OXFORD MONOGRAPHS ON GEOLOGY AND GEOPHYSICS Series Editors H. Charnock J. R Dewey S. Conway Morris A. Navrotsky E. R. Oxburgh R. A. Price B. J. Skinner OXFORD MONOGRAPHS ON GEOLOGY AND GEOPHYSICS 1. DeVerle P. Harris: Mineral Resources Appraisal: Mineral Endowment, Resources, and Potential Supply: Concepts, Methods, and Cases 2. i. ]. Veevers (ed.): Phanerozoic Earth History of Australia 3. Yang Zunyi, Cheng Yuqi, and Wang Hongzhen (eds.): The Geology of China 4. Lin-Gun Liu and William A. Bassett: Elements, Oxides, and Silicates: High-Pressure Phases with Implications for the Earth's Interior 5. Antoni Hoffman and Matthew H. Nitecki (eds.): Problematic Fossil Taxa 6. S. Mahmood Naqvi and John J. W. Rogers: Precambrian Geology of India 1. Chih-Pei Chang and T. N. Krishnamurti (eds.): Monsoon Meteorology 8. Zvi Ben-Avraham (ed.): The Evolution of the Pacific Ocean Margins 9. Ian McDougall and T. Mark Harrison: Geochronology and Thermochronology by the 40Ar/39Ar Method 10. Walter C. Sweet: The Conodonta: Morphology, Taxonomy, Paleoecology, and Evolutionary History of a Long-Extinct Animal Phylum 11. H. J. Melosh: Impact Cratering: A Geologic Process (now in paperback) 12. J. W. Cowie and M. D. Brasier (eds.): The Precambrian-Cambrian Boundary 13. C. S. Hutchinson: Geological Evolution of South-East Asia 14. Anthony J. Naldrett: Magmatic Sulfide Deposits 15. D. R. Prothero and R. M. Schoch (eds.): The Evolution of Perissodactyls 16. M. Menzies (ed.): Continental Mantle 17. R. J. Tingey (ed.): Geology of the Antarctic 18. Thomas J. Crowley and Gerald R. North: Paleoclimatology (now in paperback) 19. Gregory J. Retallack: Miocene Paleosols and Ape Habitats in Pakistan and Kenya 20. Kuo-Nan Liou: Radiation and Cloud Processes in the Atmosphere: Theory, Observation and Modeling 21. Brian Bayly: Chemical Change in Deforming Materials 22. A. K. Gibbs and C. N. Barron: The Geology of the Guiana Shield 23. Peter J. Ortoleva: Geochemical Self-Organization 24. Robert G. 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Alan Holman: Pleistocene Amphibians and Reptiles in Britain and Europe PLEISTOCENE AMPHIBIANS AND REPTILES IN BRITAIN AND EUROPE J. Alan Holman New York • Oxford Oxford University Press 1998 Oxford University Press Oxford New York Athens Auckland Bangkok Bogota Bombay Buenos Aires Calcutta Cape Town Dar es Salaam Delhi Florence Hong Kong Istanbul Karachi Kuala Lumpur Madras Madrid Melbourne Mexico City Nairobi Paris Singapore Taipei Tokyo Toronto Warsaw and associated companies in Berlin Ibadan Copyright © 1998 by Oxford University Press, Inc. Published by Oxford University Press, Inc. 198 Madison Avenue, New York, New York 10016 Oxford is a registered trademark of Oxford University Press. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of Oxford University Press. Library of Congress Cataloging-in-Publication Data Holman, J. Alan, 1931- Pleistocene amphibians and reptiles in Britain and Europe / J. Alan Holman. p. cm. Includes bibliographical references and index. ISBN 0-19-511232-6 1. Amphibians, Fossil—Europe. 2. Reptiles, Fossil—Europe. 3. Amphibians, Fossil—Great Britain. 4. Reptiles, Fossil—Great Britain. 5. Paleontology—Pleistocene. I. Title. QE867.H63 1998 567'.S'094— dc21 97-30058 1 3 5 7 9 8 6 42 Printed in the United States of America on acid-free paper PREFACE The Pleistocene epoch, or Ice Age, an extended period of advancing and re- treating continental ice sheets, is characterized by striking climatic oscillations and sea level fluctuations. This age saw the rise and spread of humans and a great extinction of large mammals by the end of the epoch. In fact, the world today is essentially the product of dramatic changes that took place in the Pleis- tocene. Major works have dealt with Pleistocene mammals in North America and Europe and the "why" of the worldwide mammalian extinction. Moreover, a recent book (Holman, 1995c) has dealt with North American Ice Age am- phibians and reptiles. Yet no comparable work deals with British and European Pleistocene hcrpetofaunas. This book attempts to address this situation by fo- cusing on the Pleistocene amphibians arid reptiles in Britain and the European continent eastward through Poland, the Czech Republic, Hungary, Yugoslavia, and Greece. The book begins with an overview of the Pleistocene epoch and its signifi- cance, followed by a general discussion of the Pleistocene in Britain and Europe in Chapter 2, with emphasis on regional terms used to define Pleistocene chron- ological events. Chapter 3 looks at the pre-Pleistocene herpetofauna of the study area to set the stage for a discussion of the Pleistocene herpetofauna. A large portion of the book consists of a "Bestiary" (Chapter 4), a series of annotated taxonomic accounts of Pleistocene herpetological taxa from the region. Illustra- tions of bones that are especially useful in the identification of Pleistocene her- petological species arc included in many of these accounts, along with a discussion of important characters used in the identification of Pleistocene spe- cies. A list of locality numbers for each taxon in the Bestiary correlates with those of localities detailed in Chapter 5. These accounts contain lists of herpe- tological species, literature references, and. when appropriate, remarks about the site. Following this is the interpretive section. It begins with a discussion of her- petological species as paleoenvironmental indicators in Chapter 6. Next, Chapter 7 analyzes herpetological population adjustments to Pleistocene events in Britain and Europe. Chapter 8 follows with a discussion of extinction patterns in the vi PREFACE region, including a dialogue about the apparent evolutionary stasis of the Pleis- tocene herpetofauna compared with avian and mammalian populations. Finally, Chapter 9 discusses the differences and similarities between Pleistocene herpe- tological events in Britain and Europe and North America. This book is meant to be a college-level introduction to the study of Pleis- tocene amphibians and reptiles of Britain and Europe and a companion volume to Pleistocene Amphibians and Reptiles in North America (Holman, 1995c). It does not pretend to be a definitive taxonomic revision of Pleistocene herpeto- logical taxa or claim to cite every reference for each Pleistocene herpetological species found in the vast area covered by the book. It should, however, be a standard reference for anyone interested in Pleistocene flora or fauna as well as Pleistocene events in general. It should be especially useful for modern herpe- tologists, ecologists, and evolutionary biologists. I have tried to keep the ap- proach to the subject simple and direct and have purposely tried not to get involved in esoteric and convoluted arguments about Pleistocene events and patterns. East Lansing, Mich. J.A.H. ACKNOWLEDGMENTS I gratefully acknowledge the persons who have been especially helpful in the preparation of this book. Many have been hosts for my field and museum stud- ies, and others have loaned herpetological fossils or modern comparative skel- etons. All have supplied technical correspondence and/or publications: N. Ashton (London), W. Auffenberg (Gainesville), S. Balcon (Tunbridge-Wells), M. Benton (Bristol), G. Bohme (Berlin), S. Chapman (London), J. Clayden (East Runton-Cromer), J. Clack (Cambridge), A. Currant (London), R. Estes (San Diego), E. Fritz (Dresden), D. Harrison (Seven Oaks), M. Ivanov (Brno), T. Kemp (Oxford), T. v. Kolfschoten (Leiden), L. Kordos (Budapest), A. Milner (London), M. Mlynarski (Krakow), S. Parfitt (London), M. Paunovic (Zagreb), J. C. Rage (Paris), M. Roberts (London), Z. Rocek (Prague), B. Sanchiz (Ma- drid), A. Stuart (Norwich), Z. Szyndlar (Krakow), and M. Warren (Cromer). Diane Baclawski of the Geological Sciences Library at Michigan State Uni- versity helped gather Pleistocene herpetological literature. I thank Joyce Berry, Nancy Hoagland, Jessica A. Ryan, and the other editorial staff of Oxford University Press, New York, for their efforts in the production of this book. The National Geographic Society provided recent grants for field- work and museum studies for this work. I especially thank Teresa Petersen for the preparation of figures. Other artists are acknowledged in the figure legends. This page intentionally left blank CONTENTS 1 Introduction 3 2 The Pleistocene in Britain and Europe 6 3 The European Herpetofauna: Paleocene Through Pliocene 10 4 A Bestiary—Annotated Taxonomic Accounts 20 5 Pleistocene Herpetological Localities 130 6 Herpetological Species as Paleoenvironmental Indicators 201 7 Herpetological Population Adjustments in the Pleistocene of Britain and Europe 207 8 Extinction Patterns in the Herpetofauna of the Pleistocene of Britain and Europe 217 9 Pleistocene Events in the Herpetofaunas of Britain and Europe, and North America Compared 224 References 228 General Index 249 Taxonomic Index 251 Site Index 252
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