DEFINING TURTLE DIVERSITY PROCEEDINGS OF A WORKSHOP ON GENETICS, ETHICS, AND TAXONOMY OF FRESHWATER TURTLES AND TORTOISES CAMBRIDGE, MASSACHUSETTS, 8–12 AUGUST 2005 EDITED BY H. BRADLEY SHAFFER, NANCY N. FITZSIMMONS, ARTHUR GEORGES, AND ANDERS G.J. RHODIN CHELONIAN RESEARCH MONOGRAPHS Number 4 – December 2007 Published by Chelonian Research Foundation in association with National Science Foundation, University of California at Davis, Harvard Museum of Comparative Zoology, Conservation International, and IUCN/SSC Tortoise and Freshwater Turtle Specialist Group CHELONIAN RESEARCH MONOGRAPHS Contributions in Turtle and Tortoise Research Series Editor ANDERS G.J. RHODIN Chelonian Research Foundation 168 Goodrich Street Lunenburg, Massachusetts 01462 USA [[email protected]] CHELONIAN RESEARCH MONOGRAPHS (ISSN 1088-7105) is an international peer-reviewed scientific publica- tion series for monograph-length manuscripts, collected proceedings of symposia, edited compilations, and other longer turtle-related research documents. The series accepts contributions dealing with any aspects of chelonian research, with a preference for conservation or biology. Manuscripts or edited compilations dealing with conservation biology, systematic relationships, chelonian diversity, geographic distribution, natural history, ecology, reproduction, morphology and natural variation, population status, husbandry, and human exploitation or conservation management issues are of special interest. Bibliographic and other reference materials are also of interest. Submit proposals for publications directly to A.G.J. Rhodin at the e-mail address above. Instructions for manuscript submission and detailed guidelines for authors are available at the same address.The series is published on an occasional basis by Chelonian Research Foundation. The publication of the current issue has benefited from the institutional support and association of National Science Foundation, University of California at Davis, Harvard Museum of Comparative Zoology, Conservation International, and IUCN/SSC Tortoise and Freshwater Turtle Specialist Group, for which the publisher and editors are grateful. Published Issues in Series 1. The Galápagos Tortoises: Nomenclatural and Survival Status. 1996. By PETER C.H. PRITCHARD. 85 pp. ISBN: 0-9653540- 0-8 (hard cover); 0-9653540-1-6 (soft cover). 2. Asian Turtle Trade: Proceedings of a Workshop on Conservation and Trade of Freshwater Turtles and Tortoises in Asia. 2000. Edited by PETER PAUL VAN DIJK, BRYAN L. STUART, AND ANDERS G.J. RHODIN. 164 pp. ISBN: 0-9653540-2-4 (hard cover); 0-9653540-3-2 (soft cover). 3. Biology and Conservation of Florida Turtles. 2006. Edited by PETER A. MEYLAN. 376 pp. ISBN: 0-9653540-4-0 (hard cover); 0-9653540-5-9 (soft cover). 4. Defining Turtle Diversity: Proceedings of a Workshop on Genetics, Ethics, and Taxonomy of Freshwater Turtles and Tortoises. 2007. Edited by H. BRADLEY S HAFFER, NANCY N. FITZSIMMONS, ARTHUR G EORGES, AND A NDERS G.J. RHODIN. 200 pp. ISBN 0-9653540-7-5 (hard cover); 0-9653540-8-3 (soft cover). Issue in Preparation 5. Conservation Biology of Freshwater Turtles and Tortoises. Edited by PETER PAUL VAN DIJK, PETER C.H. PRITCHARD, AND ANDERS G.J. RHODIN. CHELONIAN RESEARCH MONOGRAPHS are available for purchase from Chelonian Research Foundation. No overall series subscription rate is available, as individual monograph issues are priced separately. Contact Chelonian Research Foundation, 168 Goodrich Street, Lunenburg, MA 01462 USA (Phone: 978-582-9668; Fax: 978-582-6279; E-mail: [email protected]; Web Site: www.chelonian.org/crm) for prices, titles, and to place orders. Chelonian Research Foundation, founded in 1992, is a private, nonprofit tax-exempt organization under section 501(c)(3) of the Internal Revenue Code. Copyright © 2007 by Chelonian Research Foundation Date of Publication: 31 December 2007 DEFINING TURTLE DIVERSITY PROCEEDINGS OF A WORKSHOP ON GENETICS, ETHICS, AND TAXONOMY OF FRESHWATER TURTLES AND TORTOISES CAMBRIDGE, MASSACHUSETTS, 8–12 AUGUST 2005 EDITED BY H. BRADLEY SHAFFER, NANCY N. FITZSIMMONS, ARTHUR GEORGES, AND ANDERS G.J. RHODIN DEFINING TURTLE DIVERSITY PROCEEDINGS OF A WORKSHOP ON GENETICS, ETHICS, AND TAXONOMY OF FRESHWATER TURTLES AND TORTOISES CAMBRIDGE, MASSACHUSETTS, 8–12 AUGUST 2005 EDITED BY H. BRADLEY SHAFFER1, NANCY N. FITZSIMMONS2, ARTHUR GEORGES2, AND ANDERS G.J. RHODIN3 1University of California at Davis, Davis, California, USA; 2University of Canberra, Canberra, ACT, Australia; 3Chelonian Research Foundation, Lunenburg, Massachusetts, USA CHELONIAN RESEARCH MONOGRAPHS Number 4 December 2007 Chelonian Research Foundation COVER ILLUSTRATIONS Front Cover: Radiated tortoise, Astrochelys radiata (previously in Geochelone) (Testudinidae), in the wild at Cap Ste. Marie Special Reserve, southern tip of Madagascar. Photo by ANDERS G.J. RHODIN. Frontispiece: Mating radiated tortoises, Astrochelys radiata (previously in Geochelone) (Testudinidae), in captivity at Berenty Nature Reserve, southern Madagascar. Photo by ANDERS G.J. RHODIN. Back Cover (Upper): Big-headed Amazon River turtle, Peltocephalus dumerilianus (Podocnemididae), in captivity at Villavicencio, Colombia. Photo by RUSSELL A. MITTERMEIER. Back Cover (Lower): Musk turtle, Sternotherus odoratus (Kinosternidae), in the wild on the South Branch of the Potomac River, West Virginia, USA, showing particularly striking facial markings. Photo by PETER PAUL VAN DIJK. EDITORS FOR THIS VOLUME H. BRADLEY SHAFFER, Professor of Evolution and Ecology, and Center for Population Bioloy, University of California, Davis, CA 95616 USA [[email protected]]; Director, Center for Population Biology (UC Davis); Member, Steering Committee, IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. NANCY N. FITZSIMMONS, Senior Lecturer, Institute for Applied Ecology, University of Canberra, ACT 2601 Australia [[email protected]]; Member, Board of Directors, International Sea Turtle Society; Member, IUCN/ SSC Tortoise and Freshwater Turtle Specialist Group; Member, IUCN/SSC Marine Turtle Specialist Group; Member, IUCN/ SSC Crocodile Specialist Group. ARTHUR G EORGES, Professor of Applied Ecology, Institute for Applied Ecology, University of Canberra, ACT 2601 Australia [[email protected]]; Chair, ACT Flora and Fauna Committee; Member of Executive, ARC Environmental Futures Network; Member, Steering Committee, IUCN/SSC Tortoise and Freshwater Turtle Specialist Group; Member, Australian Biodiversity Information Facility (ABIF) Steering Committee; Member, ACT Natural Resource Management Committee. ANDERS G.J. RHODIN, Director, Chelonian Research Foundation, 168 Goodrich St., Lunenburg, MA 01462 USA [[email protected]]; Chair, IUCN/SSC Tortoise and Freshwater Turtle Specialist Group; Co-Chair, Turtle Conservation Fund; Member and Financial Officer, IUCN/SSC Marine Turtle Specialist Group; Member, Chairman’s Council, Conser- vation International; Editor, Chelonian Conservation and Biology; Editor, Chelonian Research Monographs. RECOMMENDED CITATION FORMATS For entire volume: SHAFFER, H.B., FITZSIMMONS, N.N., GEORGES, A., AND RHODIN, A.G.J. (Eds.). 2007. Defining Turtle Diversity: Proceedings of a Workshop on Genetics, Ethics, and Taxonomy of Freshwater Turtles and Tortoises. Chelonian Research Monographs No. 4, 200 pp. For included chapters: TURTLE TAXONOMY WORKING GROUP [BICKHAM, J.W., IVERSON, J.B., PARHAM, J.F., PHILIPPEN, H.D., RHODIN, A.G.J., SHAFFER, H.B., SPINKS, P.Q., AND VAN DIJK, P.P.]. 2007. An annotated list of modern turtle terminal taxa with comments on areas of taxonomic instability and recent change. In: Shaffer, H.B., FitzSimmons, N.N., Georges, A., and Rhodin, A.G.J. (Eds.). Defining Turtle Diversity: Proceedings of a Workshop on Genetics, Ethics, and Taxonomy of Freshwater Turtles and Tortoises. Chelonian Research Monographs No. 4, pp. 173-199. CHELONIAN RESEARCH MONOGRAPHS. Series Edited by ANDERS G.J. RHODIN. VOLUME NUMBER 4. Defining Turtle Diversity: Proceedings of a Workshop on Genetics, Ethics, and Taxonomy of Freshwater Turtles and Tortoises. Volume Edited by H. BRADLEY S HAFFER, NANCY N. FITZSIMMONS, ARTHUR G EORGES, AND A NDERS G.J. RHODIN. ISSN (monograph series): 1088-7105 ISBN (this volume): 0-9653540-7-5 (hard cover); 0-9653540-8-3 (soft cover). Published and Copyright © 2007 by Chelonian Research Foundation, Lunenburg, Massachusetts, USA. Printed by MTC Printing, Inc., Leominster, Massachusetts, USA. Defining Turtle Diversity: Proceedings of a Workshop on Genetics, Ethics, and Taxonomy of Freshwater Turtles and Tortoises H. Bradley Shaffer, Nancy N. FitzSimmons, Arthur Georges, and Anders G.J. Rhodin, Eds. Chelonian Research Monographs 4:5-6 • © 2007 by Chelonian Research Foundation TABLE OF CONTENTS Preface H. BRADLEY SHAFFER, ARTHUR GEORGES, PHILLIP Q. SPINKS, AND NANCY N. FITZSIMMONS ..........................................7 Participants and Contributors ..........................................................................................................................................9 Executive Summary H. BRADLEY SHAFFER, NANCY N. FITZSIMMONS, ARTHUR GEORGES, AND ANDERS G.J. RHODIN ...................................13 Genetic Studies of Freshwater Turtles and Tortoises: A Review of the Past 70 Years NANCY N. FITZSIMMONS AND KRISTEN M. HART ........................................................................................................15 From Molecules to Organisms: Research Applications of Modern Genetic Tools for Turtle Biology and Conservation