Bull. Natn . Sci. Mus., Tokyo, Ser. B, 22(3) , pp. 113- 134, September 22, 1996

A Checklist of the Mosses of Vanuatu
By

Masanobu HIGUCHI
Department of Botany, National Science Museum,4- 1- 1 Amakubo, Tsukuba, lbaraki 305, Japan

Abstract A checklist of the mosses of Vanuatu is given based on published reports.
Totals of 33 families, 84 genera and 236 species and infraspecific taxa are recorded.
The chromosome number, n = 11 , for Acanthorrhynchium papilla tu m is reported forthe first time by the re-examination of the reported specimens.

Vanuatu, previously known as New Hebrides, is situated in the southwest
region of the Pacific Ocean. Its neighbors are New Caledonia, ea. 400 km to thesouthwest, Fiji, ea. 800 km to the east, and the Solomon Islands, ea. 800 km to thenorthwest. The irregular Y -shaped archipelago in a northwest/southeast orientation consists of about 80 islands, and stretches 900 km between 13 °S and 22 °S
latitude, and between 166°E and 175 °E longitude (Fig. 1). The highest peak of1879 m is Mt. Tabwemasana on Espiritu Santo which is the largest island in thearchipelago. Geologically the islands of Vanuatu originated from volcanic
activity and coral formation, and date from the Tertiary (Vitt, 1991). The total
land mass is approximately 12,930 square kilometers (Whyte, 1990) . From
November to April the weather is warm and humid.
The mosses of Vanuatu have been reported by Mitten (1868), Brotherus &Watts (1915), Theriot (1938), Dixon (1948), and Tixier (1972, 1973, 1974). The
present state of knowledge of the moss flora of Vanuatu is far from complete.
There is no checklist of mosses of Vanuatu except that of Schultze-Motel ( 1973)
who compiled a catalog of mosses of Melanesia. In January 1992 I had theopportunity to do field research at several localities on Efate and Espiritu Santo,and collected more than 500 bryophyte specimens. Based on this collection,
recent reports were published by Matsui & Iwatsuki (1993), Inoue & Higuchi(1994), and Iwatsuki & Suzuki (1995) .
Checklist
The present list includes 33 families, 84 genera, and 236 species and infraspecific taxa (Table 1) . The nomenclature used follows Index Muscorum(Wijk et al., 1959-1969) except for that found in more recent publications. The

Masanobu H IGUCH I

114

1b Torres Islands
<;)

0

a

Vanua Lava

0""

-Q. c

Banks Islands

0

<\, o

Efate ~

Erromango~

100 km0

FutunaAnatomFig. I.

Q

Outline map of Vanuatu.

arrangement of the families follows Crosby & M a gill ( 1978). Genera and species
are arranged alphabetically in families.
Fissidentaceae
Fissidens angustifolius Sull. - Iwatsuki & Suzuki ( 1995; Efate, Espiritu San to) .
Fissidens consociatus Ther. - Iwatsuki & Suzuki (1995; Espiritu Santo).
Fissidens dealbatus Hook. f. & Wilson - lwatsuki & Suzuki (1995; Efate).
Fissidens ftabellulus Thwaites & Mitt. - Iwatsuki & Suzuki (1995; Efate) .
Fissidens hollianus Dozy & Molk. - Iwatsuki & Suzuki (1995; Efate, Espiritu
Santo).
Fissidens subscabrisetus Broth. & Watts - Brotherus & Watts ( 1915, sp.

A Checklist of the mosses of Vanuatu
Table I.

A summary of the taxa of mosses reported from Vanuatu.

FamilyFissidentaceae
Ditrichaceae
Dicranaceae
Dicnemonaceae
Leucobryaceae
Calymperaceae
Pottiaceae
Splachnaceae
Bryaceae
Mniaceae
Calomniaceae
Rhizogoniaceae
Hypnodendraceae
Bart ramiaceae
Spiridentaceae
Orthotrichaceae
Racopilaceae
Cyrtopodaceae
Myuriaceae
Pterobryaceae
Meteoriaceae
Phyllogoniaceae
Daltoniaceae
Neckeraceae
Lembophyllaceae
Hookeriaceae
Hypopterygiaceae
Thuidiaceae
Brachytheciaceae
Entodontaceae
Sematophyllaceae
Hypnaceae
Polytrichaceae
Total

115

GeneraSpecies and infraspecific taxa
17

4

12

I

2
6
3

13
23

6

2

2

2

4

2

4
4
8

2

6

7
5

I
12
11
I

7

2

2

7
4
13
4
8
6
4

10
7

22
28

84

236

4
I
3

2
3
3

nov. ; Paama). cf. Iwatsuki & Suzuki (1995).
Fissidens hyalinus Hook. & Wilson - Iwatsuki & Suzuki (1995; Espiritu Santo) .
Fissidens lagenarius Mitt.- Iwatsuki & Suzuki (1995; Efate, Espiritu Santo).
Fissidens lautokensis Dixon - lwatsuki & Suzuki ( 1995; Espiritu San to).
Fissidens laxus Sull. & Lesq.- Iwatsuki & Suzuki ( 1995; Efate, Espiritu San to) .
Fissidens linearis Brid. var. obscurirete (Broth. & Paris) I. G. Stone- lwatsuki& Suzuki (1995; Efate).
Fissidens maceratus Mitt. - lwatsuki & Suzuki (1995; Efate, Espiritu Santo).
Fissidens oblongifolius Hook. f. & Wilson- Iwatsuki & Suzuki (1995; Efate).

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Masanobu HIGU C HI

Fissidens subbryoides Hampe ex Gangulee- I watsuki & Suzuki ( 1995; Efate).
Fissidens sylvaticus Griff. - Tixier ( 1974; Espiritu San to).
Fissidens zippelianus Dozy & Molk. var. zippelianus - Theriot (1938; Tanna),
Iwatsuki & Suzuki ( 1995; Espiritu San to) .
Fissidens zippelianus Dozy & Molk. var. robinsonii (Broth.) Z. lwats. & Suzuki- Iwatsuki & Suzuki (1995 ; Espiritu Santa) .
Fissidens zollingeri Mont. - Iwatsuki & Suzuki (1995; Efate, Espiritu Santa).
Ditrichaceae
Ceratodon purpureus (Hedw.) Brid. - Tixier (1973; Tanna) .
Dicranaceae
Campylopus caudatus (Miili. Hal.) Mont. - Tixier (1974; Espiritu Santa) .
Campylopus comosus (Schwagr.) Bosch & San de Lac. var. auberti Ther. Theriot (1938, var. nov.; Tanna) .
Campylopus introflexus (Hedw.) Brid. - Brotherus & Watts (1915; Anatom) .
Campylopus umbellatus (Arnold) Paris
Pilopogon blumii (Dozy & Molk.) Broth. - Brotherus & Watts (1915;
Anatom).
Campylopus blumii (Dozy & Molk.) Bosch & San de Lac. - Theriot ( 1938;
Aoba) .
Campyopus richardii Brid. - Tixier (1974; Espiritu Santa) .
Campylopus verrucosus Besch. - Tixier (1974; Espiritu Santa) .
Campylopus sp.- Brotherus & Watts (1915; Anatom) .
Dicranodontium sp.- Brotherus & Watts (1915; Futuna).
Dicranoloma blumii (Nees) Paris - Tixier (1974; Espiritu Santa).
Dicranoloma brachysteleum (Miill. Hal.) Paris - Miilier (1897, sp. nov.;
Anatom), Brotherus & Watts (1915; New Hebrides).
Dicranoloma braunii (Miill. Hal.) Paris - Brotherus & Watts (1915 ; Anatom).
Leucoloma molle (Miili. Hal.) Mitt. - Tixier (1972; Anatom).
Leucoloma subtenuifolium Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Anatom), Tixier (1973; Anatom).
Dicnemonaceae
Dicnemon aneitensis (Broth. & Watts) B. H. Alien - Alien (1989; Anatom).
Synodontia aneitensis Broth. & Watts- Brotherus & Watts (1915, sp. nov. ;Anatom).

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117

Leucobryaceae
Leucobryum aduncum Dozy & Molk. - Dixon (1948; Anatom).
Leucobryum aneitense Broth. & Watts - Brotherus & Watts (1915, sp. nov. ;Anatom), Dixon (1948; Anatom).
Leucobryum bowringii Mitt. - Tixier (1973; Anatom), Tixier (1974; Espiritu
Santa) .
Leucobryum candidum (P. Beauv.) Wilson - Brotherus & Watts (1915;
Anatom, Epi, Futuna) , Theriot (1938; Erromango), Tixier (1974; Efate).
Leucobryum chlorophyllosum Mull. Hal.
Leucobryum stenophyllum Besch. - Brotherus & Watts (1915; Anatom). cf.
Yamaguchi & Iwatsuki ( 1987) .
Leucobryum conocladon Besch. - Brotherus & Watts (1915 ; Anatom), Dixon(1948; Anatom) .
Leucobryum fragile Mtill. Hal. , nom. nud. - Muller (1901 ; New Hebrides).
Leucobryum gunnii Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Anatom, Futuna) .
Leuobryum neo-caledonicum Besch. - Brotherus & Watts (1915; Futuna) ,Dixon (1948 ; Anatom), Yamaguchi & Iwatsuki (1987; New Hebrides) .
Leuobryum sanctum (Brid.) Hampe - Mitten (1873 ; Anatom), Fleischer ( 1904;
New Hebrides), Brotherus & Watts (1915 ; Espiritu Santa) .
Octoblepharum sanctum (Brid.) Mitt. ex Paris - Mitten (1882; Anatom).
Leucophanes albescens Mull. Hal.- Theriot (1938; Futuna) .
Leucophanes candid urn (Schwagr.) Lindb. - Theriot ( 1938; Futuna) .
Leucophanes glauculum Mull. H al. ex M. Fleisch. - Brotherus & Watts (1915;
Espiritu Santa).
Leucophanes octoblepharioides Brid. - Brotherus & Watts ( 1915; Epi, Paama,
Espiritu San to).
Calymperaceae
Arthrocormus schimperi (Dozy & Molk.) Dozy & Molk. - Brotherus ( 1909;
Espiritu Santa, Tongoa) , Brotherus & Watts (1915 ; Espiritu Santa), Tixier(1973, Anatom), Tixier (1974, Erromango) .
Calymperes aongstroemii Besch. - Brotherus & Watts (1915 ; Espiritu Santa) .
Calymperes chamaeleonteum Mtill. Hal. - Tixier (1973; Anatom) .
Calymperes serratum A. Braun ex Mull. Hal. - Brotherus & Watts (1915;
Anatom, Espiritu Santa).
Calymperes lorifolium Mitt. - Brotherus & Watts (1915; Anatom).
Calymperes tahitense (Sull.) Mitt. - Mitten (1873; Anatom) , Mitten (1882;
Anatom) .

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Masanobu HIG UCHI

Exodictyon dentatum (Mitt.) Cardot - Brotherus & Watts (1915; Futuna,
Espiritu Santo), Theriot (1938; Erromango, Futuna, Pentecost) .
Exodictyon subscabrum (Broth.) Cardot - Brotherus & Watts (1915; Espiritu
Santo).
Mitthyridium constrictum (Sull.) H. Rob.- Nowak (1980; Espiritu Santo,
Tongoa) .
Syrrhopodon constrictus (Sull.) Mitt. - Brotherus & Watts (1915; Espiritu
Santo) .
Mitthyridium cuspidatum (M. Fleisch.) H . Rob.
Thyridium cuspidatum M. Fleisch. - Tixier ( 1974; Erromango, Espiritu
Santo) .
Mitthyridium fasciculatum (Hook. & Grev.) H. Rob.
Syrrhopodon fasciculatus Hook. & Grev. - Brotherus & Watts (1915;
Anatom, Epi, Espiritu Santo).
Thyridiumfasciculatum (Hook. & Grev.) Mitt. - Tixier (1973; Anatom).
Mitthyridium obtusifolium (Lindb.) H. Rob.
Thyridium obtusifolium (Lindb.) M. Fleisch. - Theriot (1938; Erromango).
Mitthyridium samoanum (W. Schultze-Motel) H . Rob. - Nowak (1980;
Anatom, Erromango, Espiritu Santo, Futuna).
Octoblepharum albidum Hedw. - Brotherus & Watts (1915; Espiritu Santo).
Syrrhopodon albo-vaginatus Schwagr.- Brotherus & Watts (1915; Anatom).
Syrrhopodon japonicus (Besch.) Broth.
Syrrhopodon aneitensis Broth. & Watts - Brotherus & Watts ( 1915, sp. nov.;
Anatom). cf. Reese et al. (1986).
Syrrhopodon tenuinervis Broth. & Watts - Brotherus & Watts (1915, sp.
nov.; Futuna). cf. Reese et al. (1986).
Syrrhopodon diversiretis Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Anatom).
Syrrhopodon ftammeonervis Mull. Hal. subsp. lilliei (Broth.) Tixier
Syrrhopodon lilliei Broth. - Brotherus & Watts (1915, sp. nov. ; Anatom) .
Syrrhopodon kindelii Broth. & Paris - Dixon ( 1948; Erromango, Espiritu San to,
Tongoa).
Syrrhopodon laevigatus Mitt.
Syrrhopodon graeffeanus Mull. Hal. - Theriot (1938; Vanua Lava) . cf.
Schultze-Motel (1973) .
Syrrhopodon mammillosus Miill. Hal. - Theriot (1938; Tanna) .
Syrrhopodon muelleri (Dozy & Molk.) Sande Lac. - Brotherus & Watts (1915;
Anatom, Espiritu Santo) , Theriot (1938; Futuna).
Syrrhopodon perarmatus Broth. - Brotherus & Watts (1915, sp. nov.; Anatom,
Futuna, Espiritu Santo) .
Syrrhopodon tristichus Nees ex Schwagr. - Brotherus & Watts (1915; Espiritu

A Checklist of the mosses of Vanuatu

119

Santa) , Tixier (1972; Anatom), Tixier (1973; Tanna), Tixier (1974; Tanna).
Pottiaceae
Barbula indica (Hook.) Spreng. - Theriot (1938; Tanna) .
Hydrogonium aneitense (Broth. & Watts) W. Schultze-Motel
Barbula aneitensis Broth. & Watts - Brotherus & Watts ( 1915, sp. nov.;
Ana tom).
Hydrogonium arcuatum (Griff.) Wijk & Margad.
Hydrogonium comosum (Dozy & Molk.) Hilp. - Tixier ( 1973; Espiritu
Santa).
Hydrogonium consanguineum (Thwaites & Mitt.) Hilp. - Tixier ( 1974; Espiritu
San to).
Hydrogonium inflexum (Duby) P. C. Chen - Tixier (1973; Espiritu Santa).
Hyophila microphylla Broth. & Watts- Brotherus & Watts (1915, sp. nov.;
Ana tom).
Splachnaceae
Tayloria braithwaitiana Tixier - Tixier (1974, sp. nov. - Espiritu Santa).
Bryaceae
Brachymenium indicum (Dozy & Molk.) Bosch & Sande Lac. - Brotherus &Watts (1915; Anatom).
Epipterygium vanuatuicum H . A. Mill. - Miller (1988, sp. nov. ; Ambrym).
Mniaceae
Plagiomnium succulentum (Mitt.) T. J. Kop. (?)
Mnium 'esculentum' Mitt. - Tixier ( 1974; Espiritu San to).
Tixier (1974) reported Mnium 'esculentum' Mitt. (p. 35) from Espiritu
Santa. However, this name is not listed on "Index Muscorum (Wijk et al., 1964,1969)", "Index of Mosses 1963- 1989 (Crosby et al., 1992)" and "Indices to theSpecies of Mosses and Lichens Described by William Mitten (Thiers, 1992)."
Judging from the note of the range, "Nepal, Assam, lndes, Ceylan, Malaisie,Japan, lndochine, Sumatra, Java, Borneo, Nouvelle-Guinee," it may be theorthographic error of Mnium succulentum Mitt.

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Masanobu HIGUCHI

Calomniaceae
Calomnion melanesicum H . A. Mill.- Miller (1987; Ambrym, Aoba, Efate,
Espiritu Santo, Pentecost, Tanna).
Rhizogoniaceae
Pyrrhobryum medium (Besch.) ManuelRhizogonium medium Besch. - Brotherus & Watts (1915; Anatom).
Pyrrhobryum spiniforme (Hedw.) Mitt. - Inoue & Higuchi ( 1994; Espiritu
Santo).
Rhizogonium spiniforme (Hedw.) Bruch - Brotherus & Watts (1915 ;Anatom, Espiritu Santo, Futuna), Tixier (1973; Espiritu Santo).
Rhizogonium setosum (Mitt.) Mitt.- Mitten (1882; Anatom), Brotherus &Watts (1915; Anatom, Epi, Futuna), Theriot (1938; Erromango, Futuna),
Tixier (1973; Anatom, Tanna), Tixier (1974; Anatom, Erromango, Espiritu
Santo).
Rhizogonium setosum (Mitt.) Mitt. f. crispulum Ther.- Theriot (1938, f. nov.;
Erromango, Tanna, Vanua Lava).
Hypnodendraceae
Hypnodendron dendroides (Brid.) Touw - Touw ( 1971; Anatom, Espiritu
Santo, Vanua Lava).
Mniodendron densirameum Broth.- Theriot ( 1938; Vanua Lava) . cf. Touw
(1971).
Mniodendron hellwigii Broth. ex Broth. - Brotherus & Watts (1915;
Anatom), Dixon (1948; Anatom). cf. Touw (1971).
Hypnodendron ftagelliferum Broth. & Watts - Brotherus & Watts (1915, sp.
nov.; Anatom), Touw (1971; Anatom), Tixier (1972; Anatom).
Hypnodendron fusco-mucronatum (Mull. Hal.) A. Jaeger subsp. chalmersii(Mitt.) Touw-Touw (1971; Espiritu Santo).
Hypnodendron milnei Mitt. subsp. milnei - Mitten (1873, sp. nov.; Anatom),Mitten (1882; Anatom), Touw (1971; Anatom).
Mniodendron milnei (Mitt.) Mitt.- Brotherus & Watts (1915; Anatom).
Hypnodendron spininervium (Hook.) A. Jaeger - Brotherus & Watts (1915;
Anatom, Futuna, Espiritu Santo), Theriot (1938; Tanna).
Hypnodendron subspininervium (MUll. Hal.) A. Jaeger subsp. subspininervium- Brotherus & Watts (1915; Anatom), Theriot (1938; Futuna), Touw
(1971; Anatom, Futuna, Tongoa).
Braithwaitea arborescens Miit. - Mitten ( 1882; Anatom) .

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121

Hypnodendron vitiense Mitt. subsp. vitiense - Touw ( 1971; Ana tom, Espiritu
Santo, Futuna, Tanna, Tongoa), Tixier (1973; Espiritu Santo), Tixier (1974;
Espiritu San to) .
Hypnodendron arcuatum (Hedw.) Lindb. ex Mitt.- Tixier ( 1972;
Ana tom).
Bartramiaceae
Breutelia arundinifolia (Duby) M. Fleisch. -Tixier (1974; Espiritu Santo).
Philonotis hastata (Duby) Wijk & Margad.
Philonotis laxissima Mitt. nom. illeg.- Brotherus & Watts (1915; Anatom) .
Philonotis obtusifolia (Mitt.) Paris - Brotherus & Watts ( 1915; Espiritu San to).
Philonotis turneriana (Schwagr.) Mitt.- Tixier (1974; Espiritu Santo) .
Spiridentaceae
Spiridens aristifolius Mitt. - Theriot ( 1938; Futuna) .
Spiridens aristifolius Mitt. var. pallidus Ther. - Theriot ( 1938, var. nov.;
Futuna) .
Spiridens ftagellosus Schimp. - Mitten (1873; Anatom), Mitten (1882;
Anatom), Brotherus & Watts (1915; Anatom, Futuna, Espiritu Santo),
Theriot (1938; Pentecost), Tixier (1972; Anatom), Tixier (1973; Anatom,
Espiritu Santo), Tixier (1974; Efate).
Spiridens reinwardtii Nees - Brotherus & Watts (1915; Anatom, Espiritu
Santo) .
Orthotrichaceae
Macromitrium orthostichum Nees ex Schwagr. - Brotherus & Watts (1915;
Futuna).
Macromitrium orthostichum Nees ex Schwagr. subsp. micropoma M. Fleisch. Tixier (1974; Anatom).
Macromitrium pulchrum Besch.- Dixon (1948; Anatom).
Macromitrium reinwardtii Schwagr. - Brotherus & Watts ( 1915; Ana tom),Tixier (1973; Anatom).
Macromitrium salakanum Mtill. Hal. -Tixier (1972; Anatom), Tixier (1973;
New Hebrides), Tixier (1974; Anatom, Erromango).
Macromitrium semi-pellucidum Dozy & Molk.- Tixier (1974; Efate) .
Macromitrium subtile Schwagr.- Brotherus & Watts (1915; Futuna, Espiritu
Santo) .
Macromitrium sulcatum (Hook.) Brid. -Tixier (1973; Anatom).

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Masanobu HIGUC HI

Racopilaceae
Powellia involutifolia Mitt. - Inoue & Higuchi (1994; Espiritu Santa).
Racopilum cuspidigerum (Schwagr.) Angstr. - Brotherus & Watts (1915;
Espiritu Santa).
Racopilum gracillimum Mi.ill.Hal. - Muller (1896, sp. nov.; Erromango) .
Racopilum pacificum Besch. - Brotherus & Watts (1915; Futuna), Fleischer(1923; New Hebrides).
Racopilum pacificum Besch. var. gracilescens Besch. - Theriot (1938; Tanna).
Racopilum spectabile Reinw. & Hornsch. - Brotherus & Watts (1915; Anatom,
Futuna, Espiritu Santa), Theriot (1938; Aoba), Tixier (1974; Erromango),
Inoue & Higuchi ( 1994; Espiritu San to).
Cyrtopodaceae
Bescherellia elegantissima Duby- Brotherus & Watts ( 1915; Anatom), Bartram(1953; Anatom).
Myuriaceae
Myurium rufescens (Reinw. & Hornsch.) M . Fleisch. subsp. purpuratum (Mitt.)
Maschke
Oedicladium purpura tu m Mitt. - Mitten ( 1882; Ana tom).
Myurium purpuratum (Mitt.) Dixon & Ther. - Brotherus & Watts (1915;
Anatom) .
Pterobryaceae
Calyptothecium urvilleanum (Mi.ill. Hal.) Broth. - Brotherus & Watts (1915 ;Espiritu Santa), Tixier (1974; Espiritu Santa) .
Euptychium dumosum (Besch.) Broth.- Bartram (1953; New Hebrides).
Euptychium setigerum (Sull.) Broth. subsp. setigerum- During (1977; Anatom,
Efate, Futuna) .
Euptychium assimile Broth. & Watts - Brotherus & Watts ( 1915, sp. nov.;
Futuna), Tixier (1972; Anatom) . cf. During (1977).
Euptychium gunnii Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Anatom), Tixier (1973; Anatom), Tixier (1974; Erromango). cf. During(1977).
Euptychium setigerum (Sull.) Broth. subsp. austro-caledonicum (Besch.) During
Euptychium austro-ca/edonicum (Besch.) Broth. - Tixier (1974; Anatom).
Garovaglia elegans (Dozy & Molk.) Hampe ex Bosch & San de Lac. - During

A Checklist of the mosses of Vanuatu

123( 1977; An atom, Espiritu San to, Futuna).
Endotrichella elegans (Dozy & Molk.) M. Fleisch. - Dixon ( 1948;
Anatom) .
Endotrichella campbelliana Hampe-- Hampe (1874, sp. nov.; Anatom),Mitten (1882; Anatom), Brotherus & Watts (1915; Anatom) . cf. During(1977) .
Endotrichella samoana (Mitt.) Mtill. Hal.- Theriot (1938; Futuna). cf.
During ( 1977).
Pterobryella rigida (Mitt.) Touw- Touw (1971; Anatom).
Hypnodendron rigidum Mitt. - Mitten (1873, sp. nov.; Anatom), Mitten(1882; Anatom), Brotherus & Watts (1915; Anatom).
Pterobryella vagapensis Mtill. Hal. - Brotherus & Watts (1915; Anatom),Tixier ( 1972; Ana tom). cf. Schultze-Motel ( 1973).
Symphysodon gunnii Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Anatom, Epi, Futuna), Tixier (1974; Erromango).
Symphysodon neckeroides Dozy & Molk.
Symphysodon micholitzii Broth. - Brotherus & Watts (1915 ; Espiritu
Santo) .
Symphysodon vitianus (Sull.) Broth. - Brotherus & Watts (1915; Anatom, Epi,
Espiritu Santo), Tixier (1973; Anatom).
Symphysodontella cylindracea (Mont.) M. Fleisch. - Brotherus & Watts (1915;
Anatom).
Symphysodontella cylindracea (Mont.) M. Fleisch. var. attenuata Ther. Theriot (1910, var. nov.; Anatom).
Trachyloma tahitense Besch. - Tixier ( 1974; Espiritu San to).
Meteoriaceae
Aerobryopsis longissima (Dozy & Molk.) M. Fleisch.- Brotherus & Watts(1915; Anatom, Epi, Espiritu Santo), Tixier (1972; Anatom), Tixier (1973;
Tanna).
Aerobryopsis striatula (Mitt.) Broth. - Mitten (1873, sp. nov.; Anatom),Brotherus & Watts (1915; Anatom, Epi, Futuna).
Aerobryopsis vitiana (Sull.) M. Fleisch. - Brotherus & Watts (1915; Espiritu
Santo).
Barbella futunensis Ther. -Theriot (1938, sp. nov.; Futuna).
Floribundaria floribunda (Dozy & Molk.) M. Fleisch. - Brotherus & Watts(1915; Espiritu Santo) , Theriot (1938; Futuna), Tixier (1973; Anatom,
Banks, Efate, Espiritu San to), Tixier ( 1974; Espiritu San to).
Floribundaria floribunda (Dozy & Molk.) M. Fleisch. var. brevifolia (Ren . &Cardot) M. Fleisch. - Theriot ( 1938; Efate, Pentecost) .

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Masanobu HI GUC HI

Floribundaria floribunda (Dozy & Molk.) M. Fleisch. var. latifolia Cardot Theriot (1938; Futuna, Pentecost) .
Floribundaria pseudo-floribunda M. Fleisch. - Brotherus & Watts (1915 ;Ana tom, Futuna, Paama, Espiritu Santo).
Meteorium miquelianum (Mtill. Hal.) M. Fleisch. - Brotherus & Watts (1915;
Anatom, Epi, Futuna), Theriot (1938 ; Futuna) ., Tixier (1973; Tanna).
Papillaria helictophylla (Mont.) Broth. - Brotherus & Watts (1915; Futuna) ,Theriot (1938; Futuna) .
Papillaria pellucida Broth. & Watts- Brotherus & Watts (1915, sp. nov.;
Anatom, Futuna) .
Phyllogoniaceae
Orthorrhynchium cylindricum (Lindb.) Broth. - Brotherus & Watts (1915;
Espiritu Santo) , Theriot (1938; Foutuna, Pentecost).
Daltoniaceae
Cyathophorella spinosa (Mtill. Hal.) M. Fleisch. - Brotherus & Watts (1915;
Espiritu Santo).
Cyathophorella tahitensis (Besch.) M. Fleisch.- Brotherus & Watts (1915;
Ana tom), Tixier ( 1974; Espiritu San to) .
N eckeraceae
Himantocladium flagelliferum (Broth.) Broth. - Theriot ( 1938; Pentecost) .
Himantocladium loriforme (Bosch & Sande Lac.) M . Fleisch. - Brotherus &Watts (1915; Epi), Tixier (1974; Espiritu Santo) .
Himantocladium plumula (N ees) M. Fleisch. - Tixier ( 197 4; Espiritu San to).
Homaliodendron flabellatum (Sm.) M . Fleisch. - Tixier (1973; New Hebrides) .
Homaliodendron dendroides (Hook.) M. Fleisch. - Brotherus & Watts(1915; Anatom, Futuna) .
Homaliodendron microdendron (Mont.) M. Fleisch. - Tixier ( 1973; Espiritu
Santo).
Neckeropsis lepineana (Mont.) M. Fleisch. - Brotherus & Watts (1915 ;Anatom, Epi, Futuna), Theriot (1938; Pentecost) , Touw (1962; Anatom,
Futuna, Tanna), Tixier ( 1973; Anatom) .
Thamnobryum aneitense (Mitt.) W. Schultze-Motel
Porotrichum aneitense Mitt. - Mitten (1873 ; Anatom).
Thamnium aneitense (Mitt.) Mitt. - Mitten (1882; Anatom), Brotherus &Watts (1915 ; Anatom) .

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125

Lembophyllaceae
Camptochaete arbuscula (Sm.) Reichardt var. deflexa (Wilson) Dixon
Camptochaete dejiexa (Wilson) A. Jaeger - Tixier ( 1972; Anatom).
Camptochaete leichhardtii (A. Jaeger) Broth. - Brotherus & Watts (1915;
Anatom) .
Thamnium leichhardtii (A. Jaeger) Kindb. - Kindberg (1902; New Hebrides) .
Camptochaete porotrichoides (Besch.) Broth. - Brotherus & Watts (1915;
Anatom), Bartram (1953; Anatom), Tixier (1973; Anatom) .
Camptochaete prolongata Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Ana tom) .
Hookeriaceae
Callicostella campbelliana (Hampe) A. Jaeger- Brotherus & Watts ( 1915; New
Hebrides) .
Hookeria campbelliana Hampe- Hampe (1874, sp. nov. ; Anatom).
Callicostella frateri Broth. & Watts - Brotherus & Watts (1915 , sp. nov. ;Paama).
Callicostella oblongifolia (Sull.) A. Jaeger- Brotherus & Watts ( 1915; Espiritu
Santa) .
Callicostella papillata (Mont.) Mitt. - Matsui & Iwatsuki (1993; Efate, Espiritu
Santa) .
Chaetomitrium aneitense Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Anatom).
Chaetomitrium aneitense Broth. & Watts f. brachyphylla Dixon - Dixon ( 1948,
f. nov. ; Anatom).
Chaetomitrium hebridense Dixon - Dixon (1948, sp. nov.; Efate).
Chaetomitrium tahitense (Sull.) Mitt. var. deplanchei (Besch.) Wijk & Margad.
Chaetomitrium geheebii Broth. - Brotherus & Watts ( 1915; Espiritu Santa) .
Chaetomitrium torquescens Bosch. & Sande Lac. - Tixier ( 1973; New Hebrides).
Distichophyllum capillatum Mitt. - Mitten (1882, sp. nov. - Anatom) .
Distichophyllum graeffeanum (M till. Hal.) Broth. - Matsui & Iwatsuki ( 1993;
Espiritu San to).
Distichophyllum mittenii Bosch & Sande Lac.- Brotherus & Watts (1915;
Anatom) , Matsui & lwatsuki (1993; Espiritu Santa).
Distichophyllum vitianum (Sull.) Mitt. - Brotherus & Watts (1915 ; Futuna) .

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Masanobu HIGUCHI

Hypopterygiaceae
Hypopterygium bowiei Broth. & Watts- Brotherus & Watts (1915, sp. nov.;
Espiritu Santa).
Hypopterygium micholitzii Paris- Brotherus & Watts (1915; Espiritu Santa) .
Hypopterygium tenellum Mull. Hal.
Hypopterygium neo-caledonicum Besch.- Brotherus & Watts (1915;
Anatom, Futuna), Tixier (1972; Anatom). cf. Schultze-Motel (1973).
Lopidium struthiopteris (Brid.) M. Fleisch.
Hypopterygiumjavanicum (Hampe) A. Jaeger-Brotherus & Watts (1915;
Anatom).
Thuidiaceae
Herpetineuron toccoae (Sull. & Lesq.) Cardot - Brotherus & Watts (1915; Epi),
Theriot (1938; Ambrym).
Pelekium velatum Mitt. - Brotherus & Watts (1915; Epi, Espiritu Santa), Tixier(1973; Espiritu Santa).
Thuidium campbellianum (Hampe) A. Jaeger- Mitten (1882; New Hebrides),Brotherus & Watts (1915; New Hebrides).
Cryto-hypnum campbellianum Hampe- Hampe (1874, sp. nov.; Erromango).
Thuidium cymbifolium (Dozy & Molk.) Dozy & Molk. - Brotherus & Watts(1915; Anatom, Espiritu Santa), Tixier (1974; Espiritu Santa, Efate).
Thuidium ramentosum (Mitt.) Mitt. - Brotherus & Watts (1915; Anatom,
Epi).
Thuidium glaucinoides Broth.- Brotherus & Watts (1915; Espiritu Santa),
Fleischer (1923; New Hebrides), Theriot (1938; Futuna), Tixier (1974;
Espiritu Santa).
Thuidium glacinum (Mitt.) Bosch & Sande Lac. - Brotherus & Watts (1915 ;Anatom, Epi, Espiritu Santa), Theriot (1938; Tanna), Tixier (1973 ;Ana tom, Espiritu San to).
Thuidium meyenianum (Hampe) Dozy & Molk. - Htirlimann (1965; New Hebrides), Tixier (1973; Espiritu Santa), Tixier (1974; Mallicolo).
Thuidium plumulosum (Dozy & Molk.) Dozy & Molk. - Brotherus & Watts(1915; Espiritu Santa), Theriot (1938; Aniwa, Pentecost, Tanna).
Brachytheciaceae
Eurhynchium savatieri Besch.
Oxyrrhynchium savatieri (Besch.) Broth. -Theriot (1938; Tanna) .

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127

Rhynchostegiella menadense (San de Lac.) E. B. Bartram
Rhynchostegium menadense (Sande Lac.) A. Jaeger - Brotherus & Watts(1915; Epi).
Rhynchostegiella vitiensis Dixon - Theriot ( 1938; Efate, Tanna).
Rhynchostegium javanicum (Bel.) Besch. - Brotherus & Watts (1915; Espiritu
Santo).
Rhynchostegium oblongifolium Broth. & Watts- - rotherus & Watts (1915, sp.
nov. ; Futuna, Espiritu Santo).
Rhynchostegium trachypelma (Mtill. Hal.) A. Jaeger
Rhynchostegium javanicum (Bel.) Besch. var. majus Besch. - Tixier ( 1974;
Espiritu San to, Mallicolo ) .
Entodontaceae
Entodon aneitensis Mitt. - Mitten ( 1882, sp. nov. ; Ana tom).
Entodon pallidus Mitt. - Mitten (1873 , sp. nov.; Anatom) , Brotherus & Watts(1915; Epi, Futuna).
Entodon plicatiformis Dixon - Dixon (1948, sp. nov.; Efate) .
Plagiotheciopsis oblonga (Broth.) Broth. - Brotherus & Watts (1915; New
Hebrides).
Sematophyllaceae
Acanthorrhynchium papillatum (Harv.) M. Fleisch. - Theriot (1938; Erromango) .
Taxithelium papillatum (Harv.) Broth. - Brotherus & Watts (1915;
Anatom) .
Inoue & Higuchi ( 1994) reported the chromosome number of Trichosteleum
gunnii based on the plants (Higuchi 21141 , 21145) from Espiritu Santo Island.
However these are the misidentification of Acanthorrhynchium papillatum. The
chromosome number, n = 11, for A. papillatum is reported for the first time.
Acanthorrhynchium substigmosum (Mull. Hal.) M. Fleisch. - Theriot ( 1938;
Futuna, Pentecost).
Taxithelium substigmosum (Mtill. Hal.) Broth. - Brotherus & Watts (1915 ;Espiri tu San to).
Acroporium hamulatum (M. Fleisch.) M. Fleisch. - Tixier ( 1974; Espiritu
Santo) .
Acroporium hermaphroditum (Mull. Hal.) M . Fleisch. -Theriot (1938; Aoba) .
Acroporium oxyporum (San de Lac.) M. Fleisch. - Dixon ( 1948; Ana tom,
Efate) .
Sematophyllum serricalyx Broth. & Watts - Brotherus & Watts ( 1915, sp.

128

Masanobu HI GUC HI

nov.; Anatom, Epi, Futuna, Espiritu Santa).
Acroporium sigmatodontium (Mtill. Hal.) M. Fleisch. - Theriot (1938; Futuna) .
Sematophyllum sigmatodontium (M till. Hal. ) A . Jaeger - Brotherus &Watts (1915; Espiritu Santa) .
Acroporium turgidum (Mitt.) Mitt. - Tixier ( 1973; Tanna), Tixier ( 1974;
Erromango, Espritu Santa) .
Sematophyllum turgidum (Mitt.) Mitt. - Brotherus & Watts (1915;
Anatom) .
Aptychella glabrifolia (Broth. & Watts) Broth.
Sematophyllum glabrifolium Broth. & Watts - Brotherus & Watts (1915,
sp. nov. ; Anatom) .
Clastobryum hebridense Broth. - Brotherus & Watts ( 1915, sp. nov.; An atom) .
Rhaphidostichum bunodicarpum (M till. Hal.) M . Fleisch.
Sematophyllum bunodiocarpum (Mtill. Hal.) Broth. - Brotherus & Watts(1915 ; Anatom, Epi).
Sematophyllum microcladiellum M . Fleisch. - Tixier (1974; Erromango) .
Taxithelium annandii Broth. & Watts - Brotherus & Watts (1915 , sp. nov. ;Espiritu San to).
Trichosteleum gunnii Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Anatom) .
Trichosteleum hamatum (Dozy & Molk.) A . Jaeger var. semi-mammillosum(Mtill. Hal.) Paris - Brotherus & Watts (1915; Anatom) , Tixier (1973;
Anatom, Tanna) .
Trichosteleum hebridarum W . Schultze-Motel
Trichosteleum subtile Broth. & Watts, horn. illeg. - Brotherus & Watts(1915, sp. nov.; Paama) .
Trichosteleum neocaledonicum Ther. - Dixon ( 1948; Erromango) .
Trichosteleum pickeringii (Sull.) A . Jaeger - Brotherus & Watts ( 1915;
Anatom, Espiritu Santa), Inoue & Higuchi (1994; Espiritu Santa) .
Trichosteleum subrhinophyllum (Mtill. Hal.) A. Jaeger - Theriot (1938 ; Vanua
Lava).
Trismegistia complanatula (Mtill. Hal.) Mtill. Hal. - Brotherus & Watts (1915;
Anatom, Espiritu Santa) , Tixier (1972; Anatom) , Tixier (1973; Anatom) .
Trismegistia pedunculata (Mitt.) Broth. - Brotherus & Watts (1915; Anatom) .
Acanthocladium pedunculatum Mitt. - Mitten (1882, sp. nov.; Anatom) .
Trismegistia ridida (Mitt.) Broth.
Sematophyllum rigidum (Mitt.) Mitt. - Mitten ( 1873; Anatom).
Wijika tanytricha (Mont.) H. A. Crum
Acanthocladium tanytrichum (Mont.) Broth. - Tixier (1972; Anatom).

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129

Hypnaceae
Ectropothecium aneitense Broth. & Watts - Brotherus & Watts ( 1915, sp. nov.;
Ana tom).
Ectropothecium aubertii Ther.- Theriot (1938, sp. nov.; Vanua Lava), Tixier(1973; Tanna), Tixier (1974; Erromango).
Ectropothecium bowiei Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Espiritu San to).
Ectropothecium buitenzorgii (Bel.) Mitt.
Ectropothecium brachyphyllum Broth. & Watts - Brotherus & Watts (1915,
sp. nov.; Anatom) . cf. Schultze-Motel (1973) .
Ectropothecium gunnii Broth. & Watts- Brotherus & Watts (1915, sp. nov.;
Anatom), Theriot (1938; Vanua Lava), Dixon (1948; Anatom).
Ectropothecium ichnotocladum (M till. Hal.) A. Jaeger- Tixier ( 1974; Espiritu
Santo).
Ectropothecium laticuspis Broth. - Theriot (1938; Vanua Lava).
Ectropothecium malacoblastum (M till. Hal.) A. Jaeger - Theriot ( 1938;
Futuna).
Ectropothecium micholitzii Broth. - Brotherus & Watts (1915; Epi, Futuna),
Theriot (1938; Pentecost).
Ectropothecium pacificum Mitt. - Mitten (1868, sp. nov.; Erromango), Muller(1874; Erromango), Brotherus & Watts (1915; Anatom), lnoue & Higuchi( 1994; Espiritu San to) .
Ectropothecium percomplanatum Broth. - Brotherus & Watts (1915; Epi,
Espiritu San to).
Ectropothecium sodale (Sull.) Mitt. - Brotherus & Watts (1915; Espiritu
San to), Inoue & Higuchi ( 1994; Espiritu San to) .
Ectropothecium tannaense Ther.- Theriot (1938, sp. nov.; Tanna), Tixier(1974; Espiritu Santo).
Ectropothecium tannaense Ther. f. inundata Ther. - Theriot ( 1938, f. nov.;
Tanna).
Ectropothecium zollingeri (M till. Hal.) A. Jaeger
Glossadelphus obscurum (Broth. & Paris) Broth. - Tixier (1974; Espiritu
Santo). cf. Higuchi & Iwatsuki (1994).
lsopterygium albescens (Hook.) A. Jaeger - Tixier ( 1974; Efate ), Inoue &Higuchi (1994; Espiritu Santo).
Isopterygium gunnii Broth. & Watts - Brotherus & Watts (1915, sp. nov.;
Futuna).
Isopterygium minutirameum (M till. Hal.) A. Jaeger
Isopterygium austro-pusillum (M till. Hal.) A. Jaeger - Theriot ( 1938;
Tanna).

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Masanobu HIG UC HI

Isopterygium sarasinii Ther. - Theriot (1938; Futuna) .
Mittenothamnium nano-operculatum (Ther.) W. Schultze-Motel
Microthamnium nano-operculatum Ther. - Theriot (1938, sp. nov.; Pentecost).
Pseudotaxiphyllum pohliaecarpum (Sull. & Lesq.) Z. Iwats.
Isopterygium pohliaecarpum (Sull & Lesq.) A. Jaeger -Tixier (1973;
Espiritu Santo).
Taxiphyllum taxirameum (Mitt.) M. Fleisch. - Inoue & Higuchi ( 1994; Espiritu
Santo).
Isopterygium taxirameum (Mitt.) A. Jaeger - Brotherus & Watts (1915;
Epi, Futuna, Espiritu San to).
Taxiphyllum taxirameum (Mitt.) M. Fleisch. f. gracilis Ther. - Theriot ( 1938;
Tanna) .
Trachythecium hebridense (Dixon) W. Schultze-Motel
Ectropothecium hebridense Dixon - Dixon (1948, sp. nov. ; Anatom, Efate,
Erromango) .
Vesicularia aperta (Sull.) Ther. -Theriot (1938; Futuna), Tixier (1974; Mallicolo).
Vesicularia inflectens (Brid.) Miill. Hal. - Brotherus & Watts (1915; Epi),
Theriot (1938; Futuna) , Inoue & Higuchi (1994; Espiritu Santo) .
Vesicularia subinflectens Miill. Hal. - Tixier ( 1974; Espiritu San to).
Vesicularia perviridis ( Angstr.) M till. Hal. - Dixon (1948; Efate).
Vesicularia pinnatula M till. Hal. ex Broth. - Inoue & Higuchi ( 1994; Espiritu
Santo).
Polytrichaceae
Pogonatum neesii (Miill. Hal.) Dozy - Hyvonen (1989; Vanuatu) .
Pogonatum circinatum Besch. - Brotherus & Watts (1915 ; Anatom), Tixier(1973 ; Espiritu Santo) , Tixier (1974; Espiritu Santo) . cf. Hyvonen (1989).
Polytrichum collieanum Mtill. Hal. - MUller (1897, sp. nov.; Anatom). cf.
Hyvonen (1989).
Acknowledgments
I wish thank to Dr. T . Yamaguchi of Hiroshima University for sending mesome literature and the curator of the Botanical Museum, University of Helsinki
for the loan of specimens, and Dr. W. R. Buck of the New York Botanical
Garden for reading the manuscript and giving some comments.

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131

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473- 491.

Appendix

Synonyms
Acanthocladium pedunculatum = Trismegistia pedunculata
Acanthocladium tanytrichum = Wijika tanytricha
Barbula aneitensis = Hydrogonium aneitense
Braithwaitea arborescens = Hypnodendron subspininervium
Calymperes lorifolium = Calymperes serratum
Camptochaete deflexa = Camptochaete arbuscula var. deftexa

A Checklist of the mosses of Vanuatu

133

Campylopus blumii = Campylopus umbellatus
Campylopus richardii = Campylopus umbellatus
Chaetomitrium geheebii = Chaetomitrium tahitense
Cryto-hypnum campbellianum = Thuidium campbellianum
Ectropothecium brachyphyllum = Ectropothecium buitenzorgii
Ectropothecium hebridense = Trachythecium hebridense
Endotrichella campbelliana = Garovaglia elegans
Endotrichella elegans = Garovaglia elegans
Endotrichella samoana = Garovaglia elegans
Euptychium assimile = Euptychium setigerum subsp. setigerum
Euptychium austro-caledonicum = Euptychium setigerum subsp. austro-caledonicum
Euptychium gunnii = Euptychium setigerum subsp. setigerum
Fissidens subscabrisetus = Fissidens hollianus
Glossadelphus obscurum = Ectropothecium zollingeri
Homaliodendron dendroides = Homaliodendron fiabellatum
Hookeria campbelliana = Callicostella campbelliana
Hydrogonium comosum = Hydrogonium arcuatum
Hypnodendron rigidum = Pterobryella rigida
Hypnodendron arcuatum = Hypnodendron vitiense subsp. vitiense
Hypopterygium javanicum = Lopidium struthiopteris
Hypopterygium neo-caledonicum = Hypopterygium tenellum
Isopterygium austro-pusillum = Isopterygium minutirameum
Isopterygium pohliaecarpum = Pseudotaxiphyllum pohliaecarpum
Isopterygium taxirameum = Taxiphyllum taxirameum
Leucobryum stenophyllum = Leucobryum chlorophyllosum
Microthamnium nano-operculatum = Mittenothamnium nano-operculatum
Mniodendron densirameum = Hypnodendron dendroides
Mniodendron hellwigii = Hypnodendron dendroides
Mniodendron milnei = Hypnodendron milnei subsp. milnei
Mnium succulentum = Plagiomnium succulentum
Myurium purpuratum = Myurium rufescens subsp. purpuratum
Octoblepharum sanctum = Leuobryum sanctum
Oedicladium purpuratum = Myurium rufescens subsp. purpuratum
Oxyrrhynchium savatieri = Eurhynchium savatieri
Philonotis laxissima = Philonotis hastata
Pilopogon blumii = Campylopus umbellatus
Pogonatum circinatum = Pogonatum neesii
Polytrichum collieanum = Pogonatum neesii
Porotrichum aneitense = Thamnobryum aneitense
Pterobryella vagapensis = Pterobryella rigida

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Rhizogonium medium = Pyrrhobryum medium
Rhizogonium spiniforme = Pyrrhobryum spiniforme
Rhynchostegium javanicum var. majus = Rhynchostegium trachypelma
Rhynchostegium menadense = Rhynchostegiella menadense
Sematophyllum bunodiocarpum = Rhaphidostichum bunodicarpum
Sematophyllum glabrifolium = Aptychella glabrifolia
Sematophyllum rigidum = Trismegistia ridida
Sematophyllum serricalyx = Acroporium oxyporum
Sematophyllum sigmatodontium = Acroporium sigmatodontium
Sematophyllum turgidum = Acroporium turgidum
Symphysodon micholitzii = Symphysodon neckeroides
Synodontia aneitensis = Dicnemon aneitensis
Syrrhopodon aneitensis = Syrrhopodon japonicus
Syrrhopodon constrictus = Mitthyridium constrictum
Syrrhopodon fasciculatus = Mitthyridium fasciculatum
Syrrhopodon grae.ffeanus = Syrrhopodon laevigatus
Syrrhopodon lilliei = Syrrhopodon flammeonervis subsp. lilliei
Syrrhopodon tenuinervis = Syrrhopodon japonicus
Taxithelium papillatum = Acanthorrhynchium papillatum
Taxithelium substigmosum = Acanthorrhynchium substigmosum
Thamnium aneitense = Thamnobryum aneitense
Thamnium leichhardtii = Camptochaete leichhardtii
Thuidium ramentosum = Thuidium cymbifolium
Thyridium cuspidatum = Mitthyridium cuspidatum
Thyridium fasciculatum = Mitthyridium fasciculatum
Thyridium obtusifolium = Mitthyridium obtusifolium
Trichosteleum subtile = Trichosteleum hebridarum
Vesicularia subinflectens = Vesicularia inflectens