~RenA/Duel /'% 0' 1 /1 William E Editor · Duellman Museum ofNaturaI U· e Universi •. lstory Th versfty of 1979 Monog.-aph N0.7 Kansas THE SOUTH AMERICAN HERPETOFAUNA: ITS ORIGIN, EVOLUTION, AND DISPERSAL WILLIAM E. DUELLMAN EDITOR Museum of Natural History and Department of Systematics and Ecology The University of Kansas Lawrence, Kansas 66045, USA .. MONOGRAPH OF THE MUSEUM OF NATURAL HISTORY, THE UNIVERSITY OF KANSAS NUMBER 7 1979 MONOGRAPH OF THE MUSEUM OF NATURAL HISTORY, THE UNIVERSITY OF KANSAS Number 7, pages 1-485, 172 figures in text Issued December 28, 1979 © 1979by The Museum of Natural History, The University of Kansas, Lawrence, Kansas. All rights reserved. No part of this book may be reproduced in any form or by any means without permission in writing from the publisher. ISBN Number: 0-89338-008-3 Cover design by Linda Trueb PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS, USA PREFACE This volume is the result ofa symposium obvious omissions in this volume are chapters of the same title held on 11-13 August 1977 on the South American-North American her­ in conjunction with the joint annual meetings petofaunal relationships and the herpetofau­ of the Herpetologists' League and the Society nas of the Brasilian Highlands, the Atacama for the Study of Amphibians and Reptiles at Desert, and the caatinga and campos cerrados Lawrence, Kansas. I originally conceived the of Brasil. idea for such a symposium in August 1975 This volume is organized in much the same while returning from a 15-month sojourn in way as was the symposium, except that my South America. My interactions with many introductory chapter provides an overview of South American biologists during that trip the South American herpetofauna. Chapter 2 had convinced me that the time was appropri­ deals with the fossil record of amphibians and ate for a thorough discussion of ideas and reptiles in South America, .and Chapters 3 and presentation of our existing knowledge of 4 are concerned with the relationships of the the South American herpetofauna. The initial South American herpetofauna with those of response from colleagues was heartening, so Africa and Australia. The Quaternary bio­ during the following year the symposium was geography of the continent is the subject of organized. Unfortunately, owing to various Chapters 5-7. Treatments of regional herpeto­ circumstances not all subjects were covered; faunas are found in Chapters 8-15, and the Participants in. the Symposium on the South American Herpetofauna held in Lawrence, Kansas, 11-13 August 1977. Front row (left to right): Alberto Veloso M.,Beryl B. Simpson, Jaime E. Pefaur, Ana Maria Baez, Jose M. Cei, Second row: Lars Brundin, Thomas E. Lovejoy, Donn E. Rosen, jfirgen Haffer, Thomas H. Fritts, Ramon Formas, Raymond F. Laurent. Back row: William E. Duellman, James R. Dixon, Marinus S. Hoogmoed, John D. Lynch, W. Ronald Heyer, Michael J. Tyler, Jose M. Gallardo. final chapter is devoted to the conservation of and style. Rose Etta Kurtz retyped many the herpetofauna. pages of manuscript, and Rebecca A. Pyles I am grateful to the contributors to this painstakingly worked on the index. To all of volume for their scholarly efforts and for their these persons lowe a debt of gratitude for patience and understanding while it was their endeavors in behalf of this volume. being produced. For their participation in Throughout the early phases of develop­ the symposium, I thank the contributors and ment and organization of the symposium, as Lars Brundin, Thomas H. Fritts, W. Ronald well as during the production of this volume, Heyer, Jaime E. Pefaur, Donn E. Rosen, and Philip S. Humphrey, Director of the Museum Alberto Veloso M. Their enthusiastic partici­ of Natural History, has provided advice, en­ pation contributed a high level of scholarly couragement and support. Ronald K. Cal­ interaction, as well as much good cheer. gaard, Vice Chancellor for Academic Affairs, During the editing of this volume I called and George R. Waggoner, Associate Vice upon many colleagues to review manuscripts. Chancellor for International Programs, The The quality of the papers included herein University of Kansas, gave enthusiastic sup­ benefited from reviews by Avelino Barrio, port for the symposium. Richard F. Treece Lars Brundin, Richard Estes, Thomas H. of the Bureau of Conferences and Institutes Fritts, Steven Gorzula, W. Ronald Heyer, coordinated the logistics of the meetings. Philip S. Humphrey, Jean Lescure, Alan E. Without their interest and aid the symposium Leviton, John D. Lynch, Larry D. Martin, and this volume would not have been pos­ Braulio Orejas-Miranda, Jaime E. Pefaur, sible. Alan H. Savitzky, Beryl B. Simpson, Linda Financial support for bringing together Trueb, T. van der Hammen, Alherto Veloso the participants in the symposium was gen­ M. and Richard G. Zweifel. The drawings for erously provided by the National Science many of the papers were executed by Debra Foundation (DEB 76-16767), the World K. Bennett, Staff Illustrator of the Museum of Natural History at The University of Kansas. Wildlife Fund (WWF-US-71) and the Office Jaime E. Pefaur translated many of the sum­ of Academic Affairs, The University of Kansas. maries and edited the Spanish of others. Lin­ Support for the preparation of the index was da Trueb's competent editorial review of the provided by a grant from the General Re­ manuscripts is evident in their consistency search Fund of The University of Kansas. William E. Duellman Lawrence, Kansas September 6, 1979 CONTENTS 1. The South American Herpetofauna: A Panoramic View. William E. Duellman ~ --------_--_---_---_-_~ --__ --_--~_---------_---~_--_---~--_~--~_--- 1 2. The South American Herpetofauna: An Evaluation of the Fossil Record. Ana Maria Bdez and Zulma B. de Gasparini.:--------------~--------------------------~-- --~-------_- 29 3. Herpetofaunal Relationships Between Africa and South America. Raymond F. Laurent ~ ~----------------_--------- ----_---_--_---_---_--_-_-----~_-~_- 55 4. Herpetofaunal Relations of South America with Australia. Michael J. Tyler ~ ~ - ------. -_-------------_---__ 73 5. Quaternary Biogeography of Tropical Lowland South America. Jiirgen Haffer ~__~ __~ ----- ~ - ------_---------_--------------- -- 107 6. Late Cenozoic Environmental Changes in Temperate Argentina. Ana Maria Bdez and Gustavo Juan Scillato Yane -_---~_--_--_------ 141 7. Quaternary Biogeography of the High Montane Regions of South America. BerylB. Simpson __ ~ __ ~_~_~ .- ~_- --_---_--_--_-_ 157 8. The Amphibians of the Lowland Tropical Forests. John D. Lynch ~ ~ ~ ~ ~ ~_.______________________________________________________________________189 9. Origin and Distribution of Reptiles in Lowland Tropical Rainforests of South America. James R. Dixon -~---------~_~----.--------~------~ ~__ ~_~ ~ ~ ~___________________________ 217 10. The Herpetofauna of the Guianan Region. Marinus S. Hoogmoed -~----_~--~----~-_-- ~ --------_----------~-~--------------~----241 11. Origin and Distribution of the Herpetofauna of the Dry Lowland Regions of Northern South America. Carlos Bioero-Blanco and James R. Dixon ~ ~__ ~ ~ ~ ~ " ~ ~ 281 12. Composici6n, Distribuci6n y Origen de la Herpetofauna Chaquefia. Jose M. Gallardo ~ ~__ -----~---.------~---------_~_~ ~_~ ~ __ --_---~--------"-~------~-~----~----____ 299 13. ThePatagonian Herpetofauna. Jose M. Cei . ~ ~-_---------"----- ~ ~ -----"--__" ~__"__ ~____ 309 14. La Herpetofauna de los Bosques Temperados de Sudamerica. J. RamOn Formas --_---~~--_-----~-------~--~------_~---------------_-------------~--------------------~------~-----------_ 341 15. The Herpetofauna of the Andes: Patterns of Distribution, Origin, Differentiation and Present Communities. William E. Duellman ~ ~ ~ ~ ~ ~ ~ ~ ~__ ~___ 371 16. Refugia, Refuges and Minimum Critical Size: Problems in the Conservation of the Neotropical Herpetofauna. Thomas E. Looeioy ~ ~ ~ ---~------_-----~-_-~- ~-------------~----~-----------------~--- 461 Subject Index ~__ ~ ~ -_--~_--_-_-~------------_~-- ------------_---_-__ -_-_---_--_---------------~---- ~--_--_-_ 465 Taxonomic Index. ~ ----~ ~_~ -____________________________ 470 10. The Herpetofauna of the Guianan Region Marinus s. Hoogmoed Rijksmuseum van Natuurli;ke Historie Postbus 9517 2300 RA Leiden, The Netherlands Although this paper deals with a highland Guyana, several of which even at the present fauna, it is not limited to the reptiles and day remain unvisited. The most renowned of amphibians that occur at elevations of more these tepuis is Roraima with an altitude of than 1000 m, One of the main reasons for 2810 m, discovered in 1838 by Robert Schorn­ this is that "highlands" above 1000 m in the burgk and climbed for the first time in 1884 Guiana area are few and occupy only a very by 1m Thurn and Perkins. small part of the total area of the Guiana The first zoological collection ever made Shield. Another reason is that our knowledge near any tepui was assembled there in of the herpetofauna at higher elevations in 1842 by Richard Schomburgk. Other collec­ Guiana is still very fragmentary. These facts tions were made at the foot in the 1880's. prompted me to deal with the herpetofauna The first herpetological specimens from the of the entire Guiana Shield. summit were secured by 'Quelch and McCon­ The coast of the Guianas was discovered nell in 1894. In 1898 they made a second in 1499 by Alonso de Ojeda and Amerigo expedition to the summit plateau. The ma­ Vespucci. After the discovery of the so-called terial of these expeditions