The Conservation Biology of Tortoises Edited by Ian R. Swingland and Michael W. Klemens IUCN/SSC Tortoise and Freshwater Turtle Specialist Group and The Durrell Institute of Conservation and Ecology Occasional Papers of the IUCN Species Survival Commission (SSC) No. 5 IUCN—The World Conservation Union IUCN Species Survival Commission Role of the SSC 3. To cooperate with the World Conservation Monitoring Centre (WCMC) The Species Survival Commission (SSC) is IUCN's primary source of the in developing and evaluating a data base on the status of and trade in wild scientific and technical information required for the maintenance of biological flora and fauna, and to provide policy guidance to WCMC. diversity through the conservation of endangered and vulnerable species of 4. To provide advice, information, and expertise to the Secretariat of the fauna and flora, whilst recommending and promoting measures for their con- Convention on International Trade in Endangered Species of Wild Fauna servation, and for the management of other species of conservation concern. and Flora (CITES) and other international agreements affecting conser- Its objective is to mobilize action to prevent the extinction of species, sub- vation of species or biological diversity. species, and discrete populations of fauna and flora, thereby not only maintain- 5. To carry out specific tasks on behalf of the Union, including: ing biological diversity but improving the status of endangered and vulnerable species. • coordination of a programme of activities for the conservation of biological diversity within the framework of the IUCN Conserva- tion Programme. Objectives of the SSC • promotion of the maintenance of biological diversity by monitor- 1. To participate in the further development, promotion and implementa- ing the status of species and populations of conservation concern. tion of the World Conservation Strategy; to advise on the development of IUCN's Conservation Programme; to support the implementation of • development and review of conservation action plans and priorities the Programme; and to assist in the development, screening and monitor- for species and their populations. ing of projects for conservation action. • promotion of implementation of species-oriented conservation 2. To maintain an international network of independent volunteer members action plans and response to related issues. selected for their expertise in species conservation and to provide a forum • periodic evaluation of the status of species and biological diversity for the exchange of views and scientific information on species and conservation initiatives. populations of conservation concern. The publication is produced by IUCN—The World Conservation Union, in collaboration with The Durrell Institute of Conservation and Ecology at the University of Kent, The World Wide Fund for Nature—International, The American Association of Zoological Parks and Aquariums, and Wildlife Conservation International. © 1989 International Union for Conservation of Nature and Natural Resources Reproduction of this publication for educational and other non-commercial purposes is authorized without permission from the copyright holder, provided the source is cited and the copyright holder receives a copy of the reproduced material. Reproduction for resale or other commercial purposes is prohibited without prior written permission of the copyright holder. ISBN 2-88032-986-8 Published by: IUCN, Gland, Switzerland Printed by: Kelvyn Press, Inc., Broadview, Illinois, United States of America Cover photo: Hatching Aldabran tortoise (Geochelone gigantea). (Photo by I.R. Swingland.) The Conservation Biology of Tortoises Edited by Ian R. Swingland and Michael W. Klemens IUCN/SSC Tortoise and Freshwater Turtle Specialist Group and The Durrell Institute of Conservation and Ecology The results of "Operation Tortoise," a worldwide project conducted by the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group Occasional Papers of the IUCN Species Survival Commission (SSC) No. 5 IUCN—The World Conservation Union Contents Page Page Preface iii Homopus boulengeri 78 The Methodology of Conservation 1 Richard C. Boycott Michael W. Klemens Homopus femoralis 80 North America William Branch Gopherus agassizi 5 Homopus signatus 82 Kristin H. Berry Richard C. Boycott Gopherus berlandieri 8 Psammobates geometricus 85 Francis L. Rose and Frank W. Judd Ernst H.W. Baard Gopherus flavomarginatus 10 Psammobates oculifer 88 DJ. Morafka, G. Aguirre, and G.A. Adest Richard C. Boycott and William Branch Gopherus polyphemus 14 Psammobates tentorius 91 Joan E. Diemer William Branch South America Madagascar Geochelone carbonaria 17 Acinixys planicauda 94 Paul Walker Lee Durrell, Brian Groombridge, Geochelone chilensis 20 Simon Tonge, and Quentin Bloxam Paul Walker Geochelone radiata 96 Geochelone denticulata 22 Lee Durrell, Brian Groombridge, Paul Walker Simon Tonge, and Quentin Bloxam Geochelone elephantopus 24 Geochelone yniphora 99 Ian R. Swingland Lee Durrell, Brian Groombridge, Mediterranean Basin and Western Asia Simon Tonge, and Quentin Bloxam Regional Introduction 29 Pyxis arachnoides 103 David Stubbs Lee Durrell, Brian Groombridge, Testudo graeca 31 Simon Tonge, and Quentin Bloxam David Stubbs Aldabra Island Testudo hermanni 34 Geochelone gigantea 105 David Stubbs Ian R. Swingland Testudo horsfieldi 37 South and Southeast Asia David Stubbs Regional Introduction 111 Testudo kleinmanni 39 Edward O. Moll David Stubbs Geochelone elegans 113 Testudo marginata 41 Edward O. Moll David Stubbs Geochelone platynota 115 Sub-Saharan Africa Exclusive of South Africa Edward O. Moll Geochelone pardalis 43 Indotestudo elongata 116 Donald G. Broadley Edward O. Moll Geochelone sulcata 47 Indotestudo forstenii 118 Donald G. Broadley Edward O. Moll Kinixys belliana 49 Manouria emys 119 Donald G. Broadley Edward O. Moll Kinixys erosa 56 Manouria impressa 121 Donald G. Broadley Edward O. Moll Kinixys homeana 58 Appendix 1. List of editors and contributors 123 Donald G. Broadley Appendix 2. Classification of the Testudinidae 124 Kinixys natalensis 60 Bibliography Donald G. Broadley Kent R. Beaman, Russell L. Burke, Malacochersus tornieri 62 and Pamela K. Harpel-Burke Donald G. Broadley Introduction 126 South Africa North America 126 Regional Introduction 65 South America 139 William Branch Galapagos 145 Chersina angulata 68 Africa, Mediterranean, Indian Ocean 153 William Branch Asia 195 Homopus areolatus 72 Fossil 200 William Branch Homopus bergeri 75 William Branch ii Preface This contribution to the Occasional Papers of the IUCN Species and wisdom. In economic terms, the tortoise is an important Survival Commission (SSC) on the status and distribution of part of rural dynamics, being used for food in most parts of the the Testudinidae, or terrestrial tortoises, is the result of five world, as a musical instrument (maracas and banjo), as a scoop years of work by members of the IUCN/SSC Tortoise Specialist or water bail in boats, and canned as meat in parts of the Group (which has since become the Tortoise and Freshwater Mediterranean. The adults are often kept in village pens for Turtle Specialist Group) and is published on the occasion of the food and as a source of hatchlings, which are becoming a new First World Congress of Herpetology at the University of Kent, economic product of this traditional habit. These hatchlings September 1989. This work was stimulated by our lack of often fetch very high prices and are easily smuggled. Since knowledge regarding the forty species of tortoises which be- tortoises are capable of eating almost anything, they are the came apparent at our inaugural meeting at Oxford in October equivalent of long-lived pigs that do not require feeding or 1981 and our inability to answer many of the basic questions watering daily, and like pigs, every part of their body can be relating to their conservation. It was executed under the title used. "Operation Tortoise." The downside of the economics of tortoises is their rapidly We have provided the latest information on each species, escalating value to private and public collectors (which the including the Latin name, common names, description, taxon- European ban on bulk trade in 1984 and their general scarcity omy, geographic variation, status and distribution, habitat and has exacerbated) and the ways in which they are used as key ecology (particularly behaviour, reproduction, and feeding), species to defend areas programmed for development. In threats to survival, conservation reserves and recommenda- certain areas, the presence of tortoises has stopped the contin- tions, and current research. We have also provided the most ued destruction of habitats by off-road vehicles and bikes by the complete bibliography on the Testudinidae ever published. enforcement of conservation legislation. In parts of the Medi- In drawing up this report we have been concerned that the terranean, local villagers have been impeded from developing information is as up-to-date as possible, but we expect, indeed seaside areas for profit. The use of CITES regulations to defend hope, that it will be redundant in a few years as more and more these very vulnerable animals is of minimal value as long as people work on these fascinating animals. We have also been those charged with monitoring and policing the areas or coun- conscious of the needs of the local people and their interests, a tries where tortoises occur remain ignorant of the problem. vital part of any successful conservation