Bull. Natl. Mus. Nat. Sci., Ser. B, 36(3), pp. 91–105, August 22, 2010

Moss Flora of Lanyu, Taiwan
Masanobu Higuchi1,*, Naoki Nishimura2, Kuei-Yu Yao3 and Shan-Hsiung Lin4
1

Department of Botany, National Museum of Nature and Science,
Amakubo 4–1–1, Tsukuba, 305–0005 Japan
2
Botanical Garden, Okayama University of Science,
Ridai-cho 1–1, Okayama, 700–0005 Japan
3
Endemic Species Research Institute, Council of Agriculture,
1 Ming-Shen E. Road, Chi-Chi, Nantou, Taiwan
4Department of Life Science, Tunghai University,
No 181, Sec. 3, Taichung-Kan Road, Taichung, Taiwan*
E-mail: higuchi@kahaku.go.jp(Received 21 May 2010; accepted 23 June 2010)

Abstract The moss flora of Lanyu, Taiwan, was investigated in 2006. The mosses recognizedcomprise 23 families, 39 genera and 62 species. Twenty seven species are new additions to themoss flora of Lanyu. Calymperes moluccense, Ectropothecium sodale and Radulina elegantissima
are reported as new to the moss flora of Taiwan. The moss flora of Lanyu comprising 29 families,
60 genera and 123 species is characterized by richness of tropical species as compared with that of
the main island. For each species recognized here, locality, substrate, specimen number and taxonomic note are provided.
Key words : Botel Tobago, Lanyu, mosses, Taiwan.

Introduction
This study deals with the moss flora of Lanyu,
Taiwan. In 2006 we made field researches andcollected bryophytes mainly from the areas of
Mt. Hongtoushan and Datienchi. Taiwan consistsof the main island and many smaller islands inthe northwest region of the Pacific Ocean and locates 400 km between 21º50 and 25º20N latitude. Lanyu is located off the southeastern coastof the main island and is the second largest island(45 km2). This is a volcanic island and the geology of the island is mainly andesite conglomerate.
Most parts of the island are mountainous andsteep. The tallest mountain is Mt. Hongtoushan
at 552 m above sea level. The climate is subtropical and it is characterized by strong wind andplenty of precipitation. The natural vegetation remains only around higher mountainous regions.
Lanyu has been also called Orchid Island or
known as Tabaco Xima, Botel Tobago, Hong-touyu or Kôtôsho.
The mosses of Lanyu have been reported by
Horikawa (1935), Noguchi (1937, 1939), Herzog
and Noguchi (1955), Yang and Lee (1964), Wang
(1969), Shin (1971, 1973). However, no bryophyte flora of Lanyu has been compiled. The purpose of this study is to investigate the moss floraof Lanyu and compile it based on the specimens
collected and references.
Materials and Methods
Field studies were carried out in November
2006 and a total of ca. 130 specimens were collected. The specimens cited here are deposited in
the herbarium of the Department of Botany,
National Museum of Nature and Science (TNS) andduplicates in the herbarium of the Department ofLife Science, Tunghai University, Taichung(TUNG).
The sites investigated are divided as follows.

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I: Mt. Hongtoushan, 150–552 m alt., 22°03N,
121°32E, November 9, 2006. (Fig. 1: A, C)
II: Datienchi, 180–360 m alt., 22°01N, 121°34E,
November 10, 2006. (Fig. 1: B)
III: Central Cross Road, 170 m alt., 22°01N,
121°33E, November 10, 2006
IV: Hongtou, 10 m alt., 22°01N, 121°33E,
November 10, 2006

V: Xiaotienchi, 280 m alt., 22°04N, 121°31E,
November 11, 2006
Results and Discussion
The mosses recognized in this study comprise
23 families, 39 genera and 62 species. Twenty
seven species are new additions to the moss flora

Fig. 1. Photographs showing the area investigated and some species growing there. A. Southern slope of Mt.
Hongtoushan. B. Datienchi. C. Forest floor near the summit of Mt. Hongtoushan. D. Leucobryum bowringii
growing on tree-trunk mixed with Leucophanes angustifolium. E. Garovaglia elegans growing on low tree. F.
Distichophyllum mittenii growing on tree-trunk mixed with Hookeria acutifolia.

Moss Flora of Lanyu, Taiwan

of Lanyu. Among them Calymperes moluccense
Schwägr., Ectropothecium sodale (Sull.) Mitt.
and Radulina elegantissima (M.Fleisch.) W.R.
Buck & B.C. Tan are new to Taiwan. In total themoss flora of Lanyu comprise 29 families, 60
genera and 123 species.
In Taiwan, the moss flora of the lowlands is
related to that of the Philippines, southern China
and Southeast Asia, and has suffered much morehuman activities (Higuchi and Lin, 2004). The
affinities of the flora and fauna of Lanyu and thePhilippines have been indicated (e.g., Kano,
1931a, 1931b; Kanehira, 1935).
The 123 species known from Lanyu mainly
consist of tropical species, but also include several other elements of distribution, such as East
Asiatic (e.g., Eurhynchium savatieri, Neckeropsis
nitidula, etc.), cosmopolitan (e.g., Fissidens taxifolius, Weissia controversa, etc.) or circumboreal
species (e.g., Eurhynchium hians, Ditrichum pallidum, etc.). They are classified roughly into thefollowing floristic groups: palaeotropic species(74 spp., 60%), East Asiatic species (21 spp.,17%), pantropical species (12 spp., 9%), cosmopolitan (6 spp., 5%), circumboreal species (5spp., 4%) and others (5 spp., 4%). While, themosses of Taiwan consists of East Asiatic species(44%), palaeotropic species (27%), circumboreal
species (12%), cosmopolitan (7%), pantropical
species (2%) and others (8%) (Higuchi and Lin,
2004). The result shows that the moss flora of
Lanyu is characterized by richness of tropical
species as compared with that of Taiwan.
Enumeration of the moss species
known from Lanyu
The families, genera and species are arrangedalphabetically. The generic position follows
Goffinet et al. (2009). In the following enumeration an asterisk (*) preceding a species indicates“new to Lanyu.” The species that were recognized in this study are in italics, and the species
that are cited from the references are in Roman
type. Fissidens splachnobryoides Broth. f. subbrachyneuron (Thér. & P.de la Varde) Herzog,

93

which was reported from the island by Herzog
and Noguchi (1955), is not available here because of the lack of information. Each species isreferred by collecting site (I–V), substrate andspecimen number, and some species by taxonomic note.
Anomodontaceae
Herpetineuron toccoae (Sull. & Lesq.) Cardot,
Beih. Bot. Centralbl., Abt. 2, 19(2): 127 (1905).
Note. This species has been reported from theisland by Shin (1971, 1973).
Bartramiaceae*Philonotis hastata (Duby) Wijk & Marg.,
Taxon 8: 74 (1959).
Specimen examined. IV, on stone wall
(Higuchi 48398Brachymenium exile, Hyophila
involuta).
Note. This species is new to Lanyu.
Philonotis lancifolia Mitt., J. Proc. Linn. Soc.,
Bot., 8: 151 (1864).
Note. This species has been reported from theisland by Shin (1973).
Philonotis mollis (Dozy & Molk.) Mitt., J. Proc.
Linn. Soc., Bot., Suppl. 1: 60 (1859).
Note. This species has been reported from theisland by Wang (1969).
Philonotis thwaitesii Mitt., J. Proc. Linn. Soc.,
Bot., Suppl. 1: 60 (1859).
Note. This species has been reported from theisland by Shin (1973).
Philonotis turneriana (Schwägr.) Mitt., J. Proc.
Linn. Soc., Bot., Suppl. 1: 62 (1859).
Specimen examined. V, on soil (Higuchi
48409).
Note. This species has been reported from theisland by Shin (1973).

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Masanobu Higuchi et al.

Brachytheciaceae
Eurhynchium celebicum (Sande Lac.) E.B.Bartram, Bernice P. Bishop Mus. Bull., 101: 217(1933).
Note. This species has been reported from theisland by Wang (1969).
Eurhynchium hians (Hedw.) Sande Lac., Ann.
Mus. Bot. Lugduno-Batavi, 2: 299 (1866).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as
Oxyrrhynchium hians).
*Eurhynchium savatieri Schimp. ex Besch.,Ann. Sci. Nat. Bot., sér. 7, 17: 378 (1893).
Specimen examined. V, on soil (Higuchi
48407).
Note. This species is new to Lanyu.
Oxyrrhynchium vagans (A.Jaeger) Ignatov &Huttunen, Arctoa, 11: 273 (2002).
Note. This species has been reported from theisland by Wang (1969 as Rhynchostegium
vagans).
*Rhynchostegium contractum Cardot, Bull. Soc.
Bot. Genève, 4: 381 (1912).
Specimen examined. I, on basal part of tree
(Higuchi 48300).
Note. This species is new to Lanyu.
Rhynchostegium riparioides (Hedw.) Cardot,
Bull. Soc. Bot. France, 60: 231 (1913).
Note. This species has been reported from theisland by Noguchi (1939 as Platyhypnidium
schottmulleri).
Bryaceae
Brachymenium exile (Dozy & Molk.) Bosch &Sande Lac., Bryol. Jav., 1: 139 (1860).
Specimens examined. IV, on stone-wall(Higuchi 48398Hyophila involuta, Philonotis
hastata, 48399).
Note. This species has been reported from theisland by Shin (1973).
Bryum apiculatum Schwägr., Sp. Musc. Frond.,
Suppl. 1, 2: 102 (1816).
Note. This species has been reported from theisland by Shin (1973 as B. nitense).
Bryum atrovirens Brid., Muscol. Recent., 2(3):48 (1803).
Note. This species has been reported from theisland by Shin (1973 as B. erythrocarpum).
Bryum cellulare Hook., Sp. Musc. Frond.,
Suppl. 3, 1(1): 214 (1827).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as B. japonense) and Shin (1973).
Pohlia flexuosa Harv., Icon. Pl. 1: pl. 19: f. 5
(1836).
Note. This species has been reported from theisland by Shin (1973).
Calymperaceae*Calymperes afzelii Sw., Jahrb. Gewächsk., 1: 3
(1818).
Specimens examined. I, on tree-trunk (Higuchi
48317Fissidens taxifolius, 48399). II, on rockcliff (Higuchi 48363Barbula subcomosa).
Note. This species is new to Lanyu.
*Calymperes graeffeanum Müll.Hal., J. Mus.
Godeffroy, 3(6): 64 (1874).
Specimens examined. I, on tree-trunk (Higuchi
48296). II, on tree-trunk (Higuchi 48365Haplohymenium pseudo-triste).
Note. This species is new to Lanyu.
*Calymperes moluccense Schwägr., Sp. Musc.
Frond., Suppl. 2, 1: 99 (1824). (Fig. 2)
Specimen examined. II, on basal part of treetrunk (Higuchi 48368).
Note. This species is new to Taiwan. Calymperes moluccense is characterized by thickwalled laminal cells around shoulders of hyalineMoss Flora of Lanyu, Taiwan

Fig. 2. Calymperes moluccense Schwägr. (Higuchi 48368). A. Plants. B. Nongemmiferous leaf. C, D. Gemmiferous leaves. E. Lower part of gemmiferous leaf showing cancellinae and laminal cells. F. Leaf apex of gemmiferous leaf with gemmae. G. Gemmae. H, I. Cross-sections of nongemmiferous leaf (H: middle part, I:
lower part). Scale in A1 mm. Scale bars in B–D1 mm; E–G0.5 mm; H & I 0.2 mm.

95

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Masanobu Higuchi et al.

base (Fig. 2: E) and broad “collar” of the chlorophyllose lamina at apices of gemmiferous leaves(Fig. 2: F) (Ellis and Tan, 1999). This species is
distributed widespread in the Indo-Pacific regions, and Lanyu is the northern limit of the distribution.

ma molle, 48358Exostratum blumii). II, on
basal part of tree (Higuchi 48387).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as L. octoblepharioides), Wang (1969 as L. octoblepharioides) and Shin (1973 as L. octoblepharioides).

Calymperes serratum A.Braun ex Müll.Hal.,
Syn. Musc. Frond., 1: 527 (1849).
Specimen examined. I, on tree-trunk (Higuchi
48326).
Note. This species has been reported from theisland by Noguchi (1939 as C. cristatum).

Leucophanes candidum (Schwägr.) Lindb.,
Öfvers. Förh. Kongl. Svenska Vetensk.-Akad.,
21: 602 (1865).
Note. This species has been reported from theisland by Shin (1973).

Calymperes strictifolium (Mitt.) G.Roth, Hedwigia 51: 127 (1911).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as C. tuberculosum) and Wang (1969).
Calymperes tahitense (Sull.) Mitt., J. Linn.
Soc., Bot., 10: 172 (1868).
Note. This species has been reported from theisland by Herzog and Noguchi (1955).
Calymperes tenerum Müll.Hal., Linnaea 37: 174(1872).
Note. This species has been reported from theisland by Herzog and Noguchi (1955) and Wang
(1969).
Exostratum blumii (Nees ex Hampe) L.T.Ellis,
Lindbergia, 11: 25 (1985).
Specimens examined. I, on decaying log(Higuchi 48342, 48358Leucophanes angustifolium).
Note. This species has been reported from theisland by Horikawa (1935 as Exodictyon blumii),
Herzog and Noguchi (1955 as Exodictyon blumii)
and Shin (1973 as Exodictyon blumii).
Leucophanes angustifolium Renauld & Cardot,
Rev. Bot. Bull. Mens., 9: 395 (1891).
Specimens examined. I, on root (Higuchi
48298); on tree-trunk (Higuchi 48308,
48316Leucobryum bowringii, 48323Leucolo-

Leucophanes glaucum (Schwägr.) Mitt., J. Proc.
Linn. Soc., Bot., Suppl. 1: 25 (1859).
Specimen examined. II, on root (Higuchi
48375).
Note. This species has been reported from theisland by Shin (1973 as L. albescens).
*Mitthyridium flavum (Müll.Hal.) H.Rob., Phytologia, 32: 434 (1975).
Specimens examined. II, on root (Higuchi
48366); on basal part of tree (Nishimura 12079).
Note. This species is new to Lanyu.
Daltoniaceae
Calyptrochaeta japonica (Cardot & Thér.) Z.
Iwats. & Nog., J. Hattori Bot. Lab., 46: 236(1979).
Note. This species has been reported from theisland by Yang and Lee (1964 as Eriopus mollis).
Distichophyllum cuspidatum (Dozy & Molk.)
Dozy & Molk., Musci Frond. Ined. Archip. Ind.,
4: 101 (1846).
Specimen examined. I, on tree-trunk (Higuchi
48340).
Note. This species has been reported from theisland by Noguchi (1937).
Distichophyllum jungermannioides (Müll.Hal.)
Bosch & Sande Lac., Bryol. Jav. 2: 22 (1861).
PAL (As 2, 4)Note. This species has been reported from theMoss Flora of Lanyu, Taiwan

island by Horikawa (1935), Herzog and Noguchi(1955) and Yang and Lee (1964).
*Distichophyllum maibarae Besch., J. Bot.
(Morot), 13: 40 (1899).
Specimens examined. I, on boulder (Higuchi
48319, 48354); on decaying log (Higuchi 48325
Trichosteleum boschii); on tree-trunk (Higuchi
48344, 48353). II, on soil (Higuchi 48378,
48379).
Note. This species is new to Lanyu.
Distichophyllum mittenii Bosch & Sande Lac.,
Bryol. Jav., 2: 25 (1861).
Specimens examined. I, on root (Higuchi
48322Hookeria acutifolia); on decaying log(Higuchi 48351Hookeria acutifolia).
Note. This species has been reported from theisland by Noguchi (1939).
Distichophyllum osterwaldii M.Fleisch., Musci
Fl. Buitenzorg, 3: 994 (1908).
Note. This species has been reported from theisland by Yang and Lee (1964).
Dicranaceae
Leucoloma molle (Müll.Hal.) Mitt., J. Proc.
Linn. Soc., Bot., Suppl. 1: 13 (1859).
Specimens examined. I, on tree-trunk (Higuchi
48323Leucophanes angustifolium, 48327,
48335, 48356); on branch (Higuchi 48341
Hypnodendron reinwardtii).
Note. This species has been reported from theisland by Noguchi (1939).
Trematodon longicollis Michx., Fl. Bor.-Amer.
2: 289 (1803).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as T.
drepanellus, T. flaccidisetus and T. paucifolius).
Ditrichaceae*Ditrichum difficile (Duby) M.Fleisch., Musci
Fl. Buitenzorg, 1: 300 (1904).

97

Specimen examined. II, on soil at roadsidebank (Higuchi 48393).
Note. This species is new to Lanyu.
Ditrichum pallidum (Hedw.) Hampe, Flora, 50:
182 (1867).
Note. This species has been reported from theisland by Wang (1969).
Fissidentaceae
Fissidens bryoides Hedw., Sp. Mus. Frond., 153(1801).
Note. This species has been reported from theisland by Shin (1971, 1973).
Fissidens ceylonensis Dozy & Molk., Ann. Sci.
Nat. Bot., sér. 3, 2: 304 (1844).
Note. This species has been reported from theisland by Shin (1971, 1973).
Fissidens crispulus Brid., Muscol. Recent.
Suppl., 4: 187 (1819 [1818]).
Specimens examined. II, on soil (Higuchi
48391). III, on soil (Higuchi 48396).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as F. sakourae) and Shin (1973 as F. incrassatus).
Fissidens gardneri Mitt., J. Linn. Soc., Bot., 12:593 (1869).
Note. This species has been reported from theisland by Shin (1971, 1973 as F. garberi).
*Fissidens hollianus Dozy & Molk., Bryol. Jav.,
1: 4 (1855).
Specimen examined. II, on boulder (Higuchi
48372Taxithelium parvulum).
Note. This species is new to Lanyu.
Fissidens nobilis Griff., Calcutta J. Nat. Hist., 2:505 (1842).
Specimen examined. II, on soil (Higuchi
48369).
Note. This species has been reported from theisland by Noguchi (1939 as F. japonicus), Herzog

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Masanobu Higuchi et al.

and Noguchi (1955) and Wang (1969 as F. filicinus, F. nobilis).
*Fissidens oblongifolius Hook.f. & Wilson,
London J. Bot., 3: 547 (1844).
Specimen examined. II, on soil (Higuchi
48382).
Note. This species is new to Lanyu.
Fissidens taxifolius Hedw., Sp. Musc. Frond.,
155 (1801).
Specimen examined. I, on tree-trunk (Higuchi
48317Calymperes afzelii).
Note. This species has been reported from theisland by Shin (1971, 1973).
*Fissidens teysmannianus Dozy & Molk., Pl.
Jungh., 317 (1854).
Specimens examined. II, on soil (Higuchi
48361, 48370).
Note. This species is new to Lanyu.
Fissidens zollingeri Mont., Ann. Sci. Nat. Bot.,sér. 3, 4: 114 (1845).
Note. This species has been reported from theisland by Shin (1971, 1973 as F. xiphioides).
Hookeriaceae
Hookeria acutifolia Hook. & Grev., Edinburgh J.
Sci., 2: 225 (1825).
Specimen examined. I, on boulder (Higuchi
48310); on root (Higuchi 48321Homaliodendron microdendron, Vesicularia flaccida, Higuchi
48322Distichophyllum mittenii); on decayinglog (Higuchi 48351Distichophyllum mittenii);
on soil (Higuchi 48381); on basal part of tree
(Nishimura 12073).
Note. This species has been reported from theisland by Noguchi (1937), Herzog and Noguchi(1955), Yang and Lee (1964) as H. nipponensis
and Wang (1969).
Hypnaceae
Ectropothecium buitenzorgii (Bél.) Mitt., J.

Linn. Soc., Bot., 10: 180 (1868).
Note. This species has been reported from theisland by Herzog and Noguchi (1955) and Wang
(1969).
Ectropothecium dealbatum (Reinw. & Hornsch.)
A.Jaeger, Ber. S. Gall. Naturw. Ges., 1877–78:
264 (1880).
Specimen examined. I, on boulder (Nishimura
12061). II, on soil (Nishimura 12085).
Note. This species has been reported from theisland by Wang (1969).
*Ectropothecium moritzii (Müll.Hal.) A.Jaeger,
Ber. S. Gall. Naturw. Ges., 1877–78: 262 (1880).
Specimens examined. I, on tree-trunk (Higuchi
48336); on root (Nishimura 12065Vesicularia
montagnei, 12066, 12067, 12068, 12069, 12071,
12075, 12076, 12077). II, on humus (Higuchi
48374); on soil (Nishimura 12080); on rock
(Nishimura 12084).
Note. This species is new to Lanyu.
Ectropothecium perminutum Broth. ex E.B.
Bartram, Philipp. J. Sci., 68: 356 (1939).
Note. This species has been reported from theisland by Herzog and Noguchi (1955).
*Ectropothecium sodale (Sull.) Mitt., J. Linn.
Soc., Bot., 10: 180 (1868).
Specimen examined. II, on rock (Nishimura
12078).
Note. This species is new to Taiwan. Ectropothecium sodale is characterized by small
plants, strongly falcate and acuminate leaves andlaminal cells distinctly prorate at upper ends.
Ectropothecium zollingeri (Müll.Hal.) A.Jaeger,
Ber. S. Gall. Naturw. Ges., 1877–78: 272 (1880).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as Taxiphyllum formosanum Herzog & Nog. sp. nov.).
Taxiphyllum taxirameum (Mitt.) M.Fleisch.,Musci Fl. Buitenzorg, 4: 1435 (1923).
Specimens examined. III, on soil (Higuchi

Moss Flora of Lanyu, Taiwan

48397, Nishimura 12086).
Note. This species has been reported from theisland by Wang (1969).
*Vesicularia flaccida (Sull. & Lesq.) Z.Iwats., J.
Hattori Bot. Lab., 26: 70 (1963).
Specimens examined. I, on root (Higuchi
48321Homaliodendron microdendron, Hookeria acutifolia). II, on decaying log (Higuchi
48384Isopterygium minutirameum, Trichosteleum boschii).
Note. This species is new to Lanyu.
*Vesicularia montagnei (Bel.) Broth. in Engler& Prantl, Nat. Pfl., I(3): 1094 (1908).
Specimens examined. I, on decaying log(Higuchi 48297, 48309, 48350, 48360Callicostella papillata, Nishimura 12064, 12065
Ectropothecium moritzii); on boulder (Nishimura
12062). II, on soil (Higuchi 48389, Nishimura
12083). V, on soil (Higuchi 48406).
Note. This species is new to Lanyu.
Vesicularia reticulata (Dozy & Molk.) Broth. inEngler & Prantl, Nat. Pfl., I(3): 1094 (1908).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as Vesicularia shimadae).
Hypnodendraceae*Hypnodendron reinwardtii (Schwägr.) A.
Jaeger, Ber. S. Gall. Natur. Ges., 1877–78: 358(1880).
Specimens examined. I, on branch (Higuchi
48341Leucoloma molle); on decaying log(Higuchi 48343).
Note. This species is new to Lanyu.
Hypopterygiaceae*Hypopterygium flavolimbatum Müll.Hal., Syn.
Musc. Frond., 2: 10 (1850).
Specimen examined. I, on tree-trunk (Higuchi
48349).
Note. This species is new to Lanyu.

99

Leucobryaceae
Leucobryum bowringii Mitt., J. Proc. Linn. Soc.,
Bot., Suppl. 1: 26 (1859).
Specimens examined. I, on humus (Higuchi
48312); on root (Higuchi 48315); on tree-trunk(Higuchi 48316Leucophanes angustifolium);on decaying log (Higuchi 48324). II, on decayinglog (Higuchi 48385); on humus (Higuchi 48367).
Note. This species has been reported from theisland by Herzog and Noguchi (1955), Wang
(1969) and Shin (1973).
*Leucobryum javense (Brid.) Mitt., J. Proc.
Linn. Soc., Bot., Suppl. 1: 25 (1859).
Specimen examined. I, on basal part of tree
(Higuchi 48347).
Note. This species is new to Lanyu.
Leucomiaceae
Leucomium strumosum (Hornsch.) Mitt., J.
Linn. Soc., Bot., 12: 502 (1869).
Note. This species has been reported from theisland by Wang (1969 as Leucomium aneurodictyon).
Meteoriaceae
Aerobryopsis parisii (Cardot) Broth. in Engler &Prantl, Nat. Pfl., I (3): 820 (1906).
Specimen examined. I, on tree-trunk (Higuchi
48328).
Note. This species has been reported from theisland by Shin (1971, 1973).
Aerobryopsis wallichii (Brid.) M.Fleisch.,Musci Fl. Buitenzorg, 3: 789 (1908).
Note. This species has been reported from theisland by Wang (1969 as Aerobryopsis longissima).
Meteorium miquelianum (Müll.Hal.) M.
Fleisch. in Engler & Prantl, Nat. Pfl., I (3): 818(1906).
Note. This species has been reported from the100

Masanobu Higuchi et al.

island by Wang (1969).
Trachypodopsis serrulata (P.Beauv.) M.Fleisch.
var. crispatula (Hook.) Zanten, Blumea 9: 521(1959).
Note. This species has been reported from theisland by Wang (1969).
Mniaceae
Plagiomnium succulentum (Mitt.) T.J.Kop.,
Ann. Bot. Fenn., 5: 147 (1968).
Specimen examined. I, on boulder (Higuchi
48303).
Note. This species has been reported from theisland by Horikawa (1935 as Mnium formosicum), Herzog and Noguchi (1955 as Mnium
succulentum var. integrum), Wang (1969 as
Mnium succulentum var. integrum) and Shin
(1973 as Mnium succulentum var. integrum).
Neckeraceae
Himantocladium cyclophyllum (Müll.Hal.) M.
Fleisch., Musci Fl. Buitenzorg, 3: 887 (1908).
Note. This species has been reported from theisland by Noguchi (1937 as H. loriforme) and
Shin (1973 as H. loriforme).
Himantocladium formosicum Broth. & Yasuda,Rev. Bryol., 53: 2 (1926).
Note. This species has been reported from theisland by Shin (1973).
Himantocladium plumula (Nees) M.Fleisch.,Musci Fl. Buitenzorg, 3: 889 (1908).
Specimens examined. I, on tree-trunk (Higuchi
48313, 48330, Nishimura 12072).
Note. This species has been reported from theisland by Wang (1969).
Homaliadelphus targionianus (Mitt.) Dixon &P.de la Varde, Rev. Bryol. Lichénol., 4: 142(1932).
Note. This species has been reported from theisland by Herzog and Noguchi (1955).

Homaliodendron exiguum (Bosch & Sande
Lac.) M.Fleisch., Musci Fl. Buitenzorg, 3: 897(1908).
Specimen examined. I, on basal part of tree
(Higuchi 48301).
Note. This species has been reported from theisland by Noguchi (1937), Herzog and Noguchi(1955), Wang (1969) and Shin (1973).
Homaliodendron microdendron (Mont.) M.
Fleisch., Hedwigia, 45: 78 (1906).
Specimens examined. I, on root (Higuchi
48321Hookeria acutifolia, Vesicularia flaccida). II, on basal part of tree (Higuchi 483767).
Note. This species has been reported from theisland by Noguchi (1937), Wang (1969) and Shin
(1973).
Neckeropsis lepineana (Mont.) M.Fleisch.,Musci Fl. Buitenzorg, 3: 879 (1908).
Note. This species has been reported from theisland by Noguchi (1937), Herzog and Noguchi(1955), Wang (1969) and Shin (1973).
Neckeropsis nitidula (Mitt.) M.Fleisch., Musci
Fl. Buitenzorg, 3: 882 (1908).
Specimens examined. I, on boulder (Higuchi
48295). V, on boulder (Higuchi 48403).
Note. This species has been reported from theisland by Herzog and Noguchi (1955), Wang
(1969) and Shin (1973).
Neckeropsis obtusata (Mont.) M.Fleisch. inEngler & Prantl, Nat. Pfl. ed. 2, 11: 187 (1925).
Specimens examined. II, on tree-trunk(Higuchi 48364); on basal part of tree (Higuchi
48373Taxithelium oblongifolium).
Note. This species has been reported from theisland by Shin (1971) and Shin (1973 as Neckera
obtusata). Plants from Lanyu (Higuchi 48364,
48373) are smaller than those of other areas.
Thamnobryum incurvum (Nog.) Nog. & Z.
Iwats., Misc. Bryol. Lichenol., 6: 33 (1972).
Note. This species has been reported from theisland by Noguchi (1937 as Thamnium incurvum)

Moss Flora of Lanyu, Taiwan

and Shin (1973 as Thamnium incurvum).
Orthotrichaceae
Macromitrium angustifolium Dozy & Molk.,
Ann. Sci. Nat. Bot., sér. 3, 2: 311 (1844).
Note. This species has been reported from theisland by Herzog and Noguchi (1955) and Shin
(1973).
Macromitrium foxworthyi Broth., Philipp. J.
Sci., 3: 16 (1908).
Specimen examined. I, on tree-trunk (Higuchi
48305Calymperes afzelii).
Note. This species has been reported from theisland by Shin (1971, 1973).
Macromitrium holomitrioides Nog., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot., 3: 135 (1938).
Note. This species has been reported from theisland by Shin (1971, 1973).
Macromitrium japonicum Dozy & Molk., Ann.
Sci. Nat. Bot., sér. 3. 2: 311 (1844).
Note. This species has been reported from theisland by Wang (1969) and Shin (1971, 1973).
Pilotrichaceae
Callicostella papillata (Mont.) Mitt., J. Proc.
Linn. Soc., Bot., Suppl. 1: 136 (1859).
Specimens examined. I, on boulder (Higuchi
48304, 48359); on decaying log (Higuchi 48360
Vesicularia montagnei, Nishimura 12063).
Note. This species has been reported from theisland by Noguchi (1937 as C. paillata f. longifolia) and Herzog and Noguchi (1955 as C. paillata
f. longifolia).
Pottiaceae
Barbula consanguinea (Thwaites & Mitt.) A.
Jaeger, Ber. S. Gall. Naturw. Ges., 1877–78: 409(1880).
Note. This species has been reported from theisland by Herzog and Noguchi (1955) and Wang

101

(1969), but is not listed in the checklist of Taiwan
mosses by Chiang et al. (2001).
Barbula indica (Hook.) Spreng., Nomencl. Bot.,
2: 72 (1824).
Specimen examined. II, on concrete (Higuchi
48394Hyophila involuta).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as Hymenostomum malayense) and Wang (1969).
Barbula subcomosa Broth., Hedwigia, 38: 211(1899).
Specimen examined. II, on rock-cliff (Higuchi
48363Calymperes afzelii).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as Hydrogonium subcomosum).
Bryoerythrophyllum recurvirostre (Hedw.)
P.C.Chen, Hedwigia, 80: 5 (1941).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as Didymodon recurvirostre).
Gymnostomum recurvirostrum Hedw., Sp.
Musc. Frond., 33 (1801).
Note. This species has been reported from theisland by Shin (1973).
Hydrogonium arcuatum (Griff.) Wijk & Margad., Taxon, 7: 289 (1958).
Note. This species has been reported from theisland by Wang (1969 as Barbula subulata), but
is not listed in the checklist of Taiwan mosses by
Chiang et al. (2001).
*Hyophila involuta (Hook.) A.Jaeger, Ber. S.
Gall. Naturw. Ges., 1871–72: 354 (1873).
Specimens examined. II, on concrete (Higuchi
48394Barbula indica, Higuchi 48395). IV, on
stone wall (Higuchi 48398Brachymenium exile,
Philonotis hastata).
Note. This species is new to Lanyu.
*Weissia controversa Hedw., Sp. Musc. Frond.,

102

Masanobu Higuchi et al.

67 (1801).
Specimen examined. V, on soil (Higuchi
48400). II, on decaying log (Higuchi 48384
Isopterygium minutirameum.
Note. This species is new to Lanyu.
Weissia exserta (Broth.) P.C.Chen, Hedwigia,
80: 158 (1941).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as Hymenostomum edentulum).
Ptychomniaceae
Garovaglia elegans (Dozy & Molk.) Hampe ex
Bosch & Sande Lac., Bryol. Jav., 2: 81 (1863).
Specimens examined. I, on tree-trunk (Higuchi
48320, 48357).
Note. This species has been reported from theisland by Noguchi (1939 as Endotrichella elegans var. brevicuspis).
Pylaisiadelphaceae*Isocladiella surcularis (Dixon) B.C.Tan &Mohamed, Cryptog. Bryol. Lichénol., 11: 357(1990).
Specimen examined. I, on fallen log (Higuchi
48318).
Note. This species is new to Lanyu.
Isopterygium albescens (Hook.) A.Jaeger, Ber.
S. Gall. Naturw. Ges., 1876–77: 433 (1878).
Note. This species has been reported from theisland by Shin (1973).
Isopterygium minutirameum (Müll.Hal.) A.
Jaeger, Ber. S. Gall. Naturw. Ges., 1876–77: 434(1878).
Specimens examined. I, on decaying log(Higuchi 48348, 48352, Nishimura 12074). II, on
decaying log (Higuchi 48384Trichosteleumboschii, Vesicularia flaccida).
Note. This species has been reported from theisland by Wang (1969).

Racopilaceae
Racopilum aristatum Mitt., J. Proc. Linn. Soc.,
Bot., 8: 155 (1864).
Note. This species has been reported from theisland by Noguchi (1937), Herzog and Noguchi(1955) and Shin (1973).
Racopilum cuspidigerum (Schwägr.) Ångström,Öfvers. Förh. Kongl. Svenska Vetensk.-Akad.,29(4): 10 (1872).
Note. This species has been reported from theisland by Shin (1971, 1973 as Racopilum ferriei).
Racopilum spectabile Reinw. & Hornsch., Nova
Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat.
Cur. 14(2): 721 (1829).
Note. This species has been reported from theisland by Wang (1969).
Rhabdoweisiaceae
Amphidium papillosum E.B.Bartram, Philipp.
J. Sci., 68: 48 (1939).
Note. This species has been reported from theisland by Wang (1969).
Symblepharis reinwardtii (Dozy & Molk.)
Mitt., Trans. Roy. Soc. Edinburgh, 31: 331(1888).
Note. This species has been reported from theisland by Wang (1969).
Rhizogoniaceae*Pyrrhobryum spiniforme (Hedw.) Mitt. var.
ryukyuense (Z.Iwats.) Manuel, Cryptog. Bryol.
Lichénol. 1: 70 (1980).
Specimens examined. I, on root (Higuchi
48314); on decaying log (Higuchi 48337).
Note. This species is new to Lanyu.
Sematophyllaceae*Radulina elegantissima (M.Fleisch.) W.R.
Buck & B.C.Tan, Acta Bryol. Asiat., 1: 10 (1989

Moss Flora of Lanyu, Taiwan[1990]).
Specimen examined. I, on root (Higuchi
48339).
Note. This species is new to Taiwan. The
genus Radulina is characterized by pluripapillose
leaf cells, a basal row of much inflated, hyaline
alar cells and strongly collenchymatous exothecial cells (Tan and Jia, 1999). R. elegantissima isdistinguished from R. hamata by its small plants,
leaves less than 1 mm long and 0.2 mm wide andshort setae less than 5 mm long.
Radulina hamata (Dozy & Molk.) W.R.Buck &B.C.Tan, Acta Bryol. Asiat., 1: 10 (1989 [1990]).
Specimens examined. I, on decaying log(Higuchi 48302); on tree-trunk (Higuchi 48307).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as Trichosteleum hamatum).
Rhaphidostichum piliferum (Broth.) Broth. inEngler & Prantl, Nat. Pfl., ed. 2, 11: 434 (1925).
Specimens examined. I, on tree-trunk (Higuchi
48345); on decaying log (Higuchi 48334).
Note. This species has been reported from theisland by Herzog and Noguchi (1955). R. piliferum is often confused with Wijikia tanytricha. Tan
and Jia (1999) pointed out, “It is possible that all
Chinese reports of R. piliferum are based on
misidentification.” R. piliferum is characterized
by stout plants, leaves broadly ovate, more than
2.5 mm long and 1 mm wide and abruptly constricted into a filiform acumen of leaves.
Taxithelium batanense E.B.Bartram, Philipp. J.
Sci., 68: 345 (1939).
Note. This species has been reported from theisland by Herzog and Noguchi (1955) and Shin
(1973).
Taxithelium instratum (Brid.) Broth., Rev.
Bryol. 28: 110 (1901).
Note. This species has been reported from theisland by Herzog and Noguchi (1955) and Wang
(1969).

103*Taxithelium oblongifolium (Sull. & Lesq.) Z.
Iwats., J. Hattori Bot. Lab., 29: 60 (1966).
Specimens examined. I, on root (Higuchi
48294); on tree-trunk (Higuchi 48306Taxithelium parvulum). II, on basal part of tree (Higuchi
48373Neckeropsis obtusata, Nishimura 12082).
Note. This species is new to Lanyu.
*Taxithelium parvulum (Broth. & Paris) Broth.
in Engler & Prantl, Nat. Pfl., I (3): 1092 (1908).
Specimens examined. I, on tree-trunk (Higuchi
48306Taxithelium oblongifolium). II, on boulder (Higuchi 48372Fissidens hollianus).
Note. This species is new to Lanyu.
Trichosteleum boschii (Dozy & Molk.) A.
Jaeger, Ber. S. Gall. Naturw. Ges., 1876–77: 421(1878).
Specimens examined. I, on decaying log(Higuchi 48325Distichophyllum maibarae). II,
on decaying log (Higuchi 48383, 48384
Isopterygium minutirameum, Vesicularia flaccida,
Nishimura 12081).
Note. This species has been reported from theisland by Wang (1969) and Shin (1973).
Symphyodontaceae
Chaetomitriopsis glaucocarpa (Reinw. ex
Schwägr.) M.Fleisch., Musci Fl. Buitenzorg, 4:1372 (1923).
Note. This species has been reported from theisland by Shin (1971, 1973).
Trachythecium micropyxis (Broth.) E.B.Bartram, Philipp. J. Sci., 68: 363 (1939).
Note. This species has been reported from theisland by Herzog and Noguchi (1955).
Trachythecium verrucosum (A.Jaeger) M.
Fleisch. var. binervulum Herzog, J. Hattori Bot.
Lab., 14: 70 (1955).
Note. This varietas was described based on thespecimens from Lanyu and Lutao (Herzog andNoguchi, 1955).

104

Masanobu Higuchi et al.

Thuidiaceae
Claopodium aciculum (Broth.) Broth. in Engler& Prantl, Nat. Pfl., I (3): 1009 (1908).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as C. aciculum var. brevifolium).
Claopodium gracillimum (Cardot & Thér.)Nog., J. Hattori Bot. Lab., 27: 33 (1964).
Note. This species has been reported from theisland by Shin (1973).
Haplohymenium pseudo-triste (Müll.Hal.) Broth.
in Engler & Prantl, Nat. Pfl., I (3): 986 (1907).
Specimen examined. II, on tree-trunk (Higuchi
48365Calymperes graeffeanum).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as H. filiforme and H. submicrophyllum) and Shin (1973).
Pelekium bifarium (Bosch & Sande Lac.) M.
Fleisch., Musci Fl. Buitenzorg 4: 1513 (1923).
Note. This species has been reported from theisland by Herzog and Noguchi (1955) and Wang
(1969).
Pelekium bonianum (Besch.) Touw, J. Hattori
Bot. Lab., 90: 203 (2001).
Note. This species has been reported from theisland by Herzog and Noguchi (1955 as Thuidium lejeuneoides) and Shin (1973 as Thuidium
lejeuneoides).
Pelekium velatum Mitt., J. Linn. Soc., Bot., 10:
176 (1868).
Note. This species has been reported from theisland by Herzog and Noguchi (1955) and Shin
(1973).
Pelekium versicolor (Müll.Hal.) Touw, J. Hattori
Bot. Lab., 90: 205 (2001).
Note. This species has been reported from theisland by Wang (1969 as Thuidium minutulum).
*Thuidium pristocalyx (Müll.Hal.) A.Jaeger,

Ber. S. Gall. Naturw. Ges., 1876–77: 257 (1878).
Specimen examined. II, on soil (Higuchi
48380).
Note. This species is new to Lanyu.
Acknowledgements
This study was supported in part by a Grantin-Aid (no. 18570097 to M. Higuchi) from the
Ministry of Education, Culture, Sports, Scienceand Technology, Japan and Japan Society for the
Promotion of Science.

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