Taxonomy and Biogeography of the Genus Colura

Taxonomy and Biogeography of the Genus Colura Jiroat Sanngrattanaprasert   English: Taxonomy and biogeography of the genus Colura (Dumort.) Dumort. (Lejeuneaceae, Marchantiophyta) in Tropical East Asia Introduction Colura (Dumort.) Dumort. was coined by Dumortier (1835) in Recueil dObservations sur les Jungermanniacà ©es 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 Pà ³cs (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. (Pà ³cs, 1996; Heinrichs et al., 2012). Two critically endangered Colura species (C. heimii Jovet-Ast and C. obesa Jovet-Ast) found in Rà ©union 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 Chit tagong 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 (Pà ³cs, 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 Expect Results 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. Literature review 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. Pà ³cs 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. Sà ¶derstrà ¶m, 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. Pà ³cs (2012) have small collected epiphyllous liverworts from Laos, which Colura tenuicornis (Evans) Steph. was reported with this list. Pà ³cs and Ninh (2012) reported epiphyllous liverworts from Cà ¡t Tià ªn 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. Pà ³cs (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). Pà ³cs, Luong and Ho (2013) examined epiphyllous liverworts in Bidoup-Nà ºi Bà   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. Pà ³cs 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. Pà ³cs 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 Pà ³cs 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. Pà ³cs 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. Sà ¶derstrà ¶m 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. Pà ³cs (1993) found new Colura from Comoro Archipelago and coin it as C. hattoriana Pà ³cs 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 Pà ³cs, 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 Pà ³cs (2003) treated Colura zoophaga Fischer (1999) to be a synonym of C. kilimanjarica Pà ³cs 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 Pà ³cs, 2003; Fischer, 1999). Mà ¼ller and Pà ³cs (2007) investigated epiphyllous bryophyte of Bioko Island (Equatorial Guinea), biogeography of three species of Colura (viz. C. calderae Pà ³cs, C. digitalis (Mitt.) Steph. and C. hattoriana Pà ³cs, 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, Pà ³cs 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. Pà ³cs (2010) reported new bryophyte taxa in Mayotte Island, among them, Colura heimii Jovet-Ast was included. Pà ³cs (2011) reported four species of Colura, viz. C. calderae Pà ³cs, C. digitalis (Mitten) Steph., C. tenuicornis (Evan) Steph. and C. thomeensis Pà ³cs in Sà £o Tomà © island. Among them, C. calderae was a new to Sà £o Tomà © and C. thomeensis was a new to science. Morphological characters Pà ³cs (2012) investigated morphological character of leafy liverworts and found endogenous gemma in Cololejeunea bifalcata Pà ³cs, C. crateris Pà ³cs, Colura calderae Pà ³cs and Co. vitiensis Pà ³cs et J. Eggers Research Method 1. Literature to which the previous investigations releveant will be revised. 2. 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