Conservation Biology of Freshwater Turtles and Tortoises: ExtinctA Compilation Pleistocene Project of and the IUCN/SSCHolocene Tortoise Turtles and of Freshwater the World Turtle Checklist Specialist – Group2015 000e.1 A.G.J. Rhodin, P.C.H. Pritchard, P.P. van Dijk, R.A. Saumure, K.A. Buhlmann, J.B. Iverson, and R.A. Mittermeier, Eds. Chelonian Research Monographs (ISSN 1088-7105) No. 5, doi:10.3854/crm.5.000e.fossil.checklist.v1.2015 © 2015 by Chelonian Research Foundation • Published 16 April 2015 Turtles and Tortoises of the World During the Rise and Global Spread of Humanity: First Checklist and Review of Extinct Pleistocene and Holocene Chelonians TURTLE EXTINCTIONS WORKING GROUP* *Authorship of this article is by this joint Working Group of the IUCN SSC Tortoise and Freshwater Turtle Specialist Group and specialists in Chelonian Paleontology, which for the purposes of this document consisted of the following contributors: ANDERS G.J. RHODIN1, SCOTT THOMSON2, GEORGIOS L. GEORGALIS3,4, HANS-VOLKER KARL5, IGOR G. DANILOV6, AKIO TAKAHASHI7, MARCELO S. DE LA FUENTE8, JASON R. BOURQUE9, MASSIMO DELFINO4,10, ROGER BOUR11, JOHN B. IVERSON12, H. BRADLEY SHAFFER13, AND PETER PAUL vaN DIJK14 1Chairman Emeritus, IUCN SSC Tortoise and Freshwater Turtle Specialist Group, Chelonian Research Foundation, 168 Goodrich St., Lunenburg, Massachusetts 01462 USA [[email protected]]; 2Museu de Zoologia da Universidade de São Paulo, Divisão de Vertebrados (Herpetologia), Ipiranga, 04263-000 São Paulo, Brazil [[email protected]]; 3Department of Geosciences, University of Fribourg/Freiburg, Chemin du Musée 6, 1700 Fribourg, Switzerland [[email protected]]; 4Dipartimento di Scienze della Terra, Università di Torino, Via Valperga Caluso 35, 10125 Torino, Italy [[email protected]]; 5Department of Prehistoric and Early Historic Archaeology, Friedrich-Schiller-University Jena, Löbdergraben 24a, 07743 Jena, Germany [[email protected]]; 6Department of Herpetology, Zoological Institute of the Russian Academy of Sciences, Universitetskaya Emb. 1, St. Petersburg, 199034, Russia [[email protected]]; 7Department of Zoology, Faculty of Science, Okayama University of Science, Ridaicho 1-1, Kitaku, Okayama 700-0005, Japan [[email protected]]; 8Departamento de Paleontología, Museo de Historia Natural de San Rafael, Parque Mariano Moreno s/n, (5600) San Rafael, Provincia de Mendoza, Argentina [[email protected]]; 9Division of Vertebrate Paleontology, Florida Museum of Natural History, Dickinson Hall, University of Florida, 1659 Museum Rd., Gainesville, Florida 32611 USA [[email protected]]; 10Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICP, Campus de la UAB s/n, 08193 Cerdanyola del Vallès, Barcelona, Spain; 11Laboratoire des Reptiles et Amphibiens, Muséum National d’Histoire Naturelle, Paris, France [[email protected]]; 12Department of Biology, Earlham College, Richmond, Indiana 47374 USA [[email protected]]; 13Department of Ecology and Evolutionary Biology and Institute of the Environment and Sustainability, University of California, Los Angeles, California 90095 USA [[email protected]]; 14Co-Chair, IUCN SSC Tortoise and Freshwater Turtle Specialist Group, Conservation International, 2011 Crystal Drive, Suite 500, Arlington, Virginia 22202 USA [[email protected]] ABSTRACT. – We provide a first checklist and review of all recognized taxa of the world’s extinct Pleistocene and Holocene (Quaternary) turtles and tortoises that existed during the early rise and global expansion of humanity, and most likely went extinct through a combination of earlier hominin (e.g., Homo erectus, H. neanderthalensis) and later human (H. sapiens) exploitation, as well as being affected by concurrent global or regional climatic and habitat changes. This check- list complements the broader listing of all modern and extant turtles and tortoises by the Turtle Taxonomy Working Group (2014). We provide a comprehensive listing of taxonomy, names, synonymies, and stratigraphic distribution of all chelonian taxa that have gone extinct from ap- proximately the boundary between the Late Pliocene and Early Pleistocene, ca. 2.6 million years ago, up through 1500 AD, at the beginning of modern times. We also provide details on modern turtle and tortoise taxa that have gone extinct since 1500 AD. This checklist currently includes 100 fossil turtle and tortoise taxa, including 84 named and apparently distinct species, and 16 ad- ditional taxa that appear to represent additional valid species, but are only identified to genus or family. Modern extinct turtles and tortoises include 8 species, 3 subspecies, and 1 unnamed taxon, for 12 taxa. Of the extinct fossil taxa, terrestrial tortoises of the family Testudinidae (including many large-bodied island forms) are the most numerous, with 60 taxa. When the numbers for fossil tortoises are combined with the 61 modern (living and extinct) species of tortoises, of the 121 tortoise species that have existed at some point since the beginning of the Pleistocene, 69 (57.0%) have gone extinct. This likely reflects the high vulnerability of these large and slow terrestrial (often insular) species primarily to human exploitation. The other large-bodied terrestrial turtles, the 000e.2 Conservation Biology of Freshwater Turtles and Tortoises • Chelonian Research Monographs, No. 5 giant horned turtles of the family Meiolaniidae, with 7 taxa (also often insular), all went extinct by the Late Holocene while also exploited by humans. The total global diversity of turtles and tortoises that has existed during the history of hominin utilization of chelonians, and that are currently recognized as distinct and included on our two checklists, consists of 336 modern spe- cies and 100 extinct Pleistocene and Holocene taxa, for a total of 436 chelonian species. Of these, 109 species (25.0%) and 112 total taxa are estimated to have gone extinct since the beginning of the Pleistocene. The chelonian diversity and its patterns of extinctions during the Quaternary inform our understanding of the impacts of the history of human exploitation of turtles and the effects of climate change, and their relevance to current and future patterns. KEY WORDS. – Reptilia, Testudines, turtle, tortoise, chelonian, taxonomy, distribution, extinction, fossils, paleontology, archaeology, humanity, hominin, exploitation, chelonophagy, megafauna, island refugia, climate change, Pliocene, Pleistocene, Holocene, Anthropocene, Quaternary As an addition to the annual checklist of extant and A. grandidieri, that went extinct in about 1200 modern turtle taxa (Turtle Taxonomy Working Group AD and 884 AD, respectively, not long after humans [TTWG] 2014), we here present an annotated checklist reached Madagascar ca. 2000 years ago (Pedrono 2008). of extinct Pleistocene and Holocene turtle and tortoise Additionally, some large insular species of Chelonoidis species that existed in relatively recent times, prior to from the Bahamas region of the Caribbean West Indies 1500 AD, during the history of the rise and global spread were eaten into extinction by pre-Columbian natives as of humanity and concurrent global climatic and habitat late as ca. 1170–1400 AD (Carlson 1999; Franz et al. changes. These species, recorded from archaeological 2001; Hastings et al. 2014). and paleontological sites from the Pleistocene and Holo- Among the Meiolaniidae, we have the remarkable cene epochs (Quaternary period), approximately the last giant terrestrial horned turtle, Meiolania damelipi from 2.6 million years, are currently considered to be valid, Vanuatu in the southern Pacific Ocean, also eaten into and not synonymous with modern (post-1500 AD) taxa. extinction by humans by about 810 BC (White et al. These fossil species, including some unnamed taxa 2010), as well as an unnamed giant horned turtle from of indeterminate or undescribed generic or specific al- nearby New Caledonia, that went extinct as recently location, represent the majority of the chelonian diver- as about 531 AD (Gaffney et al. 1984). This unnamed sity that has gone extinct relatively recently. Many of and vanished species was apparently the last surviving these taxa were likely extirpated by anthropogenic ex- member of this most impressively distinct and ancient ploitation over the relatively long prehistory of earlier family of giant horned terrestrial turtles. Several recent hominin (e.g., Homo erectus, H. neanderthalensis, and phylogenies suggest that the Meiolaniidae branched others) and later human (H. sapiens) exploitation of tur- off as a separate clade of turtles before the Cryptodira– tles and tortoises. In addition, many were also likely af- Pleurodira split (e.g., Joyce 2007; Sterli and de la Fuente fected by global and regional climate change and cycles 2013), but others (e.g., Gaffney 1996; Gaffney et al. of warming and cooling and habitat alterations, such as 2007; Gaffney and Jenkins 2010) place them among the those associated with glacial and interglacial periods Cryptodira. In either case, their recent extinction was and sea level changes and aridification, or stochastic indeed major, not just for their disparate and bizarre events such as volcanism. As such, these recently ex- morphology, but also because had they persisted, tinct fossil species and taxa are eminently relevant to they would have been one of the most evolutionarily our understanding of distribution and extinction patterns and phylogenetically distinct lineages