cryptogamie

Bryologie 2021 ● 42 ● 19
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Rédacteurs associés / Associate Editors

Biologie moléculaire et phylogénie / Molecular biology and phylogeny
Bernard GOFFINET
Department of Ecology and Evolutionary Biology, University of Connecticut (United States)
Mousses d’Europe / European mosses
Isabel DRAPER
Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid (Spain)
Francisco LARA GARCÍA
Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid (Spain)
Mousses d’Afrique et d’Antarctique / African and Antarctic mosses
Rysiek OCHYRA
Laboratory of Bryology, Institute of Botany, Polish Academy of Sciences, Krakow (Pologne)
Bryophytes d’Asie / Asian bryophytes
Rui-Liang ZHU
School of Life Science, East China Normal University, Shanghai (China)
Bioindication / Biomonitoring
Franck-Olivier DENAYER
Faculté des Sciences Pharmaceutiques et Biologiques de Lille, Laboratoire de Botanique et de Cryptogamie, Lille (France)
Écologie des bryophytes / Ecology of bryophyte
Nagore GARCÍA MEDINA
Department of Biology (Botany), and Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de Madrid (Spain)

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 Data to the Malaysian liverwort flora, III: New Lejeuneaceae
records from Sabah and Peninsular Malaysia

Muhammad Syafiq SARIMI

Faculty of Science and Marine Environment, 21030 Kuala Nerus,
Universiti Malaysia Terengganu, Terengganu (Malaysia)

Tamás PÓCS
Eszterházy Károly University, Institute of Biology,
Botany Department, Eger, Pf. 43, H-3301 (Hungary)

Gaik Ee LEE
Eco-Innovation Research Interest Group, Faculty of Science and Marine Environment,
21030 Kuala Nerus, Universiti Malaysia Terengganu, Terengganu
and Institute of Tropical Biodiversity and Sustainable Development, 21030 Kuala Nerus,
Universiti Malaysia Terengganu, Terengganu (Malaysia)
[email protected] (corresponding author)

Submitted on 30 April 2021 | Accepted on 26 October 2021 | Published on 20 December 2021

Sarimi M. S., Pócs T. & Lee G. E. 2021. — Data to the Malaysian liverwort flora, III: New Lejeuneaceae records from Sabah and
Peninsular Malaysia. Cryptogamie, Bryologie 42 (19): 249-267. https://doi.org/10.5252/cryptogamie-bryologie2021v42a19.
http://cryptogamie.com/bryologie/42/19

ABSTRACT
New Malaysian records from the family Lejeuneaceae are presented, including four species new to
Malaysia and seven new to Peninsular Malaysia (Kelantan or Terengganu). The new records include
KEY WORDS Lejeunea convexiloba M.L.So & R.L.Zhu, which was only known from China and Radula grandilobula
Lejeuneaceae, Promma & Chantanaorr. , which was only known from its type from southern Thailand and Borneo.
liverworts,
Malaysia, All species are illustrated, their diagnostic characteristics and morphological comparison between their
new records. allies are provided and discussed, habitat and distribution are given.

RÉSUMÉ
Données sur la flore d’hépatiques malaisienne, III: Nouveaux enregistrements de Lejeuneaceae de Sabah et
de la Malaisie péninsulaire.
De nouveaux signalements malais de la famille des Lejeuneaceae sont présentés, dont quatre espèces
nouvelles pour la Malaisie et sept nouvelles pour la Malaisie péninsulaire (Kelantan ou Terengganu).
Les nouveaux enregistrements incluent Lejeunea convexiloba M.L.So & R.L.Zhu, qui n’était connu
MOTS CLÉS qu’en Chine et Radula grandilobula Promma & Chantanaorr., qui n’était connu que par son type dans
Lejeuneaceae, le sud de la Thaïlande et à Bornéo. Toutes les espèces sont illustrées, leurs caractéristiques diagnos-
hépatiques,
Malaisie, tiques et la comparaison morphologique avec les taxons proches sont fournies et discutées, l’habitat
signalements nouveaux. et la distribution sont donnés.

CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.cryptogamie.com/bryologie 249
 Sarimi M. S. et al.

INTRODUCTION on fresh materials although some older collections made

by the last author were also incorporated. Oil bodies were
Lejeuneaceae is the largest family of the liverworts and widely studied within one day (less than 12 hours) after collect-
distributed in the tropical regions. An estimated 30% of the ing of the material. Names of species new to Malaysia are
liverwort species in the tropics and up to 50% of the bryophytes preceded by an asterisk. Collecting localities are indicated
in a lowland rain forest are members of this family (e.g., Cor- in Figure 1. All specimens are deposited in the Herbarium
nelissen & Ter Steege 1989; Zartman 2003; Gradstein 2006). of Universiti Malaysia Terengganu (UMTP), with selected
Since the publication of the catalogue of Lejeuneaceae duplicates in Sandakan Herbarium (SAN).
of Malaysia (Lee et al. 2013), new genera and species of
Lejeuneaceae have been added to the Malaysian liverwort
flora, e.g., Colura cristata Jovet-Ast (Sangrattanaprasert RESULTS AND DISCUSSION
et al. 2019), Leptolejeunea tripuncta (Mitt.) Steph. (Shu
et al. 2016), and Thiersianthus silamensis R.L.Zhu & L.Shu Family LEJEUNEACEAE Rostovzev
(Zhu et al. 2017), names of species and genera have been Genus Ceratolejeunea (Spruce) J.B.Jack & Steph.
reduced to synonymy or transferred to other genera, e.g.,
Drepanolejeunea brunnea Mizut. (Mohamedia brunnea Ceratolejeunea minor Mizut.
(Mizut.) R.L.Zhu & L.Shu), Lepidolejeunea borneensis (Fig. 2)
(Steph.) R.M.Schust. (Mohamedia borneensis (Steph.)
R.L.Zhu & L.Shu (Zhu et al. 2019), Lejeunea tamaspocsii Journal of the Hattori Botanical Laboratory 49: 311 (1981).
G.E.Lee (synonym of Lejeunea pulchriflora G.E.Lee et al. Specimen examined. — Malaysia. Terengganu: Mt. Tebu Forest
(Lee & Gradstein 2013; Lee et al. 2016a), Leptolejeunea Reserve, 15 km S of Jerteh town. Primary lowland dipterocarp forest
serrulata Herzog (synonym of Leptolejeunea tripuncta), Tax- with small waterfalls and slow stream. Along the forest trail at the
ilejeunea compressiuscula Steph. (Lejeunea compressiuscula Lata Belatan Recreational Forest and waterfall, situated at the base
of Mt. Tebu at 100 m alt., on living leaves, 16.IV.2019, G.E. Lee
(Steph.) G.E.Lee & Heinrichs (Lee et al. 2016b, 2020), et al. 19013, 19014, ibid, E. Pesiu et al. 19157, 19182.
and genus Mastigolejeunea (Spruce) Steph. (synonym of
Thysananthus Lindenb.; Sukkharak & Gradstein 2017), or
reinstated (genus Allorgella Tixier; Bechteler et al. 2016; Remarks
several Leptolejeunea species, Bechteler et al. 2017; Shu & This species is recognized by the weakly keeled perianth
Zhu 2019). In the present study, we report four species of without horn-like projections, the small, distant underleaves,
Lejeuneaceae new to Malaysia and seven new to Peninsular and the obovate perianth. Previously known from the type
Malaysia, collected in the state of Sabah and on the East in Sabah, Mt. Templer (Mizutani 1981) and subsequently
Coast of Peninsular Malaysia (Kelantan, Terengganu). Sabah reported from China (Zhu et al. 2005). New to Peninsular
is the most intensively inventoried state of Malaysia for Malaysia (Terengganu). The specimens from Terengganu
liverworts (Lee et al. 2018) but large parts of Kelantan and are similar to the Chinese material in the entire margins of
Terengganu remain virtually unexplored. Only 12 species underleaf lobes and the absence of weak Allorgella-typed den-
(Frullania intermedia (Reinw. et al.) Dumort., 11 spp. of ticulations in the margins of leaf lobes; in the type of Sabah
Lejeunea Lib.) have been recorded from Kelantan (Chuah- the underleaf lobes have 1-2 marginal teeth and the margin
Petiot 2011; Lee 2013; Lee et al. 2013) and two species denticulation is distinct. This species grows on sandstone, tree
(Radula retroflexa Taylor, Plagiochila arbuscula (Brid. ex trunks or living leaves (this study) in the understorey and
Lehm. & Lindenb.) Lindenb.) from Terengganu (Lee et al. canopy of lowland and montane rainforests at an elevation
2019). This paper aims to contribute to the knowledge of of 100-1000 m (Mizutani 1981; Zhu et al. 2005).
liverwort diversity, particularly Lejeuneaceae, in Malaysia.

Ceratolejeunea singapurensis (Lindb.) Schiffn.

MATERIAL AND METHODS (Fig. 3)

This study is based on the intensive liverwort explorations Conspectus Hepaticarum Archipelagi Indici 273 (1898).
carried out by the authors during 2018-2020 in Sabah and Specimen examined. — Malaysia. Terengganu: Mt. Tebu Forest
the East Coast of Peninsular Malaysia (Kelantan, Tereng- Reserve, 15 km S of Jerteh town. Primary lowland dipterocarp for-
ganu). The collections yielded several species new to the est with small waterfalls and slow stream. Along the forest trail at
country, to Sabah or to Peninsular Malaysia, in spite of the the Lata Belatan Recreational Forest and waterfall, situated at the
base of Mt. Tebu at 100 m alt., on living leaves, 16.IV.2019, G.E.
fact that only part of the epiphyllous specimens from Sabah Lee et al. 19012.
were identified. A first paper on the epiphyllous communi-
ties of Sabah has already been published (Pócs et al. 2020).
The present paper included new floristic records. This is the Remarks
continuation of a series on new Malaysian records (Pócs The distinctive features are the presence of 1-6 basal
et al. 2014; Pócs & Lee 2016). The study was largely based ocelli, the approximate to distantly arranged and orbicu-

250 CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19)

 Data to the Malaysian liverwort flora, III

BHUTAN CHINA

BANGLADESH TAIWAN

MYANMAR Philippine Sea

LAOS
Bay of Bengal
South
THAILAND China Sea

CAMBODIA PHILIPPINES
Andaman
Sea VIETNAM

Peninsular Sabah
Malaysia
Sarawak Celebes Sea

SUMATERA KALIMANTAN
Bismarck Sea

INDONESIA
PAPUA NEW GUINEA

Solomon Sea

AUSTRALIA

B C

THAILAND

   Mt. Kinabalu

Kelantan
Terengganu 
Mt. Alab

Sabah


Mt. Silam

SUMATERA KALIMANTAN

Fig. 1. — A, Map of Malaysia showing two separate regions; Peninsular Malaysia and Malaysian Borneo (Sabah, Sarawak); B, C, the stars indicate the locality
where the specimens were collected.

lar underleaves, the entire leaf margin, the absence of (Mizutani 1981; Udar & Shaheen 1985) and in Sabah,
utricles, and the oblong leaf lobe with rounded apices. Malaysia (Herzog 1952). New to Peninsular Malaysia
Only known from the type in Singapore and from India (Terengganu).

CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19) 251

 Sarimi M. S. et al.

A B C

D E

I J

F G H

Fig. 2. — Ceratolejeunea minor Mizut.: A, part of plant in ventral view; B, C, leaves; D, upper part of leaf lobule when flattened (hyaline papilla shown in gray);
E, marginal leaf lobe cells; F, G, underleaves; H, cross-section of stem; I, median leaf lobe cells; J, basal leaf lobe cells and ocelli. From E. Pesiu et al. 19157.
Scale bars: A, 0.2 mm; B-C, 0.5 mm; F, G, 0.1 mm; D, E, H-J, 50 µm.

Genus Cololejeunea (Spruce) Steph. Remarks

Cololejeunea appressa is a member of subg. Taeniolejeunea and
Cololejeunea appressa is distinguished from other members in this subgenus by the
(A.Evans) Benedix single row of 3-4 ocelli, the unicellular first lobule tooth and
(Fig. 4) first and second lobule teeth clearly visible, not overlapping
each other. A pantropical species, preferably growing on barks
Feddes Repertorium Specierum Novarum Regni Vegetabilis. Beiheft or on living leaves at low elevations, reported in Asia from
134: 31 (1953). China, Indonesia (Java, Sumatra), Indonesian New Guinea,
Japan, and recently from Laos (Mizutani 1961; Piippo 1990;
Specimen examined. — Malaysia. Terengganu: Mt. Tebu Forest Pócs & Kovács 2019; Söderström et al. 2020). New to Pen-
Reserve, 15 km S of Jerteh town. Primary lowland dipterocarp for-
est with small waterfalls and slow stream. Along the forest trail at insular Malaysia (Terengganu). In Malaysia the species is
the Lata Belatan Recreational Forest and waterfall, situated at the only known from Mt. Kinabalu, Sabah (Mizutani 1966). In
base of Mt. Tebu at 100 m alt., on living leaves, 16.IV.2019, G.E. view of its wide world distribution, however, this species is
Lee et al. 19011. expected to occur in many lowland forest areas in Malaysia.

252 CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19)

 Data to the Malaysian liverwort flora, III

A B C

D E

H I J K

Fig. 3. — Ceratolejeunea singapurensis (Lindb.) Schiffn.: A, part of plant in ventral view; B, C, leaves; D, E, upper part of leaf lobule when flattened (hyaline papilla
shown in gray); F, H, I, underleaves; G, cross-section of stem; J, K, basal leaf lobe cells and ocelli. From G.E. Lee et al. 19012. Scale bars: A-C, 0.2 mm; D, E,
G, 50 µm; F, H-J, K 100 µm.

Mt. Silam from the telecommunication towers to the main summit.

*Cololejeunea equialbi Tixier Lower montane rainforest with Shorea tenuiramulosa and Borneoden-
(Figs 5; 6A, B) dron enigmaticum at elevation of 700-800 m alt., on living leaves,
04°57’12”N, 118°9’39”E, 9.VIII.2018, J. Havasi & T. Pócs 1811/AX.
Annales de la Faculté des Sciences, Université de Phnom Penh 3:
178 (1970). Remarks
This species is characterized by the small leaf lobule usually
Specimen examined. — Malaysia. Terengganu: Mt. Tebu Forest with 1-celled first tooth and obsolete second tooth, unicellular
Reserve, 15 km S of Jerteh town. Hill dipterocarp forest, area around
the summit of Mt. Tebu at 940 m alt., on living leaves, 16.IV.2019, E. stylus, leaf cells with small trigones and indistinct intermedi-
Pesiu et al. 1929, 1948, 1954, 1955, 1956; Sabah: Mt. Silam, Sapagaya ate thickenings, absence of ocelli and vitta, and the widely
Forest Reserve, 22 km WSW of Lahad Datu town. Summit ridge of spreading to retrorsed leaf lobe, 90-160° from the stem. A

CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19) 253

 Sarimi M. S. et al.

widespread Indopacific species (Dey & Singh 2016); new to Specimen examined. — Malaysia. Kelantan: Gunung Basor For-
Malaysia. Because of its wide distribution, its occurrence in est Reserve, halfway between Dabong and Jeli towns near Bkt.
Malaysia was predictable. The species grows exclusively on Gerongan village at Renyok No. 3 water intake, at 670-680 m alt.,
on living leaves, submontane rainforest at cataracts. 05°31.171’N,
living leaves (Zhu & So 2001; Dey & Singh 2016), therefore 101°46.940’E, 7.XI.2013, T. Pócs et al. 13180/R.
can be regarded as an obligate epiphyll.
Remarks
Cololejeunea gottschei is close to C. longifolia and C. stellaris
*Cololejeunea ensifera (subg. Leptocolea sect. Salebrosae) but differs from the latter
Tixier ex L.Söderstr., Pócs, Váňa & A.Hagborg two species in the apiculate, sometimes denticulate leaf lobes,
(Figs 6C-E; 7) and furthermore from C. longifolia in the obovate perianth
and sexangular, multicellular gemmae with four adhesive
Phytotaxa 220 (2): 199 (2015). — Cololejeunea ensifera Tixier, cells and from C. stellaris by its smooth, not papillose cells.
Revue Bryologique et Lichénologique 36 (3/4): 562 (1969) nom. inval.
A widespread epiphyllous Indomalesian species, in Malaysia
Specimen examined. — Malaysia. Sabah: Crocker Range National known only from Sabah (Mizutani 1966, under Cololejeunea
Park NNW of Tambunan. Gunung Alab Substation at elevation of paroica Mizut.). New to Peninsular Malaysia (Kelantan).
1900-1940 m. Mossy cloud (elfin) forest, about 6 m high canopy of
Phyllocladus hypophyllus, Rhododendron, Dacrydium and Nepenthes,
17.VIII.2018. J. Havasi & T. Pócs 1823D, 1823L, 1823O, 1823AC,
1823AH, 1823AL, 1823AN. Genus Colura (Dumort.) Dumort.

Remarks Colura ari (Steph.) Steph.

A species hitherto known from Vietnam, Cambodia and Thai- (Figs 10; 11C, D)
land (Tixier 1969; Chantanaorrapint & Pócs 2014). New to
Malaysia. Typical features are the acute, sword-like first lobule Species Hepaticarum 5: 936 (1916).
tooth crossing the triangular second tooth, the pluricellular Specimen examined. — Malaysia. Terengganu: Mt. Tebu Forest
stylus and the biseriate vitta. On leaves of shrubs and ferns. Reserve, 15 km S of Jerteh town. Primary lowland dipterocarp for-
est with small waterfalls and slow stream. Along the forest trail at
the Lata Belatan Recreational Forest and waterfall, situated at the
base of Mt. Tebu at 100 m alt., on living leaves, 16.IV.2019, G.E.
Cololejeunea schmidtii Steph. Lee et al. 19018, 19019, 19020, 19021.
(Fig. 8)
Botanisk Tidsskrift 24 (3): 278 (1902). Remarks
Specimen examined. — Malaysia. Terengganu: Mt. Tebu Forest Distinctive features of Colura ari include the strongly serrate
Reserve, 15 km S of Jerteh town. Primary lowland dipterocarp for- leaf margin, the position of lobule and sac more or less parallel
est with small waterfalls and slow stream. Along the forest trail at to the stem, truncate and apiculate sac (apex usually pointing
the Lata Belatan Recreational Forest and waterfall, situated at the downward), and leaf cells with large trigones and conspicu-
base of Mt. Tebu at 308 m alt., on living leaves, 16.IV.2019, G.E.
Lee et al. 19015, 19016, 19017. ous intermediate thickenings (2 per adjacent trigones in basal
leaf cells). A common and widespread Indopacific epiphyl-
lous species. New to Peninsular Malaysia (Terengganu). In
Remarks Malaysia, only known from Sabah (Mizutani 1966, 1970).
Cololejeunea schmidtii is unlikely to be confused with other
members of Malaysian Cololejeunea by its 2-celled first tooth
of leaf lobule, large papillose-tuberculate cuticle (1 per cell, *Colura inuii Horik.
usually almost occupying its whole surface), denticulate (Figs 11A, B; 12)
margin of the dorsal leaf base and the 4-keeled, papillose
perianth. A widespread, epiphyllous Indopacific species Journal of Science of Hiroshima University, Series B, Division 2,
Botany 1: 68 (1931).
known from Sri Lanka, Indochina, China, Japan, Philip-
pines, Papua New Guinea and Fiji (Zhu & So 2001; Pócs Specimen examined. — Malaysia. Terengganu: Mt. Tebu Forest
et al. 2011). New to Peninsular Malaysia (Terengganu). In Reserve, 15 km S of Jerteh town. Primary lowland dipterocarp for-
Malaysia, the species was only known from Sabah (Kinabalu est with small waterfalls and slow stream. Along the forest trail at
the Lata Belatan Recreational Forest and waterfall, situated at the
area) (Mizutani 1966). base of Mt. Tebu at 80 m alt., on living leaves, 16.IV.2019, G.E.
Lee et al. 19018, 19019, 19020, 19021, ibid., 13.III.2021, G.E. Lee
21008, 21009, 21010, 21011, 21012.
Cololejeunea gottschei
(Steph.) Pandé, K.P.Srivast. & Ahmad Remarks
(Fig. 9) Colura inuii is very similar to the widespread Indopacific
C. conica (see Zhu & So 2001) but in the latter species, the sac
Journal of the Indian Botanical Society 36 (3): 345 (1957). always has a crested, toothed apiculus whereas in C. inuii, it

254 CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19)

 Data to the Malaysian liverwort flora, III

A B C

F G

Fig. 4. — Cololejeunea appressa (A.Evans) Benedix: A, part of plant in ventral view; B, C, F, leaves; D, stem portion, leaf lobule and stylus (hyaline papilla shown in
gray); E, marginal leaf lobe cells; G, median leaf lobe cells; H, basal leaf lobe cells and ocelli. From G.E. Lee et al. 19011. Scale bars: A, 0.2 mm; B, C, F, 0.1 mm;
D, H, 50 µm; E, G, 25 µm.

does not occur. The perianth keels of C. conica have 1-3 celled into indistinct granules) in C. inuii while granular (faintly
sharp teeth at the end of the wings (Jovet-Ast 1953, fig. 51: rough externally) in C. conica (see Yang et al. 2013: figs 2g, 3e).
1-3) whereas in C. inuii, the perianth keels are often strongly Furthermore, both species can be distinguished by the presence
mamillose but teeth are never produced. Moreover, the oil of 18-20 median valve cells and 17-19 surrounding hyaline
bodies are homogenous (smooth externally, rarely segmented cells in C. inuii while 13-17 and 15-16 in C. conica. Colura

CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19) 255

 Sarimi M. S. et al.

A B C

D
F

Fig. 5. — Cololejeunea equialbi Tixier: A, part of plant in ventral view; B, C, leaves; D, marginal leaf lobe cells; E, median leaf lobe cells; F, androecial shoot;
G, stem portion, leaf lobule and stylus (hyaline papilla shown in gray); H, perianth; I, female bracts. From E. Pesiu et al. 1955. Scale bars: A, H, I, 0.2 mm; B-D,
G, 100 µm; E, 50 µm; F, 0.5 mm.

inuii was known from China (Taiwan), Japan and Vietnam species usually grows on living leaves, rarely on bark and can
(Horikawa 1931; Zhu & So 2001; Shu et al. 2017). New to be considered, as typical epiphyll; in Malaysia it was found
Malaysia, which record in Peninsular Malaysia represents its on living and dry fallen leaves in a lowland dipterocarp forest
southernmost locality, extending to the Malesian region. The and also in montane forest, above 750 m alt.

256 CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19)

 Data to the Malaysian liverwort flora, III

A C

Fig. 6. — Cololejeunea equialbi Tixier: A, part of plant in ventral view and gemmae; B, part of leaf lobe and lobule. From J.Havasi & T.Pócs 1811/AX. Cololejeunea
ensifera Tixier ex L.Söderstr., Pócs, Váňa & A.Hagborg: C, part of plant in ventral view; D, E, part of leaf lobe, lobule and vitta. From J. Havasi & T. Pócs 1823AL.
Scale bars: 50 µm.

Genus Lejeunea Lib. Remarks

The distinguishing characters of Lejeunea convexiloba include
*Lejeunea convexiloba M.L.So & R.L.Zhu 1) the autoicous plant; 2) the relatively large (to ½ the length
(Fig. 13) of leaf lobe) leaf lobules with flat free margin; 3) cross-section
of stem with less than 10, usually 5-6 medullary cells; 4) the
The Bryologist 101 (1): 137 (1998). indistinct trigones and absence of intermediate thickenings;
Specimen examined. — Malaysia. Sabah: Mt. Kinabalu: Paka Cave, 5) the distantly arranged and orbicular underleaves; 6) the
along the trail between Villosa shelter and Laban Rata resthouse, on rough cuticle with finely punctate to verrucose-small papil-
tree trunk, at 2900 m alt., 13.XI.2010, G.E. Lee 1940. lae; and 7) the 5-keeled, obovoid perianth with 2 long ventral

CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19) 257

 Sarimi M. S. et al.

A B

D H

Fig. 7. — Cololejeunea ensifera Tixier ex L.Söderstr., Pócs, Váňa & A.Hagborg: A, part of plant in ventral view; B, C, leaves; D, median leaf lobe cells; E, stem
portion, leaf lobule and stylus (hyaline papilla shown in gray); F, perianth; G, female bracts; H, basal leaf lobe cells and vitta. From J. Havasi & T. Pócs 1823AL.
Scale bars: A-C, F, G, 0.2 mm; D, E, H, 50 µm.

258 CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19)

 Data to the Malaysian liverwort flora, III

A B C

E F

Fig. 8. — Cololejeunea schmidtii Steph.: A, part of plant in ventral view; B, C, leaves; D, perianth, female bracts; E, median leaf lobe cells; F, marginal leaf lobe cells;
G, stem portion, leaf lobule and stylus (hyaline papilla shown in gray), H, gemma. From G.E. Lee et al. 19015. Scale bars: A, D, 0.2 mm; B, C, 0.1 mm; E-H, 50 µm.

keels (more than ⅔ length of perianth) (So & Zhu 1998). L. konosensis and L. pallide-virens but differs by the large leaf
By its autoicy, verrucose cuticle, and less than 10 medullary lobules with free margin and the long ventral keels of perianth.
cells in cross-section of stem, L. convexiloba most resembles The species was previously known only from China, new to

CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19) 259

 Sarimi M. S. et al.

A B

G
H

I J K

Fig. 9. — Cololejeunea gottschei (Steph.) Pandé, K.P.Srivast. & Ahmad: A, part of plant in ventral view; B, C, leaves; D, stem portion and leaf lobule; E, gemma;
F, leaf lobule when flattened (hyaline papilla shown in gray); G, median leaf lobe cells; H, stem portion, lower part of leaf lobule and stylus; I, female bracts; J, peri-
anth and female bracts; K, marginal leaf lobe cells; L, upper part of perianth. From T. Pócs et al. 13180/R. Scale bars: A-C, 0.2 mm; D-F, H-L, 100 µm; G, 50 µm.

260 CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19)

 Data to the Malaysian liverwort flora, III

A B

G H

Fig. 10. — Colura ari (Steph.) Steph.: A, part of plant in ventral view; B, C, leaves; D, underleaf; E, marginal leaf lobe cells; F, valve; G, median leaf lobe cells; H,
apex of lobule sac; I, perianth. From G.E. Lee et al. 19020. Scale bars: A-C, I, 0.5 mm; D, 0.2 mm; E, G, 100 µm; F, H, 50 µm.

Malaysia. It occurs on bark at high elevations and was found in tree trunk at 1500 m in a nature reserve. It is expected that
China at 1500 m and in Malaysia (Mt. Kinabalu) at 2900 m. L. convexiloba may be distributed more widely in montane
Moreover, the holotype in China also found growing on a rainforests of Malesia.

CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19) 261

 Sarimi M. S. et al.

A B

C D

E F

Fig. 11. — Colura inuii Horik.: A, habit; B, oil bodies. From G.E.Lee 21010. Colura ari (Steph.) Steph.: C, habit; D, oil bodies. From G.E. Lee et al. 19020. Radula
grandilobula Promma & Chantanaorr.: E, habit; F, oil bodies. From G.E. Lee 21003. Scale bars: A, C, E, 2 mm; B, D, 50 µm; F, 25 µm.

262 CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19)

 Data to the Malaysian liverwort flora, III

B C
A

D E

G I
H

M N

L
K

Fig. 12. — Colura inuii Horik.: A, part of plant and perianth in ventral view; B, C, leaves; D, cross-section of stem; E, underleaf; F, oil bodies; G, marginal leaf lobe
cells; H, L, valves; I, median leaf lobe cells; J, K, apex of lobule sacs; M, N, perianths and female bracts. A, E, G, D from G.E. Lee et al. 19019; others from G.E.
Lee 21011. Scale bars: A-C, 0.5 mm; D, G, H, L, 50 µm; E, I-K, 100 µm.

CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19) 263

 Sarimi M. S. et al.

A C
D

I
J

Fig. 13. — Lejeunea convexiloba M.L.So & R.L.Zhu: A, B, underleaves; C, part of plant and perianth in ventral view; D, E, leaves; F, G, upper part of leaf lobules
when flattened (hyaline papilla shown in gray); H, perianth; I, median leaf lobe cells; J, cross-section of stem; K, female bracts and bracteole. From G.E. Lee
1940. Scale bars: A, B, D, E, 0.1 mm; C, H, K, 0.2 mm; F, G, I, J, 50 µm.

264 CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19)

 Data to the Malaysian liverwort flora, III

B C
A

D E

Fig. 14. — Radula grandilobula Promma & Chantanaorr.: A, part of plant in ventral view; B, C, leaves; D, median leaf lobe cells; E, leaf lobule and rhizoids (hyaline
papilla shown in gray); F, perianth and female bracts; G, cross-section of stem; H, gemma. From G.E. Lee et al. 19022. Scale bars: A, F, 0.5 mm; B, C, 0.2 mm;
D, G, 50 µm; E, H, 100 µm.

Genus Radula Dumort. the Lata Belatan Recreational Forest and waterfall, situated at the
base of Mt. Tebu at 100 m alt., on living leaves, 16.IV.2019, G.E.
Lee et al. 19022, 19023, 19024, 19025, ibid., 13.III.2021, G.E. Lee
Radula grandilobula Promma & Chantanaorr. 21001, 21002, 21003, 21004.
(Figs 11E, F; 14)
Cryptogamie, Bryologie 36 (3): 219 (2015). Remarks
Specimen examined. — Malaysia. Terengganu: Mt. Tebu Forest Radula grandilobula is close to the widespread Indopacific
Reserve, 15 km S of Jerteh town. Primary lowland dipterocarp for- R. acuminata (Zhu & So 2001); differs in having triangular-
est with small waterfalls and slow stream. Along the forest trail at ovate leaf lobes, frequently recurved free margin of leaf lobules,

CRYPTOGAMIE, BRYOLOGIE • 2021 • 42 (19) 265

 Sarimi M. S. et al.

the leaf lobule apices pointing towards the leaf lobe apices or ‒ a liverwort hotspot. Cryptogamie, Bryologie 27 (1): 141-152.
directed away from the stem, and the finely segmented oil bod- H erzog T. 1952. — Nachtrag zu ‘Hepaticae Borneense
ies, 2-4 per cell (Promma & Chantanaorrapint 2015). Radula (Oxford University Expedition to Sarawak, 1932)’. Transac-
tions of the British Bryological Society 2: 71-73. https://doi.
grandilobula was first described in Nakhon Si Thammarat, org/10.1179/006813852804878435
southern Thailand (Promma & Chantanaorrapint 2015), and Horikawa Y. 1931. — Studies on the hepaticae of Japan. V. Jour-
its occurrence in northern Malaysia was therefore expected. nal of Science of the Hiroshima University: Series B, Divison 2
New to Peninsular Malaysia (Terengganu). The species has (Botany) 1: 55-76.
exclusively been found on living leaves. Jovet-Ast S. 1953. — Le genre Colura, Hépatiques, Lejeuneaceae,
Diplasiae. Revue Bryologique et Lichénologique 22: 206-312.
Lee G. E. 2013. — A systematic revision of the genus Lejeunea Lib.
(Marchantiophyta : Lejeuneaceae) in Malaysia. Cryptogamie,
Acknowledgements Bryologie 34 : 381-484.
This study was supported by the Ministry of Education Lee G. E. & Gradstein S. R. 2013. — Distribution and habi-
(MOE) Malaysia through Fundamental Research Grant tat of the Malaysian species of Lejeunea (Marchantiophyta:
Lejeuneaceae), with description of Lejeunea tamaspocsii sp. nov.
Scheme (FRGS/1/2018/WAB13/UMT/03/1) awarded to Polish Botanical Journal 58: 59-69. https://doi.org/10.2478/
G.E. Lee. Special thanks are due to our companions Ms pbj-2013-0007
Judit Havasi, for helping in collecting and preparing the Lee G. E., Gradstein S. R., Söderström L. & Latiff A. 2013. —
epiphyllous samples and to Mr Danial Tang for taking Catalogue of the Lejeuneaceae of Malaysia. Malayan Nature
care of all logistics and helping in the fieldwork in Sabah. Journal 65: 81-129.
Lee G. E., Bechteler J., Pócs T., Schäfer-Verwimp A. & Hein-
Thanks also to Ms Elizabeth Pesiu and Ms Nor Aishah richs J. 2016a. — Molecular and morphological evidence for
Shafie for their support and assistance during the fieldwork an intercontinental range of the liverwort Lejeunea pulchriflora
in Mt. Tebu Forest Reserve, to Sabah Biodiversity Council, (Marchantiophyta: Lejeuneaceae). Organisms, Diversity and Evo-
Sabah Parks, and Sabah Forestry Department (Ms. Andi lution 16: 13-21. https://doi.org/10.1007/s13127-015-0243-5
Maryani A. Mustapeng) for their support in obtaining Lee G. E., Bechteler J. & Heinrichs J. 2016b. — A revision of
unrevised names in the former genus Taxilejeunea (Marchantio-
permits and research permissions [SaBC access license: phyta: Lejeuneaceae) from Asia. Phytotaxa 358: 26-48. https://
JKM/MBS.1000-2/2 JLD.7 (107)]. Finally, we are very doi.org/10.11646/phytotaxa.358.1.2
thankful to Prof S. Robbert Gradstein for the very care- Lee G. E., Pócs T., Gradstein S. R., Damanhuri A. & Latiff A.
ful corrections, to Prof R. L. Zhu for sending important 2018. — Abundant but neglected past and present of liverwort
literature, and to the anonymous reviewer for the helpful (Marchantiophyta) studies in Malaysia. Cryptogamie, Bryologie
39 (1): 83-91. https://doi.org/10.7872/cryb/v39.iss1.2018.83
remarks on the manuscript. Lee G. E., Damanhuri A. & Norhazrina N. 2019. — Diver-
sity of bryophytes of Terengganu and their ecological roles in
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Submitted on 30 April 2021;

accepted on 26 October 2021;
published on 20 December 2021.

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