A checklist of the bryophyte flora of Mount Tebu Forest Reserve in Terengganu, Peninsular Malaysia, is presented. A total of 189 taxa in 71 genera and 26 families were enumerated. This figure represents 63% of the 298 bryophyte species recorded so far for the State of Terengganu. Out of 189 taxa of bryophytes, 26 liverworts are new additions to the bryoflora of Terengganu. The most prominent liverwort family is represented by Lejeuneaceae, with 54 species from 17 genera, while the moss family is the Sematophyllaceae, with 34 taxa in 13 genera. The majority of the species are epiphytes, either corticolous or ramicolous. Almost half of the bryophyte species have wider elevational ranges and occur from the lowlands to the summit of Mount Tebu.

Biodiversity, bryophytes, checklist, Malaysia, Marchantiophyta, taxonomy

Mount Tebu (1039 m) is the second-highest mountain after Mount Lawit (1519 m) in the northernmost part of Terengganu (Fig. 1). It is located within one of the primary mountain ranges of Peninsular Malaysia, known as the Timur Range (Banjaran Timur). The mountain comprises undulating lowlands, hill and upper hill dipterocarp forest. It has been gazetted as one of the state forest reserves, including the lowlands of Lata Belatan Recreational Forest at the base of Mount Tebu. Geologically, Mount Tebu is composed of unconsolidated alluvium, metasedimentary and igneous rocks in the lowlands to the summit of the mountain (Mohamed and Ali 2014). The unique landscape feature provides ample habitat for a diverse flora and fauna community with high conservation value (see Abdul Rahim et al. (2014) for several extensive floristic and ecological studies). It also offers a variety of vegetation and habitats favourable to the growth and diversity of bryophytes. The history of bryophyte exploration in Terengganu has been reviewed by Lee et al. (2019). The early investigation was conducted by British and Japanese bryologists and yielded only a few bryophyte species, nine being mosses and two were liverworts (Dixon 1926; Yamada 1979; Inoue 1984). Subsequently, more recent collections of bryophytes from this region have been carried out, of which 11 species of bryophyte have been reported for the first time in Peninsular Malaysia and 77 taxa are new records to Terengganu (Lee et al. 2018, 2022; Pesiu et al. 2021; Sarimi et al. 2021).

Figure 1. 

The map of Peninsular Malaysia shows the study area, Mt. Tebu Forest Reserve. Map modified from Dr Blofeld - http://www.maps-for-free.com, CC BY 3.0.

This study is based on the authors’ intensive bryophyte explorations from April 2019–November 2021 in Terengganu and a re-examination of previous moss collections of A. Damanhuri was made during the Mount Tebu scientific expedition in 2012. All the bryophyte samples were collected from various microhabitats along the trails within the study area, including tree trunks and branches, rocks, soils, fallen logs, rotten wood and leaves. Liverwort specimens were deposited in the Herbarium of Universiti Malaysia Terengganu (UMTP) and moss specimens were deposited in the Herbarium of Universiti Kebangsaan Malaysia (UKMB). About 1000 samples of bryophytes were collected from the study area and were examined by light microscopy. The drawing of the specimen was produced using an Olympus BX43 microscope, equipped with a drawing tube.

A total of 189 taxa in 71 genera and 26 families were found in the Mount Tebu Forest Reserve, of which 109 are mosses and 80 are liverworts (Figs 2–4). This represents 63% of the 298 bryophyte species recorded so far for the State of Terengganu (Pócs and Lee 2016; Pesiu et al. 2021; Sarimi et al. 2021; Lee et al. 2022). Out of 80 species of liverworts, 26 are reported for the first time for Terengganu. The largest liverwort family found is the Lejeuneaceae, with 54 species, followed by Lepidoziaceae (eight species) and Radulaceae (seven species). The largest moss family is the Sematophyllaceae, with 34 taxa, followed by Calymperaceae (32 taxa) and Hypnaceae (seven taxa). The smallest liverwort and moss families were represented by only one species, for example, liverworts: Calypogeiaceae, Pallaviciniaceae, Plagiochilaceae, Solenostomataceae and Schistochilaceae and mosses: Diphysciaceae, Myuriaceae, Neckeraceae and Thuidiaceae. As expected, the distinct dominance of species is from the family Lejeuneaceae and mosses Sematophyllaceae and Calymperaceae, representing about 60% of all the bryophyte species found in Mount Tebu. They are the most common bryophyte families in the lowland tropical rainforests with high light intensity, dense canopy, high temperatures and many evergreen tree species.

Figure 2. 

Mosses and their habit A Diphyscium mucronifolium Mitt B Leucobryum sanctum (Schwägr.) Hampe C Fissidens ceylonensis Dozy & Molk D Pyrrhobryum latifolium (Bosch & Sande Lac.) Mitt E Arthrocormus schimperi (Dozy & Molk.) Dozy & Molk F Octoblepharum albidum Hedw G Mitthyridium fasciculatum (Hook. & Grev.) H. Rob H Ectropothecium buitenzorgii (Bél.) Mitt. I Syrrhopodon muelleri (Dozy & Molk.) Sande Lac.

Figure 3. 

Liverworts and their habit A Bazzania uncigera (Reinw., Blume & Nees) Trevis B Pycnolejeunea grandiocellata Steph C Caudalejeunea reniloba (Gottsche) Steph D Leptolejeunea epiphylla (Mitt.) Steph E Bazzania densa (Sande Lac.) Schiffn F Pallavicinia lyellii (Hook.) Gray G Drepanolejeunea pentadactyla (Mont.) Steph.

Figure 4. 

Liverworts from Mount Tebu Forest Reserve, all in ventral view A Frullania gracilis (Reinw. et al.) Nees B Frullania trichodes Mitt C. Cololejeunea wightii Steph D Bazzania longicaulis (Sande Lac.) Schiffn E Bazzania albifolia Horik F Ptychanthus striatus (Lehm. & Lindenb.) Nees G Thysananthus spathulistipus (Reinw. et al.) Lindenb H Heteroscyphus coalitus (Hook.) Schiffn I Radula formosa (Spreng.) Nees J Spruceanthus polymorphus (Sande Lac.) Verd K Drepanolejeunea vesiculosa (Mitt.) Steph L Lejeunea sordida (Nees) Nees M Lepidolejeunea integristipula (J.B. Jack & Steph.) R.M. Schust N Pycnolejeunea grandiocellata Steph. (Scale = 0.5 mm).

Our study found that the diversity of moss species was higher than that of liverworts, a scenario similar to all the states in Peninsular Malaysia (Fig. 5). Reasons may be lower liverwort collecting, difficulty identifying liverwort species and lack of comprehensive field guides and local bryologists dealing with liverwort. The moss flora of Peninsular Malaysia has been well-studied taxonomically, in which exploration and species inventory of mosses have been more intensive and detailed. Thus far, 524 moss species have been reported from Peninsular Malaysia and all but Perlis and Malacca are well-represented with above 100 species (Yong et al. 2013; Ellis et al. 2019a, b). In comparison, only 491 taxa of liverworts are known from Peninsular Malaysia, suggesting that several States, particularly the northern regions, such as Perlis, Kedah and the east coast (Kelantan), have been under-collected and understudied (Lee and Gradstein 2021; Lee et al. 2022). The State of Pahang seems to be the centre of bryophyte diversity in Peninsular Malaysia (Fig. 5). The presence of major highlands and montane forests in Pahang often provides more favourable and more varied microhabitats for a rich bryophyte flora.

Figure 5. 

The number of bryophyte species reported from the States of Peninsular Malaysia.

Most of the bryophyte species in Mount Tebu are epiphytic, growing on the bark of tree trunks, on branches or tree stumps and the base of trees (Fig. 6). About half (49%) of ca. 1000 specimens examined were collected on trees (trunks, branches, twigs), while 22% were from leaves, 14% from rocks, 9% from soil or humus and 6% from rotten logs. About 18 species had broad substrate preferences and occurred on bark and branches of trees, leaves, soils and decaying logs. Others had more narrow preferences and occurred on only one substrate type, for example, Pallavicinia lyellii and Solenostoma comatum were always found on soil, Ephemeropsis tjibodensis and Leptolejeunea epiphylla occurred exclusively on leaves and Diphyscium mucronifolium grew only on rock (Appendix 1).

Figure 6. 

Habitats of bryophyte species of Mount Tebu Forest Reserve A lowland dipterocarp forest B area around the summit C–E bryophytes on tree bases, branches, trunks F on rocks G, H on leaves.

The distribution of the bryophyte species in Mount Tebu shows a distinct elevational differentiation from sea level to the mountain’s summit (Fig. 7). About half of the moss species have wide elevational ranges and occur from the lowlands to the summit of Mount Tebu. The remaining half of the species have more narrow elevational ranges and are restricted to a lower range, below 500 m. Liverwort species have wider elevational ranges and occur in all elevation belts. However, both groups show a similar trend where most of the species are elevational generalist species, occurring in most rainforest belts and lowland specialists, being found only below 500 m. Of 189 taxa, only 29 species are restricted to the submontane rainforest and occur exclusively at 700–1000 m a.s.l. For example, Acroporium condensatum and Mastopoma uncinifolium are obligate highland species known only from Cameron Highlands, Mount Jerai and Mount Tebu (this study) (Tixier 1980; Yong et al. 2006). Other moss species typical of high elevations found in Mount Tebu are Campylopus exasperatus, Leucoloma molle, Pogonatum cirratum subsp. macrophyllum, Acroporium stramineum and Trichosteleum saproxylophilum and liverworts are Frullania gracilis, F. trichodes, Cheilolejeunea ceylanica, C. trifaria, Cololejeunea aequabilis, C. appressa, C. equialbi, C. falcata, C. inflectens, C. metzgeriopsis, C. obliqua, C. ocelloides, C. sigmoidea, C. stephanii, Drepanolejeunea dactylophora, Ptychanthus striatus, Schistochila aligera, Spruceanthus polymorphus and Tuyamaella molischii.

Figure 7. 

The elevational distribution of bryophyte taxa found in Mount Tebu Forest Reserve.

We want to thank Mr Syamsul Bahri Mahammud and Mr Mat Rafi Daud, our local nature guides, for their invaluable assistance during the field sampling in Mount Tebu Forest Reserve and to Mr Baizul Hafsyam Badli Sham, Ms Noor Shahirah Ibrahim and Mr Muhammad Fatihah Syafiq for helping and support during the fieldwork. We extend our gratitude to Matt von Konrat, the subject editor, as well as Anders Hagborg and two anonymous reviewers, whose invaluable comments greatly improved earlier drafts of the manuscript.