Cover Page. The handle holds various files of this Leiden University dissertation.

Transcription

1 Cover Page The handle holds various files of this Leiden University dissertation. Author: Kok, Philippe Jacques Robert Title: Islands in the sky : species diversity, evolutionary history, and patterns of endemism of the Pantepui Herpetofauna Issue Date:

2 ISLANDS IN THE SKY: SPECIES DIVERSITY, EVOLUTIONARY HISTORY, AND PATTERNS OF ENDEMISM OF THE PANTEPUI HERPETOFAUNA PHILIPPE J. R. KOK

3 KOK, PHILIPPE JACQUES ROBERT ISLANDS IN THE SKY: SPECIES DIVERSITY, EVOLUTIONARY HISTORY, AND PATTERNS OF ENDEMISM OF THE PANTEPUI HERPETOFAUNA THESIS LEIDEN UNIVERSITY COVER PHOTO: KAMARKAWARAI-TEPUI, VENEZUELA PHILIPPE J.R. KOK. PRINTED BY: GILDEPRINT DRUKKERIJEN THE NETHERLANDS ISBN: All rights reserved. No parts of this book may be reproduced or transmitted in any form or by any means, electronical, mechanical, photocopying, recording, or otherwise, without the prior written permission of the author.

4 ISLANDS IN THE SKY: SPECIES DIVERSITY, EVOLUTIONARY HISTORY, AND PATTERNS OF ENDEMISM OF THE PANTEPUI HERPETOFAUNA Proefschrift ter verkrijging van de graad van Doctor aan de Universiteit Leiden, op gezag van de Rector Magnificus Prof. Mr. C.J.J.M. Stolker, volgens besluit van het College voor Promoties te verdedigen op dinsdag 28 mei 2013 klokke 16:15 uur door Philippe Jacques Robert Kok geboren te Watsa, Democratische Republiek Congo, in 1970

5 PROMOTIECOMMISSIE Promotors: Co-promotor: Overige leden: Prof. Dr. E. Gittenberger (Universiteit Leiden) Prof. Dr. E.F. Smets (Universiteit Leiden) Dr. M.S. Hoogmoed (Museu Paraense Emílio Goeldi, Belém) Prof. Dr. F. Bossuyt (Vrije Universiteit, Brussel) Prof. Dr. C.J. ten Cate (Universiteit Leiden) Dr. D.B. Means (Florida State University, Tallahassee) Prof. Dr. M.K. Richardson (Universiteit Leiden) Prof. Dr. M. Schilthuizen (Universiteit Leiden)

6 TABLE OF CONTENTS ACKNOWLEDGEMENTS 11 NEDERLANDSE INLEIDING EN SAMENVATTING 13 CHAPTER 1. GENERAL INTRODUCTION AND SUMMARY 19 The tepuis 21 Pantepui, the tepuis, and their geomorphological evolution 22 The Lost World 32 PhD objectives 32 Brief introduction to the selected taxa 36 Literature cited 45 MORPHOLOGY CHAPTER 2. A NEW HIGHLAND SPECIES OF ARTHROSAURA BOULENGER, 1885 (SQUAMATA: GYMNOPHTHALMIDAE) FROM MARINGMA TEPUI ON THE BORDER OF GUYANA AND BRAZIL 53 Zootaxa 1909: 1 15 Introduction 55 Material and methods 56 Results 58 Acknowledgments 70 Literature cited 70 Appendix 1. Comparative material examined 72 CHAPTER 3. LIZARD IN THE CLOUDS: A NEW HIGHLAND GENUS AND SPECIES OF GYMNOPHTHALMIDAE (REPTILIA: SQUAMATA) FROM MARINGMA TEPUI, WESTERN GUYANA 73 Zootaxa 1992: Introduction 75 Material and methods 75 Results 76 Acknowledgments 90 Literature cited 90 Appendix 1. Additional comparative material examined 92

7 CHAPTER 4. A NEW SPECIES OF OREOPHRYNELLA (ANURA: BUFONIDAE) FROM THE PANTEPUI REGION OF GUYANA, WITH NOTES ON O. MACCONNELLI BOULENGER, Zootaxa 2071: Introduction 95 Material and methods 95 Results 96 Acknowledgments 109 Literature cited 110 Appendix 1. Additional material examined 113 CHAPTER 5. A NEW SPECIES OF ANOMALOGLOSSUS (ANURA: AROMOBATIDAE) FROM THE PAKARAIMA MOUNTAINS OF GUYANA 115 Zootaxa 2660: Introduction 117 Material and methods 118 New species description 119 Acknowledgments 131 Literature cited 131 Appendix. Additional material examined 134 CHAPTER 6. A NEW HIGHLAND SPECIES OF PRISTIMANTIS JIMÉNEZ DE LA ESPADA, 1871 (ANURA: STRABOMANTIDAE) FROM THE PANTEPUI REGION, NORTHERN SOUTH AMERICA 137 Zootaxa 2934: 1 19 Introduction 139 Material and methods 141 New species description 142 Acknowledgments 159 Literature cited 160 Appendix. Additional material examined 163 CHAPTER 7. A NEW SPECIES OF ANADIA (REPTILIA, SQUAMATA) FROM THE VENEZUELAN LOST WORLD, NORTHERN SOUTH AMERICA 165 European Journal of Taxonomy 3: 1 18 Introduction 167 Material and methods 168 Results 169 Acknowledgements 182 References 183 Appendix. Additional material examined 186

8 CHAPTER 8. A REDESCRIPTION OF ANOMALOGLOSSUS PRADERIOI (LA MARCA, 1998) (ANURA: AROMOBATIDAE: ANOMALOGLOSSINAE), WITH DESCRIPTION OF ITS TADPOLE AND CALL 187 Papéis Avulsos de Zoologia 50: Introduction 189 Materials and methods 190 Results 191 Acknowledgments 211 Literature cited 211 Appendix. Specimens examined 214 CHAPTER 9. A NEW DIAGNOSIS AND DESCRIPTION OF ANOMALOGLOSSUS RORAIMA (LA MARCA, 1998) (ANURA: AROMOBATIDAE: ANOMALOGLOSSINAE), WITH DESCRIPTION OF ITS TADPOLE AND CALL 215 In press in South American Journal of Herpetology Introduction 217 Materials and methods 218 Results 220 Acknowledgments 240 References 241 Appendix. Specimens examined 245 MOLECULES CHAPTER 10. LOW GENETIC DIVERSITY IN TEPUI SUMMITS VERTEBRATES 247 Current Biology 22(15): R589 R590 Main text 249 References 252 Acknowledgments 252 Supplemental Information Area of study and taxon sampling Choice of markers DNA extraction, PCR, sequencing and sequence alignment Phylogenetic analyses Estimation of divergence time Supplemental Acknowledgments Supplemental References Supplemental Tables and Figures 258

9 CHAPTER 11. GENERAL CONCLUSION: THE LOST WORLD CONUNDRUM, INSIGHTS FROM PANTEPUI AMPHIBIANS AND REPTILES 269 Neotropical diversification, timing and evolutionary causes 271 Diversification in the Pantepui, current hypotheses 272 Lessons from Pantepui amphibians and reptiles 275 Test of hypotheses: towards a phylogenetically supported diversification model? 278 Conclusion 287 Literature cited 288 FUTURE PERSPECTIVES 295 CURRICULUM VITAE AND PUBLICATIONS 297 GLOSSARY OF GEOLOGICAL TERMS 303