Bull. Natl. Mus. Nat. Sci., Ser. B, 46(1), pp. 9–15, February 21, 2020

A Contribution to the Knowledge of Moss Flora of Koh-Kong andMondulkiri Provinces, Cambodia
Masanobu Higuchi1,*, Tadashi Suzuki2,†, William R. Buck3 and Vadim A. Bakalin4
1

Department of Botany, National Museum of Nature and Science,
4–1–1 Amakubo, Tsukuba, Ibaraki 305–0005, Japan
2
Hattori Botanical Laboratory, Shimada Branch,
6480–3 Takasago-cho, Shimada-shi, Shizuoka 427–0054, Japan
3
Institute of Systematic Botany, New York Botanical Garden, Bronx, NY 10458–5126, U.S.A.
4
Botanical Garden-Institute, Far Eastern Branch of the Russian Academy of Sciences,
Makovskogo Street 142, Vladivostok, 690024, Russia*E-mail: higuchi@kahaku.go.jp(Received 6 December 2019; accepted 25 December 2019)

Abstract The moss ﬂora of Koh-Kong and MondulKiri provinces, Cambodia, was investigatedin 2011. The mosses recognized comprise 20 families, 30 genera and 52 species. Among them 24
species are reported as new to the moss ﬂora of Cambodia. For each species recognized here,locality, substrate, specimen number are provided and taxonomic notes are given for some species.
Key words : bryophytes, Cambodia, Koh-Kong Province, MondulKiri Province, mosses.

This study deals with the moss ﬂora of Cambodia (Kampuchea) based on a ﬁeld trip made by
Bakalin in 2011. He made ﬁeld researches andcollected bryophytes mainly from Koh-Kong and
Mondulkiri provinces, Cambodia. The main purpose of Bakalinʼs trip to Cambodia was the collecting of liverworts, whereas mosses were collected only occasionally and, therefore, thepresent list does not pretend to be exhaustive inany sense. However, taking into account thepoorness of data available on Cambodian mosses
we think the publication of the list of specimen
identiﬁcations would be a small contribution to
the knowledge of moss taxonomic diversity inthis still poorly investigated country.
The ﬁeld studies took 10 days from the 17th tothe 26th of December 2011, in two provinces,both of which belong to the tropical zone. KohKong Province is located in the southwestern
ﬂank of the Cambodia and bordered by the Siam
Bay of the South China Sea. The mean annual© 2020 National Museum of Nature and Sciencetemperature is 27.1°C with the annual amount ofprecipitation over 4,000 mm per year with summer maximum (where from April to Octoberalmost 3,800 mm are fallen). Mondulkiri Province is located in the northeastern part of thecountry and borders with southern Vietnam in theEast. The mean annual temperature of Senmonorom city (the capital of the province) is 22.9°C
and annual precipitation is 2,200 mm, with a distinct summer maximum, when from April toOctober over 2,000 mm are fallen (https://en.
climate-data.org for Sen Monorom and KohKong). In the both provinces the annual temperature variations are very low: within 1°C in KohKong Province and 2°C in Mondulkiri Province.
Therefore the seasonal variation is evident inmoisture conditions but not in temperature
regime, this feature shows the tropical characterof the climate of the studied areas. The basic
habitats for the mosses that were gathered weredecaying or living tree trunks and moist cliffs
near waterfalls (Figs. 1–3), other habitats wererare.

10

Masanobu Higuchi et al.

Figs. 1–3. 1. Tropical forest with dense bamboo thickets in stream valley in Mondulkiri Province (S. S. Choi,2011). 2. Small waterfall—a habitat for peculiar bryophytes in Koh Kong Province (S. S. Choi, 2011).
3. Small stream valley in Koh Kong Province (S. S. Choi, 2011).

Moss Flora of Koh-Kong and Mondulkiri Provinces, Cambodia

The mosses of Cambodia have been reported
by Tixier (1966, 1967, 1975, 1979, 1980). Tanand Iwatsuki (1993) compiled a checklist ofIndochinese mosses, and listed 73 genera and152 species from Cambodia. Later Higuchi(2009) added nine species to the moss ﬂora ofCambodia. The purpose of this study is to investigate the moss ﬂora of Koh-Kong and Mondulkiri provinces and compile it based on thespecimens collected.
Materials and Methods
Field studies were carried out in December
2011 and a total of 78 moss specimens were collected. The sites investigated are divided as follows. The collections are preserved in the herbarium of the Botanical Garden-Institute (VBGI)
and the duplicates in the herbarium of National
Museum of Nature and Science (TNS).
Main collecting sites are as follows.
A. Mondulkiri Province, broad-leaved evergreen forest, 285 m alt., 12°11′04″N,
106°59′28″E, December 17, 2011.
B. Mondulkiri Province, broad-leaved evergreen forest on slope, 880 m alt., 12°22′19″N,107°18′33″E, December 18, 2011.
C. Mondulkiri Province, broad-leaved evergreen forest on slope, 860 m alt., 12°21′22″N,107°18′54″E, December 18, 2011.
D. Mondulkiri Province, broad-leaved evergreen forest near waterfall, 800 m alt.,
12°24′35″N, 107°18′46″E, December 18,
2011.
E. Mondulkiri Province, broad-leaved evergreen forest on slope, 900 m alt., 12°32′40″N,107°30′31″E, December 19, 2011.
F. Mondulkiri Province, broad-leaved evergreen forest near waterfall, 530 m alt.,
12°26′31″N, 107°09′37″E, December 20,
2011.
G. Koh-Kong Province, broad-leaved evergreen
forest on slope, 700 m alt., 11°48′19″N,103°30′39″E, December 22, 2011.
H. Koh-Kong Province, agricultural land nearvillage, 430 m alt., 11°41′06″N, 103°26′23″E,

11December 23, 2011.
I. Koh-Kong Province, broad-leaved evergreen
forest on slope to waterfall, 360 m alt.,
11°39′43″N, 103°23′55″E, December 23,
2011.
J. Koh-Kong Province, broad-leaved evergreen
forest on slope, 400 m alt., 11°41′15″N,103°25′38″E, December 23, 2011.
K. Koh-Kong Province, broad-leaved evergreen
forest on slope, 200 m alt., 11°37′33″N,103°16′07″E, December 24, 2011.
L. Koh-Kong Province, broad-leaved evergreen
forest on slope, 200 m alt., 11°35′15″N,103°11′29″E, December 24, 2011.
M. Koh-Kong Province, wet broad-leaved evergreen forest at the edge of grass land, 380 malt., 11°35′49″N, 103°13′32″E, December 24,2011.
N. Koh-Kong Province, broad-leaved evergreen
forest on slope, 700 m alt., 11°48′56″N,103°28′34″E, December 25, 2011.
O. Koh-Kong Province, area near waterfall inevergreen forest, 43 m alt., 11°35′11″N,103°05′47″E, December 26, 2011.
Results and Discussion
The mosses recognized in this study comprise
20 families, 30 genera and 52 species. Twenty
four species are new additions to the moss ﬂoraof Cambodia. Among them ten taxa, Fissidens
albolimbatus, F. autoicus, F. axilliﬂorus, F.
crispulus. var. crispulus, F. crispulus var. robinsonii, F. ﬁrmus, F. kalimpongensis, F. kamiyamae, F. pellucidus and F. pseudoceylonensis, are
new to Indochina including Cambodia, Laos,Myanmar, Thailand and Vietnam.
Enumeration of Species
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 “newto Cambodia.” Each species is referred by collecting site (A–O), substrate and specimen num-Masanobu Higuchi et al.

12

ber of Bakalin, and some species taxonomicnotes.
Bartramiaceae (by Higuchi)*Philonotis roylei (Hook. f.) Mitt.
Specimen examined. D, on wet cliff, splashedwith water near waterfall (#78-11-11).
Note. This species is reported from Myanmar(Tanaka et al., 2003) and Thailand (cf. Tan andIwatsuki, 1993) in Indochina.
Calymperaceae (by Higuchi)
Calymperes semiliber Mitt.
Specimen examined. I, on decaying wood(#83-4-11).
Leucophanes octoblepharoides Brid.
Specimen examined. G, on tree trunk (#81-2311).
Octoblepharum albidum Hedw.
Specimen examined. B, on tree trunk (#76-4011).
Syrrhopodon confertus Sande Lac.
Specimen examined. K, on decaying wood(#85-16-11).
Syrrhopodon muelleri (Dozy & Molk.) SandeLac.
Specimen examined. K, on tree trunk (#85-711).
Syrrhopodon trachyphyllus Mont.
Specimen examined. G, on tree trunk (#81-2511).
Dicranaceae (by Higuchi)*Campylopus comosus (Reinw. & Hornsch.)Bosch & Lac.
Specimens examined. J, on forest ﬂoor (#84-111); M, on forest ﬂoor (#87-6-11).
Note. This species is reported from Thailandand Vietnam in Indochina (cf. Tan and Iwatsuki,1993).
Leucoloma molle (Müll.Hal.) Mitt.
Specimens examined. G, on tree trunk (#81-311, #81-54-11).

Fissidentaceae (by Suzuki)*Fissidens albolimbatus Dixon
Specimen examined. F, on more or less wet
ﬁne-grained ground (Cam-80-24-11).
Note. This species is new to Indochina.
*F. autoicus Thér. & Dixon
Specimen examined. G, on stone in forest(Cam-81-100-11).
Note. This species is new to Indochina.
*F. axilliﬂorus Thwaites & Mitt.
Specimens examined. D, on wet stone nearwaterfall in the spray zone (Cam-78-32-11); on
wet cliff, splashed with water near waterfall(Cam-78-15-11).
Note. This species is new to Indochina.
F. ceylonensis Dozy & Molk.
Specimen examined. B, on bare soil on slope
in grass land near forest (Cam-76-6-11).
*F. crispulus Brid. var. crispulus
Specimens examined. D, on wet cliff, splashedwith water near waterfall (Cam-78-10-11); O, on
crevice behind waterfall (Cam-89-3-11).
Note. This species is new to Indochina.
*F. crispulus Brid. var. robinsonii (Broth.) Z.
Iwats. & Z.H.Li
Specimen examined. I, on wet cliff near waterfall (Cam-83-35-11).
Note. This species is new to Indochina.
*F. ﬁrmus Mitt.
Specimen examined. N, on stone in streambed(Cam-88-2-11).
Note. This species is new to Indochina.
*F. geminiﬂorus Dozy & Molk.
Specimen examined. F, on wet cave in the cliff
behind waterfall (Cam-80-49-11).
Note. This species is reported from Vietnam inIndochina (cf. Tan and Iwatsuki, 1993).
F. hollianus Dozy & Molk.
Specimen examined. F, on roadside in partshade (Cam-80-4-11).
*F. javanicus Dozy & Molk.
Specimens examined. G, on stone in forest(Cam-81-86-11, Cam-81-98-11).
Note. This species is reported from Myanmar,
Thailand and Vietnam in Indochina (cf. Tan andMoss Flora of Koh-Kong and Mondulkiri Provinces, Cambodia

Iwatsuki, 1993).
*F. kalimpongensis GanguleeSpecimen examined. F, on more or less wet
ﬁne-grained ground in steep slope to watercourse(Cam-80-6-11).
Note. This species is new to Indochina.
*F. kamiyamae Tad.Suzuki & Z.Iwats.
Specimen examined. A, on bark of tree (Cam75-8-11).
Note. This species is new to Indochina.
F. nobilis Griff.
Specimens examined. F, on wet cave in thecliff behind waterfall (Cam-80-46-11, Cam-8048-11).
*F. pellucidus Hornsch.
Specimen examined. L, on stone side (Cam86-6-11).
Note. This species is new to Indochina.
*F. pseudoceylonensis B.C.Tan & M.S.Choy
Specimen examined. C, on soil in open place(Cam-77-4-11).
Note. This species is new to Indochina.
*F. virens Thwaites & Mitt.
Specimen examined. B, on bare soil in grassland near forest (Cam-76-5-11).
Note. This species is reported from Thailandin Indochina (cf. Tan and Iwatsuki, 1993).
Hypnaceae (by Higuchi)
Pseudotaxiphyllum pohliaecarpum (Sull. &Lesq.) Z.Iwats.
Specimen examined. G, on ﬁne-grained soil onslope (#81-119-11).
Hypopterygiaceae (by Higuchi)
Lopidium struthiopteris (Brid.) M.Fleisch.
Specimen examined. E, on tree base (#79-3011).
Leucobryaceae (by Higuchi)
Leucobryum aduncum Dozy & Molk. var. scalare (M.Fleisch.) A.Eddy
Specimens examined. B, on forest ﬂoor (#76-137-11); E, on decaying wood (#79-8-11), on stone
in part to full shade (#79-47-11); I, on more orless dry cliff near waterfall (#83-66-11); K, on
decaying wood (#85-19-11).
Leucobryum javense (Brid.) Mitt.
Specimen examined. G, on decaying wood(#81-130-11).
Leucobryum sanctum (Brid.) Hampe
Specimen examined. I, on more or less dry
cliff near waterfall (#83-77-11).
Note. Cambodia is not listed in this species byTan and Iwatsuki (1993), but Yamaguchi (1993)
reported the species from Cambodia.
Meteoriaceae (by Higuchi)*Trachycladiella sparsa (Mitt.) M.Menzel
Specimen examined. E, hanging down fromshrub branch (#79-6-11).
Note. This species is reported from Myanmar
and Thailand in Indochina (cf. Tan and Iwatsuki,1993).
Mniaceae (by Higuchi)*Orthomnion bryoides (Griff.) Norkett
Specimen examined. E, on tree trunk (#79-6911).
Note. This species is reported from Myanmar
and Thailand in Indochina (cf. Tan and Iwatsuki,1993).
Neckeraceae (by Higuchi)
Himantocladium plumula (Nees) M.Fleisch.
Specimen examined. G, on tree trunk (#81-3911).
*Homaliodendoron exiguum (Bosch & SandeLac.) M.Fleisch.
Specimen examined. E, on stone in part to fullshade (#79-55-11).
Note. This species is reported from Myanmar
and Thailand in Indochina (cf. Tan and Iwatsuki,1993).
*Homaliodendoron ﬂabellatum (Sm.) M.
Fleisch.

Masanobu Higuchi et al.

14

Specimens examined. G, on tree trunk (#8124-11, #81-32-11, #81-56-11).
Note. This species is reported from Laos,
Myanmar, Thailand and Vietnam in Indochina(cf. Tan and Iwatsuki, 1993).
*Neckeropsis exserta (Hook. ex Schwägr.)Broth.
Specimens examined. I, on tree trunk (#83-1411, #83-19-11); on more or less dry cliff nearwaterfall (#83-72-11).
Note. This species is reported from Myanmar
and Thailand in Indochina (cf. Tan and Iwatsuki,1993).
*Neckeropsis semperiana (Hampe ex Müll.
Hal.) Touw
Specimen examined. K, on thin branches (#8534-11).
Note. This species is reported from Thailandand Vietnam in Indochina (cf. Tan and Iwatsuki,1993).

Specimens examined. D, on wet stone nearwaterfall in the spray zone (#78-30-11); F, on
wet cave in the cliff behind waterfall (#80-3111).
Note. This species is reported from Myanmar,
Thailand and Vietnam in Indochina (cf. Tan andIwatsuki, 1993).

Orthotrichaceae (by Higuchi)

Rhizogoniaceae (by Higuchi)*Macromitrium fasciculare Mitt.
Specimen examined. C, on tree trunk (#77-3111).
Note. This species is reported from Vietnam in
Indochina (cf. Tan and Iwatsuki, 1993).

Pyrrhobryum spiniforme (Hedw.) Mitt.
Specimens examined. B, on tree trunk (#7646-11, #76-54-11); E, on decaying wood (#7915-11), on stone in part to full shade (#79-34-11);
I, on more or less dry cliff near waterfall (#8365-11).

Polytrichaceae (by Higuchi)*Pogonatum camusii (Thér.) Touw
Specimen examined. G, on ﬁne-grained soil onslope (#81-113-11).
Note. This species is reported from Thailand
in Indochina (cf. Tan and Iwatsuki, 1993).
Pottiaceae (by Higuchi)
Barbula subcomosa Broth.
Specimen examined. H, on stone in part shadein agricultural land near village (#82-2-11).
Hyophila involuta (Hook.) A.Jaeger
Specimen examined. D, on wet cliff, splashedwith water near waterfall (#78-11-11).
*Timmiella anomala (Bruch & Schimp.) Limpr.

Pterobryaceae (by Higuchi)
Pterobryopsis crassicaulis (Müll.Hal.) M.
Fleisch.
Specimen examined. G, on tree trunk (#81-211).
Racopilaceae (by Higuchi)
Racopilum cuspidigerum (Schwägr.) Ångstr.
Specimen examined. D, on stone in forest(#78-3-11).

Sematophyllaceae (by Higuchi)
Acanthorrhynchium papillatum (Harv.) M.
Fleisch.
Specimens examined. B, on tree trunk onslope (#76-60-11); I on more or less dry cliff
near water fall on slope to waterfall (#83-65-11).
Papillidiopsis macrosticta (Broth. & Paris)
W.R.Buck & B.C.Tan
Specimen examined. I, on more or less dry
cliff near water fall (#83-68-5-11).
Sphagnaceae (by Buck)
Sphagnum perichaetiale Hampe
Specimens examined. M, on forest ﬂoor (#87-Moss Flora of Koh-Kong and Mondulkiri Provinces, Cambodia

7-11, #87-12-11, #87-13-11, #87-16-11, #87-1811, #87-19-11).
Stereophyllaceae (by Higuchi)
Entodontopsis nitens (Mitt.) W.R.Buck & Ireland
Specimen examined. E, on stone in part to fullshade (#79-42-11).
Thuidiaceae (by Higuchi)*Herpetineuron toccoae (Sull. & Lesq.) Cardot
Specimens examined. I, on more or less dry
cliff near waterfall (#83-73-11); N, on stone side(#88-25-11).
Note. This species is reported from Laos,
Thailand and Vietnam in Indochina (cf. Tan and
Iwatsuki, 1993).
Pelekium investe (Mitt.) Touw
Specimen examined. A, on stone in part shade(#75-1-11).
Thuidium plumulosum (Dozy & Molk.) Dozy& Molk.
Specimens examined. D, on wet stone nearwaterfall in the spray zone (#78-23-11); G, on
stone in forest (#81-94-11).
Thuidium pristocalyx (Müll.Hal.) A.Jaeger var.
samoanum (Mitt.) Touw
Specimens examined. G, on tree trunk (#8116-11); L, on stone side (#86-11-11).
Acknowledgments
The collecting of bryophytes and travel
expenses were supported by Prof. B. Y. Sun(Chonbuk National University, Republic of15Korea) and the photographs of habitat were provided by Dr. S. S. Choi from National Institute ofEcology (Republic of Korea) to both of whom V.
Bakalin is sincerely grateful.
References
Gofﬁnet, B., Buck, W. R. and Shaw, A. J. 2009. Morphology, anatomy, and classiﬁcation of the Bryophyta. In:
Gofﬁnet, B. and Shaw, A. J. (eds.), Bryophyte Biology.
2nd ed., pp. 55–138. Cambridge University Press, NewYork.
Higuchi, M. 2009. Moss ﬂora of Angkor, Cambodia. Bulletin of the National Museum of Nature and Science,Series B (Botany) 35: 141–150.
Tan, B. C. and Iwatsuki, Z. 1993. A checklist of Indochinese mosses. Journal of Hattori Botanical Laboratory
74: 325–405.
Tanaka, A., Zaw, K. M., Ngai, S. G. and Akiyama, H.
2003. Mosses of Natma Taung (Mt. Victoria) National
Park, Myanmar. Makinoa 3: 1–84.
Tixier, P. 1966. Bryophytae Indosinicae. A preliminary
contribution to the bryoﬂora of Angkhor (Cambodia).
Natural History Bulletin of the Siam Society 21: 129–133.
Tixier, P. 1967. Bryophytae Indosinicae. Inventaire bryologique de massif de Kirirom (Cambodge). Annales
de la Faculté des Sciences, Université de Phnom Penh
1: 41–68.
Tixier, P. 1975. Bryophytae Indosinicae.―XXIII. A preliminary contribution to the knowledge of the coastal
southern bryoﬂora of Cambodia. Natural History Bulletin of the Siam Society 26: 11–24.
Tixier, P. 1979. Bryogéographie du Mont Bokor (Cambodge) (Bryophyta Indosinicae XXIV). Bryophytorum
Bibliotheca 18: 1–113.
Tixier, P. 1980. Bryophytae Indosinicae.―XXVI. Cambodian bryoﬂora. Collections from Phnom Kulen. NovaHedwigia 32: 369–376.
Yamaguchi, T. 1993. A revision of the genus Leucobryum(Musci) in Asia. Journal of Hattori Botanical Laboratory 73: 1–123.