (1832).
[Fig. 4]

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Atsushi Ebihara and Kunio Iwatsuki

Figs. 2–6. 2a–b. Hymenophyllum peltatum (Ebihara 000223-011 [TNS]). 3a–b. H. multifidium (a. Ebihara
011217-06 [TNS]; b. Ebihara 011217-10 [TNS]). 4. H. minimum (Ebihara 011225-03 [TNS]). 5a–b. H. marginatum (a. Fraser s. n. [K, holotype]; b. Ebihara 010915-03 [TI]). 6a–b. H. armstrongii (Ebihara 011219-07[TI]). Scale1 cm for whole leaves, and 0.4 mm for sori.

— Meringium minimum (A. Rich.) Copel.,Philipp. J. Sci. 73: 457 (1941).
Type: ? (“Nova-Zeelandia”New Zealand) (n.
v.).
Hymenophyllum pygmaeum Colenso, Trans.
Proc. New Zealand Inst. 13: 376 (1881).
Syntypes: Enys s. n. (Bealey, New Zealand);Collector unknown (Preservation Inlet; Resolution Island; Lyttelton Harbour; south of Hokitika,
New Zealand) (n. v.).
Hymenophyllum pumilio Rosesnt., Repert.
Spec. Nov. Regni Veg. 9: 72 (1910).
Type: Franc 1433, 1910 (Mt. Tao, New Caledonia) [isotypes in BM MICH* NY* P UC*
US*].
Distribution: New Caledonia (one collection),
New Zealand.
Chromosome number: n26 (Lovis in Dawson et al., 2000, New Zealand).
Note: This species has very minute fronds
with spinulose teeth of involucre. In New
Zealand, it appears to be a dwarf form of H. revolutum, but it is clearly distinguishable by its
glabrous rachis and stipe. This species usually
grows on the coast where sometimes spindrift

comes, which is an exceptional habitat for Hymenophyllaceae.
The identity of H. pygmaeum was carefully
studied by Lovis (1982) who concluded that the
species is a synonym of H. minimum.
Hymenophyllum pumilio, once collected innorthern New Caledonia, has little obvious difference from H. minimum.
4. Hymenophyllum marginatum Hook. et
Grev., Icon. Filic. 2: t. 34 (1831).
[Fig. 5]
— Pachyloma marginata (Hook. et Grev.)
Bosch, Verslagen Meded. Afd. Natuurk. Kon.
Akad. Wetensch. 11: 318 (1861) — Craspedophyllum marginatum (Hook. et Grev.) Copel.,Philipp. J. Sci. 67: 27 (1938).
Type: Fraser s. n. (Port Jackson, “Nova Hollandia”Australia) [K; E* L NSW].
Distribution: Australia (QLD, NSW and TAS).
Chromosome number: 2n24 (Tindale andRoy 2002, Australia).
Note: This and the following species havefronds bordered by black cells, and were formerly separated as a genus Craspedophyllum.

Pacific Hymenophyllaceae I

5. Hymenophyllum armstrongii (Baker) Kirk,
Trans. Proc. New Zealand Inst. 10: t. 21, fig. A(1878).
[Fig. 6]
— Trichomanes armstrongii Baker in Hook. et
Baker, Syn. Fil. 452 (1868) — Hymenophyllum
cheesemanii Baker var. armstrongii (Baker)
Cheesem., Man. N. Z. Fl. 938 (1906) — Microtrichomanes armstrongii (Baker) Copel.,Philipp. J. Sci. 73: 457 (1941) — Craspedophyllum armstrongii (Baker) Rae ex Copel., Gen. Fil.
33 (1947).
Type: Armstrong s. n., 1868 (New Zealand)[K].
Hymenophyllum cheesemannii Baker, Hooker’s Icon. Pl. 12: 30, t. 1132 (1873) — Craspedophyllum cheesemannii (Baker) N. A. Wakef.,
Victorian Naturalist 66: 59 (1949).
Type: Cheesemann s. n., Oct. 1871 (Titiranga
range, New Zealand) [K; GH* US].
Hymenophyllum
melanocheilos
Colenso,Trans. Proc. New Zealand Inst. 17: 255 (1885).
Type: Rowson s. n., 1884 (Whangaroa, Mongonui, New Zealand) [K; W*].
Distribution: New Zealand (endemic).
Chromosome number: n13 (Lovis in Bronwnsey, 1991, New Zealand).
Note: In contrast to Australian H. marginatum
whose frond margin are entire, New Zealand H.
armstrongii shows spinulose-serrate margins.
6. Hymenophyllum pumilum C. Moore in
Hook. et Baker, Syn. Fil. (ed. 2) 464 (1874).
[Fig. 7]
Type: Moore s. n. (Mill Creek, Mt. Tomah,
NSW, Australia) [K].
Hymenophyllum moorei Baker in Hook. et
Baker, Syn. Fil. (ed. 2) 464 (1874).
Type: Moore 5, Dec. 1871 (Lord Howe Isl.)[K; BM US].
Hymenophyllum minimum auct. non A.Rich.:
Benth., Fl. Austral. 706 (1878).
Distribution: Australia (NSW and Lord Howe
Isl.)
Chromosome number: n34 (Tindale andRoy, 2002, Australia).
Note: The Lord Howe Island form H. moorei,

59

having larger sori, needs further comparison with
continental plants.
7. Hymenophyllum gracilescens Domin, Biblioth. Bot. 20(85): 23, t. 1, figs. 2–3 (1913).
[Fig. 8]
Type: Domin Iter Austral. 37, Dec. 1909 (Mt.
Bellenden Ker, QLD, Australia) [PR*] (examinedby Croxall, 1975).
Distribution: Australia (QLD) (endemic).
Chromosome number: unknown.
Note: This species, known from limited collections at Mt. Bellenden Ker range, resembles H.
peltatum, but is distinguished by the crenulate
lips of the involucre.
8. Hymenophyllum bivalve (G. Forst.) Sw., J.
Bot. (Schrader) 1800(2): 99 (1801).
[Fig. 9]— Trichomanes bivalve G. Forst., Fl. Ins. Austr.
84 (1786)—Sphaerocionium bivalve (G. Forst)
C. Presl, Hymen. 33 (1843)—Meringium bivalve(G. Forst.) Copel., Philipp. J. Sci. 67: 44 (1938).
Lectotype: Forster Herb. 301 (New Zealand ?)[BM]—designated by Tindale (1963).
Hymenophyllum spathulatum Colenso, Tasman. J. Nat. Sci. 2: 184 (1846).
Type: Colenso s. n., Dec. 1841 (Shores of
Waikare Lake, New Zealand) (n. v.).
Hymenophyllum pyriforme Bosch, Ned.
Kruidk. Arch. 5(3): 173 (1863).
Type: D’Urville s. n. (New Zealand) [B*] (seeTindale, 1963).
Distribution: Australia (QLD and NSW), New
Zealand.
Chromosome number: n22 (Brownlie 1958,
New Zealand; Vessey and Barlow 1963, Australia); n211b (Tindale and Roy 2002, Australia).
9. Hymenophyllum revolutum Colenso, Tasman. J. Nat. Sci. 2: 186 (1846).
[Fig. 10]
Type: Colenso 276, Dec. 1841 (shores of
Waikare Lake, New Zealand) [K].
Hymenophyllum zeelandicum Bosch, Ned.
Kruidk. Arch. 5(3): 175 (1863).
Syntypes: J. D. Hooker s. n. (Bay of Islands,

60

Atsushi Ebihara and Kunio Iwatsuki

Figs. 7–14. 7. Hymenophyllum pumilum (Moore s. n. [K, holotype]). 8. H. gracilescens (Dockrill 1083 [K]).
9a–b. H. bivalve (Ebihara 011216-05 [TNS]. 10a–b. H. revolutum (a. Colenso 276 [K, holotype]; b. Ebihara
011222-11 [TNS]). 11a–b. H. cupressiforme (Ebihara 010905-01 [TNS]). 12. H. baileyanum (Bailey s. n. [K,isotype]). 13a–b. H. deplanchei (a. Deplanche 174 [K, isotype]; b. Ebihara 001228-05 [TI]). 14a–b. H. feejeense (a. Wilkes Expedition 10 [K, isotype]; b. Kokubo Fiji-5 [TNS]). Scale1 cm for whole leaves, and 0.4
mm for sori.

Pacific Hymenophyllaceae I

New Zealand) [K]; Wilkes s. n. (New Zealand)(n.v.).
Hymenophyllum pusillum Colenso, Trans.
Proc. New Zealand Inst. 12: 365 (1880).
Syntypes: Colenso ? (Te Ranga mountain,
head-waters of Waikare, Bay of Islands, 1836;
head of the Wairarapa Valley, 1852; west slopesof Ruahine mountain range, near the head-watersof the River Manawatu, 1878–1880, New
Zealand) (n. v.).
Hymenophyllum tunbridgense auct. non (L.)
Sm.: Cheesem., Man. N. Z. Fl. 13 (1925).
Hymenophyllum unilaterale auct. non Bory:
Dobbie, N. Z. Ferns 2nd. ed., 1921.
Distribution: New Zealand (endemic).
Chromosome number: n22 (Brownlie 1958,
New Zealand).
Note: The difference between this and the subsequent species is delicate. See the note for H.
cupressiforme.
10. Hymenophyllum cupressiforme Labill.,
Nov. Holl. Pl. 2: 102, t. 250, fig. 2 (1806).
[Fig. 11]Hymenophyllum tunbridgense (L.) Sm. var. cupressiforme (Labill.) Hook.f., Fl. N. Zel. 2: 11,
1852.
Type: Labillardiere s. n. (“Nova Hollandia et
Terra Diemen”Tasmania, Australia) [FI*] (examined by Tindale, 1963).
Hymenophyllum antarcticum C. Presl, Hymen.
31, 50, t. 12A (1843).
Syntypes: Sieber syn. fil. no. 134 (Port Jackson, “Nova Hollandia”Australia) [isotypes in
E* L* W*]; Sieber flora mixia no. 254 [PR*](see Croxall, 1975).
Hymenophyllum tunbridgense auct. non (L.)
Sm.: R.Br., Prodr. Fl. Nov. Holl. 159 (1810);
Benth., Fl. Austral. 7: 706 (1878); Domin, Biblioth. Bot. 20(85): 22 (1913).
Distribution: Australia (QLD, NSW, VIC and
TAS), New Zealand.
Chromosome number: n21 (Tindale and Roy2002); 2n42 (Tindale and Roy, 2002); n22
(Vessey and Barlow, 1963; Laws in Tindale andRoy, 2002); nca. 24 (Tindale, 1963). All mate-61

rials are from Australia.
Note: This is one of the commonest Hymenophyllum in southern Australia, and apparently
rare in New Zealand (Parris and Croxall, 1972).
The difference between H. revolutum and this is
not so clear as noted by Parris and Croxall(1972); the distinguishing character of H. cupressiforme is entirely winged rachis, but occasionally H. revolutum has wings toward the base of therachis (personal observation). Whether the New
Zealand distribution of this species is true or notstill needs further investigation.
The species name Hymenophyllum antarcticum was given to the form that has near-entire
lips of involucres. The denticulation of the lips of
H. cupressiforme tends to be less distinct thanthat of H. revolutum.
11. Hymenophyllum baileyanum Domin,
Biblioth. Bot. 20(85): 21, t. 2, figs. 2–3 (1913).
[Fig. 12]Hymenophyllum trichomanoides F. M. Bailey,Rep. Govt. Sci. Exp. Bellenden-Ker 74 (1889),
non Bosch — Hemicyatheon baileyanum (Domin)
Copel., Philipp. J. Sci. 67: 28 (1938).
Type: Bailey s. n., 1889 (summit of Mt. Bellenden Ker, QLD, Australia) [BRI*; BM MEL*P] (see Croxall, 1975).
Distribution: Australia (QLD) (endemic).
Chromosome number: unknown.
Note: Hymenophyllum baileyanum and the following species were separated as the genus
Hemicyatheon by Copeland (1938), typified bythis species. Copeland (1937) thought Hemicyatheon occupied an intermediate position between his Mecodium and Meringium, the twolargest groups in Hymenophyllum s. l. Recent
studies (Hennequin et al., 2003; 2006) suggested
both Mecodium and Meringium to be polyphyletic, and the two “Hemicyatheon” species to form aprobable monophyletic group (Hennequin et al.,2006).
12. Hymenophyllum deplanchei Mett. ex
Kuhn, Linnaea 35: 393 (1868).
[Fig. 13]
Hemicyatheon deplanchei (Mett. ex Kuhn)

62

Atsushi Ebihara and Kunio Iwatsuki

Copel., Philipp. J. Sci. 67: 28 (1938).
Type: Deplanche 174 (Mt. Mou, New Caledonia) [B*; K P] (see Brownlie, 1969).
Distribution: New Caledonia (endemic).
Chromosome number: nca. 22 (Braithwaite,
unpublished, New Caledonia).
Note: Though the chromosome number of thisspecies has not been published, Braithwaite labeled “nca. 22” in his specimen deposited in K.
13. Hymenophyllum feejeense Brack., U.S.
Expl. Exped., Filic. 16: 266, t. 37, fig. 2 (1854).
[Fig. 14]
— Meringium feejeense (Brack.) Copel.,Philipp. J. Sci. 67: 44 (1938).
Type: Wilkes Expedition 10, 1838–1842(Ovolau, “Feejee Islands”Fiji) [US; GH* K].
Distribution: Fiji and Samoa.
Chromosome number: n22 (Braithwaite,
1975, Fiji).
Note: This species resembles H. bivalve except
for its dentate lips.
14. Hymenophyllum serrulatum (C. Presl) C.
Chr., Index Filic. 367 (1905).
[Fig. 15]
— Didymoglossum serrulatum C. Presl, Hymen.
23, 48 (1843) — Leptocionium serrulatum (C.
Presl) Bosch, Ned. Kruidk. Arch. 4: 383 (1859).
Type: Cuming 221 (Luzon, Philippines) [PRC;
B BM GH K L P].
Leptocionium violaceum Meyen ex Bosch,
Ned. Kruidk. Arch. 5(3): 147 (1863) —Meringium meyenianum C. Presl, Hymen. 24, t.
8 (1843) — Trichomanes meyenianum (C. Presl)
Bosch, Ned. Kruidk. Arch. 4: 379 (1859) — Hymenophyllum meyenianum (C. Presl) Copel.,Philipp. J. Sci. 64: 25, t. 8 (1937).
Type: Cuming 264 (Philippines) [PRC; B BM
GH K L P].
Hymenophyllum bougainvillense Jeff W.
Grimes, Kew Bull. 34: 233, t. 12 (1978).
Type: Schodde and Craven 3846A, 15 Aug
1964 (alt. 700 m, Maide River Gorge, lower
slopes of L. Loloru crater, c. 24 km north ofBuin, Bougainville Isl.) [CANB*].
Distribution: Solomon Isl., Vanuatu, Fiji, Mar-quesas Isls.; Malay Peninsular, Malesia.
Chromosome number: n21 (Braithwaite,
1969, Solomon Isl.; Braithwaite, 1975, Vanuatu).
Note: This is a typical Malesian element. In
New Guinea this and its allies (e. g., plants
named “H. rubellum”, “H. firmum” and “H. ovatum”) still remain an issue of great complexity. In
Solomon Islands, under a heavy influence from
New Guinea, there are various forms provisionally assigned to this species. In Vanuatu and Fiji,
the southernmost part of the range, the form isuniform and resembles typical H. serrulatum.
15. Hymenophyllum edentulum (Bosch) C.
Chr., Index Filic. 360 (1905).
— Leptocionium edentulum Bosch, Ned.
Kruidk. Arch. 5(3): 148 (1863) — Meringium
edentulum (Bosch) Copel., Philipp. J. Sci. 67: 41(1938).
Syntypes: Griffith s. n. (Assam) [K; BO L];
Lobb s. n. (Sarawak) [K; L].
Trichomanes denticulatum Baker, Syn. Fil. 82(1867), non Houtt., nec Poir — Hymenophyllum
bakeri Copel., Sarawak Mus. J. 2: 309 (1917) —Meringium bakeri (Copel.) Copel., Philipp. J.
Sci. 67: 42 (1938).
Type: Motley s. n. (Borneo) [K].
Distribution: Micronesia; Malesia, South Asia.
Chromosome number: unknown.
Note: The difference between this and H. serrulatum is delicate—H. edentulum has less hairy
axis, more slender rachis and less thickened internal cell walls than H. serrulatum.
16. Hymenophyllum holochilum (Bosch) C.
Chr., Index Filic. 362 (1905).
[Fig. 16]
— Didymoglossum holochilum Bosch, Pl.
Jungh.
1:
561(1853) — Leptocionium
holochilum (Bosch) Bosch, Ned. Kruidk. Arch.
4: 383 (1859) — Meringium holochilum (Bosch)
Copel., Philipp. J. Sci. 67: 41 (1938).
Type: Hasskarl s. n. (Java) [L].
Hymenophyllum affine Brack., U.S. Expl.
Exped., Filic. 16: 265 t. 37 fig. 1 (1854).
Type: Wilkes (Brackenridge) 7 (Ovolau, “Feejee Islands”Fiji) [US*].

Pacific Hymenophyllaceae I

Hymenophyllum subdimidiatum Rosesnt.,
Meded. Rijks-Herb. 11: 1 (1912).
Type: Schlechter 14799, 24 Sept. 1902 (mountains near Yaouhe, New Caledonia) [isotypes in
BM K L MICH* NSW P].
Hymenophyllum tunbridgense (L.) Sm. var.
exsertum F. M. Bailey, Rep. Govt. Sci. Exp. Bellenden-Ker
74(1889) — Hymenophyllum
praetervisum Christ var. australiense Domin.,
Biblioth. Bot. 20(85): 21 (1913).
Type: Bailey s. n. (Mt. Bellenden Ker, QLD,
Australia) [BRI*; BM K NSW*] (see Croxall,
1975).
Hymenophyllum babindae Watts, Proc. Linn.
Soc. New South Wales 39: 766, t. 87, fig. 5(1915).
Type: Watts s.n., July 1913 (Frenchman’s
Creek, near Babinda, QLD, Australia) [NSW;BRI* MEL?*].
Hymenophyllum pseudotunbridgense Watts,
Proc. Linn. Soc. New South Wales 39: 766(1915).
Type: Waller [P1813], 1908 (Evelyn Scrub,
QLD, Australia) [NSW; BRI?* MEL?*].
Hymenophyllum viride Rosesnt. ex Copel.,Philipp. J. Sci. 64: 59, t. 24 (1937).
Type: same as the type of H. subdimidiatum.
Hymenophyllum macgillivrayi auct. non(Baker) Copel.: Copel., Philipp. J. Sci. 64: 60, t.
25 (1937).
Distribution: Micronesia, Solomon Isl., Vanuatu, New Caledonia, FIji, Australia (QLD); Taiwan, Thailand, Malesia.
Chromosome number: n21 (Braithwaite
1975, Vanuatu).
Note: As already pointed out by Ebihara et al.
(2003), H. subdimidiatum and H. pseudotunbridgense, which have been applied to collections
from Oceania, fall in the range of H. holochilum.
17. Hymenophyllum praetervisum Christ,
Bot. Jahrb. Syst. 23: 338 (1896).
[Fig. 17]
Meringium praetervisum (Christ) Copel., Philipp.
J. Sci. 67: 44 (1938).
Syntypes: Reinecke s. n. (Savaii, Samoa); Reinecke 63 (Upolu, Samoa) [isosyntype in K]; Rei-

63

necke 88 (Upolu, Samoa) [isosyntype in BM K
P]; Reinecke 88a (Upolu, Samoa) (n. v.); Reinecke 88b (Tutuila, Samoa) [isosyntype in K];
Reinecke 88c (Manua-Inseln, Samoa)
Distribution: Samoa (endemic?).
Chromosome number: unknown.
Note: This is apparently similar to widespread
H. holochilum, but characterized by terminal soriand receptacles often exerted from involucres.
18. Hymenophyllum dimidiatum Mett. ex
Kuhn, Linnaea 35: 393 (1868).
[Fig. 18]
Meringium dimidiatum (Mett.) Copel., Philipp. J.
Sci. 67: 43 (1938).
Type: Deplanche s.n. (New Caledonia) [B*](see Brownlie, 1969).
Hymenophyllum piliferum C. Chr., Vierteljahrsschr. Naturf. Ges. Zurich 70: 221 (1925).
Type: Franc 1450, 1913 (Baie du sud, New
Caledonia) [P; BM K MICH*].
Distribution: New Caledonia (endemic).
Chromosome Number: n21 (Braithwaite,
1975, New Caledonia).
Note: In New Caledonia, this species resembles H. holochilum; both species has ‘dimidiate
pinna’ (i. e., the upper and lower parts of the
pinna divide unevenly). But H. dimidiatum iseasily distinguishable by the serrate lips of the involucre. The incorrect original description“folia ...glaberrima,...” has confused subsequent
taxonomists; the type collection actually has
hairs on the underside of the axis (cf. Brownlie,1960).
19. Hymenophyllum rolandi-principis Rosens.,
Repert. Spec. Nov. Regni Veg. 9: 72 (1910). (Fig.
19)— Rosenstockia rolandi-principis (Rosesnt.)
Copel., Gen. Fil. 36 (1947).
Type: Franc 1422, Jan. 1910 (Mt. Tao, New
Caledonia) [isotypes in P UC* US*].
Distribution: New Caledonia (endemic).
Chromosome number: n21 (Ebihara et al.,2002, New Caledonia).
Note: Copeland (1947) established the 34th
genus Rosenstockia for this peculiar plant. The

64

Atsushi Ebihara and Kunio Iwatsuki

Figs. 15–20. 15a–b. Hymenophyllum serrulatum (a. Braithwaite RSS 4748 [BM]; b. Bernardi 13363 p.p. [K]).
16a–b. H. holochilum (a. Balansa 1642 [P]; b. Ebihara 001226-01 [TI]). 17. H. praetervisum (Reinecke 88[K, isosyntype]). 18a–b. H. dimidiatum (Ebihara 001229-01 [TI]). 19a–b. H. rolandi-principis (Ebihara
001225-11 [TI]). 20a–b. H. affine (Braithwaite 691 [K]). Scale1 cm for whole leaves, and 0.4 mm for sori.

Pacific Hymenophyllaceae I

study by Ebihara et al. (2002) suggested this isactually a result of drastic morphological specialization from H. dimidiatum.
20. Hymenophyllum macgillivrayi (Baker)
Copel., Philipp. J. Sci. 64: 60. t. 25 (1937).
[Fig. 20]
— Trichomanes macgillivrayi Baker, Ann. Bot.
(London) 5: 195 (1891) — Meringium macgillivrayi (Baker) Copel., Philipp. J. Sci. 67: 44(1938).
Type: Macgillivray s. n., 1854 (“Feejee
Island”Fiji) [K].
Hymenophyllum affine auct non Brack.: E.
Fourn., Ann. Sci. Nat., Bot. V 18: 264 (1873);
Brownlie, Beih. Nova Hedw. 10: 532 (1977) p.p.;
Ebihara, Ann. Tsukuba Bot. Gard. 21: 9 (2003).
Distribution: Vanuatu, Fiji.
Chromosome Number: n21 (Braithwaite
1975, Fiji).
Note: There is much confusion concerning theidentity of this species. Though Brownlie (1960)
once recognized two species (H. affine with campanulate lips and H. pseudotunbridgense withlonger lips) in Fiji, later he reduced them to one
species H. affine (Brownlie, 1977). The name H.
macgillivrayi has often been treated as a synonym of H. affine, but campanulate involucres of
the type material of H. macgillivrayi are clearly
distinguishable from those of H. affine. Consequently, H. affine should be a synonym of H.
holochilum, and the correct name for the plants
with campanulate sori is H. macgillivrayi.
21. Hymenophyllum gorgoneum Copel.,Philipp. J. Sci. 64: 60, t. 26 (1937).
[Fig. 21]
— Meringium gorgoneum (Copel.) Copel.,Philipp. J. Sci. 67: 44 (1938).
Type: Brass 3304 (Tiratona, Santa Ysabel,
Solomon Islands) [PNH* (now lost?); GH
MICH].
Distribution: Solomon Islands; New Guinea
Chromosome number: n21 (Braithwaite,
1969, Solomon Isl.).

65

22. Hymenophyllum denticulatum Sw., J.
Bot. (Schrader) 1800(2): 100 (1801).
[Fig. 22]
— Trichomanes denticulatum (Sw.) Poir. in
Lamarck et Poir., Encycl. 8: 75 (1808) — Didymoglossum denticulatum (Sw.) Hassk., Obs. Bot.
2: 16 (1857) — Leptocionium denticulatum (Sw.)
Bosch, Ned. Kruidk. Arch. 4: 382 (1859) —Meringium denticulatum (Sw.) Copel., Philipp. J.
Sci. 67: 42 (1938).
Type: Thunberg s. n. (Java) (n. v.).
Distribution: Fiji; Asia to Malesia.
Chromosome number: unknown.
Note: The Hymenophyllum denticulatum—H.
acanthoides complex is one of the typical Malesian Hymenophyllum. It is somewhat enigmaticthat this species was repeatedly collected in Fiji
though there is no record from either Solomon
Islands or Vanuatu adjacent to Fiji.
23. Hymenophyllum kerianum Watts, Proc.
Linn. Soc. New South Wales 39: 767, t. 87, fig. 6(1915).
[Fig. 23]
— Meringium kerianum (Watts) Copel., Philipp.
J. Sci. 67: 43 (1938).
Lectotype: Watts s. n. [NSW P1240], July
1913 (Frenchman’s Creek, QLD, Australia)[NSW; BRI*]; designated by Croxall, Austral. J.
Bot. 23: 515 (1975).
Distribution: Australia (QLD) (endemic).
Chromosome number: unknown.
Note: This species is undoubtedly a close relative of H. denticulatum, and needs further careful
comparison.
24. Hymenophyllum brassii C. Chr., Brittonia
2: 273 (1937).
[Fig. 24]
— Meringium brassii (C. Chr.) Copel., Philipp.
J. Sci. 73: 463 (1941) — Myriodon brassii (C.
Chr.) Copel., Gen. Fil. 36 (1947).
Type: Brass 4114, 27 May 1933 (New Guinea)[BM; BO GH MICH].
Hymenophyllum
odontophyllum
Copel.,
Philipp. J. Sci. 64: 73, t. 35 (1937) — Myriodon
odontophyllum (Copel.) Copel., Philipp. J. Sci.
67: 47 (1938).
Type: Ledermann 10357 (New Guinea)

66

Atsushi Ebihara and Kunio Iwatsuki

Figs. 21–26. 21a–b. Hymenophyllum gorgoneum (Braithwaite RSS 4316 [BM]). 22. H. denticulatum (Herald1263 [K]). 23. H. kerianum (Watts P1241 [BM, possible syntype]. 24. H. brassii (Braithwaite RSS 4415 [K]).
25. H. walleri (Croxall and Parris 3247 [K]). 26. H. francii (Franc 163 [P; holotype]). Scale1 cm for whole
leaves, and 0.4 mm for sori.

Pacific Hymenophyllaceae I[MICH; B BM].
Distribution: Solomon Isl.; New Guinea.
Chromosome number: n21 (Braithwaite,
1969, Solomon Isl.).
Note: This species is characterized by the lessexpanding lamina cells, and Copeland (1947)
separated it as a distinct genus Myriodon, alongwith the type species Myriodon odontophyllum(Copel.) Copel. It is clear that this is a specialized form of a species of “H. acanthoides clade”(Ebihara et al., 2002; Hennequin et al., 2006).
25. Hymenophyllum walleri Maiden et
Betche, Proc. Linn. Soc. New South Wales 35:
802 (1910).
[Fig. 25]
— Mecodium walleri (Maiden & Betche)
Copel., Philipp. J. Sci. 67: 21 (1938).
Type: Waller s. n., 1908 (Evelyn Scrub, near
Herberton, QLD, Australia) [NSW; BRI*
MEL?*] (see Croxall, 1975).
Distribution: Australia (QLD) (endemic).
Chromosome number: unknown.
Note: This species, having entire margins andlips, belonged to Mecodium by the definition of
Copeland’s system (1938). However, since its
rachis and stipe underneath are densely covered
with multicellular hairs, this species falls into thecategory of “Hymenophyllum sect. Pseudomecodium” sensu Iwatsuki (1984), now attributed tosubgen. Hymenophyllum, the same as H. exsertum Wall. ex Hook., H. oligosorum Makino andH. pachydermicum Ces.
26. Hymenophyllum francii (Christ) Ebihara
et K.Iwats., Taxon 53: 945 (2004).
[Fig. 26]
— Trichomanes francii Christ, Bull. Herb.
Boissier II 7: 648 (1907) — Microtrichomanes
francii (Christ) Copel., Philipp. J. Sci. 67: 37(1938).
Type: Franc 163, Feb. 1906 (Mt. Mou, New
Caledonia) [P; B BO NSW NY SING UC US](see Ebihara et al., 2005).
Trichomanes cuneatum Christ, Bull. Herb.
Boissier II 7: 649 (1907).
Type: Franc 165, July 1906 (Mt. Mou, New
Caledonia) [P].

67

Distribution: New Caledonia (endemic).
Chromosome number: unknown.
Note: This is the only species whose systematic position we could not assume from the morphological characters. Although it was formerly
treated as a member of Microtrichomanes, its
wiry root strongly suggests that it be included in
Hymenophyllum, contrary to the proposition by
Iwatsuki (1975).
Doubtful species
Hymenophyllum lobbii T. Moore ex Bosch, Ned.
Kruidk. Arch. 5(3): 176 (1863).
— Meringium lobbii (T. Moore ex Bosch)
Copel., Philipp. J. Sci. 67: 43 (1938).
Note: This dwarf species has been recorded
from Queensland only once, but it is quite unnatural for this Bornean species to have an isolated
distribution in Australia. It is possibly a mislabeling of the collections by the collector (P. Bostock, personal communication).
Acknowledgments
The authors are grateful to curators of theherbaria BM, CHR, K, KYO, L, MO, NOU,
NSW, NTBG, P and TI.
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