TAXONOMY AND ECOLOGY OF AMPHIBIAN COMMUNITIES IN SOUTHERN VIETNAM: LINKING MORPHOLOGY AND BIOACOUSTICS TRAN, THI ANH DAO Taxonomy and ecology of amphibian communities in Southern Vietnam: Linking morphology and bioacoustics Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn vorgelegt von TRAN, THI ANH DAO aus Ho Chi Minh City, Vietnam Bonn, 2013 Angefertigt mit Genehmigung der Mathematisch- Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn 1. Gutachter Prof. Dr. Wolfgang Böhme 2. Gutachter Prof. Dr. Wolfgang Wägele Tag der Promotion:03/12/2013 Erscheinungsjahr: 2014 LIST OF CONTENTS LIST OF CONTENTS ........................................................................ i LIST OF TABLES .............................................................................. vi LIST OF FIGURES ............................................................................ viii LIST OF PLATES .............................................................................. xiv ABBREVIATIONS ............................................................................ 1 CHAPTER I. INTRODUCTION .......................................................... 3 1. Background ....................................................................... 3 2. Study area ......................................................................... 8 2.1. Topography ............................................................... 8 2.2. Climate ...................................................................... 10 2.3. Vegetation types ....................................................... 11 2.4. Vertebrate fauna in brief .......................................... 11 2.5. Threats to the biodiversity ........................................ 13 CHAPTER II. METHODS AND MATERIALS ...................................... 15 1. Sampling and field data collecting .................................... 15 1.1. Survey sites ............................................................... 15 1.2. Survey efforts ............................................................ 18 1.3. Sampling .................................................................... 19 1.4. Call recording ............................................................ 20 2. Analyses ............................................................................ 20 2.1. Morphological analyses ............................................ 20 2.2. Acoustic analyses ...................................................... 22 2.3. Statistical analyses .................................................... 24 CHAPTER III. RESULTS .................................................................... 25 i Bufonidae ................................................................... 26 Duttaphrynus melanostictus (Schneider, 1799) ... 26 Ingerophrynus galeatus (Günther, 1864) ............. 27 Megophryidae............................................................ 29 Brachytarsophrys intermedia (Smith, 1921) ........ 29 Leptobrachium leucops Stuart, Rowley, Tran, Le, and Hoang, 2011 ............................................ 32 Leptobrachium pullum (Smith, 1921) .................. 35 Leptolalax bidoupensis Rowley, Le, Tran, and Hoang, 2011 ......................................................... 38 Ophryophryne hansi Ohler, 2003 ......................... 41 Xenophrys major (Boulenger, 1908) .................... 43 Microhylidae .............................................................. 46 Calluella guttulata (Blyth, 1856) ........................... 46 Kaloula pulchra Gray, 1831 .................................. 47 Microhyla annamensis Smith, 1923 ..................... 48 Microhyla berdmorei (Blyth, 1856) ...................... 50 Microhyla heymonsi Vogt, 1911 .......................... 51 Microhyla pulchra (Hallowell, 1861) .................... 53 Microhyla sp. 1 ..................................................... 54 Microhyla sp. 2 ..................................................... 56 Dicroglossidae ............................................................ 58 Occidozyga lima (Gravenhorst, 1829) .................. 58 Occidozyga martensii (Peters, 1867) .................... 60 Fejervarya limnocharis (Gravenhorst, 1829) ........ 62 Limnonectes dabanus (Smith, 1922) .................... 64 Limnonectes poilani (Bourret, 1942) .................... 66 Ranidae ................................................................................ 69 Hylarana attigua (Inger, Orlov, and Darevsky, 1999) .................................................................... 69 Hylarana milleti (Smith, 1921) .............................. 71 Hylarana montivaga (Smith, 1921) ....................... 74 Hylarana nigrovittata (Blyth, 1856) ...................... 77 Odorrana gigatympana (Orlov, Ananjeva, and Ho, 2006) .............................................................. 80 ii Odorrana graminea (Boulenger, 1900) ................. 82 Rana johnsi Smith, 1921 ....................................... 84 Rhacophoridae ........................................................... 86 Chiromantis nongkhorensis (Cochran, 1927) ....... 86 Feihyla palpebralis (Smith, 1924) ......................... 88 Kurixalus baliogaster (Inger, Orlov, and Darevsky, 1999) .................................................... 90 Polypedates megacephalus Hallowell, 1861 ........ 92 Polypedates cf. mutus (Smith, 1940) .................... 94 Raorchestes gryllus (Smith, 1924) ........................ 95 Rhacophorus annamensis Smith, 1924 ................. 98 Rhacophorus calcaneus Smith, 1924 .................... 101 Rhacophorus robertingeri Orlov, Poyarkov, Vassilieva, Ananjeva, Nguyen, Nguyen, Geissler, 2012 ...................................................................... 103 Rhacophorus vampyrus Rowley, Le, Tran, Stuart, and Hoang, 2010 ....................................... 106 Theloderma asperum (Boulenger, 1886) .............. 107 Theloderma bambusicolum Orlov, Poyarkov, Vassilieva, Ananjeva, Nguyen, Nguyen & Geissler, 2012 ....................................................... 109 Theloderma gordoni Taylor, 1962 ........................ 111 Theloderma palliatum Rowley, Le, Hoang, Dau & Cao 2011 ........................................................... 113 Theloderma stellatum Taylor, 1962 ..................... 115 Theloderma truongsonense (Orlov & Ho, 2005) .. 117 2.4. Ecological niche segregation of amphibians in the Langbian Plateau ..................................................... 123 3. Biogeographic relationships of the amphibian communities among forests within and neiboring the Langbian Plateau ............................................................ 128 3.1. Biogeographic relationships of the amphibian communities among the forests within the Langbian Plateau ..................................................... 128 Brachytarsophrys intermedia ............................... 130 Leptobrachium leucops ........................................ 132 iii Leptobrachium pullum ......................................... 134 Leptolalax bidoupensis ......................................... 139 Xenophrys major .................................................. 144 Hylarana attigua ................................................... 146 Hylarana milleti .................................................... 148 Hylarana nigrovittata ........................................... 151 Chiromantis nongkhorensis ................................. 152 Kurixalus baliogaster ............................................ 154 Raorchestes gryllus .............................................. 156 Rhacophorus annamensis .................................... 158 Rhacophorus calcaneus........................................ 165 CHAPTER IV. DISCUSSION ............................................................. 167 1. Taxonomic problems and species complexes .................. 167 1.1. Brachytarsophrys intermedia ................................... 167 1.2. Hylarana nigrovittata complex ................................. 167 1.3. Polypedates leucomystax complex ........................... 168 1.4. Raorchestes gryllus ................................................... 169 1.5. Rhacophorus calcaneus and Rhacophorus robertingeri ............................................................. 170 2. Bioacoustics ..................................................................... 171 2.1. Vocal property and behavior similarities within the anuran families ................................................. 171 Megophryidae............................................................ 171 Ranidae ...................................................................... 173 Rhacophoridae ........................................................... 178 2.2. Multiple call types .................................................... 182 2.3. Chorus behavior ....................................................... 182 3. Diversity and biogeography of the amphibian fauna of the Langbian Plateau...................................................... 186 3.1. Diversity .................................................................... 186 3.2. Biogeographic relationships of the amphibian communities among the forests within and adjacent the Langbian Plateau ................................ 188 iv 4. Conservation potential ..................................................... 190 CHAPTER V. CONCLUSIONS AND OUTLOOK .................................