Reptiles of Venezuela: an updated and commented checklist by Luis Alejandro Rodriguez J.

ISSN 1175-5326 (print edition)

Monograph

ZOOTAXA ISSN 1175-5334 (online edition)

ZOOTAXA 3211

Reptiles of Venezuela: an updated and commented checklist GILSON A. RIVAS1, CÉSAR R. MOLINA2, GABRIEL N. UGUETO3, TITO R. BARROS1, CÉSAR L. BARRIO-AMORÓS4 & PHILIPPE J. R. KOK5,6,7 1

Museo de Biología, Facultad Experimental de Ciencias, La Universidad del Zulia, apartado postal 526, Maracaibo 4011, Maracaibo, Venezuela 2 Instituto de Zoología y Ecología Tropical, Facultad de Ciencias, Universidad Central de Venezuela, Caracas, Venezuela 3 11111 Biscayne Boulevard, Jockey Club, Phase III, #556, Miami, Florida 33181, USA 4 Instituto de Biodiversidad Tropical. Apartado Postal 328-1260, Escaz, Costa Rica 5 Department of Vertebrates, Royal Belgian Institute of Natural Sciences, 29 rue Vautier, B-1000 Brussels, Belgium. 6 Department of Biology, Unit of Ecology and Systematics, Vrije Universiteit Brussel, 2 Pleinlaan, B-1050 Brussels, Belgium 7 Corresponding author. E-mail: philippe.kok@naturalsciences.be

Magnolia Press Auckland, New Zealand

Accepted by S. Carranza: 7 Oct. 2011; published: 29 Feb. 2012

GILSON A. RIVAS, CÉSAR R. MOLINA, GABRIEL N. UGUETO, TITO R. BARROS, CÉSAR L. BARRIO-AMORÓS & PHILIPPE J. R. KOK Reptiles of Venezuela: an updated and commented checklist (Zootaxa 3211) 64 pp.; 30 cm. 29 Feb. 2012 ISBN 978-1-86977-867-5 (paperback) ISBN 978-1-86977-868-2 (Online edition)

FIRST PUBLISHED IN 2012 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: zootaxa@mapress.com http://www.mapress.com/zootaxa/

© 2012 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326

(Print edition)

ISSN 1175-5334

(Online edition)

RIVAS ET AL.

Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Resumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Taxonomic richness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Species endemism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Endangered species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Introduced (non-native) and displaced (native) species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Undescribed species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Species erroneously reported from Venezuela . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Order TESTUDINES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Order SQUAMATA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Taxonomic comments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Order TESTUDINES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Order SQUAMATA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Species of possible occurrence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Literature cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

Abstract We update the list of reptiles of Venezuela, reporting a total of 370 species from the country (four of these exotic), arranged in 122 genera (one exotic), 30 families and three orders. Introduced species and dubious or erroneous records are discussed. Taxonomic, nomenclatural and distributional comments are provided when required. Considering species of probable occurrence in the country (known to occur in Colombia, Brazil and Guyana at localities very close to the Venezuelan border) and still undescribed taxa, we estimate that the total number of species in Venezuela could exceed 400.

Resumen Se actualiza la lista de reptiles de Venezuela a un total de 370 especies (cuatro de ellas exóticas), pertenecientes a 122 géneros (un exótico), 30 familias y tres órdenes. Se discuten las especies introducidas, los registros dudosos o erróneos y se proporcionan comentarios taxonómicos, nomenclaturales o de distribución en casos necesarios. Después de considerar las especies de probable presencia en el país (registros conocidos cercanos a la frontera con Colombia, Brasil y Guyana) y los taxones aun no descritos, se estima que el número total de especies en Venezuela podría superar las 400 especies. Key words: Diversity, Reptiles, South America, Venezuela.

Introduction South America harbours some of the most species-rich regions of the world (e.g. the Amazon Basin and the Andes). Venezuela is located at the northernmost portion of the continent and is considered the ninth most reptile diverse country in the world (Aguilera et al. 2003). Partial lists of Venezuelan reptiles have been published using taxonomy-based (Roze 1966; Donoso-Barros 1968; Lancini 1979; Medem 1983; Pritchard & Trebbau 1984) or distribution-based approaches (Roze 1964b; Staton & Dixon 1977; Rivas & Oliveros 1997; Gorzula & Señaris 1999; Markezich 2002; Mijares-Urrutia & Arends 2000; Molina et al. 2004; Rivas Fuenmayor & Barrio-Amorós 2005), but few studies have focused on complete checklists of the Venezuelan herpetofauna (Péfaur 1992; La Marca 1997; Péfaur & Rivero 2000). Among these, only the publication by Péfaur and Rivero (2000) is considered a comprehensive analysis of all Venezuelan bioregions, but their contribution is currently outdated, especially considering all the new species that have been described over the last 10+ years. Venezuela has a considerable diverse reptile fauna. Knowledge of the country’s reptiles has improved considerably in the last two decades, thanks to research conducted by both Venezuelan and foreign herpetologists. Most recent research has been carried out in the Venezuelan Guayana, the Andean region, and the Coastal Ranges. The number of reptile species currently known from Venezuela increased to 370, which is fewer than in Ecuador (414 REPTILES OF VENEZUELA

species), Colombia (578 species), and Brazil (721 species), but more than in other neighbouring countries like Guyana (170 species), Suriname (143 species) and French Guiana (162 species) (Hoogmoed 1973; Pritchard & Trebbau 1984; Avila-Pires 1995; Métrailler & Le Gratiet 1996; Starace 1998; Abuys 2003; de Massary 2004; Bérnils 2010; Torres-Carvajal & Salazar-Valenzuela 2011; Uetz 2011). Fair comparison between these countries is difficult due to tremendous differences in countries size and varying intensity of collecting efforts. Checklists are dynamic and should be considered as a still frame in time that has no lasting value, only showing the state of knowledge at a peculiar moment. Reports of new species, synonymisations and elevation of old synonyms to specific status, clarification of prior mistakes and new data about species distributions rapidly change our knowledge of biological diversity in tropical countries. With this paper we present the current state of knowledge of the reptile diversity in Venezuela.

Materials and methods All data on species geographic distributions are based on published records and voucher specimens housed in museum collections (except in very rare occasions, see specific comments). With few exceptions we examined specimens from all species recorded in this work. No formal record based only on field observations is provided, except for Hydrops martii because a catalogued photographic voucher exists. In order to provide the most accurate species list possible we consulted the pertinent literature published, mostly until march 2011, including: Albuquerque and McDiarmid 2010; Avila-Pires 2005; Avila-Pires and Hoogmoed 2000; Bailey and Thomas 2006; Barrio 2001; Barrio and Calcaño 2001; Barrio and Navarrete 1999; Barrio et al. 1999, 2003; Barrio-Amorós 2007; BarrioAmorós and Díaz de Pascual 2008; Barrio-Amorós and Molina 2010; Barrio-Amorós and Rivas Fuenmayor 2003; Barrio-Amorós et al. 2010; Barros 2000; Barros and Pirela 2000; Barros and Rincón 1999; Barros et al. 1996, 2007; Bisbal et al. 2008; Calcaño and Barrio-Amorós 2002, 2003a, 2003b, 2010; Carvajal-Cogollo and BernalGonzález 2011; Carvalho 2002; Casado and La Marca 1996; Casado and Zoltan 1996; Castro 1994; Cole and Kok 2006; Cole et al. 1990, 1993, 1995; Cornejo and Prieto 2001; Dixon 1973, 1983a, 1983b, 1987, 1989; Dixon and Hendricks 1979; Dixon and Huey 1970; Dixon and Kofron 1983; Dixon and Lamar 1981; Dixon and Markezich 1992; Dixon et al. 1993; Doan and Castoe 2005; Donnelly and Myers 1991; Donnelly et al. 1992; Duellman 1958; Esqueda 2005; Esqueda and La Marca 1999, 2004, 2005; Esqueda et al. 2004, 2005a, 2005b; Etheridge 1970; Fernández et al. 2002; Franz et al. 1985; Fuentes and Barrio 1999a, 1999b; Fuentes and Rivas Fuenmayor 2000; Fuentes and Rodriguez-Acosta 1997; Gallagher and Dixon 1992; Galvis Peñuela et al. 2011; Gans 1962; García and La Marca 1989; González et al. 2004; González Sponga 1971; González Sponga and Gans 1971; Gorzula 1985, 1989, 1992; Gorzula and Arocha-Piñango 1975; Gutberlet and Campbell 2001; Harris 1982, 1994; Harvey et al. 2003; Harvey et al. 2004; Hellmich 1953a, 1953b; Hernández and Rada 1992; Hernández Ruz 2005; Hollis 2006; Hoogmoed and Dixon 1977; Hoogmoed et al. 1992; Horton 1973; Infante-Rivero 2009; Infante-Rivero et al. 2005, 2009; Johnson and Dixon 1984; Kaiser et al. 2003; Klaczko et al. 2010; Kok 2006; Kok and Rivas 2008, 2011; Kok et al. 2007; Kornacker 1998, 2001a, 2001b; Lamar and Medem 1982; La Marca 1995; La Marca and García 1986, 1987; La Marca and García-Pérez 1990a, 1990b; La Marca and Milano 1995; La Marca and Soriano 1995, 2000, 2004; La Marca et al. 1995, 2004; Lancini 1962a, 1962b, 1963a, 1963b, 1963c, 1964, 1965, 1967, 1968, 1969; Lemoine et al. 1998; Lotzkat et al. 2008, Mägdefrau 1991; Mägdefrau et al. 1991; Manzanilla and Natera 2003; Manzanilla and Sánchez 2004; Manzanilla et al. 1996, 1997, 1998a, 1998b, 1999, 2001; Marcuzzi 1950a, 1950b; Markezich 2001a, 2001b; Markezich and Dixon 1979; Markezich and Rossman 1992; Markezich and Taphorn 1993, 1994; Massary and Hoogmoed 2001; Mattei and Barrio 1999; Maturana 1962; Mausfeld et al. 2002; McCrystal and Dixon 1987; McDiarmid and Donnelly 2005; Mendoz and Fernández Lucero 2009; Mertens 1925, 1954; Michaud and Dixon 1987; Mijares-Urrutia and Arends 1992, 1995, 1997, 1999a, 1999b; Mijares-Urrutia et al. 1992, 2000; Miralles and Carranza 2010; Miralles et al. 2005a, 2005b, 2009; Molina 1998, 2001a, 2001b, 2001c, 2001d, 2004; Molina and Rivas 1996; Molina and Rivero 2003; Molina and Señaris 2001; Molina et al. 2002a, 2002b, 2003a, 2003b; Myers 1997; Myers et al. 1993, 2009; Natera-Mumaw 2008; Natera-Mumaw and Manzanilla 2000; Natera-Mumaw et al. 2006a, 2006b; Navarrete et al. 2005; Oliveros et al. 2000; Paolillo 1984, 1985, 1986; Passos and Fernandes 2005; Passos et al. 2009; Peloso and Avila-Pires 2010; Pérez-Bravo 1978; Rada and Hernández 1999; Rivas 2001; Rivas and Oliveros 1997; Rivas and Rojas-Runjaic 2006; Rivas and Schargel 2008; Rivas et al. 2001, 2005, 2006; Rivas Fuenmayor 2001a, 2001b, 2002a, 2002b, 2002c; Rivas and BarrioAmorós 2003, 2005; Rivas Fuenmayor and Fuentes 2000, 2002; Rivas Fuenmayor and La Marca 2001a, 2001b,

RIVAS ET AL.

2001c; Rivas and Manzanilla Puppo 1999; Rivas Fuenmayor and Molina 1998, 2001, 2003, 2004; Rivas Fuenmayor et al. 1999, 2000, 2001, 2002a, 2002b, 2005a, 2005b, 2006; Rivero-Blanco 1967; Rivero-Blanco and Barrio-Amorós 2002; Rivero-Blanco and Lancini 1967; Robinson 1989; Rodríguez and Rojas-Suárez 2008; Rodríguez-Acosta and Fuentes 1996; Rojas-Runjaic and Infante Rivero 2004a, 2004b, 2006a, 2006b, Rojas-Runjaic and Rivas Fuenmayor 2006; Rojas-Runjaic et al. 2008, 2010; Rossman 1976; Roux 1927, 1929; Roze 1952a, 1952b, 1953a, 1953b, 1954, 1955a, 1955b, 1957a, 1957b, 1958a, 1958b, 1958c, 1958d, 1959a, 1959b, 1959c, 1961, 1963, 1964a, 1964b, 1967, 1983, 1989, 1994, 1996; Roze and Bernal-Carlo 1987; Roze and da Silva 1990; Roze and Trebbau 1958; Sandner-Montilla 1985; Schargel et al. 2005, 2007, 2010; Seijas 2006; Señaris 1999; Señaris and Rivas 2006; Shreve 1947; Sturaro and Avila-Pires 2011; Suárez et al. 2000; Test et al. 1966; Ugueto and Harvey 2010; Ugueto et al. 2007, 2009a, 2009b; Uzzell 1958, 1966; Van Devender 1969; Vanzolini and Calleffo 2002; Walker and Rhoads 2003; Williams 1974, Williams et al. 1996; Wilson 1980; Wüster et al. 2001; Zaher and Caramaschi 2000; Zaher and Prudente 1999.

FIGURE 1. Map of Venezuela showing the delimitation of the bioregions considered in this study, which are: 1. Marine (Marina); 2. Insular; 3. Maracaibo Lake Basin (Depresión del Lago de Maracaibo); 4. Mountain Range of Mérida (Cordillera de Mérida); 5. Mountain Range of Perijá (Sierra de Perijá); 6. Lara-Falcón Hill System (Sistema de Colinas Lara-Falcón); 7. Coastal Area (línea costera; highlighted by a grey line); 8. Central Coastal Range (Cordillera de la costa central); 9. Eastern Coastal Range (Cordillera de la costa oriental); 10. Orinoco Delta (Delta del Orinoco); 11. Llanos; 12. Guayana; 13. Amazonia. Note: the Pantepui (fourteenth bioregion considered in this paper) corresponds to the highlands (>1500 m elevation) of the Guiana Shield in the Amazonia and Guayana bioregions.

MARN (2001) divided Venezuela into ten main bioregions based on vegetational and climatic characteristics. We consider the Mountain Range of Perijá (Sierra de Perijá) and the Mountain Range of Mérida (Cordillera de Mérida) as distinct bioregions instead of subdivisions of the Andes as proposed by MARN (2001). We also consider the Central and Eastern Coastal Ranges as separate bioregions instead of a single area named “Coastal range” by MARN (2001), and the Amazonia, Guayana and Pantepui as distinct bioregions instead of grouping them under the single term “Guayana” as proposed by MARN (2001), the latter following Hoogmoed (1979), McDiarmid and Donnelly (2005) and Rodríguez et al. (2010). Significant endemism (in flora and fauna) is in favour of considering REPTILES OF VENEZUELA

these bioregions as separate entities. We thus recognise the following 14 Venezuelan bioregions: 1) Marine; 2) Insular; 3) Maracaibo Lake Basin; 4) Mountain Range of Perijá; 5) Mountain Range of Mérida; 6) Lara-Falcón Hill System; 7) Coastal Area; 8) Central Coastal Range; 9) Eastern Coastal Range; 10) Orinoco Delta; 11) Llanos; 12) Guayana (< 1500 m); 13) Amazonia; and 14) Pantepui [≥ 1500 m above sea level following McDiarmid and Donnelly (2005), even if we agree that this implies some arbitrariness] (Fig. 1). These bioregions also differ from those proposed by Barrio (1998) for amphibians and by Péfaur and Rivero (2000) for amphibians and reptiles. Each bioregion is characterized by a number of taxonomic entities (i.e. orders, families, genera and species) and endemic species. A cluster analysis with Jaccard's coefficient and centroid clustering method was applied to the presence/absence of reptile species to reveal affinities between biogeographical regions. Introduced and/or translocated species, those of dubious occurrence or erroneously reported, and species requiring taxonomic, nomenclatural or distributional comments are also discussed. Museum acronyms follow Leviton et al. (1985), except for MHNLS (Museo de Historia Natural La Salle, Caracas, Venezuela) and MBLUZ (Museo de Biologia, la Universidad del Zulia, Maracaibo, Venezuela).

Results Taxonomic Richness We recorded three orders, 30 families, 122 genera (one exotic), and 370 reptile species (four exotics), in Venezuela (Table 1). Previous publications listed 292 (La Marca 1997) and 299 species (Péfaur & Rivero 2000). Although these taxa do occur in Venezuela, the introduced species (Anolis extremus, Ctenosaura similis, Hemidactylus frenatus and H. mabouia) are not included in our species richness analysis. The order Squamata contains 92.0% of the taxa (337 species), followed by Testudines (6.6%, 24 species) and Crocodylia (1.4%, 5 species). The most diverse families within Squamata are Dipsadidae (31.8%, 107 species), followed by Gymnophthalmidae (15.4%, 52 species), Colubridae (11.0%, 37 species), Sphaerodactylidae (7.4%, 25 species), Polychrotidae (7.1%, 24 species), Teiidae (5.0%, 17 species), Elapidae (3.9%, 13 species) and Viperidae (3.6%, 12 species), whereas the most speciose genera are Atractus (8.9%, 30 species), Anolis (6.8%, 23 species), Gonatodes (5.0%, 17 species), Liophis (4.2%, 14 species), Micrurus (3.6%, 12 species), and Anadia and Thamnodynastes with 3.0 % (10 species each). TABLE 1. Updated checklist of the Venezuelan reptiles. For each species the biogeographical regions where it has been recorded are indicated based on revision of museum specimens, pictures and records from the literature. Bioregions: 1. Marine (Marina); 2. Insular; 3. Maracaibo Lake Basin (depresión del Lago de Maracaibo); 4. Mountain Range of Mérida (cordillera de Mérida); 5. Mountain Range of Perijá (sierra de Perijá); 6. Lara-Falcón Hill System (sistema de colinas Lara-Falcón); 7. Coastal area (línea costera); 8. Central Coastal Range (cordillera de la costa central); 9. Eastern Coastal range (cordillera de la costa oriental); 10. Orinoco Delta (Delta del Orinoco); 11. Llanos; 12. Guayana; 13. Amazonia; 14. Pantepui (highlands of the Guiana shield above 1500 m). Threatened species: E, endangered; CE, critical endangered; V, vulnerable. ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ

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CLASS REPTILIA Order CROCODYLIA FAMILY ALLIGATORIDAE (Cuvier 1807) Genus Caiman Spix 1825 Caiman crocodilus Linnaeus 1758 Genus Paleosuchus Gray 1862

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Paleosuchus palpebrosus (Cuvier 1807) Paleosuchus trigonatus (Schneider 1801) FAMILY CROCODYLIDAE (Cuvier 1807) Genus Crocodylus Laurenti 1768 Crocodylus acutus (Cuvier 1807) Crocodylus intermedius Graves 1819

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E x

Order TESTUDINES FAMILY CHELONIIDAE Oppel 1811 Genus Caretta Rafinesque 1814 Caretta caretta (Linnaeus 1758) Genus Chelonia Brongniart 1800 Chelonia mydas (Linnaeus 1758) Genus Eretmochelys Fitzinger 1843 Eretmochelys imbricata (Linnaeus 1766) Genus Lepidochelys Fitzinger 1843 Lepidochelys olivacea (Eschscholtz 1829) FAMILY DERMOCHELYIDAE Fitzinger 1843 Genus Dermochelys Blainville 1816 Dermochelys coriacea (Vandelli 1761) FAMILY EMYDIDAE Rafinesque 1815 Genus Trachemys Agassiz 1857 Trachemys callirostris (Gray 1855) FAMILY GEOEMYDIDAE Theobald 1868 Genus Rhinoclemmys Fitzinger 1835 Rhinoclemmys diademata (Mertens 1954) Rhinoclemmys flammigera (Paolillo 1985)

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Rhinoclemmys punctularia (Daudin 1802) FAMILY KINOSTERNIDAE Agassiz 1857 Genus Kinosternon Spix 1824 Kinosternon scorpioides (Linnaeus 1766) FAMILY TESTUDINIDAE Batsch 1788 Genus Chelonoidis Fitzinger 1835 Chelonoidis carbonaria (Spix 1824) Chelonoidis denticulata (Linnaeus 1766) FAMILY CHELIDAE Gray 1831 Genus Batrachemmys Stejneger 1909 Batrachemys heliostemma McCord, Joseph-Ouni and Lamar 2001 Batrachemys raniceps (Gray 1855) Batrachemys zuliae (Pritchard and Trebbau 1984) Genus Chelus Duméril 1806 Chelus fimbriatus (Schneider 1783) Genus Mesoclemmys Gray 1873 Mesoclemmys gibba (Schweigger 1812) Genus Phrynops Wagler 1830 Phrynops tuberosus (Peters 1870) Genus Platemys Wagler 1830 Platemys platycephala (Schneider 1792) FAMILY PODOCNEMIDIDAE Cope 1868 Genus Peltocephalus Duméril and Bibron 1835 Peltocephalus dumerilianus (Schweigger 1812)

x x

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RIVAS ET AL.

TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ

Genus Podocnemis Wagler 1830 Podocnemis erythrocephala (Spix 1824) Podocnemis expansa (Schweigger 1812) Podocnemis unifilis Troschel 1848 Podocnemis vogli Muller 1935

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dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

V x

Order SQUAMATA FAMILY AMPHISBAENIDAE Gray 1844 Genus Amphisbaena Linnaeus 1758 Amphisbaena alba (Linnaeus 1758) Amphisbaena fuliginosa (Linnaeus 1758) Amphisbaena gracilis Strauch 1881 Amphisbaena rozei Lancini 1963 Genus Mesobaena Mertens 1925 Mesobaena huebneri Mertens 1925 FAMILY CORYTOPHANIDAE Fitzinger 1843 Genus Basiliscus Laurenti, 1768 Basiliscus basiliscus (Linnaeus, 1758) FAMILY IGUANIDAE Oppel 1811 Genus Iguana Laurenti 1768 Iguana iguana (Linnaeus 1758) Genus Ctenosaura Wiegmann 1828 Ctenosaura similis (Gray 1831) FAMILY POLYCHROTIDAE Fitzinger 1843 Genus Anolis Daudin 1802 Anolis anatoloros Ugueto, Rivas, Barros, Sánchez and GarcíaPérez 2007 Anolis annectens (Williams 1974)

x x

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Anolis apollinaris Boulenger 1919 Anolis auratus Daudin 1802 Anolis bellipeniculus (Myers and Donnelly 1995) Anolis biporcatus Wiegmann 1834 Anolis blanquillanus Humelinck 1940 Anolis bonairensis Ruthven 1923 Anolis carlostoddi (Williams, Praderio and Gorzula 1996) Anolis deltae Williams 1974 Anolis euskalerriari (Barros, Williams and Viloria 1996) Anolis extremus Garman 1887 Anolis fuscoauratus Duméril and Bibron 1837 Anolis jacare Boulenger, 1903 Anolis neblininus (Myers, Williams and McDiarmid 1993) Anolis nicefori (Dunn 1944) Anolis onca (O'Shaughnessy 1875)

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

Anolis ortonii Cope 1869 Anolis planiceps Troschel 1848 Anolis punctatus Daudin 1802 Anolis squamulatus Peters 1863 Anolis tetarii (Barros, Williams and Viloria 1996) Anolis tigrinus Peters 1863 Anolis tropidogaster Hallowell 1837 Genus Polychrus Cuvier 1817 Polychrus marmoratus (Linnaeus 1758) FAMILY TROPIDURIDAE Bell 1843

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Genus Plica Gray 1831

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RIVAS ET AL.

TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Plica lumaria Donnelly and Myers 1991 Plica pansticta Myers and Donnelly 2001 Plica plica (Linnaeus 1758) Plica umbra (Linnaeus 1758) Genus Stenocercus Duméril and Bibron 1837 Stenocercus erythrogaster (Hallowell 1856) Genus Tropidurus Wied-Neuwied 1825 Tropidurus bogerti Roze, 1958 Tropidurus hispidus (Spix 1825) Genus Uracentron Kaup 1826 Uracentron azureum Mertens 1925 Genus Uranoscodon Kaup 1825 Uranoscodon superciliosum (Linnaeus 1758) FAMILY SPHAERODACTYLIDAE Underwood 1954 Genus Coleodactylus Parker 1926 Coleodactylus amazonicus (Andersson 1918) Coleodactylus septentrionalis Vanzolini 1980 Genus Gonatodes Fitzinger 1843 Gonatodes albogularis (Duméril and Bibron 1836) Gonatodes alexandermendesi Cole and Kok 2006 Gonatodes annularis Boulenger 1887 Gonatodes antillensis (Van Lidth de Jeude 1887)

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

ŶĚĞŵŝĐ ƐƉĞĐŝĞƐ

x x

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Gonatodes astralis Schargel, Rivas, Makowsky, Señaris, Natera, Barros, Molina and Barrio-Amorós 2010 Gonatodes ceciliae Donoso-Barros 1966 Gonatodes falconensis Shreve 1947 Gonatodes humeralis (Guichenot 1855) Gonatodes infernalis Rivas and Schargel 2008 Gonatodes lichenosus Rojas-Runjaic, InfanteRivero, Cabello and Velozo 2010 Gonatodes ligiae Donoso-Barros 1967 Gonatodes petersi Donoso-Barros 1967 Gonatodes purpurogularis Esqueda 2004 Gonatodes seigliei Donoso-Barros 1966 Gonatodes superciliaris Barrio-Amorós and Brewer-Carias 2008 Gonatodes taniae Roze 1963 Gonatodes vittatus (Lichtenstein 1856) Genus Lepidoblepharis Peracca 1897 Lepidoblepharis montecanoensis Markezich and Taphorn 1994 Lepidoblepharis sanctaemartae Ruthven 1916 Genus Pseudogonatodes Ruthven 1915 Pseudogonatodes guianensis (Parker 1935) Pseudogonatodes lunulatus (Roux 1927) Pseudogonatodes manessi Avila-Pires and Hoogmoed 2000 Genus Sphaerodactylus Wagler 1830 Sphaerodactylus molei Boettger 1894 FAMILY GEKKONIDAE Gray 1825

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

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RIVAS ET AL.

TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Genus Hemidactylus Gray 1825 Hemidactylus frenatus Schlegel 1836 Hemidactylus mabouia (Moreau de Jonnés 1818) Hemidactylus palaichthus Kluge 1969 FAMILY PHYLLODACTYLIDAE Gamble, Bauer, Greenbaum and Jackman 2008 Genus Phyllodactylus Gray 1828 Phyllodactylus dixoni Rivero-Blanco and Lancini 1968 Phyllodactylus rutteni Hummerlinck 1947 Phyllodactylus ventralis O'Shaughnessy 1875 Genus Thecadactylus Oken 1817 Thecadactylus rapicauda (Houttuyn 1782) FAMILY GYMNOPHTHALMIDAE Merrem 1820 Genus Adercosaurus Myers and Donnelly 2001 Adercosaurus vixadnexus Myers and Donnelly 2001 Genus Anadia Gray 1845 Anadia bitaeniata Boulenger 1903 Anadia blakei Schmidt 1932 Anadia brevifrontalis (Boulenger 1903) Anadia escalerae Myers, Rivas and Jadin 2009 Anadia hobarti La Marca and García-Pérez 1990 Anadia marmorata (Gray 1846) Anadia mcdiarmidi Kok and Rivas 2011 Anadia pamplonensis Dunn 1944 Anadia pariaensis Rivas, La Marca and Oliveros 1999 Anadia steyeri Nieden 1914

x x

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x x

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Genus Arthrosaura Boulenger 1885 Arthrosaura montigena Myers and Donnelly 2008 Arthrosaura reticulata (O'Shaughnessy 1881) Arthrosaura synaptolepis Donnelly, McDiarmid and Myers 1992 Arthrosaura testigensis Gorzula and Señaris 1998 Arthrosaura tyleri (Burt and Burt 1931) Arthrosaura versteegii Van Lidth de Juede 1904 Genus Bachia Gray 1845 Bachia bicolor (Cope 1896) Bachia flavescens (Bonnaterre 1789) Bachia guianensis Hoogmoed and Dixon 1977 Bachia heteropa (Wiegmann 1856) Bachia pyburni Kizirian and McDiarmid 1998 Genus Cercosaura Wagler 1830 Cercosaura ampuedai (Lancini 1968) Cercosaura nigroventris (Gorzula and Señaris 1998) Cercosaura ocellata Wagler 1830 Cercosaura phelpsorum (Lancini 1968) Genus Euspondylus Tschudi 1845 Euspondylus acutirostris (Peters 1862) Euspondylus auyanensis Myers, Rivas and Jadin 2009 Euspondylus monsfumus (Mijares-Urrutia, Señaris and Arends 2001) Genus Gymnophthalmus Merrem 1820 Gymnophthalmus cryptus Hoogmoed, Cole and Ayarzaguena 1996

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

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RIVAS ET AL.

TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Gymnophthalmus speciosus (Hallowell 1861) Genus Leposoma Spix 1825 Leposoma caparensis Esqueda 2005 Leposoma hexalepis Ayala and Harris 1982 Leposoma parietale (Cope 1885) Leposoma percarinatum (Muller 1923) Genus Neusticurus Duméril and Bibron 1839 Neusticurus bicarinatus (Linnaeus 1758) Neusticurus medemi Dixon and Lamar 1981 Neusticurus racenisi Roze 1958 Neusticurus rudis Boulenger 1900 Neusticurus tatei (Burt and Burt 1931) Genus Proctoporus Tschudi 1845 "Proctoporus" cephalolineatus GarcíaPérez and Yustiz 1995 Genus Ptychoglossus Boulenger 1890 Ptychoglossus kugleri Roux 1927 Ptychoglossus brevifrontalis Boulenger 1912 Genus Riama Gray 1858 Riama achlyens (Uzzell 1958) Riama inanis (Doan and Schargel 2003) Riama luctuosa (Peters 1862) Riama rhodogaster Rivas, Schargel and Meik 2005 Genus Riolama Boulenger 1900 Riolama leucosticta (Boulenger 1900) Riolama luridiventris Esqueda, La Marca and Praderio 2004

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

x x x

x x

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Riolama uzzelli Molina and Señaris 2003 Genus Tretioscincus Cope, 1862 Tretioscincus bifasciatus Shreve 1947 Tretioscincus oriximinensis Avila-Pires 1995 FAMILY TEIIDAE Gray 1827 Genus Ameiva Meyer 1795 Ameiva ameiva (Linnaeus 1758) Ameiva bifrontata Cope 1862 Ameiva provitaae Garcia-Perez 1995 Genus Cnemidophorus Wagler 1830 Cnemidophorus arenivagus Markezich, Cole and Dessauer 1997 Cnemidophorus cryptus Cole and Dessauer 1993 Cnemidophorus flavissimus Ugueto, Harvey and Rivas 2009 Cnemidophorus gramivagus McCristal and Dixon 1987 Cnemidophorus lemniscatus (Linnaeus 1758) Cnemidophorus leucopsammus Ugueto and Harvey 2010 Cnemidophorus nigricolor Peters 1873 Cnemidophorus rostralis Ugueto and Harvey 2010 Cnemidophorus senectus Ugueto, Harvey and Rivas 2009 Genus Crocodilurus Spix 1825 Crocodilurus amazonicus (Spix 1825) Genus Kentropyx Spix 1825 Kentropyx altamazonica Cope 1876 Kentropyx calcarata Spix 1825 Kentropyx striata (Daudin 1802)

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

x x x

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RIVAS ET AL.

TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Genus Tupinambis Daudin 1803 Tupinambis teguixin (Linnaeus 1758) FAMILY SCINCIDAE Gray 1825 Genus Mabuya Fitzinger 1826 Mabuya carvalhoi ReboucasSpieker and Vanzolini 1990 Mabuya croizati Horton 1973 Mabuya falconensis Mijares-Urrutia and Arends 1997 Mabuya nebulosylvestris Miralles, Rivas, Bonillo, Schargel, Barros, GarcíaPérez and BarrioAmorós 2009 Mabuya nigropunctata Spix 1825 Mabuya meridensis Miralles, Rivas and Schargel 2005 Mabuya zuliae Miralles, Rivas, Bonillo, Schargel, Barros, García-Pérez and Barrio-Amorós 2009 FAMILY ANILIIDAE Stejneger 1907 Genus Anilius Oken 1816 Anilius scytale (Linnaeus 1758) FAMILY BOIDAE Gray 1842 Genus Boa Linnaeus 1758 Boa constrictor Linnaeus 1758 Genus Corallus Daudin 1803 Corallus caninus (Linnaeus 1758) Corallus hortulanus (Linnaeus 1758) Corallus ruschenbergerii (Cope 1876) Genus Epicrates Wagler 1830 Epicrates cenchria (Linnaeus 1758) Epicrates maurus (Gray 1849)

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Genus Eunectes Wagler 1830 Eunectes murinus Linnaeus 1758 FAMILY COLUBRIDAE Cope 1886 Genus Chironius Fitzinger 1826 Chironius carinatus (Linnaeus 1758) Chironius challenger Kok 2010 Chironius spixi (Hallowell 1845) Chironius exoletus (Linnaeus 1758) Chironius fuscus (Linnaeus 1758) Chironius monticola Roze 1952 Chironius multiventris Schmidt and Walker 1943 Chironius septentrionalis (Dixon, Wiest and Cei 1993) Chironius scurrulus (Wagler 1824) Genus Coluber Linnaeus 1758 Coluber mentovarius (Duméril, Bibron and Duméril 1854) Genus Dendrophidion Fitzinger 1843 Dendrophidion dendrophis (Schlegel 1837) Dendrophidion nuchale (Peters 1864) Dendrophidion percarinatus Cope 1893 Genus Drymarchon Fitzinger 1843 Drymarchon caudomaculatus Wüster, Yrausquin and MijaresUrrutia 2001 Drymarchon corais (Boie 1827) Drymarchon margaritae (Roze 1959) Drymarchon melanurus (Duméril, Bibron and Duméril 1854)

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

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RIVAS ET AL.

TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Genus Drymobius Fitzinger 1843 Drymobius rhombifer (Günther 1860) Genus Drymoluber Amaral 1929 Drymoluber dichrous (Peters 1863) Genus Lampropeltis Fitzinger 1843 Lampropeltis triangulum Lacépède 1788 Genus Leptophis Bell 1825 Leptophis ahaetulla (Linnaeus 1758) Leptophis cupreus (Cope 1868) Genus Mastigodryas Amaral 1843 Mastigodryas amarali (Stuart 1938) Mastigodryas bifossatus (Amaral 1931) Mastigodryas boddaerti (Sentzen 1796) Mastigodryas pleei (Duméril, Bibron and Duméril 1854) Genus Oxybelis Wagler 1830 Oxybelis aeneus (Wagler 1824) Oxybelis fulgidus (Daudin 1803) Genus Pseustes Fitzinger 1843 Pseustes poecilonotus (Peters 1867) Pseustes shropshirei (Barbour and Amaral 1924) Pseustes sulphureus (Wagler in Spix 1824) Genus Rhinobothryum Wagler 1830 Rhinobothryum bovalli Andersson 1916 Rhinobothrium lentiginosum (Scopoli 1785) Genus Spilotes Wagler 1830 Spilotes pullatus (Linnaeus 1758) Genus Stenorrhina Duméril 1853

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Stenorrhina degenhardtii (Jan 1876) Genus Tantilla Baird and Girard 1853 Tantilla melanocephala (Linnaeus 1758) Tantilla semicincta (Duméril, Bibron and Duméril 1854) FAMILY DIPSADIDAE Bonaparte 1838 Genus Atractus Wagler 1828 Atractus acheronius Passos, Rivas and BarrioAmorós 2009 Atractus duidensis Roze 1961 Atractus elaps (Gunther 1858) Atractus eriki Esqueda, La Marca and Bazó 2007 Atractus emigdioi González-Sponga 1971 Atractus erythromelas Boulenger 1903 Atractus fuliginosus (Hallowell 1845) Atractus guerreroi Myers and Donnelly 2008 Atractus insipidus Roze 1961 Atractus lancinii Roze 1961 Atractus major Boulenger 1893 Atractus matthewi Markezich and BarrioAmorós 2004 Atractus mariselae Lancini 1969 Atractus meridensis Esqueda and La Marca 2005 Atractus micheleae Esqueda and La Marca 2005 Atractus mijaresi Esqueda and La Marca 2005 Atractus multidentatus Passos, Rivas and BarrioAmorós 2009 Atractus ochrosetrus Esqueda and La Marca 2005

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

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RIVAS ET AL.

TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Atractus pamplonensis Amaral 1935 Atractus riveroi Roze 1961 Atractus steyermarki Roze 1958 Atractus tamaensis Esqueda and La Marca 2005 Atractus tamessari Kok 2006 Atractus taphorni Schargel and GarcíaPérez 2002 Atractus torquatus (Duméril, Bibron and Duméril 1854) Atractus trilineatus Wagler 1828 Atractus turikensis Barros 2000 Atractus univittatus (Jan 1862) Atractus ventrimaculatus Boulenger 1905 Atractus vittatus Boulenger 1894 Genus Clelia Fitzinger 1826 Clelia clelia (Daudin 1803) Genus Dipsas Laurenti 1768 Dipsas catesbyi (Sentzen 1796) Dipsas copei (Gunther 1872) Dipsas indica Laurenti 1768 Dipsas peruana (Boettger 1898) Dipsas pavonina Schlegel 1837 Dipsas praeornata Werner 1909 Dipsas pratti (Boulenger 1897) Dipsas variegata (Duméril, Bibron and Duméril 1854) Genus Enulius Cope 1871 Enulius flavitorques (Cope 1869) Genus Erythrolamprus Wagler 1830

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

x x x

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Erythrolamprus aesculapii (Linnaeus 1766) Erythrolamprus bizona Jan 1863 Erythrolamprus pseudocorallus Roze 1959 Genus Helicops Wagler 1830 Helicops angulatus (Linnaeus 1758) Helicops hagmanni Roux 1910 Helicops hogei Lancini 1979 Helicops pastazae Shreve 1934 Helicops scalaris Jan 1865 Genus Hydrodynastes Fitzinger 1843 Hydrodynastes bicinctus (Herrmann 1804) Genus Hydrops Wagler 1830 Hydrops triangularis (Wagler 1824) Hydrops martii (Wagler 1824) Genus Imantodes Duméril 1853 Imantodes cenchoa (Linnaeus 1758) Imantodes lentiferus (Cope 1894) Genus Leptodeira Fitzinger 1843 Leptodeira annulata (Hallowell 1845) Leptodeira bakeri Ruthven 1936 Leptodeira septentrionalis (Bocourt 1884) Genus Liophis Wagler 1830 Liophis breviceps Cope 1861 Liophis cobella (Linnaeus 1758) Liophis dorsocorallinus Esqueda, Natera, La Marca and Ilija-Fistar 2007 Liophis epinephelus (Cope 1862)

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x x

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RIVAS ET AL.

TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ

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Liophis ingeri Roze 1958 Liophis melanotus (Shaw 1802) Liophis miliaris (Linnaeus 1758) Liophis poecilogyrus Schlegel 1837 Liophis reginae (Wagler 1824) Liophis torrenicola Donnelly and Myers 1991 Liophis trebbaui Roze 1958 Liophis typhlus (Linnaeus 1758) Liophis williamsi (Roze 1958) Liophis zweifeli (Roze 1959) Genus Lygophis Fitzinger 1843 Lygophis lineatus (Linnaeus 1758) Genus Ninia Baird and Girard 1853 Ninia atrata (Hallowell 1845) Genus Oxyrhopus Wagler 1830 Oxyrhopus doliatus Duméril, Bibron and Duméril 1854 Oxyrhopus leucomelas (Werner 1916) Oxyrhopus occipitalis (Wagler 1824) Oxyrhopus melanogenys (Tschudi 1854) Oxyrhopus petolarius (Linnaeus 1758) Genus Phimophis Wagler, 1830 Phimophis guianensis (Troschel in Schomburgk, 1848) Genus Philodryas Wagler, 1830 Philodryas cordata Donnelly and Myers 1991 Philodryas olfersii (Wied-Neuwied 1825) Philodryas viridissimus (Linnaeus 1758)

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dƌĂŶƐůŽĐĂƚĞĚ ƐƉĞĐŝĞƐ

dŚƌĞĂƚĞŶĞĚ ƐƉĞĐŝĞƐ

x x

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REPTILES OF VENEZUELA

TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Genus Plesiodipsas Harvey, Rivas, Caicedo and Rueda-Almonacid 2008 Plesiodipsas perijanensis (Alemán 1953) Genus Pseudoboa Schneider 1801 Pseudoboa coronata Schneider 1801 Pseudoboa neuwiedii (Duméril, Bibron and Duméril 1854) Genus Pseudoeryx Fitzinger 1826 Pseudoeryx plicatilis (Linnaeus 1758) Pseudoeryx relictualis Schargel, Rivas, Barros, Péfaur and Navarrete 2007 Genus Sibon Fitzinger 1826 Sibon nebulata (Linnaeus 1758) Genus Siphlophis Fitzinger 1843 Siphlophis cervinus (Laurenti 1768) Siphlophis compressus (Daudin 1803) Genus Taeniophallus Cope 1895 Taeniophallus nebularis Schargel, Rivas and Myers 2005 Genus Thamnodynastes Wagler 1830 Thamnodynastes chimanta Roze 1958 Thamnodynastes corocoroensis Gorzula and Ayarzaguena 1995 Thamnodynastes dixoni Bailey and Thomas 2007 Thamnodynastes duida Myers and Donnelly 1995 Thamnodynastes ceibae Bailey and Thomas 2007 Thamnodynastes marahuaquensis Gorzula and Aryarzaguena 1995 Thamnodynastes pallidus (Linnaeus 1758) Thamnodynastes paraguanae Bailey and Thomas 2007

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Thamnodynastes ramonriveroi Manzanilla and Sánchez 2005 Thamnodynastes yavi Myers and Donnelly 1995 Genus Umbrivaga Roze 1964 Umbrivaga mertensi Roze 1964 Genus Urotheca Bibron 1843 Urotheca fulviceps Cope 1886 Urotheca multilineata (Peters 1859) Genus Xenodon Boie 1827 Xenodon merremii (Wagler 1824) Xenodon rhabdocephalus (Wierd 1824) Xenodon severus (Linnaeus 1758) Genus Xenoxybelis Machado 1993 Xenoxybelis argenteus (Daudin 1803) FAMILY ELAPIDAE Boie 1827 Genus Leptomicrurus Schmidt 1937 Leptomicrurus collaris (Schlegel 1837) Genus Micrurus Wagler 1824 Micrurus circinalis (Duméril and Bibron 1854) Micrurus dissoleucus (Cope 1860) Micrurus dumerilii (Jan 1858) Micrurus hemprichii (Jan 1858) Micrurus isozonus (Cope 1860) Micrurus lemniscatus (Linnaeus 1758) Micrurus meridensis Roze 1989 Micrurus mipartitus (Duméril, Bibron and Duméril 1854) Micrurus nattereri (Schmidt 1952)

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x x

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Micrurus psyches (Daudin 1803) Micrurus remotus (Roze 1987) Micrurus obscurus Jan 1872 FAMILY VIPERIDAE Gray 1825 Genus Bothriechis Peters 1859 Bothriechis schlegelii (Berthold 1846) Genus Bothriopsis Peters 1861 Bothriopsis bilineata (Wied 1825) Bothriopsis medusa (Sternfeld 1920) Bothriopsis taeniata (Wagler 1824) Genus Bothrops Wagler 1824 Bothrops asper (Garman 1883) Bothrops atrox (Linnaeus 1758) Bothrops brazilii Hoge 1953 Bothrops venezuelensis Sandner-Montilla 1952 Genus Crotalus Linnaeus 1758 Crotalus durissus Linnaeus 1758 Crotalus vegrandis Klauber 1941 Genus Lachesis Daudin 1803 Lachesis muta (Linnaeus 1766) Genus Porthidium Cope 1871 Porthidium lansbergii (Schlegel 1841) FAMILY ANOMALEPIDIDAE Taylor 1939 Genus Helminthophis Peters 1860 Helminthophis flavoterminatus (Peters 1857) Genus Liotyphlops Peters 1881 Liotyphlops albirostris (Peters 1881)

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TABLE 1. (continued) ŝŽƌĞŐŝŽŶƐ dĂǆŽŶŽŵŝĐ ůŝƐƚ ŽĨ ƚŚĞ sĞŶĞǌƵĞůĂŶ ƌĞƉƚŝůĞƐ Genus Typhlophis Fitzinger 1843 Typhlophis squamosus (Schlegel 1839) FAMILY LEPTOTYPHLOPIDA E Stejneger 1891 Genus Epictia Gray 1845 Epictia albifrons (Wagler 1824) Epictia goudotti (Duméril and Bibron 1844) Epictia tenella (Klauber 1939) Epictia signata (Jan 1861) Genus Rena Baird and Girard 1853 Rena affinis (Boulenger 1884) Genus Siagonodon Peters 1881 Siagonodon septemstriatus (Schneider 1801) Genus Trilepida Hedges 2011 Trilepida macrolepis (Peters 1857) FAMILY TYPHLOPIDAE Jan 1863 Genus Typhlops Oppel 1811 Typhlops brongersmianus Vanzolini 1972 Typhlops lehneri Roux 1926 Typhlops minuisquamus Dixon and Hendricks 1979 Typhlops reticulatus (Linnaeus 1766)

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x x

Among the 14 Venezuelan bioregions, the highest reptile diversity is found in Guayana (39.9%, 146 species), followed by Amazonia (35.8%, 131 species), Mountain Range of Mérida (29.6%, 108 species), Central Coastal Range (27.9%, 102 species), Eastern Coastal Range (22.1%, 81 species), Lara-Falcón Hill System (21.9%, 80 species each), Llanos (19.1%, 70 species), Orinoco Delta and Perijá (18.9%, 69 species each), Maracaibo Lake Basin (18.3%, 67 species), Insular region (13.4%, 49 species), Pantepui (10.1%, 37 species), Coastal Area (4.9%, 18 species), and Marine bioregion (1.6%, 6 species) (Table 2). However, some bioregions are inadequately surveyed (e.g. Guayana, Lara-Falcón Hill System and Perijá) and future work is needed to assess the accurate reptile diversity at these locations. Based on species composition and shared species, bioregions can be clustered in seven groups (Fig. 2; arranged in descending order according to shared species): Guayana + Amazonia; Llanos + Orinoco Delta; Moun-

REPTILES OF VENEZUELA

tain Ranges of Mérida and Perijá; Maracaibo Lake Basin + Central Coastal Range + Lara-Falcón Hill System; Coastal + Insular areas + Eastern Coastal Range. Pantepui and the Marine bioregion are independent entities. Some species (e.g. Pseudoboa neuwiedii, Tantilla melanocephala and Leptodeira annulata) are common elements of the lowlands of the Maracaibo Lake Basin and the Central Coastal Range. Other species (e.g. Anadia steyeri and Euspondylus acutirostris) are known only from the Central Coastal Range and the Lara-Falcón Hill System, whereas other taxa (e.g. Thamnodynastes paraguanae and Cnemidophorus arenivagus) occur only in the Lara-Falcon Hill System and the Maracaibo Lake Basin. Overlapping distributions are thus not consistent and are affected by many ecological factors. The great Andean Mountain chain splits into two branches in extreme northern Colombia (in a region known as the Pamplona knot), one extending northeast into Venezuela being the origin of the Cordillera de Mérida, while the other extends north being the origin of the Sierra de Perijá, which has more affinities with the Cordillera Oriental de Colombia than with the Cordillera de Mérida. Although they are considered as being the Venezuelan Andes, along with the Tamá massif, these two mountain ranges have both geological and biogeographical differences (Ataroff & Sarmiento 1994). Both regions share several Andean and Middle American species (e.g. Anolis jacare, A. biporcatus, Atractus ventrimaculatus, and Lampropeltis triangulum).

FIGURE 2. Cluster analysis, with Jaccard's coefficient and centroid clustering method showing the taxonomic similarities between the bioregions considered. Although Pantepui and Marine cluster together they do not share any similarity, these two bioregions are clearly distinct from all other bioregions.

Some aquatic species like Paleosuchus palpebrosus, Eunectes murinus, Chelus fimbriatus, Podocnemis expansa, P. unifilis, Hydrops triangularis and Helicops angulatus are distributed throughout the Llanos and the Orinoco Delta. Many of these taxa are also present throughout the Guiana Shield. Anolis onca, Gonatodes vittatus, Mabuya falconensis, and Mastigodryas pleei are all associated with habitats found along the Venezuelan coast. Such habitats are also common on many islands off the Caribbean coast of Venezuela. Endemic taxa found in the mountains of Península de Paria and Turimiquire are related to species from Trinidad, the Guiana Shield lowlands and the forests of the central portion of the Venezuelan Coastal Range (see Schargel et al. 2005).

RIVAS ET AL.

Species endemism One hundred twenty one (33.0%) reptile species are endemic to Venezuela (Table 2, see also Figs. 3 and 4). One hundred nineteen species of these endemics (98.3%) belong to the order Squamata and two species (1.7%) to Testudines. The family Dipsadidae (42 species) is the richest endemic family, followed by Gymnophthalmidae (31 species), Sphaerodactylidae (12 species) and Polychrotidae (10 species). The richest genera in endemic species are Atractus (22 species), Anolis (10 species), Gonatodes (9 species, see Fig. 3), Anadia (9 species), Liophis (6 species), Thamnodynastes (6 species), and Riama (4 species). Only one reptile genus is endemic to Venezuela, Ardecosaurus, which is monotypic and currently known only from the Yutajé-Corocoro massif in the northwestern tepui area (Amazonas State) (Myers & Donnelly 2001). The bioregion with the highest percentage of known endemic species is Pantepui (67.6%), followed by the Mountain Range of Mérida (28.7%), the Central Coastal Range (23.5%), the Insular region (18.4%), the Lara-Falcón hill system (16.3%), the Eastern Coastal Range (16.0%), Perijá (7.2%), Amazonia (6.9%) and Guayana (6.2%). The six remaining bioregions have 5.0% or less (Table 2).

Endangered Species In the third edition of the Red Book of the Venezuelan fauna (Rodríguez & Rojas-Suárez 2008), 22 species (6.0% of total species) were listed as critically endangered (CE), endangered (E) or vulnerable (V). Among them, 10 species belong to the order Testudines, five of which are marine turtles (Caretta caretta [E], Chelonia mydas [E], Dermochelys coriacea [CE], Eretmochelys imbricata [CE], and Lepidochelys olivacea [E]) and five are freshwater turtles (Batrachemys zuliae [V], Peltocephalus dumerilianus [V], Podocnemis expansa [CE], P. unifilis [V] and Rhinoclemmys diademata [V]); ten belong to the order Squamata, eight of which are lizards (Ameiva provitaae [V], Anadia blakei [V], A. pariaensis [V], Anolis euskalerriari [V], A. tetarii [V], Euspondylus monsfumus [V], Lepidoblepharis montecanoensis [CE] and Mabuya croizati [V]), and two are snakes (Atractus matthewi [V] and A. turikensis [V]); and two belong to the order Crocodylia (Crocodylus acutus [E] and C. intermedius [CE]) (see Table 1). With respect to the bioregions harbouring species included in the IUCN Red List, Marine is first with 100% of threatened species, followed by the Eastern Coastal Range (7.4%), the Llanos (5.8%), the Mountain Range of Perijá (4.3%), the Maracaibo Lake Basin (4.4%), and the Orinoco Delta (4.3%), the remaining bioregions have 3% or less of threatened species (Table 2).

TABLE 2. Orders, families, genera and species numbers of Venezuelan reptiles per bioregion, with numbers and percentages of endemic and threatened taxa. Exotic and translocated taxa have been excluded. Venezuela

Marine

Insular

Maracaibo Lake Basin

Mountain Range of Mérida

Mountain Range of Perija

Lara-Falcón Hill System

Order

Family

Genera

121

Species

366

108

% of total of species

100

1.6

13.4

18.3

29.5

18.9

21.9

Endemics

121

% endemics

33.1

18.4

4.5

28.7

7.3

16.3

Threatened 22

% threatened

100

4.5

0.9

4.4

1.3

6.0

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TABLE 2 (continued) Coastal area

Central Coastal Range

Eastern Coastal Range

Orinoco Delta

Llanos

Guayana

Amazonia

Pantepui

Order

Family

Genera

Species

102

146

131

% of total of species

4.9

27.9

22.1

18.9

19.1

39.9

35.8

10.1

Endemics

% endemics

23.5

16.1

4.4

1.4

6.2

6.9

67.6

Threatened

1.0

7.4

4.4

5.7

2.1

3.1

% threatened 0

Introduced (non-native) and displaced (native) species To date, four species of lizards are known to have been introduced to Venezuela: Anolis extremus.—The species was cited for the first time in Venezuela based on a personal communication from C. Rivero-Blanco to J. Peters (Peters & Donoso-Barros 1970). We had the opportunity to examine some specimens of that taxon (MCZ 110294−95 and MBUCV 1889), all of them collected in the city of Caracas, confirming their identification. An additional male specimen, collected in June 2011 in eastern Caracas, was recently brought to the Parque del Este Zoo in Caracas (Fig. 5). This new record confirms that the species is still present in the country, and one of us (GR) also observed A. extremus on the wall of a nursery near Macaracuay, eastern Caracas. This anole, native to the Lesser Antilles, has also been introduced to Trinidad. Ctenosaura similis.—This iguanid lizard originating from Central America has been introduced during recent decades into North America (Savage 2002; Krysko et al. 2003). The Venezuelan population is the first record of the genus in mainland South America, the species inhabits dry forest in the Coastal Area near Barcelona city (BarrioAmorós & Rivas-Fuenmayor 2008; Flores & Esqueda 2008). Date of introduction is unknown. Hemidactylus mabouia.—This species originates from Africa and has long been established in South America, including Venezuela (Kluge 1969). We here report the first record of H. mabouia from the Mountain Range of Mérida, based on a juvenile specimen (MBLUZ 1033) collected on a steel roof beam in the parking lot of a building in Ejido, Mérida. On 12 December 2010 on the Isla La Orchila, Dependencias Federales, we observed several H. mabouia (two collected, MBLUZ 1049–50) in the installations of the national military base. The geckos were very common on building walls as well as on small trees (mainly under loose bark) surrounding the area. In April 2011 we observed H. mabouia mainly on building walls (two juveniles were found on trees within an open area) on the island of Gran Roque, part of Los Roques Archipelago National Park. This species was not found on Los Roques archipelago nor on the island of La Orchila in 1950 (Roze 1954). We presume that these populations were introduced along with construction materials from mainland Venezuela (probably from La Guaira Bayard, Maiquetia International Airport). These specimens are the first records of H. mabouia on oceanic islands of the Venezuelan Caribbean. The species had been previously recorded from other Venezuelan islands, but only of continental origin (e.g. Isla Margarita). Hemidactylus frenatus.—An Asiatic species, which has only recently been recorded in Venezuela (Rivas Fuenmayor et al. 2005b) and appears to be expanding its range in South America (Jadin et al. 2009).

RIVAS ET AL.

Three Venezuelan turtles have been translocated from their natural habitats in Venezuela to other locations inside the country: Chelonoidis carbonaria.—The species was recorded by Meek (1910, as Testudo tabulata) from Los Testigos, an archipelago composed of 16 islands, located 68 km north of the eastern coast of Venezuela. The islands of Los Testigos have different climatic conditions than other archipelagos of the Venezuelan Caribbean Sea. Rainfall reaches an average of 400 mm per year, which allows a greater vegetation cover (Bisbal 2008). Meek (1910) did not mention the origin of this population, which is of special interest because of the early dates of collection (1908−1909). There are two possible origins for that population: (1) Amerindians carried specimens from the mainland as a food source prior to the arrival of Europeans or (2) turtles were brought by pirates, who frequently released goats and tortoises on different islands for food storage. Chelonoidis carbonaria is well known on many Caribbean islands. The population from Isla Margarita has also apparently been introduced (Roze 1964b). Podocnemis expansa.— That species, native to the Orinoco Delta and the most endangered turtle in Venezuela, has become rare in its natural geographic distribution (Barrio-Amorós & Narbaiza 2008). Podocnemis expansa occurs outside its original range at Lago de Valencia, a large lake in the Central Coastal Range, where it is highly endangered as well. The species was introduced in that area in the middle of the last century by the Ministerio de Agricultura y Cría (Fernández 1968). Podocnemis vogli.—Also native to the Orinoco Delta. A population of P. vogli inhabits an artificial reservoir in “Represa Burro Negro” in the Maracaibo Lake Basin. We cannot confirm when this species was introduced, but it was only recently recorded from that locality (Rojas-Runjaic et al. 2007). An adult P. vogli (EBRG 5133) was recently collected in Parroquia Río Negro, Municipio Perijá, Estado Zulia (09°30’15” N, 72°28’47”W). The specimen was collected in a stream bordered by a small gallery forest in a savannah surrounded by flat areas heavily modified by livestock (pastures). The stream is ca. 2 km from Santa Rosa River and about 1 km from the farm “Ecuador”. Apparently specimens of P. vogli were released in some ponds of the farm, which explains the origin of that population. More turtles, both adults and young specimens, were observed on the banks of two artificial lagoons (called Jagueyes) inside the farm. The impact of this population on the local wildlife needs to be evaluated.

Undescribed species To our knowledge, at least 30 new species of Venezuelan Squamata are currently being described by various authors. The lizard genera Anadia, Cnemidophorus, Gonatodes and Phyllodactylus still have several species that remain to be formally described (Rivero-Blanco 1979; McBee et al. 1984; unpubl. data from the authors). These genera are speciose and Venezuela could have more species than any other country in which these genera occur. The Guianan-Amazonian genus Neusticurus needs a taxonomic revision, especially the populations under the names N. rudis and N. racenisi (Barrio-Amorós & Brewer Carías 2008; Myers & Donnelly 2008; pers. obs. from the authors), which could lead to the description of new taxa. The family Polychrotidae and the genus Atractus in Venezuela are being revised by GU and GR, and P. Passos and colleagues, respectively, which will certainly yield some additional new species.

Species erroneously reported from Venezuela A number of species have been cited erroneously from Venezuela (e.g. in Peters & Orejas-Miranda 1970; Peters & Donoso-Barros 1970; La Marca 1997; Péfaur 1992; Péfaur & Rivero 2000). These species are listed below in alphabetical order with arguments for excluding them from the reptile fauna of Venezuela.

REPTILES OF VENEZUELA

FIGURE 3. Of the 28 species currently recognized in the genus Gonatodes, Venezuela harbors 17, more than any other country. Ten of these species are endemic, and several others remain to be described. A. Gonatodes humeralis, Reserva Forestal de Imataca, Bolívar. Photo: César L. Barrio-Amorós; B. Gonatodes petersi (endemic), Sierra de Perijá, Zulia. Photo: Tito R. Barros; C. Gonatodes antillensis, Archipelago Las Aves, Barlovento, Dependencias Federales. Photo: Pedro Velozo; D. Gonatodes seigliei (endemic), near Cueva del Guacharo, Monagas. Photo: Walter E. Schargel; E. Gonatodes infernalis (endemic), near Puerto Ayacucho, Amazonas. Photo: Tito R. Barros; F. Gonatodes ceciliae, road Güiria-Macuro, Península de Paria, Sucre. Photo: Luis A. González; G. Gonatodes astralis (endemic), near Puerto Ayacucho, Amazonas; Photo: César L. BarrioAmorós; H. Gonatodes annularis, Reserva Forestal de Imataca, Bolívar; Photo: Eric N. Smith. All specimens, except a female shown in E, are males.

RIVAS ET AL.

FIGURE 4. Some endemic squamates from Venezuela. A. Riama inanis, Los Alcaravanes, Calderas, Barinas. Photo: César L. Barrio-Amorós; B. Pseudogonatodes manessi, Rancho Grande, Aragua. Photo: Eric N. Smith; C. Anadia brevifrontalis, Paramo de Mucuchies, 3400 m, Mérida. Photo: César L. Barrio-Amorós; D. Cnemidophorus flavissimus, Puerto Real, Los Frailes Archipelago. Photo: Oscar Lasso-Alcalá; E. Cnemidophorus nigricolor, boca de Sebastopol, Los Roques archipelago. Photo: Gilson A. Rivas; F. Anadia mcdiarmidi, Abakapá-tepui, Bolívar. Photo: Philippe J. R. Kok; G. Atractus vittatus, Monumento Natural Pico Codazzi, Aragua. Photo: Marcial Quiroga; H. Atractus lancinii, Monumento Natural Pico Codazzi, Aragua. Photo: Marcial Quiroga. The species shown in B, G and H are illustrated in life for the first time.

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Order TESTUDINES Batrachemys nasuta.—The species has been cited from Venezuela, but there is no Venezuelan voucher specimen of this taxon. The Batrachemys nasuta specimens mentioned in Pritchard and Trebbau (1984) were captive animals without proper locality of occurrence. Bour and Pauler (1987) and McCord et al. (2001) do not list this species as part of the Venezuelan turtle fauna. Barrio-Amorós and Narbaiza (2008) made similar comments on this case. Podocnemis lewyana.—This species was reported from Venezuela based on a single specimen found in a bend of the Tarra River, south of Maracaibo Lake, near the town of Casigua (Pauler & Trebbau 1995). Podocnemis lewyana is endemic to the Magdalena River, Colombia, with populations decreasing due to poaching. Three of us (GR, TB & CBA) conducted field studies on Chelus fimbriatus in the Maracaibo Lake Basin (Rivas et al. 2007) and interviewed the farmer who caught the original specimen behind his house along the Tarra River. That river is often used by Colombians to bring food supplies and they might very well bring turtles as well. It was the first and only occasion the farmer saw such a turtle and interviews with other local people show no familiarity with P. lewyana. No additional specimen has ever been reported, and apparently no population established successfully in Venezuela, therefore we do not consider the species as part of the herpetofauna of Venezuela.

Order SQUAMATA Amphisbaena spurrelli.—This species was erroneously cited from Venezuela by Donoso-Barros (1968) because the author mistakenly synonymised A. rozei under A. spurrelli. Several authors followed this arrangement (see La Marca 1997; Péfaur & Rivero 2000). Amphisbaena spurrelli is restricted to the Pacific versant of Colombia and Panama, west of the Andes, while A. rozei is known from the Guayana region in Venezuela (thus well east of the Andes). Gans and Mathers (1977), and more recently Vanzolini (2002), showed substantial morphological differences between these species. Anolis frenatus.—Although this species has repeatedly been included as part of the Venezuelan herpetofauna, no voucher specimen exists to confirm its presence in the country.

FIGURE 5. Anolis extremus, an adult male from eastern Caracas, Venezuela. Photo by Luis A. Rodriguez.

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Anolis gibbiceps.—This species was described based on a single female from “Caracas” (Cope 1864). Anolis gibbiceps was subsequently reported from northeastern South America, northern Venezuela and Guyana (Burt & Burt 1933; Parker 1935; Donoso-Barros 1968; Peters & Donoso-Barros 1970), but none of these authors listed specific localities other than that of the holotype. Anolis gibbiceps is closely allied to A. fuscoauratus (Savage & Guyer 1989). The species was not considered by Williams (1976) in his grouping of South American anoles. The validity of this taxon is highly dubious. Based on the original description, the inclusion of this species in the A. fuscoauratus group does not seem appropriate. Moreover, we have been unable to find additional information about this taxon other than the original and brief description (Donoso-Barros 1968; Peters & Donoso-Barros 1970). Examination of specimens assigned to A. gibbiceps from Venezuela revealed misidentified anoles of other species (e.g. A. squamulatus). Recently, C. McCarthy (NHM) sent us photographs of the holotype (BMNH 1946.8.13.35, formerly 1846.10.28.5, Fig. 6) and informed us that the original catalogue entry (dated 1846) states that the specimen is from Jamaica. Written with the holotype is an added undated comment in the handwriting of Miss A.G.C. Grandison, who worked in the collection between 1951–1984, stating: “type locality given as Caracas in Cope’s description”. It is unclear on which basis the locality was changed from “Jamaica” to “Caracas”, but after examining the photographs of the holotype we observed similarities between that specimen and three of the seven anole species found in Jamaica (A. grahami, A. lineatopus and A. opalinus) and conclude that the holotype indeed belongs to a taxon of Caribbean or Central American origin. Thus, the type locality reported by Cope is likely in error. Because we have not examined the holotype in detail we refrain from assigning it to any particular species. It is clear, however, that this taxon does not belong to the Venezuelan herpetofauna. Anolis roquet.—This species was reported in Venezuela as Anolis roquet extremus (Peters & Donoso-Barros 1970); see our comments about A. extremus in the Introduced species section. Anolis transversalis.—Donoso-Barros (1968) mentioned this species from Venezuelan Amazonas, but there is no voucher specimen to confirm the record. Nor Williams and Vanzolini (1966) or Avila-Pires (1995) considered the species present in Venezuela. Atractus badius.—This species was erroneously recorded from the Coastal Range and the Cordillera de Mérida (Roze 1966; Lancini 1979). According to Hoogmoed (1980) this taxon probably includes several distinct species and A. badius has a limited distribution in the Guiana Shield. It is likely that the specimens cited as A. badius from Venezuela are in fact A. univittatus or A. lancinii, at least those from the Coastal Range records (W. Schargel and P. Passos, pers. comm.). Specimens from the Cordillera de Mérida could belong to an undescribed taxon. The species is not considered in our analysis, or as belonging to the Venezuelan herpetofauna. Coleodactylus meridionalis.—No voucher specimen of this species is known from Venezuela. The inclusion of this taxon in the Venezuelan herpetofauna (Rivero-Blanco 1967) is based on misidentified specimens of C. septentrionalis (Vanzolini 1980; Rivas Fuenmayor & Molina 2001). Coleodactylus meridionalis is restricted to northeastern Brazil (Geurgas et al. 2008). Gonatodes ocellatus.—This species was recorded based on individuals from Isla Margarita (Roze 1964b). Later, Rivero-Blanco (1979) examined these specimens and suggested that they belong to an undescribed species. RiveroBlanco (1979) restricted the distribution of G. ocellatus to the island of Tobago. Gymnophthalmus lineatus.—This species has been reported from Tucacas, Falcón State by Mijares-Urrutia and Arends (2000). Their record is based on a single specimen (Museo de Ciencias Naturales de Caracas 195), and the authors mentioned that its identification needs to be confirmed. Although we could not examine that specimen, we consider very likely that it represents a misidentified specimen of G. speciosus, which is present at the same locality (a possibility also suggested by Mijares-Urrutia & Arends 2000). Gymnophthalmus lineatus is thus not included in our list at this stage, even if, as mentioned below, the species could eventually be found in the Las Aves Archipelago. Micrurus lemniscatus helleri.—This subspecies has often been mentioned from Venezuela and its presence was accepted by Roze (1967, 1983). The southern Amazonas population was assigned to this subspecies, but no voucher is provided (Roze 1987). Cunha and Nascimento (1993) pointed out that meristic characters of M. l. helleri REPTILES OF VENEZUELA

are similar to those of M. l. lemniscatus, and thus that helleri should be considered a synonym of lemniscatus. Barrio-Amorós and Calcaño (2003) followed this statement, assigning the helleri population of Amazonas and a new one from Barinas to M. l. lemniscatus. Oxyrhopus trigeminus.—The species was recorded from Venezuela (Lancini 1979), but Zaher and Caramaschi (1992) re-identified the original photographs published by Lancini as O. aff. melanogenys. According to MacCulloch et al. (2009) the occurrence of O. trigeminus north to the Amazon River, and thus in Venezuela, remains highly doubtful and we follow these authors recommendation. Pliocercus psychoides.—This species was described from the border between Brazil and Venezuela with an unknown date of collection and collector (Smith & Chiszar 1996). In the original description, the authors stated that the species is probably a mimic of Micrurus p. psyches, known from southern Venezuela and the Guianas. For this reason, they concluded that the only known specimen (holotype AMNH 4433), probably came from southern Venezuela. In a subsequent publication Smith and Chiszar (2001) re-identified the same specimen as P. elapoides aequalis with a probable origin from western central Guatemala. Similarly, in the Reptile Database (Uetz 2011), P. psychoides is considered a synonym of P. elapoides according to a personal communication of V. Wallach to the website. Pliocercus elapoides inhabits southeastern México, Belize, Guatemala, Honduras and El Salvador. Curiously, in the catalogue of AMNH, the specimen catalogued 4433 remains identified as Pliocercus euryzonus, a bicolour species that ranges from Nicaragua to western Ecuador (currently placed in the genus Urotheca) and also includes the problematic locality data “Venezuela, Brazil”. Given the lack of verifiable information, we do not consider that any species or subspecies of the genus Pliocercus occurs in Venezuela. Taeniophallus brevirostris.—This species was mentioned from Venezuela as Rhadinaea brevirostris based on two specimens collected in the Sierra de Perijá, Zulia State (Alemán 1953). Dixon and Michaud (1992) correctly assigned one specimen (MHNLS 399) to Liophis melanotus lamari (see Rivas 2001c), while the second (MHNLS 1298) also proved to be L. m. lamari; see our comments below about species of possible occurrence in Venezuela.

Taxonomic, nomenclatural and distributional comments Taxa that most deserve nomenclatural/taxonomic/distributional comments are listed below and are arranged alphabetically.

Order TESTUDINES Batrachemys/Mesoclemmys spp.—Bour and Zaher (2005) synonymised the genus Batrachemys as well as two recently described genera Ranacephala and Bufocephala with the genus Mesoclemmys. We follow the proposal of McCord et al. (2001) in considering these four genera distinct. Mesoclemmys thus becomes monotypic (M. gibba). Chelus fimbriatus.—The presence of that species in the Maracaibo Lake Basin has long been controversial. The first mention of C. fimbriatus from the area is probably that in Pritchard and Trebbau (1984), who based the record on a personal communication and did not provide any voucher specimen. Rueda-Almonacid et al. (2007) “confirmed” the presence of an isolated population of this species in the Maracaibo Lake Basin—again without any voucher support—a report further mentioned by Pritchard (2008). It must be highlighted that the mention by Rueda-Almonacid et al. (2007) is also based on a personal communication. During the years 2004–2005, TB, GR and CBA conducted fieldwork in the area in search of C. fimbriatus. More than 50 localities were sampled and many interviews with local people were conducted. Although a few local people stated that the species occurs in the area, being very rarely seen, it was confirmed that no voucher specimen has ever been collected. It is possible that an isolated population inhabits the area, but a collected specimen is needed to confirm this. It should also be noted that Orinocoan turtle species are often kept in farms in the southern Maracaibo Lake area. In case of the discovery of C. fimbriatus specimens in the region, molecular analyses will be needed to determine the exact origin of the population(s) (which could have been imported from the Llanos).

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Rhinoclemmys flammigera.—Gorzula and Señaris (1999) considered the subspecies R. punctularia flammigera (Paolillo 1985) as a western extreme clinal variant or aberration of R. punctularia punctularia. Barrio-Amorós and Narbaiza (2008) elevated the subspecies R. p. flammigera to full species status based on head pattern and biogeography. Herein we recognize R. flammigera as a species. Trachemys callirostris ssp.—Formerly known as subspecies of T. scripta, two forms of the Neotropical slider turtle inhabit Venezuela, T. c. callirostris and T. c. chichiriviche (Seidel 2002). The subspecies have been differentiated using molecular data (Jackson et al. 2008).

Order SQUAMATA Ameiva ameiva.—Considered a polytypic species with 10 subspecies, four of them occurring in Venezuela. The validity of these subspecies was questioned by Vanzolini (1986). Avila-Pires (1995) followed Vanzolini noting that a thorough study of the variation in this species is necessary. Ameiva bifrontata.—Peters and Donoso-Barros (1970) recognized four subspecies of A. bifrontata, two of them from Venezuela, the nominal form on the mainland and A. b. insulana restricted to Los Testigos islands. The main difference presented by Ruthven (1924) to distinguish between the populations was the discontinuity of the brachials and antebrachials in the Los Testigos form. We do not believe that this represents more than a clinal variation and until more extensive and detailed studies of the insular form are made, we prefer to treat it as a synonym of the nominal mainland subspecies. Ameiva provitaae.—This species is a possible synonym of A. bifrontata and, until now, was not included in any taxonomic study. The characters thought to be unique to diagnose this species (such as a red snout, see García-Pérez 1995a) are present in different populations of A. bifrontata (GU, unpubl. data). Additional data are needed to clarify the status of this species.

FIGURE 6. Holotype of Anolis gibbiceps (BMNH 1946.8.13.35, formerly 1846.10.28.5). Photo by Colin McCarthy.

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Anolis species.—The taxonomic status of the genus Anolis sensu lato has been unstable and in constant change. Molecular data (Nicholson 2002) indicate that within Anolis there is a distinct clade (Norops) morphologically characterized by having the autotomy septa lying anterior to anteriorly directed transverse process (Etheridge 1960). However, the other clade (Anolis) is not monophyletic and, although the name Norops is recognized by several authors, we refrain from doing so as this would render the remaining anoles a paraphyletic group. The monophyletic status of Anolis (including Norops) has been recently reassessed based on osteology, internal anatomy, chromosomes, molecular data, allozymes, and immunology (Poe 2004). Anolis biporcatus.—The species is reported for the first time from the Maracaibo Lake Basin based on MBLUZ 992, a specimen collected in a gallery forest along the Palmar river (Km 75 on the road Maracaibo-Machiques, Zulia state, Venezuela). Previous records in Venezuela are from the Mountain Range of Perijá and the Mountain Range of Mérida (Rivas Fuenmayor & Barrio-Amorós 2003).

FIGURE 7. Adult male of Anolis bonairensis (FMNH 2684) in lateral (upper) and ventral (below) views, from an undetermined locality in Las Aves Archipelago, Venezuela. Photo by Kathleen Kelly.

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Anolis bonairensis.—Meek (1910) reported a specimen (FMNH 2684, Fig. 7) as A. alligator from Las Aves Archipelago. GU examined this specimen, which indeed agrees in most diagnostic characters with A. bonairensis. We acknowledge the presence of this species in Las Aves Archipelago, but stress that it needs to be formally confirmed by additional material. Anolis chrysolepis vs Anolis nitens.—The nomenclatural history of these species has been controversial (see Myers & Donnelly 1997, Myers 2008, Cadle 2009 for further details). We agree with these authors and no longer recognize the name Anolis nitens. D’Angiolella et al. (2011) reviewed the Anolis chrysolepis species group based on morphology and molecular data and elevated all subspecies of A. chrysolepis to species status. The species present in Venezuela is thus A. planiceps (type locality Caracas, Venezuela). Anolis eewi.—Vanzolini and Williams (1970) synonymised this species with A. chrysolepis. Gorzula and Señaris (1999) noted that the species is distinguished from A. chrysolepis by shorter legs. Myers and Donnelly (2008) synonymised the species with A. chrysolepis again. We believe that Anolis eewi is a valid species, but until new data are obtained, A. eewi is not included in the list presented herein. Atractus tamessari.—This recently described species (Kok 2006) was known only from Kaieteur National Park, western central Guyana. In the description of Atractus surucucu from Roraima State in Brazil, Prudente and Passos (2008) cited two specimens of A. tamessari from Venezuela in their list of material examined (MHNLS 15124 from Uei-tepui and MHNLS 5950 from “El Dorado-Santa Elena de Uairen”). Since Prudente and Passos (2008) erroneously reported A. tamessari as having 17 dorsal scales rows (instead of 15) the exact identity of these museum specimens is doubtful. Only one of these individuals was available to us, MHNLS 15124, which is a juvenile female of an Atractus species having 15 dorsal scales rows that fits well the description of A. tamessari. We thus formally report the presence of A. tamessari for the first time in Venezuela and expand its known distribution ca. 130 km SWW. The specimen listed as A. tamessari from “El Dorado-Santa Elena de Uairen” by Prudente and Passos (2008) should be better examined to ascertain its identity. Atractus univittatus/fuliginosus.—The original description of A. fuliginosus was based on a single specimen from “200 milles of Caracas”, Venezuela and the status of that species remains unclear (as well as the precise type locality). Atractus fuliginosus was later recorded from the northeastern part of the country (Cerro Turimiquire), but that population has since been described as A. matthewi (Markezich & Barrio-Amorós 2004). Adding some more confusion, in their description of Atractus eriki, an Andean species, Esqueda et al. (2005a) mentioned a specimen of A. fuliginosus from San Juan de Los Morros, a locality lying in the Central Coastal Range with which they compared their new taxon. Surprisingly, these authors did not compare A. eriki with A. univittatus (even if it is listed in their material examined), despite the fact that both species possess 17-17-17 dorsal scale rows and that the localities provided for the A. univittatus specimens they examined (Guanare and Barinitas) are closer to the Venezuelan Andes than San Juan de Los Morros. It is not clear whether Esqueda and colleagues considered univittatus a synonym of fuliginosus, but it should be noted that if Coluber fuliginosus Hallowell 1845 and Rhabdosoma univittatus Jan 1862 are synonyms, the name fuliginosus would have priority. We unfortunately did not examine the holotype of A. fuliginosus and cannot confirm or infirm this possibility. No additional specimen of A. fuliginosus has been collected in recent years (Roze 1966; Lancini 1979) and a revision of the type material of A. fuliginosus compared to A. univittatus is badly needed to resolve this issue. Bachia flavescens complex.—Dixon (1973) recognized two species in the B. flavescens group, each with two subspecies: B. flavescens flavescens, B. f. schlegeli, B. monodactylus monodactylus and B. m. parkeri. Hoogmoed (1973) suggested that B. monodactylus is a synonym of B. cophias. Dixon (1973) distinguished the subspecies of each species based on “weak” and variable characters. According to Dixon (1973) few characters distinguish B. flavescens from B. monodactylus (B. cophias sensu Hoogmoed 1973) and additional material should attest that the two species are conspecific. Avila-Pires (1995), concluded that B. flavescens is a monotypic, highly variable species. Kizirian and McDiarmid (1998) considered that the variation in what is known as B. flavescens is impressive and that the taxon should be considered as a species complex. We agree with Kizirian and McDiarmid (1998) and consider Bachia flavescens to be a species complex in need of revision.

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Bachia heteropa ssp.—Bachia heteropa is a polytypic species composed of five subspecies, four of which are present in Venezuela. Populations of the genus Bachia in northern Venezuela, Trinidad, Tobago, Grenada, and the Grenadines were referred to six separate taxa (B. h. heteropa, B. h. alleni, B. h. trinitatis, B. h. lineata, B. h. marcelae and B. h. anomala) (Dixon 1973). Burt and Burt (1933) synonymised B. anomala (from El Mene, Falcón State) with B. lineata. Thomas (1965) considered B. trinitatis to be a subspecies of B. alleni. Later, Donoso-Barros (1968) considered B. marcelae to be a subspecies of B. lineata, retaining B. heteropa and B. alleni as separate species. The revision of the genus by Dixon (1973) provided a better understanding of the group. Dixon noted that there seems to be a progressive reduction in size and loss of prefrontal scales as well as toes from east to west. After examining most available specimens, Dixon (1973) considered all populations in northern Venezuela, Trinidad, Tobago, Grenada and Grenadines to be B. heteropa, which was the oldest name available for this group of lizards. Dixon considered the east to west cline an adaptation to more fossorial habits. Intermediate specimens between B. h. heteropa and B. h. trinitatis are known from eastern Venezuela but intermediates between B. h. lineata and B. h. heteropa are unknown although both subspecies have been recorded from nearby areas (B. h. heteropa from la Guaira, Vargas State and B. h. lineata from Distrito Federal, Caracas). More material is definitely needed to clarify the status of these subspecies. We consider that the significant amount of variation among populations of this taxon could indicate a complex of species, but a taxonomic revision of this taxon is out of the scope of this paper and we decide to treat B. heteropa as a monotypic taxon until this situation is resolved. Bothriechis schlegelii.—This species was reported from Ureña (Pifano et al. 1950), a semi-arid area in the Táchira depression, Táchira State. Pifano et al. (1950) provided a photograph of an individual, but no catalogue number or name of a collection housing a voucher specimen. The species has been cited from a Colombian locality nearby (Campbell & Lamar 2004), and despite the absence (loss?) of a voucher specimen we consider this species as belonging to the Venezuelan herpetofauna, stressing that a new voucher should confirm its presence. Bothriechis schlegelii usually inhabits tropical and mountain rainforest. It must be noted that Sandner-Montilla (1965) apparently also mentioned the species from the Amazonas State (as stated in Campbell & Lamar 2004). Campbell and Lamar (2004) considered that these records are likely erroneous and that they must be confirmed. We were unable to obtain a copy of Sandner-Montilla (1965), but we read a copy of the second edition of that booklet (SandnerMontilla 1975), in which there is no mention of B. schlegelii from the Amazonas State. We assume that the previous records were indeed erroneous and were removed by the author in the second edition of his work. The Bothrops asper-colombiensis-isabelae problem.—For many years a great deal of controversy has arisen about the proper name to be given to the Bothrops species inhabiting the lowlands of the Venezuelan Coastal Range. One of these names is B. colombiensis, used by numerous authors for a species present north of the Orinoco River (Lancini 1979). The type locality, “Republic of Colombia, within two hundred miles of Caracas, Venezuela” could be any locality in northern Venezuela instead of Colombia because the closest border with Colombia is located 700 km SW (approximately 450 miles) from Caracas. Sandner-Montilla (1979a) synonymised B. colombiensis with B. lanceolatus, the latter currently considered endemic to the island of Martinique, Lesser Antilles (Campbell & Lamar 2004; Fenwick et al. 2009), which is 437 km off mainland Venezuela. The same author later described B. l. aidae (Sandner-Montilla 1981), based on a specimen from the Colección de ejemplares vivos de la Familia Crotalidae del Serpentario del Instituto Venezolano de Ofidiología, Caracas (Collection of living specimens of the family Crotalidae of the Serpentarium of the Venezuelan Institute of Ophidiology, Caracas) from “Selvas de Guatopo, Estado Miranda”. Sandner-Montilla (1990) also described B. l. nacaritae from “Cariprima, N. de Valencia". “Cariprima” is not found on any available map, but both Estado Miranda and Valencia are in the Coastal Range. Both subspecies were considered synonyms of B. atrox by Campbell and Lamar (2004), although they probably meant B. asper since the Coastal Range is outside the geographic distribution of B. atrox (Campbell & Lamar 2004: 377, map 57), but inside the geographic distribution of B. asper (Campbell & Lamar 2004: 372, map 56). In any case, we consider the populations from Valencia and Guatopo as B. asper rather than B. atrox, except the population from the cloud forest that is B. venezuelensis. Johnson and Dixon (1984) considered B. colombiensis a synonym of B. atrox, but at least four species were involved under what they named B. atrox (as stated by Campbell & Lamar 2004). Sandner-Montilla (1979b) described an additional species from Guanare, in the foothills of the Venezuelan Andes: Bothrops isabelae (also based on living specimens kept in captivity under number “Bi No. 1”). Markezich and Taphorn (1993) in a study on the variation of some populations of Venezuelan Bothrops concluded that B. isabelae is not valid and synonymised

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it with B. atrox. Fenwick et al. (2009) in the first attempt to use both morphological and molecular evidence to study South American pit vipers, included sequence fragments of B. isabelae obtained from Wüster et al. (2002) (no specific locality was provided) in their analyses. In all analyses by Fenwick et al. (2009), B. isabelae falls together with B. atrox, with a seemingly low genetic distance. Nevertheless, Fenwick et al. (2009) included B. isabelae in their list of Bothrops species (therefore apparently recognizing it as a valid taxon). Some authors (e.g. Lancini 1979; Navarrete et al. 2009) still use B. colombiensis to refer to the most common pit vipers that inhabit northern Venezuela without any justification. These authors also consider B. isabelae to be a valid species (see Navarrete et al. 2009). We do not consider B. colombiensis, B. isabelae, B. l. nacaritae, and B. l. aidae as valid taxa until comparative systematic analyses involving both morphological and molecular evidence from specimens from the type localities are conducted. We follow Campbell and Lamar (2004) in the use of B. asper and B. atrox for these populations of northern Venezuela. Colubridae/Dipsadidae.—Two papers on the molecular phylogeny of the Colubroidea appeared recently, both suggesting changes to the current taxonomy of this group of snakes (Hedges et al. 2009; Zaher et al. 2009). These papers support the recognition of the subfamilies Xenodontinae and Dipsadinae within the family Dipsadidae Bonaparte, 1838 (including also the new subfamily Carphophiinae). Other changes suggested by these authors involving genera occurring in Venezuela are the resurrection of the genus Lygophis and the synonymisations of Lystrophis and Waglerophis with Xenodon, and of Erythrolamprus with Liophis. Curcio et al. (2009a) mentioned that the proposal of synonymising Erythrolamprus with Liophis is premature in view of the insufficient evidence provided by Zaher et al. (2009), proposing maintenance of the traditional recognition of both names as valid until new data become available based on a more comprehensive sampling with proper technical supporting evidence (e.g. inclusion of type species). Curcio et al. (2009a) also mentioned a similar problem with the resurrection of Lygophis, pointing out that such a taxonomic change should have been guided by reference to the type species of the genus, e.g. Herpetodryas lineatus Schlegel (=Liophis lineatus), which was not sampled by Zaher et al. (2009). However, we consider the arguments provided by Zaher et al. (2009) for the recognition of Lygophis sufficient for the resurrection of that genus, which is adopted in our list. Although Zaher et al. (2009) regarded Xenoxybelis as a synonym of Philodryas, we are not convinced by their arguments and prefer to wait for more analyses involving a larger number of species of the genus Philodryas. Vidal et al. (2000) also found Xenoxybelis nested within Philodryas. However, as mentioned by these authors, a more detailed phylogenetic analysis of the newly extended genus Philodryas might show the necessity of a partition of some Philodryas and the recovered monophyletic subunits. Other results of Zaher et al. (2009) involve the allocation of most West Indian snakes to the tribe Alsophiini, a group also studied by Hedges et al. (2009) who proposed a new reclassification for these snakes. Hedges et al. (2009) supported some of the changes proposed by Zaher et al. (2009), one of them being the partitioning of Colubridae. According to their proposal, the following subfamilies and genera are now considered as belonging to the family Dipsadidae: Dipsadinae: Atractus, Dipsas, Imantodes, Leptodeira, Ninia, Plesiodipsas, Sibon and Urotheca. Xenodontinae: Thamnodynastes (Tachymenini), Taeniophallus (Echinaterini), Clelia, Oxyrhopus, Phimophis, Pseudoboa, Siphlophis (Pseudoboini), Philodryas (Philodryadini), Hydrodynastes (Hydrodynastini), Hydrops, Helicops, Pseudoeryx (Hydropsini), Liophis, Erythrolamprus, Umbrivaga and Xenodon (Xenodontini). Dipsadidae incertae sedis: Enulius. Other genera currently remain in Colubridae. Cercosaura phelpsorum.—Euspondylus phelpsi was originally described by Lancini (1968) on the basis of a single female supposedly from Cerro Jaua (corrected to Cerro Sarisariñama by Steyermark & Brewer-Carías 1976). In his revision of the genus Prionodactylus, Uzzell (1973) did not reallocate E. phelpsi to Prionodactylus because a double widened row of gular scales—one of the diagnostic characters of the genus Prionodactylus according to Uzzell—was not mentioned in the original description. In 1996, Myers and Donnelly described a new species, E. goeleti from Cerro Yavi, Amazonas State, which according to the authors appears to be closely related to E. phelpsi but differs from it by having a pale vertebral stripe distinctly broadening posteriorly (vs maintaining the same width), greenish white ventral colouration (vs slate gray with black speckling), and distinctly keeled dorsal scales (vs weakly keeled). These authors also corrected the name of the species from E. phelpsi to E. phelpsorum (Myers & Donnelly 1996). Gorzula and Señaris (1999) reallocated the species to the genus Prionodactylus without providing any argument, and proposed to synonymise P. goeleti with P. phelpsorum based on (1) the fact that all the scale

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counts given by Lancini fall within the variation given by Myers and Donnelly (1996), (2) the subjectivity of the difference between the weakly keeled scales of P. phelpsorum and the distinctly keeled scales of P. goeleti, and (3) the likely misinterpretation of the ventral colouration and shape of the middorsal stripe. Soon after, Mijares-Urrutia (2000) reported the holotype of P. phelpsorum lost and concluded that in the absence of a neotype and additional material allowing further comparisons P. goeleti should be removed from the synonymy of P. phelpsorum. Myers and Donnelly (2001) partly agreed with Gorzula and Señaris (1999) mentioning that although neither Euspondylus nor Prionodactylus have enough characters that uniquely define each genus, the allocation of E. phelpsorum in Prionodactylus “has the practical advantage of associating them in the same group with the similar-appearing Andean Prionodactylus vertebralis”. However, Myers and Donnelly (2001) disagreed with Gorzula and Señaris (1999) stating: “In allocating the name Euspondylus goeleti to the synonymy of Prionodactylus phelpsorum, Gorzula and Señaris apparently confused ontogenetic variation with the kind of extensive intraspecific variation that is not correlated with age, sex or geography”. Myers and Donnelly (2001) insisted again on the differences in width between the vertebral stripe present in P. phelpsorum and that of P. goeleti, pointing out, with the support of photographs, that the vertebral stripe of the former species does not widen posteriorly but is equally narrow, as it is not the case in all known specimens of P. goeleti (less pronounced in juveniles). Myers and Donnelly (2001) considered the difference in ventral colouration as a secondary diagnostic character and noted: “But we took Lancini at face value that the throat and venter of the female specimen of phelpsorum were slate gray in life, and assumed that if the ventral ground colour in life had been pale he would have said so. If, however, he had misleadingly extrapolated ventral colour from the preserved specimen, it would be a different matter since female goeleti became light gray in preservative...”. Myers and Donnelly (2001) also noted that the two species belong to geographically remote populations, reinforcing their arguments. In her recent revision of the genera Cercosaura, Pantodactylus, and Prionodactylus, Doan (2003) concluded that P. goeleti shows variation in the size of the gular scales. One of the specimens examined by Doan exhibited a complete double row of gulars, another had no difference in size between the midventral gulars and the ventrolateral gulars, and another one had a double row widening posteriorly only. Doan (2003) also pointed out that this can be seen in the photographs provided by Myers and Donnelly (1996, 2001). Doan (2003) concluded: “I would be hesitant to decide on its generic status except for its obviously close relationship with P. nigroventris... a species that possesses all the diagnostic characters of Prionodactylus. Based on these similarities, I retain this species in Prionodactylus”. Doan (2003) also synonymised the genera Pantodactylus and Prionodactylus with Cercosaura and considered C. goeleti a junior synonym of C. phelpsorum. We examined four museum specimens identified as Cercosaura phelpsorum, three from the type locality of C. goeleti (MHNLS 11135, MHNLS 11141−42, all collected at 2550 m elevation), and one individual from Cerro Huachamacari, a mountain located about 200 km SWW from Cerro Sarisariñama and about 230 km S from Cerro Yavi (MHNLS 12518, collected at 1700 m elevation), and an unidentified Cercosaura from Cerro Autana (Fuentes & Rivas Fuenmayor 2000), a mountain located about 220 km NW from Cerro Sarisariñama and about 180 km SW from Cerro Yavi (MBUCV 7114, collected at 1450 m elevation). We could not find any pertinent morphological difference between these five preserved specimens and the description of C. goeleti. The degree of keeling is variable among them, the middorsal-scale counts varied between 38–42 (variation not related to geographical distribution), the widening of the vertebral stripe is always present and seems variable and related with age, from 3 (one juvenile) to 4–5 scales (four adults) in the inguinal region (as also highlighted by Myers & Donnelly 2001). Unfortunately ventral colour in life is unknown, but it is similar among all preserved specimens, except the juvenile. Because none of those specimens has the vertebral stripe exactly as described by Lancini (1968) in C. phelpsorum, the state of that stripe apparently remains the only obvious morphological difference between C. goeleti and C. phelpsorum. In our opinion, this single character is not sufficient to separate species and might well be considered as interpopulational variation. This is strengthened by the fact that “C. goeleti” has a much wider range than previously thought, and that if the species is present in Cerro Autana and Cerro Huachamacari there is no reason that it would be absent from Cerro Sarisariñama, which lies in the same biogeographic area. Our new distributional data are additional arguments to agree with Gorzula and Señaris (1999) and Doan (2003) concluding that because of the absence of convincing diagnostic characters between C. goeleti and C. phelpsorum the former should be considered a junior synonym of the latter. However, new collection efforts are needed in the Jaua-Sarisariñama area in order to better understand the inter-and intrapopulational variation of C. phelpsorum since the most recent herpetological expedition to Sarisariñama (Barrio-Amorós & Brewer-Carías 2008) has been unsuccessful in finding the species.

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Cnemidophorus lemniscatus ssp.—Clearly a complex of species. Reeder et al. (2002) placed C. l. splendidus and C. arenivagus as sister taxa, suggesting that a specific status of C. l. splendidus deserves re-evaluation. Markezich et al. (1997) previously stated: “...future work may well result in elevating C. l. splendidus and other forms to recognition as species also”. C. J. Cole (pers. comm.) is currently working at the recognition of C. l. splendidus as a full species. Coluber mentovarius ssp.—A species formerly placed in the genus Masticophis, a genus that has been synonymised with Coluber several times (Savage 2002). Recent phylogenetic data corroborate that Masticophis cannot be separated from Coluber, and thus the former must be treated as a junior synonym of the latter (Utiger et al. 2005). Herein, we adopt these taxonomic changes although they have not been universally accepted (Collins & Taggart 2008). There is considerable variation within each of the Venezuelan subspecies, but they still appear to be well differentiated and separated geographically. Our data (unpublished) suggest that C. m. centralis has a same pattern of geographic distribution as other lowland colubrid species, such as Drymarchon caudomaculatus and Thamnodynastes paraguanae, which could be used as models to test biogeographic hypotheses in northern South America. The population of this subspecies in Panama should be re-evaluated, and the populations in Colombia and Venezuela assigned to C. m. centralis might deserve full species status. Crotalus maricelae.—This species was described from an intra-Andean valley in the Cordillera de Mérida (GarcíaPérez 1995b). Crotalus maricelae was diagnosed based on cephalic scutellation, colour pattern and reduced adult size (García-Pérez 1995b, Vanzolini & Calleffo 2002; Campbell & Lamar 2004). The original description is very brief and the species is virtually indistinguishable from C. durissus cumanensis. Esqueda et al. (2001) suggested that C. maricelae is a nomen nudum and that this population does not deserve recognition because all diagnostic characters fall within the variation range of C. d. cumanensis. La Marca and Soriano (2004) considered Crotalus maricelae a valid taxon, using the same characters mentioned in the original description. This species is not included in the list and we follow Campbell and Lamar (2004) in considering this taxon a synonym of Crotalus durissus. Crotalus pifanorum.—The taxonomy of this species has been controversial because of the imprecise original description and the lack of type specimens. Originally described from the Llanos in southern Guárico (SandnerMontilla 1980), C. pifanorum has been considered a subspecies of C. durissus by Campbell and Lamar (2004). In an undergraduate unpublished dissertation, Pérez-Bidó (1992) compared pifanorum and cumanensis (karyotype, protein electrophoresis and microdermatoglyphs) and supported the recognition of C. pifanorum as a valid species. Several years later, the thesis was published (Pérez et al. 1997), but without providing a list of specimens examined and their geographic provenance. Pending additional data we still consider C. pifanorum a subspecies of C. durissus. Crotalus vegrandis.—The taxonomic history of C. vegrandis has been controversial as well. Originally described as a distinct species by Klauber (1941), the same author (Klauber 1956) relegated it to a subspecies of Crotalus durissus. Nevertheless, this arrangement was not followed by all authors, and C. vegrandis is often treated as a valid species (Lancini 1967; Peters & Orejas-Miranda 1970). In their recent revision, Campbell and Lamar (2004) treated C. vegrandis as a subspecies of C. durissus. We consider C. vegrandis a full species mostly based on the following morphological differences: smaller length, peculiar colour pattern composed of gray and/or reddish brown ground colour with large white flecks distributed as irregular rings (vs rhomboidal blotches in C. durissus). Crotalus vegrandis has an isolated distribution in the lowlands of Maturín and Anzoátegui States, northeastern Venezuela. Dipsas copei.—We follow Harvey (2009) in considering that D. copei is not a synonym of Leptognathus incertus (= D. incerta) and that D. incerta could be a valid species related to D. praeornata as first noted by Amaral (1929a, 1929b) and subsequently by Harvey (2009). Dipsas cf. indica.—This species is tentatively included in the Pantepui region based on a specimen reported from La Neblina (1515 m) by McDiarmid and Paolillo (1988) and by McDiarmid and Donnelly (2005). We did not

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examine that individual and its exact identity thus remains unconfirmed. The maximum known elevation reported for D. indica is 1000 m (see Natera-Mumaw & Battiston 2008). Dipsas peruana/D. latifrontalis.—Dipsas latifrontalis was originally described from Aricagua in the Venezuelan Andes (Boulenger 1905) and later recorded from some localities in Ecuador and the Venezuelan Coastal Range (Peters 1960; Esqueda et al. 2001). Harvey and Embert (2009) resurrected the name D. praeornata for the Venezuelan Coastal Range population and relegated D. latifrontalis to a synonym of D. peruana. The status of the Ecuadorian populations assigned to this species is unclear (Cadle 2005). We provide here preliminary data on the specimen CVULA 7883 from San Isidro, Estado Barinas, 1400 m, considered the second known specimen of D. peruana (Fig. 8) from Venezuela (along with the holotype of D. latifrontalis at the BMNH). The specimen has 1515-15 dorsal scales, 192 ventral scales, 109 subcaudal scales, 8 supralabials and 11 infralabials.

FIGURE 8. Juvenile specimen of Dipsas peruana from San Isidro, Barinas (CVULA 7883). Photo: César L. Barrio-Amorós.

Dipsas pratti.—This species was described based on a female from Medellin, Colombia at an elevation of 1700 m (Boulenger 1897). Four more species of Dipsas were later described (Cope 1899; Prado 1940, 1941; Werner 1916) and were found to be juveniles of the same species (the differences reported only reflected ontogenetic changes) and therefore synonymised with D. pratti (Peters 1960). However, the species’ systematics and natural history still remain unknown. The species is currently known only from a few specimens from the Cordillera Central and the Cordillera Oriental in Colombia, where it was recently reported in the departments of Santander and Bolívar (Moreno-Arias et al. 2006). Unfortunately these authors do not provide any information on the morphological variation and natural history. Recently, several specimens assigned to D. pratti were collected on the Venezuelan side of the Sierra de Perijá (Estado Zulia) at 1600 m. A redescription of this species including the Venezuelan specimens is in press (Barros et al. in press). Epicrates maurus.—Considered a subspecies of E. cenchria by some authors (e.g. Lancini & Kornacker 1989; Perez-Santos & Moreno 1988) or full species by others (e.g. Chippaux 1987; Gorzula & Señaris 1999; BarrioAmorós & Díaz de Pascual 2008). Recently, Passos and Fernandes (2008), on the basis of external morphology, osteology and hemipenial characteristics, showed that E. maurus is a distinct species.

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Gekkonidae.—Kluge (1967) considered five subfamilies in that family. Sphaerodactylinae was recently reviewed and elevated to family status (Sphaerodactylidae) containing the New World genera Coleodactylus, Gonatodes, Lepidoblepharis, Pseudogonatodes and Sphaerodactylus, and the Old World genera Saurodactylus, Quedenfeldtia, Aristelliger, Teratoscincus, Pristurus and Euleptes (Gamble et al. 2008a). Gamble et al. (2008b) recovered another monophyletic trans-Atlantic gecko clade (Phyllodactylidae) containing, among others, the genera Phyllodactylus and Thecadactylus. The Venezuelan species of Hemidactylus (native and introduced) remain in the family Gekkonidae. Gonatodes superciliaris.—Barrio-Amorós and Brewer-Carías (2008) described G. superciliaris as a Sarisariñamatepui endemic. The new species is morphologically indistinguishable from G. alexandermendesi (Cole & Kok 2006) and G. superciliaris is likely a junior synonym of this species. However, since preliminary phylogenetic analyses do not clearly corroborate this fact (see Schargel et al. 2010) and in the absence of additional data we still include the species in our list. Gonatodes vittatus ssp.—Roze (1956) described an insular taxon, G. vittatus roquensis, from El Gran Roque, archipiélago de los Roques, Venezuela. One of us (GR) noted that the holotype (MHNLS 1031) is an adult specimen of G. antillensis (MHNLS 1031, erroneously cited as MNHLS 10800). The remaining series of paratypes deposited in MHNLS (1032–1043) and MBUCV (MBUCV 1906–1909) are G. aff. vittatus. Thus, we do not consider roquensis as a valid subspecies, but wish to stress that these insular populations should be examined using molecular data to confirm their taxonomic identity. Helicops hogei.—Here we recognize the taxon as distinct, but a reassessment of the species is suggested in order to support its validity (see Rossman 2002). Hydrops martii.—We report for the first time H. martii in Venezuela from the conjunction of the Casiquiare arm and the Siapa river in southern Amazonas State, based on photographs taken by CBA in 1998 during a fluvial expedition in search of turtles (Fig. 9) (Barrio-Amorós & Narbaiza 2008). The specimen was not collected due to the lack of permits at that time. Based on the available photographs, the specimen has 173 ventrals, and the typical pattern of the species, consisting of black rings of 1–2 scales wide surrounded by an incomplete one scale wide ring and separated from the other by a reddish ring of 2–3 scales wide. The photographs agree well with other published illustrations of the species (Bartlett & Bartlett 2003; Campbell & Lamar 2004). We consider the pictures, kept in the collection of Fundación Andigena (FA 184–85), as photographic vouchers. Lampropeltis triangulum andesiana.—The occurrence of Lampropeltis triangulum in Venezuela was reported by Roze (1966) and subsequent workers (Lancini 1979; Kornacker 1999) as the subspecies L. t. micropholis. Barrio and Navarrete (1999) clarified the subspecies identity of the Venezuelan specimens and considered them to belong to L. t. andesiana. Leptophis ahaetulla ssp.—Three subspecies of L. ahaetulla have been recorded from Venezuela: L. a. ahaetulla, L. a. coeruleodorsus, and L. a. occidentalis (Lancini 1979; Albuquerque 2009). Leptophis a. coeruleodorsus has a wide distribution throughout the lowlands of Venezuela, east of the Andes, north and south of the Orinoco River and is also present on Isla Margarita. Leptophis a. ahaetulla in Venezuela was formerly considered to represent a distinct subspecies, L. a. copei. Both taxa were shown to be identical by Albuquerque (2009), and the name ahaetulla has precedence over copei. In Venezuela Leptophis a. ahaetulla is known from few specimens collected in the southern part of the Amazonas State. Leptophis a. occidentalis is present in north and northwestern Venezuela west of the Andes. This last form is the most morphologically differentiated among the subspecies found in Venezuela. The taxonomic status of this subspecies is still poorly understood and some populations have enough unique features to deserve specific status. N. R. de Albuquerque (pers. comm.) is currently working on a systematic revision of Leptophis.

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FIGURE 9. The only known specimen of Hydrops martii from Venezuela found dead on a beach at the conjunction of the Casiquiare arm and Siapa river in southern Amazonas State in January 1998. Photo: César L. Barrio-Amorós.

Liophis miliaris.—A specimen of L. miliaris from the Amazonas State (Fuentes 2003) is herein re-identified as Liophis cf. zweifeli (see below) because, in addition to several colour pattern characteristics, it possesses a single apical pit in some dorsal scales and presents a low number of ventral scales (134). None of these characters has ever been found in any L. miliaris population (see account for L. zweifeli). Liophis miliaris is known in Venezuela from Bolívar State (Kornacker et al. 2002; Myers & Donnelly 2008). Liophis zweifeli.—We consider this species distinct from the L. reginae complex. Liophis zweifeli differs from L. reginae (meristic and colour characters taken from Dixon 1983a) in having a salt-and-pepper dorsal pattern (vs dorsum with dense pale and dark paravertebral flecking in L. reginae) and in having a higher number of subcaudal scales (69−88 vs 55−78 in L. r. reginae). Both species are forest snakes. Liophis zweifeli is found mainly throughout the Cordillera de Mérida and the Coastal Ranges in Venezuela, although some specimens of L. cf. zweifeli are known from localities in southern Venezuela. That species has been previously confused with individuals of L. miliaris from Bolívar, south of the Orinoco River (Kornacker et al. 2002). Dixon (1983a) recorded a specimen of L. zweifeli (as L. reginae zweifeli) from Cerro Yapacana in central Amazonas, Venezuela. We examined an additional specimen from the same locality (EBRG 1156, cited in Fuentes 2003), which fits generally well the description of L. zweifeli, except that it has fewer subcaudal scales, although the very tip of the tail is missing (47 vs 78–86 fide Roze [1966] and 69–88 fide Dixon [1983a]). A re-evaluation of this taxon and a revision of the L. reginae species complex are needed to properly determine the status of this population in the state of Amazonas. Oxyrhopus formosus/occipitalis.—We follow MacCulloch et al. (2009) and consider the specimens reported as O. formosus from Venezuela to be O. occipitalis. Oxyrhopus melanogenys (Fig. 10).—Lynch (2009) described Oxyrhopus vanidicus from Colombia. According to the original description, the new species only slightly differs in colour pattern from the very similar O. melanoge-

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nys. Lynch (2009) stated that some populations formerly assigned to O. melanogenys and O. aff. melanogenys from northern South America are Oxyrhopus vanidicus. We examined four specimens (EBRG 1536, 2069, MHNLS 11967, 11998) from Amazonas and Bolívar States in southern Venezuela, which fall within the variation reported for O. vanidicus. However, we could not find consistent differences between O. vanidicus and O. melanogenys. Pending additional evidence we consider all the specimens previously referred to O. aff. melanogenys and O. trigeminus in Venezuela as O. melanogenys (see also comments under Oxyrhopus trigeminus, and MacCulloch et al. 2009).

FIGURE 10. Uncollected specimen of Oxyrhopus melanogenys photographed at night at Tobogán de la Selva near Puerto Ayacucho, Amazonas State. Photo: César L. Barrio-Amorós.

Proctoporus cephalolineatus.—García-Pérez and Yustiz (1995) described Proctoporus cephalolineatus, providing a very brief and incomplete description, and suggested that it is closely related to P. achlyens and P. shrevei (P. luctuosus group). Doan and Schargel (2003) doubted that this species belongs to the P. luctuosus group and even rejected the fact that it belongs to Proctoporus because of the presence of prefrontal scales, but that it rather could belong to either Euspondylus or Pholidobolus. Including this species in the probably paraphyletic genus Euspondylus would bring more confusion. Pholidobolus is distributed far from the Venezuelan territory and is still poorly understood (Hillis 1985). The single specimen of P. cephalolineatus could be an aberrant specimen of the geographically close P. inanis, a species from the Andes of Mérida recently described and now in the genus Riama (Doan & Schargel 2003; Doan & Castoe 2005), in which case the name P. cephalolineatus would have priority. Rena dimidiata.—This species very likely occurs in Venezuela, but is not included in our list because no voucher specimen exists. The type locality of R. dimidiata has been restricted to an area within Roraima State, Brazil (close to the border with Venezuela) by Peters & Orejas-Miranda (1970). The first mention of the species in Venezuela was apparently by Lancini (1979), who based the record on Orejas-Miranda (1966, 1969) and cited the species from the country with no voucher mentioned. Based on this report the species was later reported from Venezuela by several authors (e.g. Lancini & Kornacker 1989; La Marca 1997; Péfaur & Rivero 2000). Orejas-Miranda (1966, 1969), however, did not mention R. dimidiata from Venezuela, but in 1970, Orejas-Miranda in Peters and OrejasMiranda (1970) provided the following distribution for the species: “Guianas, northern Brazil and southeastern

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Venezuela”. Hoogmoed (1977) stated: “According to Orejas-Miranda (1967) the species occurs in a restricted zone in the border area of Brazil, Guyana and Venezuela, coinciding with the Rupununi savanna and its extensions into Brazil and Venezuela. In 1970 the area is given as being slightly more extensive by the same author [=OrejasMiranda]". Hoogmoed did not explicitly report the species from Venezuela but tried to describe the general area south of Roraima given by Orejas-Miranda and also reported Orejas-Miranda´s 1970 extension of the distribution of R. dimidiata into Venezuela. Interestingly, there is no mention of R. dimidiata from Venezuela in that paper either. A revision of the family Leptotyphlopidae in Venezuela is needed, particularly to elucidate the taxonomic status of the Epictia goudotti-albifrons complex, which includes the population from Isla Margarita, currently placed in the synonymy of E. goudotti. Tantilla melanocephala.—The species is reported from the Pantepui region based on a specimen identified by GR (MHNLS 12339) collected in Cerro Jaua, between 1600-1800 m elevation. Thamnodynastes species.—Ten species of this genus are currently recognized in Venezuela. Although the taxonomy of this group has been largely confusing, resulting in occasional confrontations (e.g. Gorzula & Ayarzagüena 1996 vs Myers & Donnelly 1996) some recent contributions provide a better understanding of that difficult genus (e.g. Bailey & Thomas 2006). Tretioscincus bifasciatus.—Formerly divided into two subspecies, T. b. bifasciatus from northern Caribbean Colombia and T. b. kugleri from Venezuela and Aruba. The main difference between the subspecies appears to be the colour pattern. However, the species seems to show considerable individual variation in the intensity of dorsolateral stripes and ventral spotting. Marcuzzi (1950a) examined specimens from Bobare, Lara, and El Pilar, Sucre and questioned the status of the subspecies T. b. kugleri, arguing that colour variation in a species, which appears to be highly variable in this case, is not recommended in the recognition of subspecies. We agree with and follow Marcuzzi in not recognizing T. bifasciatus kugleri as a valid taxon.

Species of possible occurrence A considerable number of reptile species are known from non-Venezuelan localities close to the Venezuelan borders, sometimes within a same biogeographical area and they eventually could be found in Venezuela. In the Serrania de Perijá, the northernmost spur of the Andes between Colombia and Venezuela, the lizards Bachia talpa, and Leposoma rugiceps, as well as the snakes Coniophanes fissidens, Scaphiodontophis venustissimus and Urotheca lateristriga are very likely to be found on the Venezuelan slopes of the mountain range. Most of these species have been collected in Colombian localities geographically close to Venezuela. Numerous colubroid snakes like Atractus variegatus, Dendrophidion bivittatus, Geophis brachycephalus and Helicops danieli as well as the viperid Bothrocophias microphthalmus are likely to be found in the western parts of the state of Táchira, which are a prolongation of the Cordillera Oriental of Colombia. Another snake, the elapid Micrurus sangilensis has been found very close to the Venezuelan border along with the tropidurid lizards Stenocercus santander and S. trachycephalus. The southernmost part of the Venezuelan Guayana is perhaps one of the least studied areas of the country. It is likely that one or more of the following microteiids are present in the country: Gymnophthalmus leucomystax, G. underwoodi, and/or G. vanzoi. Other lizards almost certainly found in this area of Venezuela are Iphisa elegans, Kentropyx borckiana, and Anolis trachyderma. The snakes Atractus surucucu, Dipsas pakaraima and Micrurus pacaraimae also probably occur in the Venezuelan Guayana. The Venezuelan Amazonia, has been even less studied than the Venezuelan Guayana and numerous unrecorded species are suspected to occur there. Anolis transversalis has been mentioned in previous lists (Donoso-Barros 1968; Péfaur & Rivero 2000) with no supporting data (see above), but might be found in this area. Other lizard species likely to be found in the area are Cercosaura argulus, Kentropyx pelviceps, Leposoma guianense, Mabuya bistriata, and Uracentron flaviceps. Several snakes are also very likely to be found in this ecoregion, these include: Atractus poeppigi, Mastigodryas moratoi, Pseudoboa martinsi, and Umbrivaga pyburni as well as the elapid snakes Leptomicrurus scutiventris, L. renjifoi, Micrurus filiformis, M. langsdorfii, the viperid Bothrocophias hyo-

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prora, and the turtle Rhinemmys rufipes (see an unconfirmed record by Barrio-Amorós & Narbaiza 2008). One of us (CBA) saw a specimen of Dracaena guianensis swimming in the intersection of the rivers Siapa and Casiquiare in January 1998; the individual was not caught nor photographed, so the presence of this species needs confirmation. Last but not least, the numerous islands off the Venezuelan Caribbean coast might also harbour some species found on the neighbouring islands of Curaçao or Bonaire. The Gymnophthalmid lizard Gymnophthalmus lineatus could eventually be found in the Las Aves Archipelago, and the aquatic snake, Pelamis platurus could be found along the northwestern Venezuelan coast, mainly off the Península de La Guajira (Hernández-Camacho et al. 2005).

Acknowledgements We are grateful to all the people who helped in the preparation of this work. We are especially indebted to the following curators and personnel from museums who granted access to specimens under their care: F. Bisbal, J. Sanchez, E. Camargo, J. R. Rivero (EBRG), E. N. Smith, C. J. Franklin (UTA), A. Miralles, I. Ineich (MNHN), C. W. Myers, D. Kizirian, R. Pascocello (AMNH), C. McCarthy (BMNH), A. Resetar, K. Kelly (FMNH), M. Salazar, C. Ferreira (MBUCV) and C. Señaris (MHNLS). We thank C. J. Cole (AMNH), C. R. Townsend (AMNH), R. P. Reynolds (USNM), R. D. MacCulloch, A. Lathrop (ROM), and D. Calcaño for sharing unpublished data. The discovery of Hemidactylus mabouia on Los Roques archipelago, as well as on Orchila island, would not have been possible without the invitation of the Laboratorio de protección y manejo de cuencas (IVIC), as part of the development of the project: Recuperación de áreas del Caribe y/o de países miembros del ALBA, under the direction of L. E. Sánchez, to whom we are very grateful. J. Larreal (IVIC) made the map in Figure 1. Finally, W. McCord, R. C. Jadin, K. L. Krysko, W. E. Schargel and M. S. Hoogmoed made valuable suggestions improving a previous version of this paper. This work was supported in part by Instituto Bioclon (Funds to Eric Smith).

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Wilson, L.D. (1980) Systematics of the melanocephala group of the colubrid snake genus Tantilla. San Diego Society of Natural History, memoir, 11, 1–57. Wüster, W., Yrausquin, J.L. & Mijares-Urrutia, A. (2001) A new species of indigo snake from north-western Venezuela (Serpentes: Colubridae: Drymarchon). Herpetological Journal, 11, 157–165. Wüster W., da Graca Salomão, M., Quijada-Mascareñas, J.A, Thorpe R.S. & BBBSP. (2002) Origins and evolution of the South American pitviper fauna: evidence from mitochondrial DNA sequence analysis. In: Schuett G.W., Höggren, M., Douglas M.E. & Greene H.W. (Eds.). Biology of the Vipers. Eagle Mountain Publishing, Salt Lake City, pp. 111–128. Zaher, H. & Caramaschi, U. (1992) Sur le statut taxonomique d'Oxyrhopus trigeminus et O. guibei. Bulletin du Muséum national d´Histoire naturelle, Paris, 4e série, 14, 805−827. Zaher, H. & Caramaschi, U. (2000) Synonymization of Oxyrhopus venezuelanus Shreve, 1947, with Oxyrhopus doliatus Duméril, Bibron & Duméril, 1854, and revalidation of Oxyrhopus erdisii (Barbour, 1913) (Serpentes, Colubridae). Dumerilia, 4(2), 113−122. Zaher, H. & Prudente, A.L.C. (1999) Intraspecific variation of the hemipenis in Siphlophis and Tripanurgos. Journal of Herpetology, 33, 698−702. Zaher, H., Grazziotin, F.G., Cadle, J.E., Murphy, R.W., Moura-Leite, J.C. & Bonatto, S.L. (2009) Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South America xenodontines: a revised classification and description of new taxa. Papéis Avulsos de Zoologia, 49, 115–153. Note added in proof: while our manuscript was in press Townsend et al. (2011) resurrected the family Dactyloidae Fitzinger to include the lizards of the genus Anolis sensu lato that were formerly placed in the family Polychrotidae. Gamble (2011) described the genus Chatogekko to include Coleodactylus amazonicus, and Esqueda (2011) described a new Atractus species from northwestern Venezuela (bioregion 6). Finally, Ugueto and Harvey (2011) published a taxonomic revision of Ameiva ameiva in Venezuela where the authors concluded that A. ameiva (sensu lato) in Venezuela comprises at least four evolutionary species and elevated A. ameiva (sensu stricto), A. atrigularis, and A. praesignis to species. Populations from the savannahs of southern Monagas represent a new species that the authors described as A. pantherina. Thus, A. ameiva in Venezuela is distributed through bioregions 10, 12, 13 and narrowly in southern Anzoátegui (here considered part of bioregion 11); A. atrigularis is found through bioregions 8, 9 and 2 (Isla de Margarita); A. praesignis ranges through 3, 4, 6, 7 and 11; the latter bioregion is also the one inhabited by the endemic A. pantherina. According to these new arrangements and species descriptions, the number of reptiles in Venezuela increased to 374 species in 123 genera. Esqueda, L.F. (2011) A new semifossorial snake species (Dipsadidae: Atractus Wagler, 1828) from the Lara-Falcón montainous system, northwestern Venezuela. Herpetotropicos, 6, 35–41. Gamble, T., Daza, J.D., Colli, G.R., Vitt, L.J. & Bauer, A.M. (2011) A new genus of miniaturized and pug-nosed gecko from South America (Sphaerodactylidae: Gekkota). Zoological Journal of the Linnean Society, 163, 1244–1266. Townsend, T.M., Mulcahy, D.G., Noonan, B.P., Sites Jr., J.W., Kuczynski, C.A., Wiens, J.J. & Reeder, T.W. (2011) Phylogeny of iguanian lizards inferred from 29 nuclear loci, and a comparison of concatenated and species-tree approaches for an ancient, rapid radiation. Molecular Phylogenetics and Evolution, 61, 363–380. Ugueto, G.N. & Harvey, M.B. (2012 dated 2011) Revision of Ameiva ameiva Linnaeus (Squamata: Teiidae) in Venezuela: recognition of four species and status of introduced populations in southern Florida, USA. Herpetological Monographs, 25, 113–170.

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