= DIVERSITY. FOUNDER AND EDITOR - Craig Hassapakis ASSOCIATE EDITOR Jack W. Sites, Jr. | Pilagascar ted tortoises Hefpetofauna extifction Ys ‘. ‘ New Caled Rhacodactylus getkos : The Caribbeah : oe onpatote and ee (New Golimn) The trade in live reptiles and ampnitigs * mn This journal supports the programs of Partner¥ in Amphibian . %. » and Reptile Conservation (PARC) to promote e jon about | reptiles and amphibians and protection of their habitagy e G Visit www.ingenta.com to access journal Content online (full-text). Subscribers may register with ingenta for free electronic delivery via the Internet. Non-subscribers can obtain full-text articles as pay-per-views. Freely search ingenta's journal database (nearly 3.000 total journals) by keyword, title, and author (journal article abstracts available Online free through ingenta). The Iguanid Lizards of Cuba Edited by Lourdes Rodriguez Schettino “Goes far beyond a standard taxonomic handbook or catalogue treatment covering not only information on systematics, morphol- ogy, ecology, and biogeography, but also behavior, parasitology, and genetics. .. . A tremendous resource and reference.” —Kevin de Queiroz, Division of Amphibians and Reptiles, Na- tional Museum of Natural History, Smithsonian Institution “For herpetologists in general and an indispensable source for biological scientists and naturalists interested in Caribbean fauna.”—Choice Lourdes Rodriguez Schettino covers nearly every aspect of the 62 currently known iguanid species living in Cuba, including the iguana, the curly-tailed lizards, giant anoles, chameleons, and other anoline lizards. Drawing on more than 20 years of herpetological fieldwork, she summarizes existing knowledge of this dominant group of reptiles on the largest island in the West Indies. Her book contains distribution maps for every species and original, full-color illustrations for males of 50 species, females of 4 species, and juveniles of 3 species. Rodriguez Schettino also treats current problems of conservation and biodiversity resulting from urban development and a high number of species living in a comparatively small country. 58 color plates, 21 b&w drawings, 66 distribution maps, 65 tables, glossary, index. Cloth, $85.00 The 1GUANID LIZARDS of Cuba ; edited by Lourdes Rodriguez Schettino UNIVERSITY PRESS OF COMING THIS FALL! The Cuban Treefrog in Florida Life History of a Successful Colonizing Species Walter E. Meshaka, Jr. “The abundance of data cited in this study not only characterizes the Cuban Treefrog but establishes it as a model of a successful colonizer. This latter feature vastly increases the importance of this work by providing wildlife biologists with a list of characteristics that can be applied to other introduced species to determine the likelihood of successful establishment and expansion in non-native habitats—particularly those severely altered by human activity.’— Robert Powell, Carnegie Museum of Natural History Walter Meshaka discusses all facets of the natural history of the Cuban Treefrog in detail as well as the correlates of its colonization success. He also supplies a methodology for evaluating and setting priorities for the threats facing Florida’s amphibian and reptile populations, identifies the most vulnerable species, and presents a large data set associated with colonization patterns and predictions. 83 figures, 71 tables, index. September, Cloth, $69.95 AND THE CLASSIC STUDY! Amphibians and Reptiles of the West Indies Albert Schwartz and Robert W. Henderson “A definitive synopsis of West Indian herpetofauna.” —George R. Zug, National Museum of Natural History “A superb, scholarly work invaluable to faunal specialists concerned with the West Indies.”—Choice 600 maps. Cloth $75.00 Order through full-service booksellers, our website at www.upf.com or toll free with VISA or M/C: 1-800-226-3822 Gainesville, Tallahassee, Tampa, Boca Raton, Pensacola, Orlando, Miami, Jacksonville, Fort Myers HA Aj -<. 0p FOUNDER AND EDITOR Craig Hassapakis Amphibian and Reptile Conservation {aN 10 2005 Modesto, California ta = lAnaRiES..— Allison C. Alberts Center for Reproduction of Endangered Species Zoological Society of San Diego Jonathan D. Ballou National Zoological Park Smithsonian Institution Aaron M. Bauer Department of Biology Villanova University Andrew R. Blaustein Department of Zoology Oregon State University Harold G. Cogger Australian Museum Sydney, AUSTRALIA C. Kenneth Dodd, Jr. Biological Resources Division U.S. Geological Survey Lee A. Fitzgerald Department of Wildlife and Fisheries Sciences Texas A&M University Julian C. Lee Department of Biology University of Miami AMPHIBIAN ECOLOGY AND CONSERVATION Jamie K. Reaser U.S. Department of State, Bureau of Oceans and International Environmental, and Scientific Affairs ASSOCIATE EDITOR Jack W. Sites, Jr. Department of Zoology Brigham Young University ADVISORY BOARD Joseph T. Collins Natural History Museum University of Kansas Carl Gans Adjunct Professor of Zoology University of Texas at Austin Roy W. McDiarmid Herpetology, Biological Resources Division U.S. Geological Survey Russell A. Mittermeier President, Conservation International Washington, D.C. EDITORIAL REVIEW BOARD Harvey B. Lillywhite Department of Zoology University of Florida Peter V. Lindeman Department of Biology and Health Services Edinboro University of Pennsylvania Joseph C. Mitchell Department of Biology University of Richmond Henry R. Mushinsky Department of Biology University of Florida SECTION EDITORS George B. Rabb President, Chicago Zoological Society Vice chair, Communications, Species Survival Commission, The World Conservation Union Hobart M. Smith Department of Environmental, Population and Organismic Biology University of Colorado Michael Soulé Research Professor, University of California at Santa Cruz President, The Wildlands Project Jaime E. Péfaur Ecologia Animal, Facultad de Ciencias Universidad de Los Andes, Mérida, VENEZUELA Christopher J. Raxworthy Department of Herpetology American Museum of Natural History Andrew T. Storfer Wildlife Ecology and Conservation University of Florida Robert J. Wiese Assistant Director of Animal Programs Fort Worth Zoo STATISTICS AND EXPERIMENTAL DESIGN Anthony J. Krzysik School of Arts and Sciences Embry-Riddle Aeronautical University Office of Ecology and Terrestrial Conservation, Washington, D.C. COPY EDITOR Mark L. Goodwin COLUMNIST Craig M. Hoover Senior Program Officer, TRAFFIC North America, World Wildlife Fund Washington, D.C. ARGENTINA Maria E. Bridarolli Instituto Argentino de Investigaciones de las Zonas Aridas, Centro Regional de Investigaciones Cientificas y Tecnoldgicas, Mendoza INDIAAND MALAYSIA Indraneil Das Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak Sarawak, MALAYSIA ITALY AND MADAGASCAR Franco Andreone Sezione di Zoologia, Museo Regionale di Scienze Naturali, Torino, ITALY CONTRIBUTORS GRAPHIC DESIGN Kenneth W. Sholar INDEXING AND ABSTRACTING CONSULTANT/E-PUBLISHING Michael J. Tavares WEBSITE DEVELOPMENT AND CONSULTANT Jay G. Henry COUNTRY LIAISONS INDONESIA Darmawan Liswanto Executive Director, Yayasan Titian Jakarta PERU Antonio W. Salas Museo de Historia Natural Universidad Ricardo Palma SOUTH AFRICA Marius Burger National Coordinator, South African Frog Atlas Project University of Cape Town COMPUTER SUPPORT Mark Warren ZOO LIAISON Chris Banks Curator of Herpetofauna, Invertebrates & Education Animals and Co-ordinator South East Asian Conservation Programs, Melbourne Zoo, AUSTRALIA TAIWAN David McLeod National Experimental High School at Science Based Industrial Park, Huinchu U.S. VIRGIN ISLANDS Father Alejandro J. Sanchez Munoz Pastor of Saint Anne's Parish Saint Thomas VENEZUELA Jaime E. Péfaur Ecologia Animal, Facultad de Ciencias Universidad de Los Andes, Mérida DISCLOSURE: Until 2000, ARC was published irregularly with one issue being printed (premiere) and referenced as volume 1, number 1. ARC is now publishing two issues per year (Semi-annually). This issue marks number two in the serial and is listed as volume 2, number 1 (2000). With the support that ARC continues to receive we plan to progress into a quarterly serial sometime in the future. Presently, all subscribers receive four consecutively distributed issues at a minimum of two per year. Copyright © 2000 Amphibian and Re jill: Conservation. All rights reserved. ARC authorizes ph otocopying for internal or ripe ersonal use provided the appropriate fee is paid directly to the opyright Clearance Center, Inc., 222 Ros sewood Dr., Danvers, MA 01923-4599, USA. Tel: (978) 750-8400 fax (97 8) 7 50-4470; email: info(@copyright.com; website: www.copyright.com Amphibian and Reptile Conservation 2(1):4. Editorial WORDS FROM THE EDITOR—With the comple- tion and publication of this issue, Amphibian and Reptile Conservation (ARC) is now publishing semi- annually, from our previously irregular publishing schedule. We still have not given up our original plans of being published as a quarterly serial, and hope to be able to do so in the near future. We will continue to pick up the pace, and add more pages to each volume (when possible), as well as report on some of the most exciting topics in the world, from a balanced geographic perspective. ARC will con- tinue its focus on publishing research in the field of herpetology, and specifically herpetological conservation. We will try our best to publish cutting edge herpetological conservation research, but in a way that makes the subject matter accessible and enjoyable to professional herpetologists, conservation managers, naturalists, and nonprofessionals. This often requires an extra ef- fort on part of reviewers, authors, and others involved. It is to these unselfish individuals that deserve most of the credit for the success of the journal. This issue begins with an additional four pages (two in full- color) being added from that of the previous issue and volume (volume 1, number 1—premiere is- sue). It also adds new standardized graphic design throughout, important article contributions, full- color illustrated maps, country sidebars, a new col- umn titled “Herpetofauna and Humanity,” book review(s), world news, glossary, and abbreviations used. All these new improvements are imple- mented to make this science journal as accessible and interesting to as broad an audience as possible. Many of these journal elements will continue to be developed, as well as others that come to my attention, with each successive volume. I am very pleased to exhibit our latest effort and hope that you all will continue your great enthusiasm for the journal, as we work hard to establish what we think will become an important serial, specifically for the conservation and preservation of amphibians, reptiles, and their habitats worldwide. < = Craig Hassapakis Editor and Publisher Authors AARON M. BAUER received his Doctor of Philosophy (Ph.D.) degree in Zoology from the University of California, Berkeley. He is a Professor of Biology at Villanova University and a research associate of the Smithsonian Institution, Cali- fornia Academy of Sciences, and Museum of Natural Science, Loui- siana State University. He has pub- lished several books and over 200 scientific articles. His research spe- cialties are the systematics, bioge- ography, and evolutionary mor- phology of Southern Hemisphere reptiles, especially geckos. Dr. Bauer has had extensive field ex- perience in New Caledonia and with Ross Sadlier has recently written The Herpetofauna of New Caledonia, a new book on the diverse lizard fauna of this French island territory. CRAIG M. HOOVER, our columnist—Herpetofauna and Hu- manity—is a Senior Program Officer with TRAFFIC North America. Mr. Hoover has extensive experience in wildlife trade issues and wildlife law enforcement. Prior to joining TRAF- FIC in 1996, he worked for four years as a Wildlife Inspector with the United States Fish and Wildlife Service, enforcing state, federal, and foreign laws relating to the international wildlife trade. Mr. Hoover has a Bachelor of Science (B.S.) degree in Natural Resources from the University of Michigan as well as a Juris Doctorate degree from Loyola Law School. In his position with TRAFFIC, he has worked on a number of reptile trade issues, including the role of the United States in the live reptile trade, the trade in native turtles, and the trade in turtles for food in East and Southeast Asia. RONALD A. NUSSBAUM (RAN) is curator of herpetology, professor of biology, and director of the Edwin S. George re- serve at the University of Michigan in Ann Arbor. His research interests include, broadly, ecology, evolution, and systematics of amphibians and reptiles. His specific research projects in- clude evolution of parental care in salamanders, evolution and systematics of caecilians, and the systematics and biogeography of the herpetofauna of the islands of the western Indian Ocean, in particular the Seychelles archipelago and Madagascar. BRIAN A. MAURER received a B.S. in Zoology from Brigham Young University and a Ph.D. in Wildlife Ecology at Univer- sity of Arizona. After 13 years in the Department of Zoology at Brigham Young University, he recently accepted a position as associate professor in the Department of Fisheries and Wild- life and Department of Geography at Michigan State Univer- sity. He has published two books and numerous articles on population and community ecology of vertebrates, particu- larly birds. His research interests include biogeography, quan- titative ecology, and macroecology. CHRISTOPHER J: RAXWORTHY is Associate Curator in the Department of Herpetology at the American Museum of Natural History in New York, New York. His research interests focus on the evolutionary biology and ecology of reptiles, and in- clude systematics, biogeography, conservation, and behavior. Al- though he has done much of his most recent work in Madagascar, his interest in reptiles is world- wide, with ongoing fieldwork in ; ss North America, Madagasacar, West Africa, and Southeast Asia. PAUL THOMPSON ROSS A. SADLIER received his B.S. degree in Zoology from Macquarie University in Sydney, Australia. He is a Senior Scientific Officer in the Depart- ment of Herpetology at the Aus- tralian Museum. He has published extensively on the herpetofaunas of Australia and New Caledonia. His research interests are centered on the systematics of scincid liz- ards of the Pacific region. Mr. Sadlier has had extensive field ex- perience in New Caledonia and is co-author with Aaron Bauer of The Herpetofauna of New Caledonia, anew book on the lizard fauna of New Caledonia. AARON BAUER CONTENTS 15 24 Volume 2 Number 1 FEATURES Commentary on conservation of “Sokatra,” the radiated tortoise (Geochelone radiata) of Madagascar Ronald A. Nussbaum and Christopher J. Raxworthy Extinction and extinction vulnerability of amphibians and reptiles in Madagascar Christopher J. Raxworthy and Ronald A. Nussbaum New data on the distribution, status, and biology of the New Caledonian giant geckos (Squamata: Diplodactylidae: Rhacodactylus spp.) Aaron M. Bauer and Ross A. Sadlier DEPARTMENTS EDITORIAL Words from the editor Craig Hassapakis Authors COLUMN—HERPETOFAUNA AND HUMANITY The United States role in the international live ~'*? =e « i tt " . * ladagascar, ** <% ‘adjated tortoises Hefpefofauna extifiction ” “* , New Caled@hiia 7 Rhacodactylus getkos s Herpetofauna ang’Humahity (New Golumn) The trade in live reptiles gneamphibighs R Tha eral vppert the rec Perm taiass Cover Standing’s day gecko Phelsuma stand- ingi, at Zombitsy Forest, Southwest Madagascar. This species is restricted to deciduous dry forest, a habitat that is ; i d declining due to clearing for cattle graz- reptile trade ing and agriculture. Uncontrolled grass- Craig M. Hoover land fires which, every year, burn the for- BOOK REVIEWS est edge have scorched the tree stump. These day geckos have also had dra- 32 Lizards of the Caribbean: ecology, evolution and matic population declines due to over- plate tectonics collecting for the pet trade in some ar- Review by Brian A. Maurer eas of its limited distribution. Photo: C. J. NEWS AND NOTES Raxworthy. 33 World news Announcements.» 34__....Meetings ~~ Websites New literature Literature Books and literature received THE LAST PAGE 35 Glossary Abbreviation usage Errata Acknowledgments “Wi Dedication Background screen photograph (see in color on page 18): Some of the last known surviving Mantella bemhardi forest habitat cleared for cultivation between 1993-4. Photo: C. J. Raxworthy. Scope: Amphibian and Reptile Conservation (ISSN: 1083-446X) [ARC] and the accompanying online edition (ISSN: 1525-9153) is a popularly accessible. peer-reviewed scientific journal of international scope. which is devoted to the worldwide preservation and management of amphibian and reptilian diversity. Online edition: The full-text online edition is available to subscribers FREE-OF-CHARGE as PDF (Portable Document Format) files through ingenta at: www. ingenta.com. The online edition may vary slightly from the print edition due to our reducing file sizes for efficient downloading over the Internet Some background screens (photographs) are removed which are deemed not essential to the content of the article(s). There is also some loss of clarity to photographs in reducing article file sizes to a minimum If clarity of photos is a problem, the print edition of the journal should be consulted. Audience: ARC is intended for a wide readership from nonprofessional to professional herpetologists, the general public. and scientists. Frequency: ARC publishes two issues per year (semi-annually). Focus: ARC concentrates on publishing timely information in the form of feature articles, original papers and data. reviews. reports. short communications. columns, commentaries, book reviews, editorials. and news and notes. Distribution: ARC is distributed worldwide by subscription as well as quality newsstands. bookstores, and select vendors. 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Tel: (978) 750-8400; fax: (978) 750-4470: email: info@copyright.com, website: www.copyright.com Commentary on conservation of “Sokatra,” the radiated tortoise (Geochelone radiata) of Madagascar RONALD A. NUSSBAUM" ? AND CHRISTOPHER J. RAXWORTHY? Division of Amphibians and Reptiles, Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109-1079, USA 2Associate Curator, Department of Herpetology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192, USA Abstract.—The radiated tortoise Geochelone radiata of the desert regions of Southwestern Madagascar, known as “sokatra”’ among most Malagasy, has gained much attention recently as a result of increasing and highly publicized smuggling of this commercially valuable species. Sokatra have been protected by Malagasy law since 1960 and have been classified as a Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) Appendix I species since 1975. Sokatra also are protected to some extent in the four reserves where they are known to occur. In the central part of their range, on the Mahafaly and Karimbola Plateaux, sokatra are considered fady (taboo) by the Antandroy and Mahafaly people who live in this area, and they generally are not killed for food in this region. They are, however, killed and eaten by the Vezo and Antanosy people who largely occupy the northwestern and southeastern limits of the species’ range. It has been suggested that this taboo is largely responsible for the survival of the species, and there is worry that the taboo may be breaking down as a result of human famine and intermingling of tribal peoples, many of whom do not consider the flesh of sokatra to be forbidden. In addition to the possible erosion of taboo barriers, there is strong evidence of increased illegal trade in sokatra, increased deterioration of its habitat, and increased local consumption of them for food, all of which are reasons for concern. Given the conspicuousness of this species in nature, its popularity among tortoise fanciers, and concern for sokatra among conservationists, there is surprisingly little published information about them in their natural environment that would allow for objective evaluation of their status. Especially needed are intensive studies of the life history and ecology of sokatra, with special attention paid to determining their limiting environmental requirements. It will also be important to examine the degree to which zebu (cattle) and goats compete with sokatra for food and to determine the intensity of zebu and goat grazing that can be tolerated without causing local extinction of sokatra. We have no reason to believe that the sokatra is threatened with extinction over the next 20 years, just as it obviously was not threatened over the past 23 years, although classified as Appendix I during that period. We recommend that downgrading the sokatra to CITES Appendix II might be beneficial to the survival of the species if certain conditions are met, such as careful controls on the number of legally exported animals. We also strongly recommend the establishment of additional nature reserves on the Mahafaly and Karimbola Plateaux in the central part of the range of the sokatra, both for survival insurance for this species and for other rare and endemic species that occur in this area. Key words. Radiated tortoises, Geochelone radiata, Madagascar, conservation, sokatra, protection, CITES, pet trade, education, captive breeding, repatriation, monitored legal trade program Introduction The radiated tortoise, Geochelone radiata (Plate 1), is one of the attention from conservationists. Recently, awareness of radi- ated tortoises reached new heights as a result of a highly publi- most spectacular of the larger tortoise species. It grows to a maximum size of about 40 cm carapace length and may weigh up to 14 kg. Radiated tortoises are readily identified by their color pattern of bright yellow lines radiating from the center of each dorsal scute and their yellow legs and throat. Because of their beauty, and perhaps because they are members of the high pro- file Madagascan fauna, radiated tortoises are highly coveted by pet keepers. A large breeding pair may be valued up to $25,000 in the pet trade, and prices in the range of $5,000 for a single, not necessarily mature, radiated tortoise are not uncommon. Be- cause of their commercial value, and because they are killed for food by local Malagasy and served as a delicacy in some Mala- gasy restaurants, radiated tortoises have received considerable 3Correspondence. Tel: (734) 647-2201; fax: (734) 763-4080; email: nuss@umich.edu cized smuggling bust in Orlando, Florida that resulted in the confiscation of about $250,000 worth of radiated tortoises, spi- der tortoises, and Madagascan boas and in the conviction on 10 January 1997 of two smuggling partners, a German and a South African. This headline news was followed by an article (Webster 1997) in a major news magazine that described the business of smuggling rare animals and featured a color photograph of a radiated tortoise on the cover. Considering the great interest in radiated tortoises, it 1s astounding that so little of scientific merit has been published about them. Most of the sparse literature consists of anecdotal and repeated observations. Not a single in-depth study of this species in nature exists, and we are unaware of any ongoing or planned research. The most recent field survey of the species was sponsored by the World Wide Fund for Nature (WWF) and was done by Richard Lewis, whose report (1995) to WWF- RONALD A. NUSSBAUM AND CHRISTOPHER J. RAXWORTHY Aires Protégées, Madagascar, is unpublished. Juvik (1975) re- viewed the literature and presented some limited new informa- tion on radiated tortoises. Durrell et al. (1989) commented on captive breeding programs and the status of the species. Goodman et al. (1994) reported road count data for a single trip along one segment of road in the center of the range of the species. Razafindrakoto (1987) provided the only information available on food habits and other autecological‘ aspects of the species in its natural environment at Beza-Mahafaly Réserve Spéciale. Unfortunately, the population at Beza-Mahafaly has subsequently been genetically polluted and otherwise influenced by the release of numerous confiscated tortoises (Lewis 1995), so that follow-up studies would be of limited value. Distribution and habitat “Sokatra,”! or radiated tortoises, are restricted to the xeric’ re- gion of Southwestern Madagascar (Fig. 1), where they occur in a variety of habitats ranging from brushy spiny desert domi- nated by endemic Didieriaceae and euphorbs to gallery forests dominated by deciduous species such as “kily,” or the tamarind tree, Tamarindus indica. In this region, annual rainfall is low (< 400 mm) and highly unpredictable, and temperatures are very high, especially during sunny summer (November-February) days. It has been claimed that sokatra hibernate during the win- ter, but, while there are undoubtedly periods of inactivity, we have seen them active during every month of the year. Within their range, sokatra are absent from open savannahs and from forests with no understory vegetation, probably because of their need for low vegetation for grazing and for frequent shady areas to escape overheating from insolation. Historically, the eastern limit of their range was probably determined by the dense, low elevation rain forests near Télaiiaro. There are no records of the species north of the Manombo River along the western sea- board, although it is highly unlikely that the river itself poses a barrier (they occur on both sides of much larger rivers). Their restriction to a wide coastal band is somewhat mysterious, but it may be that higher inland elevations limit them to the coastal band. Before human occupation of Madagascar, the species prob- ably occurred somewhat further inland. Status and protection Theoretically, sokatra have been protected since 1960 by na- tional Malagasy law (Decree no. 60-126), which provides for fines and/or imprisonment for unauthorized collecting. Since 1975, sokatra have been classified as Appendix I species ac- cording to the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), which regulates inter- national commercial trade. The species is further protected in limited parts of its range that are set aside as nature reserves. These protected areas are Tsimanampetsotsa Réserve Naturelle Intégrale (43,200 ha), Cap Sainte Marie Réserve Spéciale (1,750 ha), Beza-Mahafaly Réserve Spéciale (580 ha), and Parcel II of Andohahela Réserve Naturelle Intégrale (12,420 ha). Two of these reserves, Beza-Mahafaly and Parcel Il of Andohahela, are at the limits of the species’ range in areas where sokatra are uncommon (Andohahela) or consist of populations genetically polluted by release of confiscated animals (Beza-Mahafaly). Tsimanampetsotsa and Cap Sainte Marie are in the center of the species’ range and are potentially important for conservation of sokatra. Both of these reserves are overgrazed by zebu (cattle) and goats and are subjected to limited woodcutting, but sokatra seem to be abundant in both reserves (pers. obs.). There.are no large, undisturbed reserves on the Mahafaly and Karimbola Pla- teaux in the region of prime sokatra habitat. A large reserve in this area is badly needed, not only for sokatra, but also for protection of many other rare species of plants and animals that occur in this unique and spectacular environment. These two plateaux have not been adequately surveyed for biodiversity. A brief survey done by us at the edge of the Mahafaly Plateau at Lac Tsimanampetsotsa yielded several rare reptiles, including two undescribed geckos of the genera Ebenavia (Malagasy leaf- toed geckos) and Paroedura (Malagasy casque-headed geckos) [Nussbaum and Raxworthy 1998]. The Antandroy and Mahafaly people of Southwestern Madagascar regard sokatra as fady (taboo), and eating sokatra flesh is forbidden. In some areas, especially where they are associated with tombs, sokatra are believed to contain spirits of ancestors and are, therefore, sacred as well as fady. It has fre- quently been suggested (e.g., Juvik 1975) that their status as forbidden and sacred animals 1s the best protection sokatra cur- rently has, and without it they might already be extinct. It is believed (Juvik 1975; Lewis 1995) that the range of sokatra 1s shrinking and the populations diminishing at the northwestern and southeastern limits of its range (Fig. 1). This may result largely from exploitation for food by the Vezo and Antanosy tribes to the northwest and southeast, respectively. Sokatra have no taboo status among the members of these two tribes, and these people eagerly seek sokatra for food. In addition to exploitation for food, habitat destruction in these two areas is advanced, especially east of Ambondro, and this undoubtedly has a negative impact on local sokatra populations. Sokatra are classified as “threatened/vulnerable” by the World Conservation Organization (IUCN) [1996 Red List of Threatened Animals}, which seems justified, given the conspicu- ousness of sokatra in their habitat, their ease of capture, their popularity for food and pets, and the ongoing degradation of their habitat. However, their classification as a CITES Appen- dix I species is highly questionable. Originally (1973), this clas- sification was reserved for species that are threatened with ex- tinction, or could be threatened with extinction within a five- year period, or have a very limited range. This has been revised so that now Appendix I species are those “threatened with extinction which are or may be affected by trade.” Because “threatened with extinction” can be interpreted very broadly (there are no guidelines), the current criterion reduces the ques- tion of status to a matter of opinion, so that almost any species of commercial value arguably could be classified as Appendix I. In our view, there is no evidence, published or otherwise, that indicates sokatra currently are threatened with extinction, al- though there are reasons for concern. There is not even any strong evidence that sokatra are less abundant or more restricted geographically now than they were in 1975 when placed on Appendix I. Clearly, more objective criteria for CITES classifi- cations are needed, and, most of all, in the case of sokatra and many other threatened/vulnerable species, intensive research is needed to determine the distribution and abundance of the spe- cies and to identify environmental factors important for main- taining viable populations. Population densities Most information on population densities of sokatra is anec- dotal or involves estimates from road counts and other rapid CONSERVATION OF THE RADIATED TORTOISE survey methods. There are no data that would provide for meaningful comparisons of past and present population densi- ties. Juvik (1975) reported that after a rain storm in 1974, sokatra were encountered at a rate of about one per km along Route National 10 between T6laiiaro and Toliara, presumably in the area around Beloha. Goodman et al. (1994) recorded road counts of about 3 sokatra per km on 30 December 1992 after heavy rain along 42.8 km of road between Ankororoka and Beloha. These counts, however, included both living and many dead sokatra, so that it is necessary to adjust for the accumu- lation of dead animals. When that is done, the encounter rate is closer to one sokatra per km. A count of sokatra in January/ February, 1995, along 15 km of road in the region of the Menarandra River, yielded 30 individuals for a rate of two sokatra per km (Lewis 1995). We have traveled extensively, logging many thousands of kilometers, over roads in this region every year and in every season since 1989, and we can confirm observations of others that sokatra activity is greatest during warm weather shortly after rain. Not a single sokatra was observed during long periods of drought along the Route National between Tsiombe and Ampanihy, but, during warm and wet weather, numerous sokatra can be observed, especially between 40 km south to 30 km north of Beloha. Normally, sokatra will not be seen during the dry, cool winters, but on 6 July 1995 we counted 0.36 sokatra/km along this stretch of road. There had been rain the day before, and the day of the count was partially cloudy with light sprinkle, and the temperature was unseasonably warm. Other than road counts, there are only two reports of den- sities. Razafindrakoto (1987) indicated a density of 1.3 sokatra per ha in Parcel I of Beza-Mahafaly Réserve Spéciale, based on a mark-recapture study. Lewis (1995) estimated densities from five transect counts in a variety of prime habitats from well within the core of the species’ distributional area. Lewis’s den- sity estimates ranged from 262 to 1,077 sokatra per km?. Using these data, Lewis gave “conservative” total population esti- mates of 1.6 to 4.0 million sokatra for the core area on the Mahafaly and Karimbola Plateaux, an area of about 10,000 km’. Juvik (1975) and Lewis (1995) believed the range of sokatra was contacting and fragmenting at the northern and eastern ends of the distributional area. While this is probably true, the his- torical records that could confirm this unequivocally do not exist. If humans are the main threat to sokatra, and if the range of sokatra has been accurately identified, then sokatra have done remarkably well over the past 2,000 years since humans colo- nized Madagascar. Most of the negative impact on sokatra popu- lations has probably occurred over that past 500 years, coinci- dent with the arrival of Europeans, and more recently with the explosion of the Malagasy population. There are reliable re- ports of large numbers of sokatra being shipped to the Mascarenes for food during the 18" and 19" centuries (Juvik 1975). Passing ships in the Western Indian Ocean regularly took on large quantities of the larger tortoises from Southern Mada- gascar and the Mascarene and Seychelles Islands for ship’s stores. This activity is thought to be responsible for the extinction of the giant tortoise (Geochelone gigantea) on the granitic Seychelles Islands. Prospect Based on his limited survey taken in 1974, Juvik (1975:145) believed that “the outlook for the radiated tortoise in Southern Madagascar is not entirely gloomy, thanks to traditional taboos on eating its flesh, improved government controls on exports, and import restrictions in other countries. Its future depends on the survival of some natural habitat; at the same time... modern agricultural developments may be indirectly beneficial.” Now, 22 years after Juvik’s optimistic report, the outlook for sokatra is still not “gloomy,” but there have been significant changes that are worrisome. Increasing human population pressures in recent years have resulted in a marked increase in habitat destruction, harvesting of sokatra for food and pets is clearly on the rise, and there is no evidence that the Malagasy government has been able to do anything about it. We agree entirely with Richard Lewis (1995) who writes, “One is left with the conclusion that there is a lack of political will to enforce the law.” Our evidence for increased habitat destruction and harvest- ing of sokatra stems mainly from our field observations, which began in the southwest in 1989. Habitat destruction is of four kinds. First, land is being cleared to establish many new agricul- tural plots, especially noticeable in the eastern part the range of sokatra, near Ambovombe and Amboasary. These new fields include small family-owned subsistence plots of corn, cassava, sweet potato, and peanuts, but also larger plots for the commer- cial growing of sisal®. Second, every year during the dry season, much of Southern Madagascar is deliberately burned, as it is widely believed that burning increases the growth of plants needed for grazing of zebu and goats. Burning grasslands inevita- bly leads to burning of spiny forest and brush, as the grass fires are not controlled. A significant but unknown amount of sokatra habitat is lost through uncontrolled burning every year, and it is our impression that the intensity of burning has increased over the past eight years. Furthermore, sokatra travel over terrain slowly compared to most animals and can easily be overcome by fire, and we suspect that large numbers are killed each year in this manner. That some tortoises are killed in this way is certain, as we have seen charred tortoise shells in several burned areas. Third, the number of zebu and goats grazed is increasing with human populations, and the intensity of grazing is exacerbated. At the same time zebu and goat populations are increasing, the amount of land available for grazing them is diminishing as a result of conversion of land to agricultural use. It seems certain that zebu, and especially goats, consume food necessary for the survival of sokatra, but the intensity of competition is unstud- ied. There is a critical need to study the effects of zebu and goats on sokatra, because virtually every square meter of sokatra range is subject to cattle and goat grazing, even in the reserves and sacred forests of the Mahafaly. Fourth, and equally worri- some, is the dramatic increase in woodcutting for firewood, char- coal making, and construction of houses. Increased woodcutting is proportional to population growth, and there is a direct effect of increased agricultural clearing on woodcutting. For example, now that the town of Amboasary is completely surrounded by sisal plantations, woodcutters who supply the cooking fires for that large village range out as far as 30 km to cut wood. Every day, a steady stream of woodcutters with bundles of firewood on their backs or in push carts can be seen along the Route National east of Amboasary, and similar scenes can be seen along the Route National east of the seaport of Toliara. Along these same two stretches of national highways, entire villages based on charcoal making and selling have sprung up. The char- coal is made deep in the bush and transported to the villages mainly on the backs of humans. MADAGASCAR ee as Mdeeca | is the antl outiifargest island wih a oul area of 587, 040 square km | (about twice the size of Arizona) located off the southeast coast of Africa (Geographic | Coordinates 20°00! S, 47°0' the Indian Ocean. Madagascar is one of the seven major 7 world centers of ee has been caved the number one ea Bee, in. the world. ea oe 3450. Species Le s human | popu- ma with a ee 3% es growth. Cli- _ MADAGASCAR deopentive ven’ “social ae pri elopment, political instability, fi habitat ise (asaresultofallthe | - and other tleast80% | | ithin 37 protected | ) fully Tepre- ee S 8 ee Madag: | pating in. pro} about what you experience and learn the process.” Maps courtesy of John W. Megahan, st, useumn no ee ey oy Midge at t Ann Arbor, ee found within villages. For example, Beloha is in the heart of Antandroy country where sokatra are supposedly fady, yet iy hundreds of discarded shells of eaten sokatra can be seen only partially hidden in the sisal plants along the roads in the village. It is not known whether these sokatra are being eaten by less traditional Antandroy who no longer consider sokatra to be fady or whether the significant harvest is done by the many non- Antandroy who live in these regions. Increasingly, except in the most remote areas of the southwest, there is a mixing of the various Malagasy tribes, and this may be one of the greatest threats to sokatra survival, as sokatra are neither fady nor sa- cred to tribes other than the Antandroy and Mahafaly. Sokatra are sold openly in restaurants in Southern Mada- gascar, and neither federal law nor local taboo influences this commerce. The Antandroy and the Mahafaly apparently place no pressure on visitors to their lands to respect the fady status of tortoises although they do insist that sacred tortoises be left alone. Seated in a small hotely (Malagasy restaurant) on a winter day, 1995, in Ejeda, a large village dominated by Mahafaly tribespeople, one of us (RAN) overheard two sol- diers and a policeman inquiring about sokatra on the menu. It was available, but they declined it because the price 2,500 Francs Malagasy (FMG) was too high, choosing instead hena kisoa (pork), which was only FMG 1,500 per plate. At that time, FMG 2,500 was worth about US$0.55. The restaurant owners made no attempt to hide their sokatra and the soldiers 200 Cape Ste. Marie (Reserve) Fig. 1. Distribution of sokatra in Southeastern Madagascar. Evidence for increased harvesting of sokatra for local con- sumption and illegal export is everywhere. Throughout coastal Southwestern Madagascar, campsites very often are cluttered with the remains of one to several sokatra that were killed and eaten (Plate 2). Sometimes these campsites are outside the range of sokatra, which means the animals were transported there for the purpose of providing meals. Remains of sokatra are also 10 and policeman obviously weren’t interested in enforcing the law. Nor, evidently, were the local Mahafaly opposed to sell- ing sokatra for food. On several occasions, we witnessed buses and other ve- hicles stopping along Routes National within the range of sokatra, so that occupants could debark to collect a hapless sokatra observed near the road. RAN once followed a bus for about 15 CONSERVATION OF THE RADIATED TORTOISE kilometers on the road south of Beloha, which stopped no less than eleven times to collect sokatra... not one was passed up. Some of these sokatra may have been destined to become pets, but most were probably eaten. Some of the passengers of these buses must have been Antandroy and Mahafaly, and yet there was a festive atmosphere about collecting the tortoises and no objections from any passengers. Plate 2. Remains of a killed and eaten sokatra near a campsite in Southeastern Madagascar. In addition to local consumption, sokatra are harvested for sale in the markets and restaurants in bigger cities, and at least limited numbers of them are killed and exported for food from the port of Toliara (Lewis 1995; pers. obs.). On 9 October 1995, about 20 km east of Ampanihy, one of us (RAN) came across five large oxcarts filled with.500-800 sokatra being transported in disturbingly inhumane conditions. The sokatra were piled on top of one another, fully exposed to the blazing sun, and were bouncing up and down on the extremely rough road. The oxcart drivers didn’t want to talk to us, but they didn’t seem worried about exposure of their illegal cargo. We were informed in Ampanihy that these tortoises were on their way to market in Toliara. We have observed, and Lewis (1995) has reported, dumpsites for sokatra shells numbering up to 300 shells per dump near Toliara. Droughts in Southern Madagascar between 1991 and the present, which caused crop failures and human starvation and death, resulted in increased killing of sokatra for food. During the drought of 1992/93, numerous small sokatra appeared in Tolafiaro where they were being sold to (mainly) the local Antanosy for food. Inquiries led one of us (RAN) to the discov- ery that nearly 3,000 sokatra were being sold openly in the markets at Ambovombe, the Antandroy capital city. The 3,000 were present on a single market day; the rate of flow of sokatra through the market is unknown. RAN was informed by many local Malagasy of both Antandroy and Antanosy nationality that during times of food shortages, nontraditional Antandroy eat sokatra regardless of the fady (some Malagasy have ceremo- nies for temporary lifting of various fadys), and of course the Antanosy openly eat sokatra with great gusto any time they are fortunate enough to have them. The danger now is that those who were forced to eat sokatra during this unprecedented fam- ine will continue to consume them after the famine. Holidays in Madagascar are especially hard times for 11 sokatra. Poor families who can’t afford meat everyday will usually find a way to provide their families with a special holiday treat. This might be a fowl, but sokatra are often available for nothing and may be more desirable than fowl for special occasions. Two other problems, perhaps of less concern, for sokatra conservation are the killing of these animals as pests in agricul- tural areas and for selling as stuffed curios to tourists, mainly in Antananarivo and Toliara (Plate 3). As land is converted to crops, sokatra are increasing obliged to forage in fields, and many sokatra are now killed because of the damage they do in fields. Many Antandroy who won’t eat them nevertheless do not hesitate to kill them if their crops are threatened. A surprising number of Malagasy citizens keep sokatra for pets, in some cases for pleasure, and in others because they believe their presence protects their poultry against diseases, especially louse infestation. Keeping sokatra as pets is not re- stricted to the south; families in the capital city (Antananarivo) and other northern villages keep numerous sokatra. In one small village near Antananarivo, about 30 sokatra are being kept by a small group of Malagasy villagers (O. Pronk, pers. comm.). The illegal harvesting and exportation of sokatra for pets is obviously on the increase, as is indicated by the numerous re- cent reports of arrests of smugglers and confiscation of these animals (e.g., Webster 1997). It is easy for tourists to purchase live sokatra in the larger cities, and customs officials confiscate significant numbers. On two occasions, we witnessed sokatra taken from the baggage of Japanese tourists at Ivato Interna- tional Airport in Madagascar, and there are regular reports of sokatra confiscated from European (mainly German) travelers in the Malagasy newspapers distributed in Antananarivo. The most recent report of which we are aware is in the 25 March 1997 issue of “Midi Madagasikara,” which told the story of 78 sokatra taken from the backpacks of two Japanese tourists at Ivato International Airport. Such is the current situation with the sokatra. Their habitat is being degraded and destroyed at an increasing rate, they are being harvested by the thousands every year, and local law enforcement does little to mitigate the situation or stop such activities. The laws protecting sokatra are well known to the Malagasy, but they have learned that these laws can be com- pletely ignored. It is equally clear that laws to prevent international trade in radiated tortoises are not working very well. Perversely, it appears that laws restricting export of radiated tortoises may do more harm than good. Banning exportation increases the value of the tortoises, both in the legal and black market trades (see Lamar 1997, for other species), which increases the deter- mination of smugglers to find ways to get the tortoises out and inevitably leads to more corruption of officials charged with enforcing the laws. With higher values, smugglers can afford to pay for illegal transport and lay out more money for bribes. Evidence that embargoes on shipments of pet trade animals from Madagascar don’t work can be seen in regard to the recent CITES embargo on most species of day geckos (Phelsuma) and chameleons from Madagascar. Since this embargo went into effect 20 January 1995, the banned species have continued to arrive in Europe and the United States in large numbers, and at least in Europe the prices have declined suggesting they are arriving in greater numbers than before (O. Pronk, pers. comm.). Economically, this may be because the smaller day geckos and RONALD A. NUSSBAUM AND CHRISTOPHER J. RAXWORTHY chameleons can be smuggled in greater numbers than tortoises, and because smugglers do not have to pay the export tax that is levied by the Malagasy government against legal animal ex- porters. Bee yi aM 4 ate 3. C. J. Raxworthy examines a stuffed sokatra for sale to tourists in the Analakely market in Antananarivo, Madagascar. An additional negative effect of banning export of radiated tortoises (and other species) is the inhumane treatment of the exported animals. Smugglers do not have to comply with Inter- national Air Transport Association (IATA) rules regarding hu- mane packing and shipment of commercial animals and, indeed, probably cannot comply with these conditions because of the need to hide the animals and use circuitous and prolonged routes for their movement. Undoubtedly, the percentage of dead, in- jured, and unhealthy animals arriving at their destinations is higher in illegal shipments than in legal shipments. It seems certain that local traditions have had more impact than federal and international laws in conserving the sokatra. But it also seems certain that the forbidden status of sokatra among the Mahafaly and Antandroy will erode as human popu- lations and the need for protein and money increases and as people from other tribes without fady constraints continue to immigrate into the range of the tortoise. Much serious thought, research, and considerable effort will be needed to insure the survival of this increasingly vulnerable species. THOMPSON PAUL 12 Solutions It has repeatedly been argued that education of the Malagasy is the key to the survival of the sokatra. This seems unlikely. Many well-educated Malagasy, who are well aware of conserva- tion issues, keep sokatra as pets and regularly eat them because they taste good. Poorly educated Malagasy, those who live in the bush and survive off the land and eat tortoises because they need food, are largely immune to education, and, in any case, all the education in the world will not stop them from doing what is necessary to survive. The dilemma these poor Malagasy face is dramatically demonstrated in Webster’s (1997) article. Education, especially in regard to resource management, may be of some value if directed at land managers and guardians of the Reserves. During a visit to Cap Sainte Marie, RAN was proudly informed by the guardians of the Reserve that they regularly collected tortoises observed on the limestone pla- teaux and placed them in one of the steep-sided canyons where the tortoise could not get out, find more food, and were pro- tected from poachers. It apparently hadn’t occurred to these guardians that the tortoises were numerous on the plateaux because the conditions there are good for them and that taking the tortoises from their familiar home ranges might be disas- trous for them. Furthermore, the guardians had not considered the negative effects of artificially concentrating tortoises in a habitat which they should have realized was suboptimal for the tortoise, otherwise there would have been more tortoises there naturally. Finally, the guardians should have realized that concentrating the tortoises in a canyon might actually make it easier for poachers to collect them. Education might also help to stop the genetic pollution caused by the irresponsible re- lease of confiscated animals. It has also been suggested that captive breeding programs are a way to ensure survival of the species. Such a program is the American Zoo and Aquarium Association’s Species Sur- vival Plan (AZA/SSP) for the radiated tortoise, underway through a consortium of zoos, in which tortoises would ulti- mately be repatriated, to the natural environment under the auspices of Malagasy guardianship. There are obvious prob- lems with this approach, not the least of which is that it will do no good to repatriate captive-bred tortoises to an environment that will not sustain them. If the ecosystem no longer supports wild-bred tortoises, then why should we expect captive-bred tortoises to do any better? If the natural environment will not support sokatra, then captive-bred animals are best kept where they are... in zoos. If natural populations still exist, then they should not be genetically polluted by release of captive-bred animals. Captive-bred animals should be used to establish popu- lations in nature only if natural populations no longer exist and only if conditions that insure protection of the released ani- mals are in place. Aside from educating resource managers and guardians, what are the solutions to conservation of sokatra? Something should be done to (1) curtail habitat degradation, (2) reduce harvesting of sokatra for local consumption, and (3) control the exporta- tion of sokatra in the pet trade. It seems highly unlikely, given the inexorable human popu- lation growth and the Malagasy local traditions, that much can be done to slow habitat degradation. The best that can be done is to establish one or more large nature reserves on the Mahafaly and Karimbola Plateaux in areas that are not currently being converted to agricultural plots and have little value for agricul- CONSERVATION OF THE RADIATED TORTOISE é = 4a ~ " 5 : oo) ee =; : soa cs Hatchling ra ture. This might be done in conjunction with the large and rela- tively undisturbed sacred forests where Mahafaly kings are bur- ied. These sacred forests are not immune to cattle grazing, and neither are the reserves. Therefore, studies are needed to learn the impact of grazing on tortoise populations and to determine the maximum allowable grazing within reserves. Control of graz- ing and poaching of tortoises on the reserves would be impos- sible without the cooperation of local and federal authorities and, more importantly, without some reward for local support of law enforcement. This would have to include compensation for lost income from reduced grazing of cattle‘and harvesting of tortoises. Threats of punishment alone won’t work. Reduction of local consumption of sokatra will be diffi- cult, especially in areas outside of the Mahafaly and Antandroy homelands. Antanosy and other Malagasy connoisseurs of sokatra will always eat them, regardless of the law. Malagasy law regarding the sokatra has been widely ignored for so long that any attempt to enact strict enforcement locally would lead to serious problems with which the Malagasy authorities are not adapted to cope. However, we note that although lemurs (family Lemuridae) are still consumed for food in Madagascar, the law protecting lemurs is both more widely enforced and respected than are the laws protecting sokatra, perhaps as a result of more intense public scrutiny related to the economy of tourism and because of greater international involvement. If the international community, through agencies such as United States Agency for International Development (US AID), can be persuaded to act more responsibly by promoting mean- ingful projects in Southern Madagascar that increase the food supply (relative to human population density) and by reacting 0. = a . Po ae = diated tortoise Geochelone radiata. Photo courtesy « 13 =, . < of R. D. Bartlett. more swiftly to drought and famine, then desperation consump- tion of sokatra might be reduced. This would mean providing food rich in protein, and not just surplus corn and bulger wheat, during times of famine. In the area around Tolaiiaro, much of the surplus food given to Malagasy by aid agencies during the recent drought, especially the dried corn, was either fed to livestock or sold for pittance so that the puzzled owners could buy real food. The problem of consumption of sokatra is probably in- tractable in peripheral areas, but creation of additional reserves and strong rewards for respecting the boundaries of the reserves would help to insure the survival of sokatra in the core area of their distribution. There are two options for reducing the impact of the illegal commercial pet trade on natural populations of sokatra. The first is to enact a monitored legal trade program that strictly limits the number of wild-caught sokatra that could be exported and would generate some income for the Malagasy government through taxation on exports of sokatra. This would be espe- cially desirable if the generated income were used to support monitoring programs and research on the tortoise. Such a solu- tion would require downgrading of the sokatra from Appendix I to Appendix II, which theoretically shouldn’t be a problem, because it is clear that the sokarra is currently misclassified by CITES, as it is not currently “threatened with extinction.” If the species was downgraded to Appendix II, then a limited number of both wild-caught and first generation, captive-bred sokatra could be legally exported as could confiscated animals (under the monitored program), a better solution than genetic adulteration of natural populations through random release into the environ- ment. Limited legal export (with controls such as internal pas- RONALD A. NUSSBAUM AND CHRISTOPHER J. RAXWORTHY Plate 4. Confiscated sokatra at a Malagasy government station in lvoloina (September 1985). sive integrated transponder [PIT] tag markers) would destroy or greatly reduce the market for smuggled animals and prevent inhumane shipping methods. Realistically, downgrading to CITES IH might be politically impossible, so a second solution would be to encourage captive breeding programs in Madagascar that would eventually yield second generation, registered, captive-bred tortoises that could be legally exported under a monitored program. This could be done under a partnership with the Malagasy government and captive breeders in which the Malagasy government retains ownership of the tortoises and in which some of the income would be returned to the Malagasy to support tortoise conser- vation. Perhaps third-party monitoring agencies would be re- quired to insure against corruption. The nucleus for a monitored, legal export program for sokatra already exists in Madagascar at Ivoloina (Plate 4), where the Malagasy government (Eaux et Foréts) keeps confiscated tortoises. Under either program, it might be possible to supply tourists with legal tortoises, under the same kind of regulation used for exporting semiprecious stones and other valued objects from Madagascar. Such pro- grams would not entirely eliminate illegal trade but might reduce it considerably, as profit from illegal activity would be greatly diminished. Acknowledgments.—We thank Richard E. Lewis for giving us access to his unpublished report on radiated tortoises and for permission to cite it, and Olaf Pronk for useful insights into the intricacies of the pet trade. Our research in Madagascar has been funded by the National Science Foundation, National Geographic Society, and Earthwatch. We have received logistic support from Conservation International, WWF, and US AID. References Durrell, L., Groombridge, B., Tonge, S., and Bloxam, Q. 1989. Geochelone radiata radiated tortoise, sokake, p. 96-98 in Swingland, I. R. and Klemens, M. W. (editors). The Conserva- tion Biology of Tortoises. Occasional Paper, IUCN No. 5, Gland, 14 Switzerland. Iv + 202 p.: illustrations, maps. Goodman, S. M., Pidgeon, M., and O’Connor, S. A. 1994. Mass mortality of Madagascar radiated tortoise caused by road con- struction. Oryx 28:115-118. IUCN. 1996. 1996 IUCN Red List of Threatened Animals. YUCN, Gland, Switzerland. 448 p. Juvik, J. O. 1975. The radiated tortoise of Madagascar. Oryx 13:145- 148. Lamar, W. W. 1997. CITES: boon or boondoggle? Herpetological Review 28:10. Lewis, R. E. 1995. Status of the radiated tortoise (Geochelone ra- diata). WWF-Madagascar, unpublished report. Nussbaum, R. A. and Raxworthy, C. J. 1998. Revision of the genus Ebenavia Boettger (Reptilia: Squamata: Gekkonidae). Herpetologica 54(1):18-34. Razafindrakoto, L. A. 1987. Contribution a l’etude bioecoetho- logique de Geochelone radiata (Shaw 1802) (famille de Testudinidae) dans la Réserve Spéciale de Beza-Mahafaly. Mémoire de Fin d’Etude, EESSA, Universite de Madagascar, Antananarivo, Madagascar. Webster, D. 1997. The looting and smuggling and fencing and hoard- ing of impossibly precious, feathered and scaly wild things: the $10 billion dollar black market in endangered animals. New York Times, February 16, 1997, Section 6, p. 26-33, 48-49, 53, 61. Footnotes The Malagasy name for the radiated tortoise varies regionally in Madagascar: “Sokatra” (Merina), “sokaka” (Antandroy and Antanosy), and ‘kotroky” (Mahafaly) are commonly used. These words are pronounced “sookot,” “sookock,”’ and “kootrook, ” respectively. 2For more information on protected areas (policy and legislation, international activities, administration and management, system reviews, addresses, protected area information, defi- nitions of protected area designations, as legislated, together with authorities responsible for their administration, maps, and thé 1993 United Nations list of national parks and protected areas), travel, health information, global biodiversity hotspots, and country information consult the following sources which were used to write this country sunimary: Protected Areas: World Conservation Monitoring Centre's Protected Areas website location: www.unep-wenic.org/ parks/index.hitml and the IUCN, 1992. Protected Areas of the World: a review of national systems. IUCN, Gland, Switzerland.and Cambridge, United Kingdom. Xx + 352 p. Also, avail- able via the Internet at: www.unep-weme.org/protected_areas/data/pa_world_text.html: Travel: United States State Department, Bureau of Consular Affairs website: travel.state. gov; Health: Shoreland’s Travel Health Online: www.tripprep.com: Global Biodiversity Hotspots: ‘wwnw.conservation. org/hotspots/default.htm: and Country information: United States Central Intelligence eney’s (CIA) World Factbook, C/A, Washington, D.C. Also, available via the Internet at: www.odei, gov/cia/publications/factbook Manuscript received: 26-April-1997 Accepted: 15-January-1998 Copyright © 2000 Amphibian and Reptile Conservation, All rights reserved. ARC authorizes Amphibian and Reptile Conservation 2(1):1 5-23 photocopying for internal or personal use provided the appropriate fee is paid directly to the Copyright Clearance Center, Inc,, 222 Rosewood Dr., Danvers, MA 01923-4599, USA. Tel: (978) 750-8400; fax: (978) 750-4470; email: info@copyright.com; website: www.copyright.com Extinction and extinction vulnerability of amphibians and reptiles in Madagascar CHRISTOPHER J. RAXWORTHY' AND RONALD A. NUSSBAUM? 1Associate Curator, Department of Herpetology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192, USA *Division of Herpetology, Museum of Zoology, University of Michigan, Ann Arbor, Michigan, 48109-1079, USA Abstract.—In Madagascar, only two herpetofaunal extinction events are well documented. Both are extinctions of subfos- sil giant tortoises, which coexisted with humans for more than 1,000 years. Modern extinctions of amphibians and reptiles are also likely, but researchers and conservationists are probably overlooking these extinction events, because the most vulnerable species, with small relict populations, are easily missed during regional surveys. To date, conservation pro- grams in Madagascar have largely ignored many relict distribution species, restricted to transitional or rare habitat types. We provide four examples of species with relict distributions that we consider vulnerable to extinction. Based on ongoing surveys and systematic revisions, many new herpetofaunal species will be described in the future, some of which may require rapid conservation efforts to prevent extinction. Conservationists frequently measure extinction vulnerability using the World Conservation Union (IUCN) Red Lists, and fifteen endemic Malagasy amphibians and reptiles are included in the IUCN Threatened List. However, this list appears to reflect a historical bias towards conserving turtles and boas in Madagascar, listing eight species in these groups, although they represent just 2 percent of the island’s actual endemic herpetofauna. Ironically, this taxonomic bias may hinder attempts to prevent herpetofaunal extinctions, by promoting some species for conservation activities that are not vulnerable (e.g., tolerant of human habitat modification, or widespread) and ignoring many species that are soon to be lost. For Malagasy amphibians and reptiles, biogeographic data appear to provide more objective criteria with which to assess extinction threats rather than suspected rates of population decline. Key words. Extinction, conservation, Madagascar, herpetology, biogeography, reptiles, amphibians Introduction Madagascar’s “megafauna” extinction event that occurred dur- Despite the considerable interest in the amphibian and reptile ing the past two thousand years (Dewar 1984). The reasons for faunas of Madagascar and the widely held view that the island their extinctions are unknown, but it is certain that they coex- represents one of the world’s top conservation priorities (e.¢., isted with humans on the island between 2000 Before Present Wright 1997), it is surprising that there has been so little discus- (B.P.), the date of earliest human occupation of the island sion regarding the patterns of extinctions (past, present, or fu- (MacPhee and Burney 1991) and 750 B.P., the youngest carbon ture) for these two highly diverse and largely endemic groups. date for giant tortoise subfossils in Madagascar (Burleigh and This is of special concern because the objective of most conser- Arnold 1986). Raxworthy and Nussbaum (1996) have suggested vation programs in Madagascar is to maintain biodiversity. There- that the modern day practice of regular (typically annual) burn- fore these programs should prevent, or at least minimize, future ing of grassland and its peripheral forests, over much of the extinctions. island, was responsible for destroying the original habitats of The purpose of this paper is to provide a summary of grazing animals such as giant tortoises, while MacPhee and Marx herpetofaunal extinctions and extinction vulnerability in Mada- (1997) consider their extinction may be due to the human intro- gascar: first by reviewing the evidence of extinction; second by duction of exotic pathogens that caused hyperdisease. providing examples of species we believe are at imminent threat The former distribution of both species of giant tortoises of extinction; and third by evaluating and summarizing methods was vast, and their remains frequently represent the most com- used to measure risks of extinctions by the conservation com- mon subfossil material recovered by paleontological excavations munity using the World Conservation Union (IUCN) Red List (Dewar 1984). Because of their former large distribution and criteria. fact that much of island of Madagascar has still been poorly surveyed, there remains a chance that either species may survive Subfossil extinctions as a small population in one of the remoter regions, although this The only well documented cases of herpetofaunal extinctions in possibility must now be considered extremely unlikely. Madagascar are for the two species of giant tortoises, No other extinct reptile or amphibian subfossil material is Dispsochelys grandidieri and Dispsochelys abrupta, which once known from Madagascar, and there are no historically docu- occupied a large area of the central and western region of the mented cases of species going to extinction. island (Bour 1984). Both species went extinct during Possible extinctions ‘Correspondence. Fax: (212) 769-5031; email: rax@amnh. One possible extinction claimed in the literature (IUCN/UNEP/ org WWE 1987) is the colubrid snake Liophidium apperti. This 15 Plate 2 Plate 3 Plate 7 Plate 8 Plate captions: 1. Antanosy day gecko Phelsuma antanosy, from a Petriky relict forest fragment that has now been lost. 2. Chamaeleo belalandaensis, at Belalanda. One of Madagascar’s rarest chameleons. 3. Angonoka tortoise Geochelone yniphora, photographed at Ampijoroa. 4. Adult big headed Madagascar side-necked turtle Erymnochelys madagascariensis, photographed at the Andranomiditra River, Bevazaha village, Ankarafantsika Reserve. Photo: Gerardo Garcia Herrero. 5. Tomato frog Dyscophus antongili, Antongil’s Bay. 6. Dumeril boa Boa dumerili. Photo courtesy of Kevin and Sue Hanley. 7. Madagascar boa Boa madagascariensis. Photo: R. D. Bartlett. 8. Sanzinia tree boa Boa manditra. Photo by Peter Stafford, courtesy of The Natural History Museum, London. Photos 1 and 2: C. J. Raxworthy. Photos 3 and 5 courtesy of Franco Andreone, Museo Regionale di Scienze Naturali, Torino, Italy. 16 Plate 12 Plate 15 Plate 16 Plate captions: 9. Campan’s chameleon Furcifer campani. 10. Labord’s chameleon Furcifer labordi (female), Kirindy. 11. Minor chameleon Furcifer minor (female), photographed at “Mandraka Breeding Centre.” 12. Standings day gecko Phelsuma standingi. 13. Golden mantella Mantella aurantiaca, from Andasibe. Photo: C. J. Raxworthy. 14. Pyxis planicauda, photographed at Ampijoroa. 15. Radiated tortoise Geochelone radiata. 16. Armored chameleon Brookesia peramata. Photos 9 and 12 courtesy of R. D. Bartlett. Photos 10, 11, and 14 courtesy of Franco Andreone, Museo Regionale di Scienze Naturali, Torino, Italy. Photos 15 and 16 courtesy of Kevin and Sue Hanley. 17 CHRISTOPHER J. RAXWORTHY AND RONALD A. NUSSBAUM species was described from a single specimen collected from a forest 7 km north of Befandrina-sud in 1968 (Domergue 1983). Domergue reported that the forest had been subsequently cleared, which prompted IUCN/UNEP/WWE (1987) to consider that “the survival of this snake must now be in question.” We think it is premature to consider this species to be extinct, because this region remains so poorly studied, and it appears that similar forest types still survive in nearby sites in the Morombe region. We have collected Liophidium c.f. apperti at several localities in southern Madagascar, and although these specimens require fur- ther taxonomic study, this also suggests this species is not con- fined to Befandrina-sud. : Possibly the best candidate we have for an extinction event occurring during the past 100 years is for the colubrid snake Pseudoxyrhopus ankafinaensis, which was not described until 1994. This very large species is represented by just a single specimen collected in 1880 from montane forest of the High Plateau. Forest of this type is now almost completely degraded in this region (Raxworthy and Nussbaum 1994), and the absence of observations of P. ankafinaensis from surviving forest of lower elevation, or other montane forest sites suggests it was endemic to this high elevational region of the High Plateau. Dubious extinctions Other species have not been found for more than one hundred years, and, therefore, could also be considered extinct. However, before this conclusion can be made with any certainty, it is important to confirm that the species were collected in Mada- gascar. During the 1800's, specimen localities were sometimes confused or lost, frequently because museums were receiving collections from throughout the world during this period. An example is the colubrid snake Pseudoxyrphopus punctatus, which was thought for a period of more than 50 years to have been collected in Madagascar. Subsequently, this was identified as a Brazilian snake in the genus Sordellina (see Raxworthy and Nussbaum 1994). Another possible example of a species that may not ever have been collected in Madagascar is Ailuronyx trachygaster. This giant gecko, known only from a single specimen MNHP 6679 (Muséum National d’ Histoire Naturelle, Paris) was col- lected by an unknown person from a locality listed in the Paris Museum as Madagascar. Interestingly, the two specimens cata- logued before A. trachygaster: MNHP 6677-8, are both Ailuronyx seychellensis collected in the Seychelles (6677 was collected by Péron and Lesueur). Possibly A. trachygaster was also collected on the same voyage. A. trachygaster 1s a valid species, with very different characters compared to A. seychellensis. Despite some uncertainty about the geographic origin, we suspect A. trachygaster is extinct because no new specimens (of what should be a very conspicuous gecko) have been found in more than 140 years. As aresult of recent surveying in Madagascar, most of the rarer species in collections have been rediscovered, such as Zonosaurus boettgeri, Phyllodactylus brevipes, Paragehyra petiti, Uroplatus alluaudi, and Pseudoxyrhopus ambreensis, but a few notable exceptions remain, especially among the most cryptic groups. For example, among the skinks, the following species have not been collected since their original description: Mabuya betsileana (possibly African), Cryptoscincus minimus, Pseudoacontias madagascariensis, and Paracontias rothschildi. Their exact distributions remain unknown at present (in Mada- 18 gascar or elsewhere) but we consider it premature to. consider any of them to be extinct. Vulnerable to extinction We consider species in the category “Vulnerable to Extinction” to have populations that are sufficiently small that near-future extinction in the wild can be considered highly likely (without conservation action). These species are restricted to primary habitat that has declined so dramatically, that they now survive only in tiny isolated patches, which are continuing to decline. As a result, the species dependent on this primary habitat are now endemic to a very small region of the island. To illustrate this type of extinction vulnerability, we have selected four spe- cies to serve as examples. However, many other herpetofaunal species exhibit similar extinction vulnerability in Madagascar. Bernard’s mantella frog Mantella bernhardi (Plate 18A and B). During a survey of Tolongoina made by Nussbaum in 1993, M. bernhardi was first discovered, in a single patch of relict forest. Within months, commercial collectors visited the locality and supplied André Peyrieras, a commercial exporter, with animals to be sold into the pet trade. Vences et al. (1994) described this species based on animals they obtained from commercial collectors working for Peyrieras. They provided no data on the exact locality, habitat requirements, or conser- vation concerns of this species, because they never saw this species in the wild. A subsequent visit to the Tolongoina region by Raxworthy in 1994 discovered that the only known M. bernhardi habitat had been further cleared, so that no more than 20 ha of forest survived (Plate 17). No other populations were discovered dur- ing this visit, and almost all other primary rain forest had been cleared from the area. Deforestation of primary forest has been so extensive in this region that almost no fragments now survive east of the Faraony River. Topographic maps indicate that this forest was almost entire during aerial photography made of the region between 1950 and 1965 (FTM 1974). Although it is likely that M. bernhardi once had a more widespread distribution, the deforestation pattern of low eleva- tion rain forest (below 800 m elevation) in this region suggests that little forest of this type now survives. The closest pro- tected forest of this type occurs in the lowest elevational areas of the Ranomafana National Park (the southern boundary limit is 18 km to the northwest). It is not known if this species occurs there, or even if the habitat is similar to that at Tolongoina. The low elevation rain forest at Manombo Reserve (140 km to the south) does not appear to have populations of M. bernhardi based on a survey by Raxworthy in 1991. Because no known populations of M. bernhardi occur within a protected area and its habitat appears to have been almost completely destroyed, we consider this species extremely vulnerable to extinction. Antanosy day gecko Phelsuma antanosy (Plate 1). This day gecko is restricted to coastal fragments of forest in the Tolagnaro region of southeastern Madagascar. Raxworthy and Nussbaum (1993) described three sites (forest fragments) in the description of this species at Petriky, Ste. Luce, and Tapera. Since then, one new site has been discovered near Manambaro, and one site (the Petriky fragment, area 81 ha in 1989) has been completely destroyed. None of the three surviving forest frag- HERPETOFAUNAL EXTINCTION IN MADAGASCAR ments is greater than 191 ha, and the Manambaro site 1s decreas- ing in area rapidly as a result of annual burning. Unlike some Phelsuma species, P. antanosy does not sur- vive in degraded or heavily modified habitats and appears to be entirely dependent on the transitional dry-humid forests that are restricted to a small region of the southeast. This habitat has now been almost completely lost from the region, having been degraded and cleared for charcoal production and agricultural land. None of the forest fragments where P. antanosy occurs is within a protected area. The surviving populations are now so small that we con- sider this species to be extremely vulnerable to extinction. Con- servation efforts are required if the last fragments of habitat are to be saved. Belalanda chameleon Chamaeleo belalandaensis (Plate 2). Very little information exists on either the distribution or habitat requirements of this chameleon. However, all data to date suggest it is endemic to a tiny region of Madagascar. The only locality is Belalanda, Southwestern Madagascar, where Raxworthy has recorded individuals as recently as 1995. How- ever, we were unable to find this species at other sites, despite intensive herpetofaunal surveys within 10-50 km of Belalanda. The habitat where this chameleon was recorded is degraded gal- lery forest, which has now been almost completely cleared. Because C. belalandaensis was not found in the other surviving primary forest habitats of the region, we suspect it is restricted to gallery forest. This chameleon is an example of a species for which bio- geographic data are urgently needed. It would be valuable to record the exact distribution limits for this chameleon, so that conservation efforts could be directed at those populations, which appear to be most viable. The species has already been subject to some commercial collecting, but the impact on the population is not known. The very localized distribution of C. belalan- daensis, in part confirmed by our survey efforts in surrounding areas, suggests that this species is vulnerable to extinction be- cause of the apparently tiny surviving populations. Angonoka tortoise Geochelone yniphora (Plate 3). This tortoise is restricted to an area of less than 1,000 km’ in the Baly Bay area of Western Madagascar. The distribution appears to be relict, because the species is now confined to two isolated areas, one to the east of the Baly Bay (Cap Sada), and the other to the west (Belambo). Dispersal across the bay is likely to be very limited or impossible. Therefore, the eastern and western popu- lations appear to be genetically isolated. Only five sites are known or suspected to have G. yniphora populations (Durrell et al. 1994). The suspected sites are based on interviews with local people and the occurrence of suitable habitat. The wild populations are thought to have declined recently due to habitat loss, predation of eggs, and juveniles by African bush pig (Potamochoerus larvatus), and collecting by people (Durrell et al. 1994). At Cap Sada the first detailed population study is now underway. The isolated eastern and western populations of G. yniphora suggest this species was previously distributed to the south of Baly Bay and that the distribution range has contracted since this time. The habitat of this tortoise is a mosaic of deciduous forest and bamboo scrub, which appears to be replaced by a palm savanna as a result of frequent burning. Between 1949- 19 1973, Curl et al. (1985) reported only minor change in tortoise habitat distribution, and suggested this habitat is no longer de- clining. However, it should be noted that during this same pe- riod, the setting of fires was both illegal and frequently enforced by local communities. The tiny isolated populations and the restricted area of surviving habitat clearly indicate that G. yniphora is vulnerable to extinction if further habitat decline continues. A major con- servation program (Project Angonoka) coordinated by the Mala- gasy Water and Forests Authority and Jersey Wildlife Preserva- tion Trust is now underway in the Cap Sada region to protect this population. IUCN Threatened species This section includes those species listed with a threatened cat- egory (Critically Endangered, Endangered, or Vulnerable) in the most recent IUCN Red List (IUCN 1996). Threatened species, as recognized by IUCN, refers specifically to the level of risk of extinction. Fifteen endemic Malagasy amphibians and reptiles are currently listed as threatened in the Red List; three species are classified as Endangered, and another 12 as Vulnerable. A significant advance with the new IUCN categories 1s that threats are presented in a quantified format. A summary of these en- demic species, as well as their perceived risks of extinction, is given in Table 1. IUCN Endangered species Big headed Madagascar side-necked turtle Erymno- chelys madagascariensis (Plate 4). The criteria used for considering this turtle as endangered is based on a 50 percent population decline in 10 years or three generations. The genera- tion time (average age of parents in the population, as used by IUCN) for this turtle is unknown, but without doubt will be greater than 10 years. For long-lived species IUCN suggests a cap of 75 years IUCN 1996). Even working with this time span, however, the problem is a lack of population data for this species, both modern and historical. There is no doubt that populations are being exploited by fishing practices, and Kuchling and Mittermeier (1993) have presented evidence that two populations have gone extinct out of a sample of nine lakes. Nevertheless, these authors recog- nized that the status of river populations have not been esti- mated because of an almost complete lack of surveys within the rivers of Western Madagascar. In addition, we still lack modern data on either lake or river populations throughout much of the species’ distribution range, especially the many remote regions of the west. Because our knowledge of the populations of E. madagascariensis is so incomplete, it is questionable if we can claim even a suspected population reduction of 50 percent. The large historical distribution area of this turtle, with an extent of occurrence of approximately 100,000 km’, does not suggest to us that this species is yet at high risk to extinction, although clearly more field work is required to determine the current distribution of this species. Angonoka tortoise Geochelone yniphora. Like E. madagascariensis, the three-generation time for G. yniphora will be greater than 10 years. We are unaware of data to support a 50 percent population decline over either 10 years or three genera- tions. However, this tortoise does qualify for Endangered status based CHRISTOPHER J. RAXWORTHY AND RONALD A. NUSSBAUM fy a Plate 17. Som Raxworthy. we pote as nee can Gi de : = Sn ea A MR LP ae s a 2 Plate 18A. Bernard’s mantella frog Mantella bernhardi. Probably Plate 18B. Mantella bernhardi (belly pattern). Photo courtesy of the rarest Mantella species. Photo courtesy of Franco Andreone, Franco Andreone, Museo Regionale di Scienze Naturali, Torino, Museo Regionale di Scienze Naturali, Torino, Italy. Ttaly. 20 HERPETOFAUNAL EXTINCTION IN MADAGASCAR on the criteria of an extent of occurrence less than 5,000 km?, and less than 5 isolated populations (see earlier). Madagascar flat tailed tortoise Pyxis planicauda (Plate 14). Adult wild P. planicauda are reported to have 10-30 growth rings (Kuchling and Bloxam 1988), which appear to reflect the growth between each period of annual aestivation’. Using a mean generation time of 20 years, three generations would represent 60 years. The Endangered category given to this tortoise is based on a 50 percent decline in the population during this period. Without question, populations of this tortoise are declin- ing due to habitat destruction for cultivation (Kuchling and Bloxam 1988; Bloxam et al. 1993; Raxworthy, pers. obs.). How- ever, our knowledge of the species distribution is actually im- proving. New localities are being discovered, including impor- tant range extensions further to the north (Behler et al. 1993; Bloxam et al. 1993). The extent of occurrence for P. planicauda is currently about 500 km?(based on the localities given in Fig. 1, Behler et al. 1993), with the species endemic to a small region of coastal, western, deciduous forest between the Morondava and Tsiribihina Rivers. This justifies P. planicauda being considered Endangered (the extent of occurrence is significantly less than 5,000 km?) based on its small distribution, rather than the crite- ria of rate of population decline. The localized distribution of this species, and its dependence on native forest, suggests this species is vulnerable to extinction. i IUCN Vulnerable species Of the 12 species classified as Vulnerable, 11 are so classifica- tion based on a criteria of a 20 percent reduction in population over 10 years or three generations. The mean wild generation time is unknown for any of these species, although a study of radiated tortoise Geochelone radiata at one site has yielded some data on the population age structure (Razafindrakoto 1987). The population size and rate of decline have never been mea- sured for these species, although populations of some species restricted to primary habitats are declining in areas subjected to habitat loss. For those species restricted to primary habitats, it may be reasonable to positively correlate rates of habitat loss (e.g., based on satellite images) to rates of population decline. Four of the Vulnerable species: the tomato frog Dyscophus antongili (Plate 5), the Dumeril boa Boa dumerili (Plate 6), the Madagascar boa Bea madagascariensis (Plate 7), and the Sanzinia tree boa Boa manditra (Plate 8), are not restricted to primary vegetation. D. antongili is found in secondary habi- tats, such as around villages and in areas of cultivation (Raxworthy 1991), and even in towns such as Maroantsetra (Glaw and Vences 1994). All three of the Malagasy boas (B. dumerili, B. madagascariensis, and B. manditra) are frequently found in heavily degraded habitats and cultivated areas, even including close proximity to villages where they are probably feeding on commensal rats. For these species, the loss of pri- mary forest does not lead to local extinction, although their population changes (negative or positive), when primary for- est is converted, are not known. The most likely source of population decline for D. antongili, and the three species of boa, is commercial collect- ing, although now all four species are listed on the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) Appendix I, which prevents legal interna- tional trade. Malagasy boas are collected in regions such as Marovoay to supply a domestic leather trade. However, boas continue to be among the most common snakes found in Mada- gascar, and the scale of commercial collecting may be compa- rable to the number of road kills that can be seen throughout Madagascar during the rainy season. The approximate extent of occurrence of each species is: D. antongili 10,000 km’, B. dumerili, 120,000 km?, B. madagascariensis 40,000 km?, and B. manditra, 100,000 km’. Based on the large distribution area, and the broad tolerance to habitat degradation, none of these species appear at risk to extinction. Of the other eight vulnerable species which are restricted to primary habitats, it is impossible for us to assess the criteria used for the following species: Campan’s chameleon Furcifer campani (Plate 9), Labord’s chameleon Furcifer labordi (Plate 10), the Minor chameleon Furcifer minor (Plate 11), Standing’s day gecko Phelsuma standingi (Plate 12), and the golden mantella Mantella aurantiaca (Plate 13), because these are based on rates of population decline (observed or suspected) for which we have been unable to find obvious supporting data. Table 1. IUCN threatened species of endemic amphibians and reptiles of Madagascar (IUCN 1996). SPECIES CATEGORY Erymnochelys madagascariensis ENDANGERED Pyxis planicauda ENDANGERED Geochelone yniphora ENDANGERED Dyscophus antongili VULNERABLE Mantella aurantiaca VULNERABLE Pyxis arachnoides VULNERABLE Geochelone radiata VULNERABLE Phlesuma standingi VULNERABLE Brookesia peramata VULNERABLE Furcifer campani VULNERABLE Furcifer labordi VULNERABLE Furcifer minor VULNERABLE SUMMARY OF CRITERIA 50% population decline in 10 years or 3 50% population decline in 10 years or 3 50% population decline in 10 years or 3 < 5,000 km? extent of occurance and < 20% population decline in 10 years or 3 generations 20% population decline in 10 years or 3 generations < 20,000 km? extent of occurance, < 10 isolated populations and decline in population 20% population decline in 10 years or 3 generations < 20,000 km? extent of occurance, < 10 isolated populations 20% population decline in 10 years or 3 generations 20% population decline in 10 years or 3 generations and < 100 km? area of occupancy or < 5 locations 20% population decline in 10 years or 3 20% population decline in 10 years or 3 20% population decline in 10 years or 3 generations generations generations, 5 isolated populations generations generations generations 21 CHRISTOPHER J. RAXWORTHY AND RONALD A. NUSSBAUM These species have been commercially traded, but the impact of this trade on wild populations is unknown or only poorly known. Two of these species have a very limited distribution: M. aurantiaca is restricted to eastern rain forest with an approxi- mate extent of occurrence of 3,000 km? and P. standingi re- stricted to deciduous western forest in the Toliara region with an approximate extent of occurrence of 2,500 km? (although an unconfirmed report suggests this species may also occur fur- ther north). In the case of P. standingi we have found this gecko is extremely rare (if not locally extinct) in forest within 5 km of major roads at Ifaty (none were seen during at least 60 man- days of searching). Commercial collecting has been occurring in this area for some time, and in many cases collecting poles could still can be found lying against baobab tree trunks, left by the local collectors in the hope of catching any remaining geckos. However, because the original. population size and density in this region is unknown, the impact of commercial collecting cannot be measured. The impact of collecting on M. aurantiaca 1s also unknown. Zimmermann and Zimmermann (1994) suggest that local popu- lation extinctions occurred in a swamp system between 1966 and 1993 as a result of habitat loss. They therefore propose that commercial collecting be prevented by placing M. aurantiaca on Appendix I of CITES. For M. aurantiaca and P. standingi, their limited known distributions, primary habitat requirements, and commercial col- lecting pressures, all suggest that both species may be vulner- able to extinction. Distribution data is used as criteria for Vulnerable status for three species: Pyxis arachnoides, G. radiata (Plate 15), and Brookesia perarmata (Plate 16). For the two tortoise species, P. arachnoides and G. radiata, the extent of occurrence is actu- ally greater than 20,000 km? (we estimate both around 30,000 km’), one of the criteria used to support the vulnerable status. In addition, although the populations of these species are also likely to becoming more fragmented, as a result of habitat loss, there are without doubt many more than 10 isolated popula- tions (another criteria used to assign these species vulnerable status, see Table 1). Therefore, the only justification that can be made for considering these tortoises, as Vulnerable, using the IUCN criteria, is the rate of observed or suspected popula- tion decline, for which we currently have very little quantita- tive information. The large distributions of P. arachnoides and G. radiata do not suggest that either species is yet threatened by extinction. B. perarmata is only known from one site: Bemaraha, in Western Madagascar. Although the surface area of the Bemaraha Reserve is large (152,000 ha), our survey of this protected area at three well separated sites, yielded this chameleon at just one locality indicating it is not continuously distributed throughout the reserve. This clearly supports the “less than five site” crite- ria for Vulnerable status. Discussion Despite the major environmental problems that have been de- veloping for so long in Madagascar, it is perhaps surprising that we have no evidence of any herpetofaunal extinction events occurring on the island over the past 500 years. Although this suggests the island’s species diversity has not declined during this period, we believe herpetofaunal extinctions have been oc- 22 curring, and are occurring now, but that researchers and conser- vationists have overlooked these events. Species most vulnerable to extinction will have small popu- lations, and therefore tiny relict distributions. This makes them difficult to survey, and, therefore, easily missed. A modern ex- ample of this is P. antanosy, distributed in a region of Madagas- car that has been subject to herpetological collecting for over 100 years. We suspect many more herpetofaunal species are on the verge of going extinct in transitional or rare habitat types. Re- gions that experience highly localized climatic conditions fre- quently have unusual transitional habitats in Madagascar. In many cases these localized habitats have been further reduced to small relict patches as a result of human exploitation and have attracted little research attention because of their small size or isolated locality. Examples of major sites we have surveyed, which offer these conditions, include Analavelona, Isalo, Bemaraha, Kelifely, Ambohijanahary, Namoroka, and Ankarana. We are in the process of describing new species from all of these sites, most of which are likely to be locally endemic, but active conservation programs are now underway in just three sites (Isalo, Bemaraha, and Ankarana). Many of these new species are reduced to such small population sizes that they are obvi- ously some of the most vulnerable herpetofaunal species in Madagascar. Other potentially vulnerable species include Phelsuma masohoala, Uroplatus malahelo, Pseudoxyrhopus kely, and Alluaudina mocquardi. Of the 15 endemic Malagasy amphibians and reptiles con- sidered Threatened by the IUCN Red List, we consider (based on distribution and habitat requirements) just three species to be obviously vulnerable to extinction: P. planicauda, G. yniphora, and B. perarmata. Another two species, M. aurantiaca and P. standingi, appear to also have small distributions, but further fieldwork is required to confirm this. The other 10 species do not appear to be vulnerable to extinction because of their much larger distributions, and in some cases, broad tolerance to human modification of primary habitats. In part, the IUCN Threatened List actually reflects a his- torical bias towards conserving turtles and boas in Madagascar, rather than actual risks to extinction. It is worth noting that 8 of the 15 Red List species are boas or turtles (53 percent), although this group represents just 2 percent of the island’s actual en- demic herpetofauna. Ironically, this conservation bias is actually hindering attempts to prevent herpetofaunal extinctions in Mada- gascar, by promoting some species for conservation activities that are not vulnerable and by ignoring many species that are about to be lost. Rates of past, present, or future population decline (ob- served, estimated, inferred, or suspected) are used as IUCN criteria for all but one of the 15 threatened species. This sur- prises us, as we are unsure what types of evidence were used to support or estimate these rates of population decline. For the herpetofauna of Madagascar, we think that it would be far more effective to base vulnerability to extinction on biogeographic data, rather than estimated population declines. The advantage is that biogeographic data is available for most species, and these data can be compared between different taxa to determine con- servation priorities. Of course, biogeographic data are never com- plete for any species, but it can be obtained with far less effort than undertaking population studies throughout a species distri- bution range. HERPETOFAUNAL EXTINCTION INMADAGASCAR Table 2 includes four biogeographic criteria that we con- sider to be the most important in determining vulnerability to extinction for amphibians and reptiles in Madagascar. Other population factors will also influence the risk of extinction (e.g., population density, generation time, reproductive output, pre- dation, and human collecting), but since it is unlikely we will ever have detailed information of this type over the entire distri- butional range on any herpetofaunal species, we think the most objective data set available to us is biogeographic. Table 2. Measuring extinction vulnerability with biogeo- graphic criteria. EXTINCTION RISK BIOGEOGRAPHIC CRITERIA HIGH LOW EXTENT OF OCCURANCE Small Large NUMBER OF KNOWN SITES Few Many DISTRIBUTION STRUCTURE Fragmented Continuous HABITS Specialist of a declining habitat Specialist of a stable habitat Provided that taxonomic biases are avoided, we believe that the analysis of biogeographic data has the potential to reveal a much more realistic picture of extinction threats to the amphib- ians and reptiles in Madagascar, compared to the criteria cur- rently being employed. This will identify those species of great- est concern, which could then become the focus of conservation programs aimed at maintaining existing levels of biodiversity in Madagascar. Acknowledgments.—This research was made possible with the help of J.-B. Ramanamanyjato, A. Raselimanana, Angelin and Angeluc Razafimanantsoa, and the cooperation of the Ministére de |’Enseignement Supérieur, the Ministére de la Production Animale et des Eaux et Foréts, and the Ministére de la Research Scientifique et Technologie pour le Developpement. This re- search was funded in part by grants from the National Science Foundation (DEB 90 24505, 93 22600), the National Geographic Society, and Earthwatch. References Behler, J. L., Bloxam, Q. M. C., Rakotovao, E. R., and Randriamahazo, H. J. A. R. 1993. New localities for Pyxis planicauda in West- central Madagascar. Chelonian Conservation and Biology 1(1):49-51. Bloxam, Q. M. C., Behler, J. L., Rakotovao, E. R., and Randriamahazo, H. J. A. R. 1993. Distribution of the Madagascar flat-tailed tortoise Pyxis planicauda. Dodo Jersey Wildlife Preservation Trust 29:149-156. Bour, R. 1984. Les tortues terrestres géantes des iles de 1’Ocean Indien Occidental: données géographiques, taxinomiques et phylogénétiques. Studia Geologica Salmanticensia Vol. Espe- cial 1 (Studia Palaeocheloniologica 1):17-76. Burleigh, R. and Arnold, E. N. 1986. Age and dietary differences of recently extinct Indian Ocean tortoises (Geochelone s. lat) revealed by carbon isotope analysis. Proceedings of the Royal Society of London B 227:137-144. Curl, D. A., Scoones, I. C., Guy, M. K., and Rakotoarisoa, G. 1985. The Madagascan tortoise Geochelone yniphora: current status and distribution. Biological Conservation 34:35-54. Dewar, R. E. 1984. Extinctions in Madagascar, p. 574-593 in Mar- tin, P. S. and Klein, R. G. (editors). Quaternary Extinctions: a prehistoric revolution. University of Arizona Press, Tucson, USA. 23 Domergue, C. A. 1983. Notes sur les Serpents de la région malgache III. Description de trois especes nouvelles rapportées au genre Liophidium Boulenger, 1896. Bulletin du Museum National de Histoire Naturelle, Paris. 4th Sér. 5, Section A (4):1109- lip. Durrell, L., Rakotonindrina, R., Reid, D., and Durbin, J. 1994. The recovery of the angonoka (Geochelone yniphora)—an inte- grated approach to species conservation, p. 384-393 in Olney, P. J. S., Mace, G. M., and Feistner, A. T. C. (editors). Creative Conservation: interactive management of wild and captive animals. Chapman and Hall, London, UK. FTM. 1974. Carte de Madagascar 1:50,000, Tolongoina, P-53 Sud. Foiben Taosarintanin | Madagasikara, Antananarivo, Mada- gascar. Glaw, F. and Vences, M. 1994. A Fieldguide to the Amphibians and Reptiles of Madagascar. Second edition. Moss Druck, Lever- kusen, Germany. 480 p. IUCN. 1996. 1996 IUCN Red List of Threatened Animals. IUCN, Gland, Switzerland. 448 p. IUCN/UNEP/WWF. 1987. Madagascar, An Environmental Profile. International Union for Conservation of Nature and Natural Resources, IUCN, Gland, Switzerland. 374 p. Kuchling, G. and Bloxam, Q. M. C. 1988. Field-data on the Madagascan flat tailed tortoise Pyxis (Acinixys) planicauda. Amphibia- Reptilia 9:175-180. Kuchling, G. and Mittermeier, R: A. 1993. Status and exploitation of the Madagascan big-head turtle, Erymnochelys madagas- cariensis. Chelonian Conservation and Biology 1(1):13-18. Macphee, R. D. E. and Marx, P. A. 1997. The 40,000 year plague: human, hyperdisease, and first-contact extinctions, p. 169- 217 in Goodman, S. M. and Patterson, P. D. (editors). Natural Change and Human Impact in Madagascar. Smithsonian In- stitution Press, Washington, USA. Macphee, R. D. E. and Burney, D. A. 1991. Dating of modified femora of extinct dwarf Hippopotamus from southern Mada- gascar: implications for constraining human colonization and vertebrate extinction events. Journal of Archaeological Sci- ence 18:695-706. Raxworthy, C. J. 1991. The reptiles and amphibians of Ambatovaky, p. 3.1-3.9 in Thompson, P. M., and Evans, M. I. (editors). A Survey of Ambatovaky Special Reserve Madagascar. Madagas- car Environmental Research Group, London, UK. Raxworthy, C. J. and Nussbaum, R. A. 1993. A new Madagascan Phelsuma, with a review of Phelsuma trilineata and comments on Phelsuma cepediana in Madagascar (Squamata: Gekkon- idae). Herpetologica 49(3):342-349. Raxworthy, C. J. and Nussbaum, R. A. 1994. A review of the Madagascan snake genera Pseudoxyrhopus and Heteroliodon (Squamata: Colubridae). Miscellaneous Publications of the Uni- versity of Michigan 182:1-46. Raxworthy, C. J. and Nussbaum, R. A. 1996. Montane amphibian and reptile communities in Madagascar. Conservation Biology 10:750-756. Razafindrakoto, L. A. 1987. Contribution a l’etude bio-eco- ethologique de Geochelone radiata (Shaw 1802) (Famille des testudinidae) dans la Réserve Spéciale de Beza-Mahafaly. Mémoire de Fin d’Etude, EESSA, Université de Madagascar, Antananarivo, Madagascar. Wright, P. C. 1997. The future of biodiversity in Madagascar: a view from Ranomafana National Park, p. 381-405 in Goodman, S. M. and Patterson, P. D. (editors). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press, Wash- ington, USA. Vences, M. F., Glaw, F., Peyrieras, A., Bohme, W., and Busse, K. 1994. Der mantella-madagascariensis-komplex: wiederent- deckung von Manatella cowani und beschreibung von Mantella bernhardi n. sp. DATZ 47(6):390-393. Zimmerman, E. and Zimmerman, H. 1994. Reproductive strategies, breeding, and conservation of tropical frogs: dart-poison frogs and Malagasy poison frogs, p. 255-266 in Murphy, J. B., Adler, K., and Collins, J. T. (editors). Captive Management and Con- servation of Amphibians and Reptiles. Society for the Study of Amphibians and Reptiles, Ithaca, USA. Manuscript received: 05-November-1997 Accepted: 22-December-1997 Copyright 2000 Amphibian and Reptile Conservation. All rights reserved. ARC authorizes photocopying for internal or personal use provided the appropriate fee is paid directly to the Copyright Clearance Center, Inc., 222 Rosewood Dr., Danvers, MA 01923-4599, USA. Tel (978) 750-8400; fax: (978) 750-4470; email: info@copyright.com; website: www.copyright.com Amphibian and Reptile Conservation 2(1):24-29. New data on the distribution, status, and biology of the New Caledonian giant geckos (Squamata: Diplodactylidae: Rhacodactylus spp.) AARON M. BAUER’ AND ROSS A. SADLIER? ‘Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085-1669, USA ?Department of Herpetology, Australian Museum, 6-8 College Street, Sydney, New South Wales 2000, AUSTRALIA Abstract.—Recent collections and observations of the New Caledonian giant geckos (Rhacodactylus) result in range extensions and new information regarding the biology of these lizards. Significant range extensions are reported for the rough-snouted giant gecko (R. trachyrhynchus) and for the recently rediscovered Guichenot’s giant gecko (R. ciliatus). Field observations confirm the association of the knob-headed giant gecko (R. auriculatus) with plants of the family Cunoniaceae and that Leach’s giant gecko (R. leachianus) feeds on fruit. Range extension of some species, and data on local abundance allows a reassessment of their conservation status. Despite implied increases in giant gecko density and range, significant threats from habitat loss, introduced predators, and illegal trade leave all species at risk. Key words. Rhacodactylus, geckos, New Caledonia, distribution, diet, conservation status Introduction Rhacodactylus is one of three carphodactyline* gecko genera occurring in New Caledonia. The genus includes the largest liv- ing species of gecko, R. leachianus (> 250 mm snout-vent length; Russell and Bauer 1986) and has attracted scientific attention because of unusual characteristics such as viviparity! (in R. trachyrhynchus; Bartmann and Minuth 1979), the possession of and specialized dentition (in R. auriculatus; Bauer and Russell 1990; Bauer and Sadlier 1994b), and prehensile tails (all species; Bauer 1990; Bauer and Russell 1994). Species of this genus have also attracted much popular attention, especially among ter- rarium keepers, because most species thrive and reproduce well in captivity (Henkel 1987, 1991, 1993; Henkel and Schmidt 1991; Tytle 1992). The systematics and morphology of Rhacodactylus have recently been the focus of significant investigation (Bauer 1990; Bauer and Russell 1990; Bauer et al. 1993; Seipp and Klemmer 1994: Good et al. 1997). However, knowledge of the distribu- tion and biology of these geckos remains largely incomplete (Bauer and Sadlier 1993). Aside from brief reports of various aspects of natural history (e.g., Mertens 1964; Meier 1979; Sameit 1985; Bauer 1990; Bauer and Vindum 1990; Henkel 1991), field-based data are limited to a few investigations of diet (Bauer and DeVaney 1987; Bauer and Sadlier 1994b) and general ac- counts of behavior and ecology in nontechnical works (de Vosjoli 1995; de Vosjoli and Fast 1995). However, increasing awareness of the uniqueness of the flora and terrestrial fauna of New Caledonia (Myers 1988; Mittermeier et al. 1996) has given new impetus to the collection of basic distributional and biological data for all members of the New Caledonian herpetofauna, and previously understudied areas are being surveyed more system- atically (e.g., Isle of Pines, Bauer and Sadlier 1994a). Observa- tions made by the authors during several recent expeditions to ‘Correspondence. Tel: (6/0) 519-4857; fax: (610) 519-7863; email: aaron.bauer@ villanova.edu 24 New Caledonia have yielded new distributional and/or dietary information for all six of the recognized species of Rhacodactylus. In addition, preliminary assessments of genetic variation within certain species (reported more fully in Good et al. 1997) were made on the basis of tissue samples accumulated over a series of trips since the mid-1980’s. We present these data here as they help to provide a more accurate picture of the geographic ranges and biological requirements of the geckos and may be useful in establishing the conservation status of Rhacodactylus species. Materials and methods Herpetological collections and observations were made on main- land New Caledonia during trips in 1994 and 1995. Specimens were collected under a series of permits issued by the conserva- tion authorities of the Province Sud (Marcel Boulet) and Prov- ince Nord (Christian Papineau) of New Caledonia to the au- thors. Preliminary estimates of genetic divergence between popu- lations were based on results derived from allozymicé data. De- tails of the electrophoretic methodology employed are presented in Good et al. 1997. Specimens cited are housed in the collec- tions of the Australian Museum (Sydney)-AMS, the Natural History Museum (London)-BMNH, and the California Acad- emy of Sciences (San Francisco)-CAS. Results and discussion Knob-headed giant gecko (Rhacodactylus auriculatus) [Plate 1]. B6hme and Henkel (1985) reported a striped color phase of R. auriculatus, now known to be common. Although polymorphisms were noted within a single population of this species, there were no fixed differences among population samples from four different localities and no suggestion of significant intraspecific genetic variation. This tends to corroborate mor- phological observations that this species is generally polymorphic" throughout its range but that there are no geo- graphically related trends in character variation (Bauer 1990). a Plate 1 Plate 5A etal oh wh Plate 5B Plate 6 Plate captions: 1. Knob-headed giant gecko, Rhacodactylus auriculatus, from Riviére Bleue. 2. Bavay’s giant gecko Rhacodactylus chahoua. Photo courtesy of R. D. Bartlett. 3A. Guichenot’s giant gecko, Rhacodactylus ciliatus, from Riviere Bleue (adult with autoto- mized tail). 3B. Rhacodactylus ciliatus, from Riviere Bleue (subadult with complete original tail and complex body patterning). 4. Leach’s giant gecko, Rhacodactylus leachianus, from Mt. Koghis. 5A. Roux’s giant gecko, Rhacodactylus sarasinorum, from Kwa Néie (adult retaining bold white dorsal markings). 5B. Rhacodactylus sarasinorum, from Riviére Bleue (adult with mottled dorsal pattern). 6. Rough-snouted giant gecko Rhacodactylus trachyrhynchus, from Mt. Aoupinié. Photos 1, 3A and B, 4, 5A and B, and 6: Ross A. Sadlier. 25 AARON M. BAUER AND ROSS A. SADLIER Plate a Geissois sp. from Mt. Do, southern New Caledonia. This plant is apparently utilized as a food source by Rhaco- dactylus auriculatus. Bavay (1869) first reported on the diet of this species, indicating that it eats the flowers of Geissois (Cunoniaceae). This was verified by the recovery of anthers, stamens, and (pos- sibly) pollen belonging to either a member of this family (or the Myrtaceae) from the stomach of one specimen (Bauer and Sadlier 1994b). On 9 January 1995 further evidence of the specific association between R. auriculatus and Geissois was obtained when geckos were found active on flowering specimens of Geissois spp. (Plate 7) 1.3-2.6 km from the summit of Mount (Mt.) Do (21°45'S, 166°00' E) in south central New Caledonia. Bavay’s giant gecko (Rhacodactylus chahoua) [Plate 2]. The known distribution of this species in central and south- ern mainland New Caledonia has been expanded by the capture of a specimen from Sarraméa (AMS R144171) and by speci- mens from unstated localities on the Isle of Pines (de Vosjoli 1995; de Vosjoli and Fast 1995) [Fig. 1]. This species is rather polymorphic with respect to coloration (Bavay 1869; Béhme and Henkel 1985; Bauer 1985), but the comparison of allozymes from individuals separated by more than 100 km suggests rela- tive genetic uniformity. Guichenot’s giant gecko (Rhacodactylus ciliatus) [Plate 3A and 3B]. This species was numerous for the first 20 years after its description (e.g., Bavay 1869) and then was not seen again for over 100 years, despite extensive searches by several researchers. It was regarded as extinct (see Bauer and Sadlier 1993). In 1994, the species was rediscovered and has since been found at a variety of localities on the Isle of Pines (Seipp and 26 Klemmer 1994; Kullmann 1995) and several smaller offshore islands (de Vosjoli 1995). De Vosjoli (1995) and de Vosjoli and Fast (1995) recorded this species as common on the Isle of Pines but stated that it was not present on the mainland of New Caledonia. Despite their claims, Bavay (1869), whose data have proved to be very accurate (see Bauer and Sadlier 1994b), re- ported collecting seven specimens of this species at several (un- specified) localities on the mainland, and the type locality given by Guichenot (1866) was at Canala (21°35'S, 165°56' E), also on the mainland. The persistence of this species on the mainland was verified by two specimens (AMS 146594-5) collected by R. A. Sadlier near Pont German, Riviere Bleue (22°06' S, 166°38' E) in the extreme south of New Caledonia (Fig. 1). It has subse- quently been taken at other localities on the mainland (Girard and Heuclin 1998; Bauer and Sadlier 2000), suggesting that it may be relatively widespread. Leach’s giant gecko (Rhacodactylus leachianus) [Plate 4]. This species has a broad distribution in the wetter areas of the New Caledonian mainland, especially along the east coast (Bauer 1990). Boulenger (1885) first recorded the species from the Isle of Pines (BMNH 53.8.16.13). Bauer and Sadlier (1994a) confirmed the presence of this species on the island with a 194 mm female (CAS 182197). Subsequently, the species has been recorded as fairly common on the Isle of Pines and nearby off- shore islands (de Vosjoli 1995; de Vosjoli and Fast 1995). Al- though most known mainland New Caledonian localities are in low- to middle-elevation forests (Bauer 1990), specimens have been recorded from up to 1100 m (Mertens 1964). Sight obser- vations in January 1995 at 540 m on Mt. Mandjélia (20°24’ 15" S, 164°31°18" E) extend the confirmed distribution of the spe- cies to the northwest, almost to the limit of the humid forest on the main island of New Caledonia. The Isle of Pines population has recently been described by Seipp and Obst (1994) as a distinctive subspecies, Rhacodactylus leachianus henkeli. The validity of this form is challenged on the basis of morphological and allozyme charac- ters by Good et al. 1997. They found the henkeli color pattern to occur among geckos in at least two regions of the New Caledonian mainland and regarded behavioral differences as at- tributable to reduced predation pressure on the insular form. Because genetic distance data’ indicated no long separation of Isle of Pines R. /eachianus from mainland populations Good et al. 1997 regarded the split of the insular population to be very recent. Indeed sea level minima of 100 m or more would have connected New Caledonia to the Isle of Pines as recently as 16,000-20,000 years ago (Stevens 1973; Holloway 1979). Although the diet in captivity of Rhacodactylus leachianus has been well documented (Mertens 1964; Bauer and De Vaney 1987; Henkel and Schmidt 1991), and a few stomach contents have been reported (Roux 1913), the natural diet remains poorly documented. At Mt. Aoupinié, in January 1995, we observed individuals feeding on fruit in humid forest trees. Examination of feces of freshly captured individuals revealed only fig seeds and partially digested fig fruit. It appears likely that this, and per- haps other Rhacodactylus species, take advantage of seasonal and local availability of figs and may play a role in seed dis- persal. Roux’s giant gecko (Rhacodactylus sarasinorum) [Plate 5A and SB]. Bauer (1990) figured the type of R. sarasinorum NEW CALEDONIA | New Caledonia is a French overseas territory, consisting of the large island of New Caledonia and the Loyalty Islands. Its location is approximately 1,200 | km east of Australia (Geographic Coordinates 21°30’ S, 165°30’ E) in the South Pacific Ocean. These islands have an extraordinary diversity of fauna and — | flora with an extreme level of endemism in many taxa including birds and reptiles. Naturally occurring plant species number 3,380 (vascular plants), birds 116, mammals 9, and reptiles 87 (71 terrestrial and 16 marine). No naturally occurring amphibian species exist on New Caledonia though a nonnative species has been introduced from Australia (green and golden bell frog Litoria aurea). Total area is 19,060 square (sq) km (land 18,575 sq km and water 485 sq km) comparatively, slightly smaller than New Jersey. The terrain is west coastal plains with interior mountains (highest point Mont Panie 1,628 m) making up two-thirds of the island. The climate is subtropical (warm and humid) modified by southeast trade winds. There is little temperature change throughout the year, averaging between 71°F and 75°F (22°C and 24°C). The natural vegetation comprises tropical evergreen rain forest up to 1,000 m and ' tropical montane rain forest above 1,000 m. Mangroves occur along western coasts. The major vegetation types are dense evergreen forest (22.8% of total land area), Niaouli savanna woodland (13.8%), maquis vegetation in mining areas (25.1%), savanna grassland (21.7%), and scrub (8.3%). New Caledonia’s human population numbers 191,003 (July 1997 estimate) with a 1.68% (1997 estimate) annual growth rate. New Caledonia’s moderately developed economy is based on mining and has more than 20% of the world’s known nickel resources as well as other natural resources as chrome, iron, cobalt, manganese, silver, gold, lead, and copper (thus mining is an important environmental issue). Only a negligible amount of the land is suitable for cultivation and food accounts for about 25% of imports. In addition to nickel, financial support from France and tourism are key to the health of the economy. The principal threats to the natural flora and fauna are mining, logging, and bushfires, reducing the forest cover from an estimated 90% cover to just 20%.' and noted variation in color pattern and body pro- portions in this species but did not elaborate. At least two color morphs have been illustrated and described by Henkel (B6hme and Henkel 1985; Henkel 1987, 1988), but there has been no sug- gestion of subspecific or specific distinction be- tween these forms. Bauer’s (1990) and Bauer and Vindum’s (1990) concept of R. sarasinorum was based in part on typical specimens and in part on an individual from Touaourou that is larger, darker, and differs from other specimens in a number of scale counts. Allozyme analysis (Good et al. 1997) revealed that this specimen differed from a typi-_ cal R. sarasinorum from Riviere Bleue (AMS R 146596) by four fixed differences. This is a greater genetic difference than that between R. ciliatus and R. chahoua. Both allozyme and morphologi- cal data thus suggest significant variation and are Fig. 1. Distribution of Rhacodactylus ciliatus ie being analyzed separately, which may result in (triangles). R. chahoua (squares), and R h “4: f Rh il i trachyrhynchus (circles) in the New Caledonia the recognition of a new Khacodactylus sara- region. Open symbols represent older records i -li i as summarized by Bauer and Henle (1994), new sinorum-like SPStles. records are indicated by solid symbols. Note especially the range extension of R. trachyrhny- chus to the west coast at Pindai and the docu- Rough-snouted giant gecko (Rhacodacty- mentation of the occurence of the other species lus trachyrhynchus) [Plate 6]. Bauer (1990) on the Isle of Pines. recorded five mainland New Caledonian localities for R. trachyrhynchus. Several of these, Coula- Borearé, Ciu, and Mt. Gouemba are in the eastern humid forest scincus festivus), New Caledonian skink (Caledoniscincus region of the island. The other two localities, La Foa and near austrocaledonicus), and a new species of elf skink (Nannoscincus Nouméa are imprecise but probably are also humid forest locali- sp.). All previous records of Rhacodactylus spp. from the New ties. All localities are at middle to low elevation. As briefly Caledonian mainland have originated from the wetter eastern noted by Bauer (1995), recent censuses have expanded the portions of the island, or from rainforest or maquis vegetation‘ known range of the species, both elevationally and geographi- in the south (Bauer 1990; Henkel 1991, 1993; Bauer and Henle cally (Fig. 1). Five specimens (AMS R146417-9, CAS 200266- 1994). The occurrence of R. trachyrhynchus at this site is thus 8) were found during rainstorms in humid forest (Fig. 2A) at intriguing and suggests a much broader habitat tolerance range approximately 520 m on Mt. Aoupinié in Central New Caledonia than previously suspected for this species. Although normally (21°09719" S, 165°19712" E), 27 km north and west of the associated with large, mature rainforest trees (Meier 1979), at previously documented range of the species. A single specimen, Pindai, this gecko was collected less than five meters from the CAS 200269, was obtained in sclerophyll forest’ at Pindai ground in the branches of a small tree. A very low genetic dis- (21°20’02" S, 164°58’21" E) at approximately 20 m elevation tance from this specimen to specimens from Mt. Gouemba and (Fig. 2B). This locality is also somewhat further north than Mt. Aoupinié (Good et al. 1997) suggests no significant differ- earlier records, but it is unique in that it is a west coastal locality entiation in the dry forest population, and the specimen is typi- in an area of low rainfall. The local vegetation is dominated by cal for the species in regard to morphology. Henkel (1991) sug- largely endemic dry forest plants and is regarded as the most gested that there are two morphs in the species, one with a threatened terrestrial habitat in New Caledonia (Jafrré et al. 1993; short, wide, robust snout, the other less so, but because these Bouchet et al. 1995). The only other reptiles collected sympat- features were noted in captive born specimens of uncertain lo- rically with R. trachyrhynchus at Pindai were Vieillard’s prehen- cality he did not imply any subspecific distinction. Our data, sile-tailed gecko (Eurydactylodes vieillardi), Giinther’s New from three widely scattered localities, do not support the recog- Caledonian gecko (Bavayia cyclura), sclerophyll forest gecko nition of any specific or subspecific subdivisions within R. (Bavayia exsuccida), festive New Caledonian skink (Caledoni- trachyrhynchus. 27 AARON M. BAUER AND ROSS A. SADLIER Fig. 2. Habitat of Rhacodactylus trachyrhynchus. Caledonia. The presence of this species on the Isle of Pines remains problematic. Boulenger (1878) recorded the species (as Chamaeleonurus trachycephalus) from this locality. Subsequent collecting activity on this island, however (Bauer and Sadlier 1994a; de Vosjoli 1995) has not verified this occurrence. The unexpected findings of this and other Rhacodactylus after many years of extensive and intensive searching, however, argue against dismissal of this early record. Bauer and Sadlier (1994a) identi- fied appropriate habitat for the species on the island. Conclusions The new dietary observations confirm earlier reports of her- bivory by Rhacodactylus species. It is especially noteworthy that R. auriculatus was found active on flowering heads of plants of the same genus on which Bavay observed them over 130 years ago. The significance of reptiles as seed dispersers and as possible pollinators was recognized by Borzi (1911) but re- mains largely unexplored. Its recent documentation for the closely related carphodactyline geckos of New Zealand (Whitaker 1987) suggests that at least some species of giant geckos are important in this regard. The importance of plant material in the diets of Rhacodactylus sp. and the significance of geckos as dispersers of pollen or seeds, however, can only be adequately addressed by a seasonal dietary study at a single site. Bauer and Sadlier (1993) reviewed the conservation status of all New Caledonian lizards on the basis of data then available. They summarized both the extent of the geographic range of the species and their apparent abundance within the appropriate B. Sclerophyll forest habitat at Pindai on the west central coast A. Overview of humid forest habitat at middle elevation on Mt. Aoupinié. 28 of New habitat types. The data reported on here necessi- tate a re-evaluation of that status report. Rhacodactylus ciliatus, previously considered possibly extinct, is now known to be common on islands of the south coast of New Caledonia and present, if somewhat less common, on the main- land. Using the terminology of Bauer and Sadlier (1993), its distribution is now regarded as re- stricted and its status as locally common. With the extension of its known range to the north and to the west coast sclerophyll forest, the distribu- tion of Rhacodactylus trachyrhynchus can now be upgraded from restricted to moderately wide- spread and its status to locally common. The pres- ence of this species, as well as a regionally en- demic Bavayia (B. exsuccida) and a new, appar- ently endemic, Nannoscincus in the sclerophyll forest in west coastal New Caldeonia (Bauer et al. 1998) adds impetus to efforts to protect the small remaining tracts of this habitat. Rhacodactylus leachianus was previously regarded as widespread and uncommon. The extension of the east coast range to the limit of humid forest underscores the fact that this is the most widely distributed of all Rhacodactylus species and observations of indi- viduals under ideal weather conditions (warm and wet) suggest that the species may best be catego- rized as common in appropriate habitats. The sta- tus of the remaining species is unchanged by the new records. Although at least several species of Rhacodactylus do appear to be locally abundant, even the most widely ranging species is endemic to the New Caledonian mainland and adjacent sat- ellite islands, an area about the size of Connecticut. Further, no species is known to be present in all native habitat types and all are excluded from agricultural or urban environments (although they may be present at the periphery of human-modified areas). Habitat destruction and the impact of introduced predators were cited as the primary threats to the herpetofauna of New Caledonia by Bauer and Sadlier in 1993. These factors remain the most significant conservation concerns, but the illegal pet trade in Rhacodactylus has increased significantly in the 1990’s and has become a potential threat to wild populations. AARON M. BAUER Acknowledgments.—We thank the permit-issuing authori- ties in New Caledonia for their continued support of our re- search. We also thank Jean Chazeau (IRD) and Alain Renevier and his family for their encouragement of our activities. Finan- cial support for this work came from Villanova University, the California Academy of Sciences (AMB), and the Australian Museum (RAS). References Bartmann, W. and Minuth, E. 1979. Ein lebendgebarender Gecko, Rhacodactylus trachyrhynchus Bocage 1873 aus Neukale- donien. Salamandra 15:58-60. Bauer, A. M. 1985. Notes on the taxonomy, morphology and be- havior of Rhacodactylus chahoua (Bavay) (Reptilia: Gekko- nidae). Bonner Zoologische Beitrdge 36:81-94. Bauer, A. M. 1990. Phylogenetic systematics and biogeography of the Carphodactylini (Reptilia: Gekkonidae). Bonner Zoolo- gische Monographien 30:1-219. NEW RHACODACTYLUS DATA Bauer, A. M. 1995. Geckos of the genus Rhacodactylus. Reptiles 3(7):32-34, 36, 38-42, 44, 46-49. Bauer, A. M. and Devaney, K. D. 1987. Comparative aspects of diet and habitat in some New Caledonian lizards. Amphibia-Rep- tilia 8:349-364. Bauer, A. M. and Henle, K. 1994. Familia Gekkonidae (Reptilia, Sauria). Part I Australia and oceania. [Das Tierreich 109 (part)]. Walter de Gruyter, Berlin, Germany. XIII + 306 p. Bauer, A. M. and Russell, A. P. 1990. Dentitional diversity in Rhacodactylus (Reptilia: Gekkonidae). Memoirs of the Queensland Museum 29:311-321. Bauer, A. M. and Russell, A. P. 1994. Is autotomy frequency reduced in geckos with “actively functional” tails? Herpetological Natu- ral History 2:1-15. Bauer, A. M., Russell, A. P., and Dollahon, N. R. 1993. Function of the mite pockets of lizards: a reply to E. N. Arnold. Canadian Journal of Zoology 71:865-868. Bauer, A. M. and Sadlier, R. A. 1993. Systematics, biogeography and conservation of the lizards of New Caledonia. Biodiversity Let- ters 1:107-122. Bauer, A. M. and Sadlier, R. A. 1994a. Terrestrial herpetofauna of the Ile des Pins, New Caledonia. Pacific Science 48:353-366. Bauer, A. M. and Sadlier, R. A. 1994b. Diet of the New Caledonian gecko Rhacodactylus auriculatus (Squamata, Gekkonidae). Russian Journal of Herpetology 1:108-113. Bauer, A. M. and Sadlier, R. A. 2000. The Herpetofauna of New Caledonia. Society for the Study of Amphibians and Reptiles, Ithaca, NY. : Bauer, A. M. and Vindum, J. V. 1990. A checklist and key to the herpetofauna of New Caledonia, with remarks on biogeogra- phy. Proceedings of the California Academy of Sciences 47:17- 45. Bauer, A. M., Whitaker, A. H., and Sadlier, R. A. 1998. Two new species of the genus Bavayia (Reptilia: Squamata: Diplo- dactylidae) from New Caledonia, southwest Pacific. Pacific Science 52:342-355. Bavay, A. 1869. Catalogue des reptiles de la Nouvelle-Calédonie et description d’espéces nouvelles. Mémoires de la Société Linnéenne de Normandie 15:1-37. Bohme, W. and Henkel, F. W. 1985. Zur Kenntnis der Herpetofauna Neukaledoniens, speziell der Gattung Rhacodactylus. Herpe- tofauna 7:23-29. Borzi, A. 1911. Ricerche sulla disseminazione delle piante per mezzo di sauri. Memorie della Societe Italiana delle Scienze, series 3, 17:97-115. . Bouchet, P., Jaffré, T., and Veillon, J.-M. 1995. Plant extinction in New Caledonia: protection of sclerophyll forests urgently needed. Biodiversity and Conservation 4:415-428. Boulenger, G. A. 1878. Description d’un genre nouveau et d’une espece nouvelle de la famille des geckotides. Bulletin de la Société Zoologique de France 3:68. Boulenger, G. A. 1885. Catalogue of the Lizards in the British Mu- seum, Volume I. Taylor and Francis, London, England. Xii + 463 p., 32 plates. de Vosjoli, P. 1995. Notes from a herpetological field trip to New Caledonia, part one: account from the daily journals of Philippe de Vosjoli and Frank Fast. The Vivarium 6(5):4-7, 12, 14-17, 36-38, 40-43. de Vosjoli, P. and Fast, F. 1995. Notes from a herpetological field trip to New Caledonia, part two: notes on three species of New Caledonian geckos of the genus Rhacodactylus. The Vivarium 6(6):26-29, 53-54. Girard, F. and Heuclin, D. 1998. Premiére mention de gecko Rhacodactylus ciliatus sur la Grande Terre (Nouvelle-Calédonie) depuis sa description en 1866. Bulletin de la Societe Herp- etologique de France 85-86:60-61. Good, D. A., Bauer, A. M., and Sadlier, R. A. 1997. Allozyme evi- 29 dence for the phylogeny of giant New Caledonian geckos (Squamata: Diplodactylidae: Rhacodactylus), with the com- ments on the status of R. leachianus henkeli. Australian Jour- nal of Zoology 45:317-330. Guichenot, A. 1866. Notice sur un nouveau genre de sauriens de la famille geckotiens, du Muséum de Paris. Mémoires de la Société Impériale des Sciences Naturelles de Cherbourg 12:248-252. Henkel, F. W. 1987. Haltung und Zucht von Rhacodactylus sarasinorum. Herpetofauna 9(50):25-26. Henkel, F. W. 1988. Rhacodactylus sarasinorum. Sauria 10(4):125- 128 [Amphibien-Reptilien/Kartei, 37-40]. Henkel, F. W. 1991. Zur Kenntnis der diplodactylinen Gecko-Gattung Rhacodactylus Fitzinger, 1843. Aspekte von Freileben, Haltung und Nachzucht. Salamandra 27:58-69. Henkel, F. W. 1993. Notes on the diplodactyline gecko genus Rhacodactylus (Fitzinger, 1843), observations in the wild as well as aspects of captive husbandry and breeding. Dactylus 1(4):22-33. Henkel, F. W. and Schmidt, W. 1991. Geckos: biologie, haltung, zucht. Eugen Ulmer, Stuttgart, Germany. 224 p. Holloway, J. D. 1979. A Survey of the Lepidoptera, Biogeography and Ecology of New Caledonia. Dr. W. Junk, The Hague, The Netherlands. Jaffré, T., Morat, P., and Veillon, J. -M. 1993. Etude floristique et phytogéographique de la forét sclérophylle de Nouvelle- Calédonie. La forét sclérophylle de la Province Sud de la Nouvelle-Calédonie. Adansonia 15:107-146. Kullmann, B. 1995. Uber die Wiederentdeckung des Kronengeckos (Rhacodactylus ciliatus) in Neu Kaledonien. Elaphe 3:68-71. Meier, H. 1979. Herpetologische Beobachtungen auf Neukaledonien. Salamandra 15:113-139. Mertens, R. 1964. Neukaledonische Riesengeckos (Rhacodactylus). Der Zoologischer Garten, N. F. 29:49-47. Mittermeier, R. A., Werner, T. B., and Lees, A. 1996. New Caledonia— a conservation imperative for an ancient land. Oryx 30:104- 112. Myers, N. 1988. Threatened biotas: “Hotspots” in tropical forests. Environmentalist 8:1-20. Roux, J. 1913. Les reptiles de la Nouvelle-Calédonie et des iles Loyalty. Nova Caledonia, Zoologie 1:77-160, plates 4-5. Russell, A. P. and Bauer, A. M. 1986. Le gecko géant Hoplodactylus delcourti et ses relations avec le gigantisme et 1’endémisme insulaire chez le Gekkonidae. Mesogéé 46:25-28. Sameit, J. 1985. Reiseziel Neukaledonien. Die Aquarien und Terrarien Zeitschrift 38:279-281. Seipp, R. and Klemmer, K. 1994. Wiederentdeckung von Rhacodactylus ciliatus Guichenot 1866 im Stiden Neukale- doniens (Reptilia: Sauria: Gekkonidae). Senckenbergiana Biologica 24:199-204. Seipp, R. and Obst, F. J. 1994. Beschreibung einer neuen Unterart des neukaledonischen Rhacodactylus leachianus Cuvier 1829. Senckenbergiana Biologica 74:205-211. Stevens, G. R. 1973. The palaeogeographic history of New Zealand. New Zealand Entomologist 5:230-239. Tytle, T. 1992. The natural history and captive husbandry of the New Caledonian lizard genus Rhacodactylus. The Vivarium 3(6):32-35. Whitaker, A. H. 1987. The roles of lizards in New Zealand plant reproductive strategies. New Zealand Journal of Botany 25:315- 328. Footnote "The research sources used in writing the country sidebar.on page 10 (Madagascar), of this volume, and footnoted on page 14 (foomote *), were also used in writing the New Caledonia summary. Manuscript received: January-1997 Accepted: April-1998 Copyright © 2000 Amphibian and Reptile Conservation. All rights reserved. ARC authorizes photocopying for internal or personal use provided the appropriate fee is paid directly to the Copyright Clearance Center, Inc., 222 Rosewood Dr., Danvers, MA 01923-4599, USA. Tel: (978) 750-8400: fax: (978) 750-4470; email: info@copyright.com; website: www.copyright.com Amphibian and Reptile Conservation 2(1):30-31. Column The United States role in the international live reptile trade CRAIG M. HOOVER" TRAFFIC North America, World Wildlife Fund (WWF), 1250 24th Street, NW, Washington, D.C. 20037-1132, USA Abstract.—In the 1990’s, the trade in live reptiles has grown substantially, and the United States (U.S.) is the world’s most significant player in the international trade in live reptiles, both as an importer of exotic species, and as an exporter of native and exotic species. In 1995, more than 2.5 million reptiles were imported into the U.S., primarily to supply the pet trade. In 1996, over 9.5 million reptiles were exported or reexported from the U.S., primarily to Europe and Asia, to supply the demand for reptiles as pets and food. Despite the large and apparertly growing number of reptiles and amphibians in trade, we have yet to quantify the impacts of this trade on the conservation of these species in the wild. Key words. Herpetofauna, import, export, reexport, live reptile, international trade By way of introduction, I would like to give you some background information about myself, TRAFFIC, and this new column in Amphibian and Reptile Conserva- tion titled, Herpetofauna and Humanity. Presently, ama Senior Program Officer for TRAFFIC North America. As a Senior Program Officer, I am responsible for the devel- opment and oversight of wildlife trade studies and the implementation of their findings and recommendations. TRAFFIC North America is a part of the worldwide TRAF- FIC Network, a program of World Wildlife Fund (WWF) and The World Conservation Union (IUCN). TRAFFIC is the world’s largest wildlife trade monitor- ing program with 21 offices covering most regions of the globe. TRAFFIC produces reports and papers documenting the find- ings of its studies and recommending mea- sures necessary to help ensure that the trade in wildlife and wildlife products are conducted in a sus- tainable and legal manner. This work is done by collecting trade data via govern- ment agencies such as the United States Fish and Wild- life Service (USFWS), customs agencies, and interna- tional bodies, carrying out market surveys, conducting literature reviews and website searches, and other means. In the three years that I have been with TRAFFIC, much of my time has been devoted to investigating various aspects of reptile and amphibian trade. Prior to coming to TRAFFIC North America, I was a wildlife inspector for the USFWS in Los Angeles, where for over four years I was able to see firsthand the scope of the trade in reptiles and amphibians. Equally important, I gained valuable knowledge of the laws that govern the trade, and the means by which these laws are implemented and enforced in the United States (U.S.). Ball python (Python regius). Photo ‘Correspondence. Tel: (202) 822-3452; fax: (202) 775- 8287; email: craig.hoover@wwfus.org . World Wildlife Fund. 30 This background, along with an education in natural resources and law, will influence the areas to be covered in this column. Among the subjects I intend to tackle in this space are reptile and amphibian trade and its implica- tions for conservation; the use of reptiles and amphibians as clothing, food and medicine; the enactment, implemen- tation and effectiveness of wildlife trade laws; illegal trade and the threat posed to reptiles and amphibians in the wild; captive breeding and the private breeder’s role in conservation; and the current events that shape the rela- tionship between herpetofauna and humankind. As an introduction to this new col- umn, Herpetofauna and Hu- manity, it would be useful to provide an overview of the U.S. role in the international live reptile trade. The follow- ing information summarizes a report released by TRAF- FIC in August, 1998, entitled The U.S. Role in the Inter- national Live Reptile Trade: Amazon tree boas to Zulu- land dwarf chameleons (Hoover 1998). The international trade in live reptiles has grown dramatically in the last de- cade. The import, export, and reexport of live reptiles sup- plies a number of markets, including zoos and aquariums, breeding facilities, research centers, private breeders and keepers, and even food markets in some segments of soci- ety. By far the most significant market for the live reptile trade is the pet market (private breeders and keepers of amphibians and reptiles). The causes of the substantial rise in the international trade in live reptiles are difficult to quantify but may include an increase in the availability and variety of species; im- proved reptile husbandry practices due to advances in tech- nology and scientific knowledge; increased restrictions on other wildlife trade; changing lifestyles that make reptiles more suitable pets than other fauna; or simply an increased popularity that has made reptiles today’s fashionable pets. Whatever the reason, and there may be some truth to all of these explanations, there can be no denying that the live reptile industry has expanded dramatically. CRAIG M. HOOVER In 1970, prior to the passage of laws such as the U.S. Endangered Species Act and adoption of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), U.S. imports of live reptiles ap- proached 2 million animals. Nearly 80 percent of this im- port volume consisted of turtles, primarily red-eared slider turtles (Jrachemys scripta elegans), with 12 percent liz- ards, 6.5 percent crocodilians, and less than 2 percent snakes (Busack 1974). Based on analysis of USFWS trade data for a number of reptile species, it is clear that the trade subsequently declined significantly and remained relatively low through- out the 1980s. However, imports increased again in the early 1990s, and in 1995 more than 2.5 million live reptiles were imported to the U.S. Yet the content of this trade differed significantly from the reptiles imported in 1970. By far the most commonly imported species was the green iguana ([guana iguana), which made up more than 45 percent of the total trade in 1995, and only 8 percent of imports in 1970. Snakes and lizards played a far more sig- nificant role in current import levels than they did in 1970, with 1995 turtle volumes greatly reduced and crocodilian imports virtually nonexistent. In fact, the dramatic increase in the total number of live reptiles imported is primarily due to fluctuations in the number of iguanas imported. However, the U.S. is not only a consumer in the international live reptile trade, but also a significant sup- plier. In fact, the U.S. presently exports or reexports more live reptiles than it imports, due largely to the export of farm-raised hatchling red-eared slider turtles, a species even more influential on overall trade volumes than the iguana. For example, in 1996, the U.S. exported or reex- ported over 9.5 million reptiles, primarily to Europe, East and Southeast Asia, yet over 88 percent of this trade consisted of the red-eared slider, at a volume of nearly 8.4 million animals. Setting aside the voluminous trade in red-eared slid- ers, there are more than one million reptiles of other spe- cies that are exported or reexported from the U.S. The North American taxa nearest to the red-eared slider in ex- port volume are the map turtles (Graptemys spp.). Ac- cording to USFWS data, map turtle exports have risen from less than 10,000 in 1990 to over 80,000 in both 1995 and 1996. Based on map turtle export data, just two of the twelve map turtle species make up 90 to 95 percent of the export trade: the common map turtle (G. geographica) and the false map turtle (G. pseudogeographica) [Ventura 1997; data provided by Weissgold 1997]. The U.S. also plays a substantial and apparently ex- panding role as an exporter of previously imported rep- tiles. For example, the U.S. reexported less than 60,000 iguanas in 1993 and more than 270,000 imported iguanas in 1996. This role as a supplier of previously imported reptiles is influenced by a number of factors. Perhaps fore- most is geography; for instance, U.S. dealers are well po- sitioned to supply Asia and Europe with Latin American reptiles. Another significant factor may be that U.S. deal- ers have long-established connections with overseas sup- pliers that may provide a competitive advantage as new markets arise. For example, most of the live reptile trade in Canada, where the trade in reptiles is less established, appears to be supplied by U.S. traders (Chatel 1998). US. trade in live reptiles also appears to make up a substantial portion of the world trade in live reptiles. A comparison of trade data for certain CITES-listed species indicates that U.S. trade constituted approximately 28 per- 31 cent of the total world trade in 1983, but constituted more than 82 percent of the world trade by 1992. However, these numbers may be artificially high given the failure of many countries to accurately report such trade. There appears to have been an increase in illegal as well as legal trade. Based on a review of press releases, wildlife trade journals, and other sources, from 1970 to 1990 there were only 11 reported investigations of inter- national live reptile smuggling, while from 1991 to 1997 there were at least 23 such cases reported. However, there are a number of alternative explanations to these results, including increased enforcement effort and better report- ing of prosecutions. Of course, all of this information on the growth in the reptile trade raises the all-important “so what” question. The overview study that TRAFFIC conducted was not meant to answer that question, but to identify areas that needed further examination so that, in some respects, the “so what” question could be addressed. That, in large part, will be the role of this column as well. In future is- sues, we will look at several “so what” questions, such as: What impact does the pet trade have on wild popula- tions of reptiles and amphibians? What is being done to monitor and protect native species found in trade? What other forms of reptile and amphibian trade may be threatening species around the world, such as the skin, food, and medicine trades? What is the impact of commercial captive breeding, farming, and ranching operations for iguanas, turtles, ball pythons, and other species on wild populations of rep- tiles and amphibians? What is the threat posed by exotic species introduc- tions that can occur with international trade, such as the red-eared slider turtle in Europe? What is being done to more effectively enforce exist- ing laws and regulations to ensure that trade is not detri- mentally affecting wild reptiles and amphibians? The U.S. is clearly the world’s largest consumer of live reptiles for the pet trade, as well as a significant sup- plier and intermediary. Yet, this is only one piece of the puzzle. The growth in popularity of reptiles and amphib- ians as pets, along with the continued enormous demand for these species for skins, meat, and medicine, present us with a broad array of subjects for discussion and debate. In coming issues, we will explore these fascinating topics in an effort to gain a better understanding of the dynamic and complex relationship between herpetofauna and hu- manity. References Chatel, K. W. 1998 (June). Personal communication to Craig Hoover, TRAFFIC North America, Washington, D.C. Busack, S. D. 1974. Amphibians and reptiles imported into the United States. U.S. Department of Interior, Washington, D.C. 36 p. Hoover, C. M. 1998. The U.S. Role in the International Live Reptile Trade: Amazon tree boas to Zululand dwarf chame- leons. TRAFFIC North America, Washington, D.C. 59 p. Ventura, J. 1997 (May). Personal communication to Craig Hoover, TRAFFIC USA, Washington, D.C. Weissgold, B. 1997 (May). Personal communication to Craig Hoover, TRAFFIC USA, Washington, D.C. Copyright © 2000 dmpitibian and Reptile Conservation, All rights reserved. AR photocopying for mntemal or personal use provided the appropriate fee is paic d Copvrizht Clearance Center, Tae., 222 Rosewood Dr. Danvers. MA 01923-4 (978) 750-8400; fax: (978) 7302 4470: ne info ruiteht co; website: wit) BRIAN A. MAU RER* Michigan 48824-1222, US. versity Pres Key words. Lizards, Au islands, history, foraging, coexistence, for ecologists, surmountable. s of study by venus Anolis, th its ability to make ae ree ane its choice of prey items, can maximize its rate of energy intake when constrained by food abundances, oe and other 1m- portant aspects of lizard biology. The abil- ity of lizards to forage profitably is re= lated tortheir fitness, which in tum, can cies of lizards that use similar resources. Using empirically based estimates of for- aging rates Roughgarden shows that it is not unreasonable to assume that much of lizard diversity, especially on small is- lands, is a consequence of constraints on foraging of different-sized species as they mteract together. Roughgarden sketches out a reasonable picture of how these in- teractions vary spatially from one island to the next, and how one might begin to make predictions of which species should be found together on which islands. Coy- ering the literature on pairs of lizard species found together in different combinations on different islands, Roughgarden builds a convincing case that resource-based competition between dif- ferent species is likely to explain many aspects of Anolis ecol- ogy in the Caribbean. But there is much more to the story, as Roughgarden goes on to show in later chapters. Lizards evolve at relatively slow rates, and Roughgarden shows how certain aspects of the his- tory of the Caribbean basin can be traced by examining the relationships among different species of lizards within the Car- ibbean dwelling species. Many aspects of the distribution of Anolis species among islands cannot be explained by rare coloni- zation events, but must bea consequence of the geological his- qeyeee tropidonotus male), Las Cuevas, Cayo istrict, Belize (April 4998). Photo hy Peter Stafford, courtesy of The Natural History Museum, London. ‘Correspondence. Tel: (517) 353- Sethe Jax: (S17) 432- 1699; email: maurerb@msu.edu ~~ _ Depariment of isheries and Wildlife, Michigan State University, East Lansing, Jonathan. 1995. Anolis Lizards of the Carib- evolution, and plate tectonics. Oxford Uni- ew York. 200 p. + xvi. (ISBN: 0-19 be related to the presence of other spe- cable. Melding together the two, how- ever, yields deep insights in to the nature Oxford Series in Ecology and Evolution Anolis Lizards of the Caribbean Ecology, Evolution, and Plate Tectonics Jonathan Roughgarden -506731-2). field experiments that a elucidate cer- tai aspects of Anolis ecology, the real meat of the story lies in weaving together evidence from a wide variety of studies. So ecology without history is incomplete, while history without ecology ts inexpli- of species assemblages that cannot be had from investigations of limited scope. Her- ~ petologists and ecologists alike have much to gain from examining Roughgarden’s broad, sweeping approach. Norops uniformis (male), eux Bani, upper Ras- ulo, Cayo District, lize (A et, 1997). Photo i by Peter Stafford, courtesy of The Natural History Mu- seum, London. Further suggested readings: Crother, Brian I. (editor). 1999. Carib- bean Amphibians and Reptiles. Academic Press, San Diego, California. 495 p. (ISBN: 0-12-197955-5). Powell, R. and Henderson, Robert W. (editors). 1996. Contribu- tions to West Indian Herpetology: a tribute to Albert Schwartz. contributions to herpetology, volume 12. Society for the Study of Amphibians and Reptiles, Ithaca, New York. 457 p. (ISBN: 0-916984-37-0). Rivero, Juan A. 1976. Los Anfibios y Reptiles de Puerto Rico. University of Puetto Rico Press. (ISBN: 0-8477-2317-8). Schettino, Lourdes Rodriguez. (editor). 1999. The Iguanid Lizards of Cuba. University Press of Florida, Gainesville, Florida. 384 p: (ISBN: 0-8130-1647-9). Availability: see inside cover ad. Schwartz, Albert and Henderson, Robert W. 1991. Amphibians and Reptiles of the West Indies: descriptions, distributions, and natu- ral history. University Press of Florida, Gainesville, Florida. 736 p. (ISBN: 0-8130-1049-7), Availability: see inside cover ad. Background screen photograph: Anolis erermanni (male), Ciales, Puerto Rico” (24-August-1999). Photo courtesy of Father Alejandro J. Sanchez Munoz, Pastor of Saint Anne’ Parish, Saint Thomas, U.S, Virgin Islands. SUBMITTING BOOKS FOR REVIEW. Please send one deskcopy to: Book Keres Daphibion and Reptile Conservation, 2525 Iowa Avenue, Modesto, California ee 467, ee Copyright Clearance Center, Inc., 222 Rosewood Dr., Danvers, MA 01923-4599, USA. Tel: (978) 750-8400; fax: (978) 750-4470: email: info@copyright.com;, website: www-.copyright.com : News and Notes WORLD NEWS Amphibian declines: unraveling the mystery. The apparent mysterious declines of amphibian populations in protected or relatively undisturbed areas was discussed recently by university scientists, government biologists, federal administrators and representatives from non-government organizations at a workshop organized by Dr. James Collins, Dr. Elizabeth Davidson, and Dr. Andrew Storfer from Arizona State University and sponsored by the National Science Foundation in Arlington, Virginia on May 28-29, 1998. On Thursday morning, May 28, strong evidence for declines of amphibian populations in different geographic locations around the world was presented. The group consensus was that there is a global amphibian decline problem, but there is no single cause. Rather, multiple factors are implicated, including: habitat loss and alteration, global change, pathogens, parasites, toxic chemicals, ultraviolet radiation, and invasive species. The potential effects of four of these factors; UV, toxic chemicals, pathogens, and global change, were discussed in detail on Thursday afternoon. On Friday, participants outlined a research and management plan and passed a resolution (below) that summarizes the plan. Follow-up workshops on specific topics were outlined, and the National Science Foundation has already funded a workshop on disease in amphibians in late July 1998. Resolution: declining amphibian populations. Whereas, there is compel- ling evidence that, over the last 15 years, there have been unusual and substantial declines in abundance and numbers of populations of various species of amphibians in globally distributed geographic regions, and Whereas, many of the declines are in protected areas or other places not affected by obvious degradation of habitats, and Whereas, these factors are symptomatic of a general decline in environmental quality, and Whereas, even where amphibian populations persist, there are factors that may place them at risk, and Whereas, some patterns of amphibian population decline appear to be linked by causative factors, and Whereas, declines can occur on multiple scales, in different phases of amphibian life cycles, and can impact species with differing ecology and behavior, and Whereas, there 1s no obvious single common cause of these declines, and Whereas, amphibian declines, including species extinctions can be caused by multiple environmen- tal factors, including habitat loss and alteration, global change, pathogens, parasites, various chemicals, ultraviolet radiation, invasive species, and stochastic events, and Whereas, these factors may act alone, sequentially, or synergistically to impact amphibian populations, and Whereas, to under- stand, mitigate and preempt the impacts of these factors, a comprehen- sive, interdisciplinary research program must be undertaken, and Whereas, this research program must be conducted in several regions around the globe, both in areas of known declines, and in areas where declines have not been docu- mented, and Whereas, this research must examine issues ranging from environmental quality of landscapes to the condition of individual animals. Now therefore be it resolved, the signatories hereto call for the establish- ment of an interdisciplinary and collaborative research program, which will specify and quantify the direct and indirect factors affecting amphibian population dynamics, and Be it further resolved, that this program will include basic research and monitoring that will test hypotheses of causative factors and examine patterns of change through historical records, field-based correla- tive data, and controlled, multi-factorial experiments, and Be it further resolved, that interdisciplinary, incident response teams should be assembled in “hot spots” of amphibian decline to identify causative factors to facilitate the mitigation of these sudden declines, and Be it further resolved, that the signatories hereto call upon both public and private agencies and institutions, to promote and support research, policies and conservation measures that will ameliorate losses and declines of amphibian populations, and Be it further resolved, that this broad-based approach to the study of amphibian population dynamics will serve as a model for study of the global biodiversity crisis. Submitted by 33 Amphibian and Reptile Conservation 2(1):33-34. Andrew Storfer, Department of Biology, Arizona State University, Tempe, Arizona 85287-1501, USA. Tel: (602) 965-5857; fax: (602) 965-2519; email: astorfer@asu.edu ANNOUNCEMENTS NARCAM. Since 1995, reports of malformed amphibians have increased in number, and public concern for the health of our environment has grown. The North American Reporting Center for Amphibian Malformations (NARCAM) was established as a central repository for information on this phenomenon. With the help of the public and scientists, NARCAM strives to convey an accurate account of the frequency and distribution of malformed amphibians. NARCAM is maintained by the Northern Prairie Wildlife Research Center in Jamestown, North Dakota, a facility of the United States Geological Survey, Biological Resources Division. The United States Environ- mental Protection Agency provides additional support. NARCAM’s worldwide website contains maps on the geographic distribution of reports in North America, along with a description of the type of malformation present at each site. The site also has photos of the different types of malformations that may be encountered. For help in identification of species, NARCAM has a growing online guide to the amphib- ians of the United States and Canada that most frequently are reported with malformations. The public and researchers can submit information directly through the Web site (http:// Www.npwic.usgs.gov/narcam) by using an online data reporting form. Individu- als who do not have Web access can phone in reports toll-free at 1 (800) 238-9801. The public is urged to report sightings of malformed amphibians. If appropriate, a local biologist will visit the site to confirm the species identity and collect additional information. Submitted by Jeff A. Jundt, Coordinator, North American Reporting Center for Amphibian Malformations, Northern Prairie Wildlife Research Center, United States Geological Service (USGS)/ Biological Resources Division, 8711 37th Street Southeast, Jamestown, North Dakota 58401, USA. Tel: (701) 253- 5580; fax: (701) 253-5553; email: Jeffrey_Jundt @ usgs.gov MEETINGS Society for the Study of Amphibians and Reptiles and the Herpetologists’ League Annual Joint Meeting, 27-31 July 2000, Indiana University Purdue University at Indianapolis, Indianapolis, Indiana. Two symposiums: amphibian popula- tion declines (organized by David M. Green and Karen Lips) and a one-day symposium on herpetological research in zoos: the academic connection. For further information contact Henry R. Mushinsky, Department of Biology, University of South Florida, Tampa, Florida 33620. Tel: (813) 974-5218; email: MUSHINSK@ CHUMAI.CAS. USF EDU 21st Annual Symposium on Sea Turtle Biology and Conservation Philadelphia, USA February 24-28, 2001. For more information reference: http://www. seaturtle.org/symposium/2001 WEBSITES AmphibiaWeb elib.cs.berkeley.edu/aw Center for North American Amphibians and Reptiles Eagle.cc.ukans.edu/~cnaar/ CNAARHomePage.html Conservation Breeding Specialist Group www.cbsg.org International Zoo News www.quantum-conservation.org/IZN/294/ IZN-294.html Kansas Herpetological Society eagle.cc.ukans.edu/~cnaar/khs/khsmain.html Online Herpetologists Directory www.smoky-hills.com/directory/search.asp Species Survival Commission www.iucn.org/themes/SSC World Conservation Monitoring Centre www.wemce.org.uk World Zoo Organization WWW.WZO.Org NEW LITERATURE Journal—A dvances in Amphibian Research in the Former Soviet Union (AARFSU) [ISSN: 1310-8840]. Volume 1, Kuzmin, S. L. and Dodd, Jr., C. K. (editors). [ISBN: 954-642-017-4], vi + 233 p. (9 color photos). Volume 2 (ISBN: 954-642-019-0), vi + 189 p., 11 color photos. Format 165 x 235 mm, paperback, color photos, numerous black and white graphs, photos, drawings, figures, maps, and tables. Text in English. US$34. Kuzmin, S. L. and Wilkinson, J. 1998. Volume 3 (ISBN: 954-642-046-8), 165 x 245 mm, graphs, maps, black and white drawings, and ~ 60 color photographs. Text in English, 245 p. US$34. Tarkhnishvili, D. N. and Gokhelashvili, R. K. 1999. Volume 4, The Amphibians of the Caucasus (ISBN: 954-642-047-6). Format 165 x 235, graphs, color and black and white drawings, photos, tables, and bibliog- raphy. Text in English. 240 p. US$34. Discounts are possible for ordering all three volumes and subscribing to future issues. Inquiries and ordering information can be obtained from PENSOPT Publishers (Sofia and Moscow-based scientific publishers and booksellers), Dr. Lyubomir D. Penev, Akad. G. Bonchev Street, Block 6, 1113 Sofia, Bulgaria. Tel: +359-2- 716451; fax: +359-2-704508; email: pensoft @ mbox.infotel.bg; website: www. pensoft.net Electronic Journal—Contemporary Herpetology (CH). CH 1s a peer- reviewed electronic journal devoted to herpetology on-line at URL: http:// vmsweb.selu.edu/~pcsd4805. CH plans to publish articles covering all aspects of herpetology, including ecology, ethnology, systematics, conservation biology, and physiology. CH also plans to publish monographs, points-of-view, and faunistic surveys of poorly known areas but will not publish herpetocultural or anecdotal papers. For more information contact the editor, Joe Slowinski, at tel:(415) 750-7041 (or) by email: jslowins @ cas. calacademy.org LITERATURE Burbrink, F. T. et al. 1998. A riparian zone in southern I]linois as a poten- tial dispersal corridor for reptiles and amphibians. Biological Conser- vation 86 (2):107-115. 34 News and Notes de Silva, Anslem. 1997. Report on the In- ternational Conference of the Biology and Conservation of the South Asian Amphibians and Reptiles. Lyrioce- phalus. 3(1):40-43. [Faculty of Medi- cine, University of Peradeniya, Sri Lanka] Gibbons, J. W. 1997. Measuring declines and natural variation in turtle popu- lations: spatial lessons from long- term studies, p. 243-246 in Abbema, J. V. (editor). Interna- tional Conference on Conservation, Restoration, and Management of Tortoises and Turtles. New York Turtle and Tortoise Society, Pur- chase, New York. Hager, H. A. 1998. Area-sensitivity of reptiles and amphibians: are there in- dicator species for habitat fragmen- tation? Ecoscience 5:139-147. Halliday, T. 1998. Ecology: a declining am- phibian conundrum. Nature 394:418- 419. Lips, K. R. 1998. Decline of a tropical montane amphibian fauna. Conser- vation Biology 12 (1):106-117. Lovich, J. E. and Gibbons, J. W. 1997. Conservation of covert species: pro- tecting species we don’t even know, p. 426-429 in Abbema, J. V. (editor). International Conference on Conser- vation, Restoration, and Management of Tortoises and Turtles. New York Turtle and Tortoise Society, Pur- chase, New York. Seigel, R. A., Sheil, C. A., and Doody, J. S. 1998. Changes in a population of an endangered rattlesnake Sistrurus catenatus following a severe flood. Biological Conservation 83:127-131. Webb J. K., and Shine R.1998. Using ther- mal ecology to predict retreat-site se- lection by an endangered snake spe- cies. Biological Conservation 86(2):233-242. BOOKS AND LITERATURE RECEIVED Points of view on contemporary education in herpetology. Herpetologica 54 (2) {Supplement]. S82 p. Laurance, W. F. and Bierregaard, Jr., R. O. (editors). 1997. Tropical Forest Rem- nants: ecology, management, and con- servation of fragmented communities. The University of Chicago Press, Chi- cago, Illinois. 616 p. (ISBN: 0-226- 46899-2). wi P. 6-14 P. 15-23 P. 24-29 Amphibian and Reptile Conservation 2(1):35. | : The Last Page GLOSSARY Definitions of words footnoted in journal articles. “qutoecological.—ts the study of the relationship of individual organisms to their environment. ’xeric.—“Dry,” of or pertaining to a habitat having a low or inadequate supply of moisture. ‘sisal.—A species of agave grown for commercial production of fiber. 4qestivation.—A state of inactivity occurring in some animals during prolonged periods of drought or heat. Diplodactylidae (In title). —Family of southwest Pacific geckos and pygopods, distributed in Australia, New Zealand, New Caledonia, and in the case of pygopods, New Guinea. ‘carphodactyline—Adjectival form of Carphodactylini, a tribe of the Diplodactylidae including Rhacodactylus. /viviparity.—Reproductive mode in which development takes place within the mother and young are born live, as opposed to oviparity, in which eggs are laid. £allozymic.—In reference to allozymes, the different forms of proteins that can be separated by electrophoresis to yield evi- dence about relationships or similarity of individuals within or between populations and species. "polymorphic.—Having many forms, usually refers to the pres- ence of more than one manifestation of particular characters within a single taxon. ‘genetic distance data.—Measurements of the relative similarity between taxa or individuals being compared, based on direct or indirect genetic information. In this instance, genetic distance data were derived from the allozyme data obtained by electro- phoresis. Jsclerophyll forest —Critically endangered dry forest found only in isolated pockets on the west coast of New Caledonia. ‘maquis vegetation.—The characteristic and highly endemic scrub vegetation of the mineral-rich soils of parts of New Caledonia, especially the south. New abbreviations appear herein. Past issues should be consulted for previous abbreviation usage. W = West N = North E = East S = South et al. = and others F = Fahrenheit e.g.= forexample C = Celsius cm =centimeter tel = telephone mm = millimeter fax = fascimile km = kilometer kg = kilogram email = electronic mail ha = hectares U.S. and US = United States USA = United States of America pers. = personal m=meter UK = United Kingdom Pp. = pages, page ~ = approximately obs. = observation(s) fig. = figure ERRATA comm. = communication(s) The following journal corrections should be noted in Volume 1, Number | (Premiere issue): Page 3. CONTENTS PAGE. The article under Next Issue: Conservation of South African’s En- demic Dwarf Chameleons has been postponed indefinitely. MASTHEAD. The premiere issue (Volume 1, Number 1) is out- of-print and no longer available, although nicely reproduced pho- tocopies are available for $10 plus shipping and handling. All back issues are available for $12 each while regular supplies last. PAGE 10. In the “Key words” section the word Viridi_ should 35 have been spelled Viridis. PAGE 16. Under the photo caption the word Plethedon should have been spelled Plethodon. PAGE 20. The third sentence of the body of the article should read “Inbreeding depression may adversely affect small populations by unmasking recessive deleterious alleles and reducing heterozy- gosity.” Pages 24-26. COLUMN. 1). The personal communica- tion on page 25 (Groombridge 1994) was not attributed to Dr. Brian Groombridge of the World Conservation Monitoring Cen- tre (WCMC), Cambridge, England. 2). The WCMC database, referred to in the column, is in fact the CITES database of trade records. 3). By comparing export records and import records, and checking against known countries of origin, it is in fact possible to get some idea of the extent of misreporting (The updated commentary for this particular article was provided by Brian Groombridge on November 8, 1996). PAGE 27. Meet- ings. The meeting date for the conference Conservation and Biodiversity of Amphibians and Reptiles of Tropical Rain For- ests was re-scheduled for June 1999 and was then subsequently cancelled. NOTE.—The most up-to-date Writer’s Guidelines and Manuscript Preparation Instructions can be located at the Am- phibian and Reptile Conservation website at: www. herpetofauna.com ACKNOWLEDGEMENTS As the journal grows and climbs towards international as well as domestic prominence, a growing list of individuals and organiza- tions must be credited with their help, assistance, and support. Individuals who have kept their enthusiasm and support for the project, as well as me individually, are Jack Sites, Harvey Lillywhite, Mark Goodwin, Jamie Reaser, Craig Hoover, Carl Gans, Peter Lindeman, Maria Bridarolli, John Megahan, Dave Stone, Chris Raxworthy, Ron Nussbaum, Aaron Bauer, Kraig Adler, Allison Alberts, Michael Soulé, George Rabb, Tony Krzysik, Jude McNally, Trevor Hare, Dave Hardy, and Dennis Caldwell. A special thanks goes to Dave Piper, Jeff Middleton, and Jose Solorzano and to all the authors, contributors, and asso- ciates of the journal. Without your generous support, help, and patience this journal would not exist. Organizations which should be acknowledged for their support and use of facilities are the Information Technology Center staff and facility (especially, Dave Piper, Jeff Middleton, and Jose Solorzano), Arizona Health Sciences Library; Science Reference, Science-Engi- neering Library; and Multimedia & Visualization Lab, Center for Computing & Information Technology; all at the University of An- zona, Tucson, Arizona; University Computing Services, Brigham Young University, Provo, Utah; F. Wayne King and the Museum of Natural History, University of Florida, Gainesville, Florida. A special tribute is extended to GRM of Tucson, Arizona, and specifically Arnell McSwain, Dennis Barreras, Albert D. Jones, Darryl A- Lacey, John Roden, Everett E. Sellers, Donald Batson, Chipper Ward, David Morrow, Tim Church, and Leland McKirryher for support, help, and friendship. A very special thanks goes to Kenneth Sholar, our graphic designer, who when I had all but given up on finding a talented graphics person to work with on the journal, took on the task regard- less of the fact he was working three jobs and attending school full- time. ARC and I extend a heart felt thanks for his commitment and fine work in helping get the journal “just nght’ in every detail. DEDICATION This issue is kindly dedicated to my family: my mother Frances, my dad Anthony, and both my brothers Steve, and Greg Hasspakis. The Best of Its Kind 3 Handbook of Reptiles and Amphibians of Florida sve. THE SNAKES This widely — acclaimed series has been mele praised © | & CROCODILIANS throughout the world. Handbooks of Reptiles and Amphibians of Florida i by Ray E. 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