English: Taxonomy and biogeography of the genus Colura (Dumort.) Dumort. (Lejeuneaceae, Marchantiophyta) in Tropical East Asia
Colura (Dumort.) Dumort. was coined by Dumortier (1835) in Recueil d’Observations sur les Jungermanniacees and C. calyptrifolia (Hook.) Dumort. was used as a holotype. This genus belongs to Lejeuneaceae, which is the largest family of the liverworts (Gradstein, 2013). The Colura was found in all Floristic regions except USA (Fig. 1), comprising 76 taxa, of which 39 endemic taxa are also reported. Tropical East Asia is the region where Pocs (1996) had reported the highest number, both total and endemic, of taxa in the area.
Species of Colura are typically recognized by their leaf morphology; the presence of lobules forming an apical sac with an aperture mechanism consisting of a valve and a hinge. This sac varies greatly in sizes and shapes, and may function a water retention and in some species as zoophagy (Barthlott et al., 2000). Most species of Colura can be considered being epiphyllous, but many of them can occur likewise on small twigs. Colura species grow in well-illuminated sites from the lowlands to about 4,000 m above sea level. A few species (e.g. C. ornithocephala) of this genus, especially those from high montane habitats, were never observed on leaves, but seem to inhabit exclusively on thin twigs or sometimes on the bark of thicker tree trunks. (Pocs, 1996; Heinrichs et al., 2012).
Two critically endangered Colura species (C. heimii Jovet-Ast and C. obesa Jovet-Ast) found in Reunion Island, where is the largest island of the Mascarene Archipelago in the western Indian Ocean, were listed in IUCN Red List (Ah-Peng et al., 2012). In addition, Gradstein, Raeymaekers and Churchill (2000) reported common taxa such C. clavigera be as an indicator of undisturbed rainforest in Tropical America (including Mexico).
Tropical East Asia region consists Myanmar, Laos, Cambodia, Vietnam, Thailand, Malaysia, Singapore, Brunei, the Philippines, Western Indonesia, Southern China (north to 30°N), plus the Ryukyu Islands of Japan and the Andaman as well as Nicobar Islands of India. (Corlett, 2009)
Fig. 1. Floristic regions where epiphyllous bryophyte occur. USA: The SE part of United States. MEA: Mesoamerica from Mexico to Panama. ANT: The Antilles. GUI: Guyanas, including the Venezuelan and Brasilian parts of Guyana Highlands. LSA: Amazonia and other lowland parts of South America, such as Choco and the Orinoco basin. SBR: The southeastern highlands of Brazil, with Paraguay and the Province Misiones in Argentina. AND: The Andes. TSA: Temperate South America. MAC: Macaronesia (excluding Cape Verde Islands). WAF: West Afirica from Guinea to the Congo. CAF: Central Africa including Zaire, Rwanda and Burundi. EAF: East Africa from Ethiopia to Mozambique. SAF: South Africa. MAD: Madagascar and the Seychelles, Comoro and Mascarene Islands. IND: India with the Himalayas, Khasia Hills (but excluding Andaman and Nicobar Islands) and lowland Bangladesh and Sri Lanka. OAS: East Asia including China, Taiwan, South Korea and Southern Japan with the Ryukyu Islands. ICH: Indochina from Chittagong Hills of Bangladesh to Vietnam. MAL: The Malesian Archipelago including the Andaman and Nicobar Islands and the Malay Peninsula but excluding West Irian. MEL: Malanesia, including the whole New Guinea, New Britain, the Bismarck and Solomon Islands, New Hebrides and New Caledonia. AUA: Australasia, including Australia, Tasmania, New Zealand and the neighboring islands. OCE: Oceania, the Pacific from Volcano and the Carolines to Hawaiian Islands and from Fiji to Easter Islands (Pocs, 1996).
The Hypothesis of Work
How many species of the genus Colura (Dumort.) Dumort. are there in Tropical East Asia?
This Present Work Mainly Aims
1. To investigate morphology, ecology and distribution of all taxa of the genus Colura on Tropical East Asia
2. To enumerate taxa account and accepted names of Colura in Tropical East Asia
1. The description, ecological data and distribution patterns of all taxa of the genus Colura on Tropical East Asia will be provided.
2. The synoptic account of Colura species will be enumerated.
Checklists in Asia
Herzog (1921) investigated liverworts in Moluccan Island, Indomalaya. Two Colura species was found, viz. C. brevistyla Herz. and C. javanica Steph., the first one was the new to science.
Mizutani (1961) revision studied of Lejeuneaceae in Japan, Colura tenuicornis and C. meijeri were found. Description, key to species and illustration were added.
Kitagawa (1969) collected hepatic plants from Penang, Malaysia. Two species of Colura was reported, viz. C. acutifolia Jovet-Ast and C. corynephora (Nees) Trev.
Pocs and Ninh (2005) collected the hepatic plants from Vu Quang Nature Reserve in central Vietnam and found Colura brevistyla Herz. as be as a new record to Indochina, furthermore, C. corynephora (Nees) Trev. as well was listed in this record.
Zhu (2006) reported a checklist of liverwort, hornwort and takakiophytes from China, of which seven taxa of Colura (C. acroloba, C. ari, C. conica, C. corynephora, C. inuii, C. karstenii and C. tenuicornis) were listed.
Lai, Zhu and Chantanaorrapint (2008) reported an updated checklist of liverwort and hornwort in Thailand. Among the account there are 6 species of Colura, viz. C. acroloba (Mont. ex Steph.) Ast, C. conica (Sande Lac.) K.I. Goebel, C. corynephora (Nees et al.) Trevis., C. siamensis Jovet-Ast, C. superba and C. tixieri.
Soderstrom, Gradstein and Hagborg (2010) reported 15 taxa of Colura in Java, including 12 taxa that be a good species or accepted except 3 taxa (C. denticulata Jovet-Ast, C. imperfecta Steph. and C. junghuhniana (Steph.) Steph.) that be probably a good species.
Chuah-Petiot (2011) listed the account and phytogeographical data of liverworts and hornworts of all states in Malaysia. Seventeen species of Colura were found and enumerated.
Wang, Lai and Zhu (2011) updated checklist and floristic accounts of liverworts as well as hornworts in Taiwan. The present list contains 512 species of liverwort belonging to 116 genera in 52 families, including three taxa of Colura (viz. C. acroloba (Mont. ex Steph.) Jovet-Ast, C. inuii Horik. and C. tenuicornis (Evans) Steph.)
Dey & Singh (2012) investigated epiphyllous liverworts of Eastern Himalaya, which Colura tenuicornis was reported in this study.
Pocs (2012) have small collected epiphyllous liverworts from Laos, which Colura tenuicornis (Evans) Steph. was reported with this list.
Pocs and Ninh (2012) reported epiphyllous liverworts from Cat Tien National Park of Southern Vietnam. They found 21 species including two species of Colura, viz. C. conica (Sande Lac.) Goebel and C. ornata Goebel, the latter was a new to Vietnam.
Pocs (2013) investigated the genus Colura in New Guinea and in neighboring areas. This paper based on the collection made in 1981 by a joint expedition of the University of Helsinki and the Humboldt State University of Arcata and on other collections, there are 25 species present. The phytogeographical evaluation of all species was provided. The description and illustration of four new to science species (C. koponenii, C. medusa, C. mizu-tanii and C. norrisii) were equipped. Furthermore, he as well treated C. simplicior Jovet-Ast (1983) to be a synonym of C. denticulata. Jovet-Ast (1954).
Pocs, Luong and Ho (2013) examined epiphyllous liverworts in Bidoup-Nui Ba National Park, Vietnam. Among the 125 samples containing 43 species belong to Lejeunea-ceae, both Colura acroloba (Mont. ex Steph.) and C. superba (Mont.) were found. These two species distribute particularly in Giang Ly station, where the altitude is 1481-1500 m a.s.l.
Checklists in Oceania
Thiers (1987) reported ten taxa of Colura in Australia, viz. C. acroloba (Mont.) Steph., C. ari Steph., C. australiensis Jovest-Ast, C. bisvoluta Herz. et Jovet-Ast, C. conica (Sande Lac.) Goeb., C. crispiloba Jovest-Ast, C. pulcherrima var. bartlettii Jovet-Ast, C. queenslandica B. Thiers, C. saccophylla Hodgs. et Herz. and C. simplicior Jovet-Ast. Among them, three species (C. acroloba, C. ari and C. conica) were first reportedly seen according to this investi-gation, especially C. queenslandica sp. nov. was new to science. A key to all ten taxa was provided. Moreover she additionally validate Colura sect. Lingua Jovet-Ast ex Thiers.
Pocs and Streimann (2006) collected five taxa of Colura, viz. C. apiculata (Schiffn.) Steph., C. leratii (Steph.) Steph., C. crispiloba Jovet-Ast, C. ornata Goebel and C. queenslan-dica Thiers in Queensland, New South Wales, and Victoria states as well as the Austral Capital Territory.
Pocs and Eggers (2007) investigated ten species of Colura from the Fiji archipelago. Three new to Fiji (C. crispiloba Jovet-Ast, C. cristata Jovet-Ast and C. queenslandica Thiers) as well as C. vitiensis Pocs et Eggers which was a new to science were added in this paper. The photograph and illustration of some species and together with dichotomous key to all species were provided.
Pocs et al. (2011) reported Colura brevistyla Herz. and C. tenuicornis (Evans) Staph. be as new record taxa for the Fiji Islands. C. acroloba also was listed in extended distribution records from new islands within Fiji.
Soderstrom et al. (2011) accounted the checklist of liverworts and hornworts of Fiji, including the islands Viti Levu, Vanua Levu, Taveuni, Kadavu, Ovalau and surrounding smaller islands. The checklist of 14 species which belong to genus Colura were provided; furthermore the synonymous data were enumerated.
Checklists in Amarica
Engel (1978) had taxonomically and phytogeographically studied of Hepaticae and Anthocerotae in Brunswick Peninsula, Chile. Reportedly, three species of Colura were found, viz. C. calyptrifolia (Hook.) Dumort., C. naumannii (Schiffn.) Steph. and C. patagonica Jovet-Ast.
Dauphin et al. (1998) reported Colura verdoornii Herz. et Jovet-Ast as new to Costa Rica, Tropical America. This species was previously known from Malaysia and Borneo, but it probably has a pantropical distribution, but not been reported from Africa.
Checklists in Africa
Jones and Harrington (1983) reported phytogeographical data of hepatic plant in Sierra Leone and Ghana, of those taxa, three species of Colura (C. digitaris (Mitten) Steph., C. dusenii (Steph.) Steph. and C. tenuicornis (Evans) Steph.) were included.
Pocs (1993) found new Colura from Comoro Archipelago and coin it as C. hattoriana Pocs in this paper, including description, type specimens, illustration and note.
Fischer (1999) reported five species of Colura from Aberdare mountain, Kenya, viz. C. berghenii Jovet-Ast, C. calyptri-folia (Hook.) Dumort., C. hedbergiana Pocs, C. tenuicornis (Evans) Steph. and C. zoophaga Fischer, the latter species was a new to science.
Wigginton (2001) investigated about habitat and location for 64 taxa that belong to Leajeuneaceae in Mulanje Mountain, Malawi. There are four Colura spp. which was reported in this investigation, viz. C. calyptrifolia , C. digitalis, C. tenuicornis and C. usambarica.
Chuah-Petiot and Pocs (2003) treated Colura zoophaga Fischer (1999) to be a synonym of C. kilimanjarica Pocs et Jovet-Ast which they was found in same place, Aber-dare Mountain, Kenya and in addition they grew together on the twig of Cliffortia nitidula (Chuah-Petiot and Pocs, 2003; Fischer, 1999).
Muller and Pocs (2007) investigated epiphyllous bryophyte of Bioko Island (Equatorial Guinea), biogeography of three species of Colura (viz. C. calderae Pocs, C. digitalis (Mitt.) Steph. and C. hattoriana Pocs, which the first one was a new to science) were provided.
Wigginton (2009) investigated liverworts and hornworts of sub-Saharan Africa, including the East African Islands. There are 22 taxa of Colura were reported; moreover, synonymous taxa as well were enumerated.
Hylander, Pocs and Nemomissa (2010) reported Colura digitalis (Mitt.) Steph. and C. tenuicornis (Mitt.) Steph. from southwest Ethiopian montane forests, in addition, ecological and biogeography were included.
Pocs (2010) reported new bryophyte taxa in Mayotte Island, among them, Colura heimii Jovet-Ast was included.
Pocs (2011) reported four species of Colura, viz. C. calderae Pocs, C. digitalis (Mitten) Steph., C. tenuicornis (Evan) Steph. and C. thomeensis Pocs in Sao Tome island. Among them, C. calderae was a new to Sao Tome and C. thomeensis was a new to science.
Pocs (2012) investigated morphological character of leafy liverworts and found endogenous gemma in Cololejeunea bifalcata Pocs, C. crateris Pocs, Colura calderae Pocs and Co. vitiensis Pocs et J. Eggers
1. Literature to which the previous investigations releveant will be revised.
2. Morphological study based on field trip specimens and herbarium specimens, description concerning vegetative part and reproductive part will be investigated in Bryology laboratory, PSU Herbarium, Department of Biology, Faculty of Science, Prince of Songkla University.
3. Illustrations of each taxa found in this investigation will be provided.
4. Hard copy format will be publicized.
The Research Period
Three years in the Ph.D. course period.
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Barthlott, W., Fischer, E., Frahm, J.-P. and Seine, R. 2000. First experimental evidence for zoophagy in the hepatic Colura. Plant Biology 2 (1): 93-97.
Chuah-Petiot, M.S. 2011. A checklist of hepaticae and anthocerotae of Malaysia. Polish Botanical Journal 56 (1): 1-44.
Chuah-Petiot, M.S. and Pocs, T. 2003. East African Bryophytes XIX. a contribution to the Bryflora of Kenya. Acta Botanica Hungarica 45 (1-2): 53-64.
Corlett, R.T. 2010. The Ecology of Tropical Asia. Oxford University Press Inc., New York.
Dauphin, G., Gradstein, S.Rob., Bernecker-Lucking, A. and Morales, M.I. 1998. Additions to the hepatic flora of Costa Rica II. Lindbergia 23: 74-80.
Dey, M. and Singh, D.K. 2012. Epiphyllous Liverworts of Eastern Himalaya. St. Joseph’s Press, Triruvananthapuram.
Dumortier, B.C. 1835. Jungermanniacees: Fascicule I.- Revision des genres. Imprimerie de J.-A. Blanquart., Tournay.
Engel, J.J. 1978. A taxonomic and phytogeographic study of Brunswick Peninsula (Strait of Magellan) Hepaticeae and Anthocerotae. Fieldiana Botany 41: 247-248.
Fischer, E. 1999. A new soecies of Colura (Lejeuneaceae) from the Aberdare Mountains/ Kenya. Tropical Bryology 16: 205-208.
Gradstein, S.R. 2013. A classification of Lejeuneaceae (Marchantiophyta) based on molecular and morphological evidence. Phytotaxa 100 (1): 6-20.
Heinrichs, J., Dong, S., Yu, Y., Schafer-Verwimp, A., Pocs, T, Feldberg, K., Hentschel, J., Schmidt, A.R. and Schneider, H. 2012. A 150 year-old mystery solved: Transfer of the rheophytic endemic liverwort Myriocolea irrorata to Colura. Phytotaxa 66: 55-64.
Herzog, Th. 1921. Die Lebermoose der 2 Freiburger Molukkenexpeditionen und einige neue Arten der engeren Indomalaya. In: Botanischen Centralblatt. G. Regierungsrat. pp. 318-332. Verlag von C. Heinrieh Dresden – N.
Hylander, K., Pocs, T. and Nemomissa, S. 2010. Liveworts of southwest Ethiopian montane forests: ecological and biogeographical notes. Journal of Bryology 32: 92-100.
Jones, E.W. and Harrington, A.J. 1983. The Hepatics of Sierra Leone and Ghana. Bulletin of the British Museum (Natural History) 11 (3): 215-289.
Kitagawa, N. 1969. A small collection of Hepaticae from Penang, Malaysia. Bulletin of Nara University of Education 18 (2): 27-43.
Lai, M.J., Zhu, R.L. and Chantanaorrapint, S. 2008. Liverworts and horworts of Thailand: an updated checklist and bryofloristic accounts. Annales Botanici Fennici 45: 321-341.
Mizutani, M. 1961. A revision of Japanese Lejeuneaceae. Journal of the Hattori Botanical Laboratory: Devoted to Bryology 24: 235-237.
Muller, F. and Pocs, T. 2007. A contribution to the knowledge of epiphyllous bryophytes of Bioko Island (Equatorial Guinea), including additional remarks on non-epiphyllous species. Journal of Bryology 29: 81-94.
Pocs, T. 1993. New or little known epiphyllous liverworts, IV. two new Cololejeuneoideae from the Comoro Archipelago. Journal of the Hattori Botanical Laboratory 74: 45-57.
Pocs, T. 1996. Epiphyllous liverwort diversity at worldwide level and its threat and conservation. Anales del Instituto de Biologia de la Universidad Nacional Autonoma de Mexico, Serie Botanica 67 (1): 109-127.
Pocs, T. 2010. East African bryophytes, XXVI. new records from Mayotte (Maore) Island (French Comoro). Acta Bryologica Asiatica 3: 105-114.
Pocs, T. 2011. New or little known epiphyllous liverworts, XIV. the genus Colura (Lejeuneaceae) in Sao Tome Island, with the description of Colura thomeensis sp. nov. The Bryologist 114 (2): 362-366.
Pocs, T. 2012. Endogenous gemmae in certain Lejeuneaceae (Marchaniophyta). The International Journal of Plant Reproductive Biology 4 (2): 101-105.
Pocs, T. 2012. New or little known epiphyllous liverworts, XVI. a small collection from Laos. Acta Biologica Plantarum Agriensis 2: 5-10.
Pocs, T. 2013. The genus Colura (Lejeuneaceae) in New Guinea and in the neighboring areas. Chenia 11: 12-38.
Pocs, T. and Eggers, J. 2007. Bryophytes from the Fiji Island, II. an account of the genus Colura, with a description of C. vitiensis sp. nov. Polish Botanical Journal 52(2): 81-92.
Pocs, T. and Ninh, T. 2012. New or little known epiphyllous liverworts, XVII. records from the Cat Tien National Park, Southern Vietnam. Acta Biologica Plantarum Agriensis 2: 11-19.
Pocs, T. and Ninh, T. Contribution to the Bryoflora of Vietnam, VI. on the liverwort flora of Vu Quang Nature Reserve. Acta Botanica Hungarica 47 (1-2): 151-171.
Pocs, T. and Streimann, H. 2006. Contributios to the Bryoflora of Australia, I. Tropical Bryology 27: 19-24.
Pocs, T., Luong, T.T. and Ho, B.C. 2013. New of little known epiphyllous liverworts, XVIII. records from the Bidoup-Nui Ba National Park, Vietnam, with the description of Drepanolejeunea bidoupensis, sp. nov. Cryptogamie, Bryologie 34 (3): 287-298.
Pocs, T., Sass-Gyarmati, A., Naikatini, A., Tuiwawa, M., Braggins, J., Pocs, S. and von Konrat, M. 2011. New liverwort (Marchantiophyta) recods for the Fiji Islands. Telopea 13 (3): 455-494.
Soderstrom, L., Gradstrin, S. and Hagborg, A. 2010. Checklist of the hornworts and liverworts of Java. Phytotaxa 9: 53-149.
Soderstrom, L., Hagborg, A., Pocs, T., Sass-Gyarmati, A. Brown, E., von Konrat, M. and Renner, M. 2011. Checklist of hornworts and liverworts of Fiji. Telopea 13 (3): 405-454.
Thiers, B.M. 1987. A preliminary accout of Colura (Hepaticae, Lejeuneaceae) in Australia. Brittonia 39 (2): 175-179.
Wigginton, M. 2009. Checklist and distribution of the liverworts and hornworts of sub-Saharan Africa, including the East African Islands. Tropical Bryology Research Reports 8: 1-116.
Wigginton, M.J. 2001. British Bryological Society expedition to Mulanje Mountain, Malawi. 15. Lejeuneaceae, and the occurrence and frequency of foliicolous taxa. Tropical Bryology 20: 83-94.
Zhu, R.L. 2006. New checklist of Chinese liverworts, hornworts, and takakiophytes. (PDF file). Bryological Laboratory, School of Life Science, East China Normal University, Shanghai.