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A Biogeographic Synthesis of the Amphibians and Reptiles of Indochina

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A Biogeographic Synthesis of the Amphibians and Reptiles of Indochina BAIN & HURLEY: AMPHIBIANS OF INDOCHINA & REPTILES & HURLEY: BAIN Scientific Publications of the American Museum of Natural History American Museum Novitates A BIOGEOGRAPHIC SYNTHESIS OF THE Bulletin of the American Museum of Natural History Anthropological Papers of the American Museum of Natural History AMPHIBIANS AND REPTILES OF INDOCHINA Publications Committee Robert S. Voss, Chair Board of Editors Jin Meng, Paleontology Lorenzo Prendini, Invertebrate Zoology RAOUL H. BAIN AND MARTHA M. HURLEY Robert S. Voss, Vertebrate Zoology Peter M. Whiteley, Anthropology Managing Editor Mary Knight Submission procedures can be found at http://research.amnh.org/scipubs All issues of Novitates and Bulletin are available on the web from http://digitallibrary.amnh.org/dspace Order printed copies from http://www.amnhshop.com or via standard mail from: American Museum of Natural History—Scientific Publications Central Park West at 79th Street New York, NY 10024 This paper meets the requirements of ANSI/NISO Z39.48-1992 (permanence of paper). AMNH 360 BULLETIN 2011 On the cover: Leptolalax sungi from Van Ban District, in northwestern Vietnam. Photo by Raoul H. Bain. BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY A BIOGEOGRAPHIC SYNTHESIS OF THE AMPHIBIANS AND REPTILES OF INDOCHINA RAOUL H. BAIN Division of Vertebrate Zoology (Herpetology) and Center for Biodiversity and Conservation, American Museum of Natural History Life Sciences Section Canadian Museum of Nature, Ottawa, ON Canada MARTHA M. HURLEY Center for Biodiversity and Conservation, American Museum of Natural History Global Wildlife Conservation, Austin, TX BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 360, 138 pp., 9 figures, 13 tables Issued November 23, 2011 Copyright E American Museum of Natural History 2011 ISSN 0003-0090 CONTENTS Abstract.......................................................... 4 Introduction . ...................................................... 4 QualityoftheData................................................. 6 Geological History of Indochina . .................................... 7 Current Geography of Indochina . .................................... 9 Northern Uplands and Lowlands . .................................... 9 Annamite Uplands and Lowlands. ................................... 10 Lower Mekong Watershed ......................................... 11 CardamomUplandsandLowlands................................... 11 CoastsandIslands.............................................. 11 CurrentClimateandVegetationofIndochina............................. 12 Northern Uplands and Lowlands . ................................... 12 Annamite Uplands and Lowlands. ................................... 13 Lower Mekong Watershed ......................................... 14 CardamomUplandsandLowlands................................... 14 CoastsandIslands.............................................. 14 Paleoclimate and Paleovegetation of Indochina ............................ 15 Review of Regional Herpetofaunal Biogeography .......................... 16 MaterialsandMethods............................................... 19 SpeciesIncludedintheAnalysis....................................... 19 SubregionsofIndochina............................................ 20 Endemism and Biogeographic Affinities to Areas Outside Indochina ............. 21 Ecological Characteristics ........................................... 21 AnalyticalMethods................................................ 24 Results.......................................................... 24 (1) Relationship of Subregional Area to Species Richness, Endemism, and Survey Effort 24 (1a) Relationship of Species Richness and Endemism to Subregional Area ....... 24 (1b) Relationship of Survey Effort to Subregional Area . ................. 25 (2)TaxonomicDistributionofSpeciesRichnessandEndemism................ 25 (3) Geographic Distribution of Richness and Endemism...................... 26 (3a)SpeciesRichness............................................. 26 (3b) Endemism ................................................. 27 (4)TheOccurrenceofNonendemicFaunas............................... 27 (4a)SouthChina................................................ 30 (4b)MainlandSoutheastAsia...................................... 30 (4c)Malaya................................................... 30 (4d)Pan-AsiaFauna............................................. 30 (5) Ecological Characteristics of Indochina’s Herpetofauna. ................. 31 (5a)AllHerpetofaunas........................................... 31 (5b) Endemism ................................................. 33 (5c)EcologicalCorrelatesoftheFauna................................ 35 (6) Affinities Among Subregions . ................................... 35 (6a)AllHerpetofaunas........................................... 35 (6b)Amphibians................................................ 35 (6c)Snakes.................................................... 35 (6d) Lizards ................................................... 36 (6e)Turtles.................................................... 36 (6f) Endemics .................................................. 37 Discussion . ..................................................... 37 Regional Patterns of Diversity ........................................ 37 Northern Uplands and Lowlands . ................................... 37 2 The Annamites, Southern Lao Uplands, and Central–South Vietnam Lowlands . 44 The Lower Mekong Watershed and Cardamom Uplands and Lowlands ........ 48 CoastsandIslands.............................................. 50 General Patterns of Distribution ...................................... 51 EndemicSpecies................................................ 51 NonendemicFauna.............................................. 53 Characterization of Indochina’s Herpetofauna. ........................... 54 DirectionsforFurtherResearch....................................... 55 Conclusions....................................................... 59 Acknowledgments.................................................. 60 References........................................................ 61 Appendix1.DefinitionsofSubregionsofIndochina.......................... 89 Appendix 2A. Diversity and Distribution of Indochinese Herpetofauna............. 90 Appendix 2B. References and Comments: Diversity and Distribution of Indochinese Herpetofauna................................................ 118 Appendix 3. Species with Unconfirmed Occurrence in Indochina ................ 138 3 ABSTRACT Indochina (Laos, Cambodia, and Vietnam) houses over 600 species of amphibians and reptiles, roughly a quarter of which has been described within the last 15 years. Herein, we undertake the first biogeographic synthesis of the regional herpetofauna since the first half of the 20th century. We review the literature to measure and map species richness and endemism, the contributions of regional faunas, and ecological characteristics of Indochina’s amphibians (Anura, Caudata), and reptiles (Serpentes, Sauria, Testudines, Crocodylia). Dividing Indochina into 19 subregions defined by topography and geology, we estimate the similarity among the regional faunas and appraise the effects of area and survey effort on these comparative analyses. Variation in species composition is broadly correlated with topography, habitat complexity, and proximity to regions outside Indochina. Indochina’s herpetofauna is dominated (in decreasing order) by endemic species, widely distributed species, a South China fauna, and a biota centered in Thailand and Myanmar. Species richness is highest in amphibians and snakes, and peaks in upland forests. Endemism, highest among amphibians and lizards, also peaks in forests of the region’s northern uplands and Annamite Range. Endemic species occupy a narrower range of habitats than nonendemics. Patterns of richness and endemism are partially explained by ecological constraints: amphibians and lizards are more restricted to forests than snakes, turtles, and crocodiles; amphibians are more restricted to uplands, turtles to lowlands. We also assess biogeography in the context of Indochina’s geology, climate, and land cover. In northern Indochina, the Red River either acts as or coincides with an apparent dispersal barrier. Herpetofauna in northeastern upland areas are closely allied with fauna of southeastern China. In southern Indochina there is little evidence that the Mekong River represents a biogeographic barrier to the regional herpetofauna. The Annamite Range is composed of at least three distinct units and its elevated species richness and endemism are also noted in adjacent lowlands. Contribution of subtropical biota to Indochina’s fauna is significantly greater than that of tropical biota and there is little other evidence for intermixing at intermediate latitudes. Our results have implications for biogeography and conservation efforts, although they must be viewed in the context of rapidly evolving systematic knowledge of the region’s amphibians and reptiles. Future survey efforts, and the phylogenetic analyses that come from them, are essential for supporting regional conservation efforts, as they will better resolve the known patterns of amphibian and reptile richness and endemism. INTRODUCTION Morice, 1875; Tirant, 1885), but the first major herpetological field investigations were Indochina, which includes the modern carried out by Smith, starting in the 1920s Southeast Asian nations of Cambodia, Laos, (e.g., Smith,
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