Reptiles of Venezuela: an updated and commented checklist

Transcripción

Zootaxa 3211: 1–64 (2012)
www.mapress.com / zootaxa/
Copyright © 2012 · Magnolia Press
ISSN 1175-5326 (print edition)
Monograph
ZOOTAXA
ISSN 1175-5334 (online edition)
ZOOTAXA
3211
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: [email protected]
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
(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
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ISSN 1175-5326
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ISSN 1175-5334
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2 · Zootaxa 3211 © 2012 Magnolia Press
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
Zootaxa 3211 © 2012 Magnolia Press ·
3
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
5
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
dĂǆŽŶŽŵŝĐůŝƐƚŽĨƚŚĞ
sĞŶĞǌƵĞůĂŶƌĞƉƚŝůĞƐ
ϭ
Ϯ
ϯ
ϰ
ϱ
ϲ
ϳ
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
x
x
x
x
x
x
x
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
dƌĂŶƐůŽĐĂƚĞĚ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
CLASS REPTILIA
Order CROCODYLIA
FAMILY
ALLIGATORIDAE
(Cuvier 1807)
Genus Caiman Spix
1825
Caiman crocodilus
Linnaeus 1758
Genus Paleosuchus
Gray 1862
x
x
......continued on the next page
RIVAS ET AL.
TABLE 1. (continued)
ŝŽƌĞŐŝŽŶƐ
Paleosuchus palpebrosus
(Cuvier 1807)
Paleosuchus trigonatus
(Schneider 1801)
FAMILY
CROCODYLIDAE
(Cuvier 1807)
Genus Crocodylus
Laurenti 1768
Crocodylus acutus
(Cuvier 1807)
Crocodylus intermedius
Graves 1819
ϭ
x
Ϯ
ϯ
x
ϰ
ϱ
ϲ
x
ϳ
x
ϴ
x
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
x
x
x
x
x
x
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
x
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
E
x
x
x
x
CE
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)
x
E
x
E
x
E
x
E
x
CE
x
x
x
x
V
x
x
7
ŝŽƌĞŐŝŽŶƐ
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)
ϭ
Ϯ
ϯ
ϰ
ϱ
x
x
x
x
x
ϲ
ϳ
ϴ
ϵ
x
x
x
x
x
x
x
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
x
x
x
x
x
x
x
x
x
x
ϭ
ϯ
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
?
x
x
x
x
x
x
x
x
V
x
x
x
x
x
x
V
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
ϭ
Ϯ
Genus Podocnemis
Wagler 1830
Podocnemis
erythrocephala (Spix
1824)
Podocnemis expansa
(Schweigger 1812)
Podocnemis unifilis
Troschel 1848
Podocnemis vogli Muller
1935
ϯ
ϰ
ϱ
ϲ
ϳ
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
CE
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
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
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
9
ŝŽƌĞŐŝŽŶƐ
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)
ϭ
Ϯ
ϯ
ϰ
ϱ
ϲ
ϳ
ϴ
ϵ
x
x
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
x
x
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
x
x
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
V
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
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
ϭ
ϰ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
V
x
x
x
x
x
Genus Plica Gray 1831
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
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)
ϭ
Ϯ
ϯ
ϰ
ϱ
ϲ
ϳ
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
ϭ
ϰ
x
x
x
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
11
ŝŽƌĞŐŝŽŶƐ
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
ϭ
Ϯ
ϯ
ϰ
ϱ
ϲ
ϳ
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
x
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
CE
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
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
ϭ
Ϯ
ϯ
ϰ
ϱ
ϲ
x
x
x
ϳ
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
x
x
x
x
x
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
V
x
x
x
x
x
x
x
x
x
x
x
x
x
x
V
x
13
ŝŽƌĞŐŝŽŶƐ
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
ϭ
Ϯ
ϯ
ϰ
ϱ
ϲ
ϳ
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
x
ϭ
ϯ
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
x
x
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
V
x
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
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
ϭ
Ϯ
ϯ
ϰ
ϱ
ϲ
ϳ
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
x
x
x
x
x
x
x
x
x
x
x
x
ϭ
ϰ
x
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
15
ŝŽƌĞŐŝŽŶƐ
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)
ϭ
Ϯ
x
ϯ
ϰ
x
ϱ
x
ϲ
x
ϳ
x
ϴ
x
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
x
x
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
V
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
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)
ϭ
Ϯ
ϯ
ϰ
x
x
x
ϱ
ϲ
ϳ
x
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
x
x
x
x
x
x
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
V
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
17
ŝŽƌĞŐŝŽŶƐ
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 1854)
ϭ
Ϯ
ϯ
ϰ
ϱ
ϲ
ϳ
ϴ
x
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
x
x
x
x
x
x
x
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
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 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
ϭ
Ϯ
ϯ
ϰ
ϱ
x
x
x
x
x
ϲ
ϳ
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
19
ŝŽƌĞŐŝŽŶƐ
Stenorrhina degenhardtii
(Jan 1876)
Genus Tantilla Baird
and Girard 1853
Tantilla melanocephala
(Linnaeus 1758)
Tantilla semicincta
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
2005
Atractus mijaresi
2005
Atractus multidentatus
Passos, Rivas and BarrioAmorós 2009
Atractus ochrosetrus
2005
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ϲ
ϳ
ϴ
x
x
x
x
x
x
x
x
x
x
x
x
ϵ
x
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
ϭ
ϰ
x
x
x
x
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
V
x
x
x
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
Atractus pamplonensis
Amaral 1935
Atractus riveroi Roze
1961
Atractus steyermarki
Roze 1958
Atractus tamaensis
2005
Atractus tamessari Kok
2006
Atractus taphorni
Schargel and GarcíaPérez 2002
Atractus torquatus
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 1854)
Genus Enulius Cope
1871
Enulius flavitorques
(Cope 1869)
Genus Erythrolamprus
Wagler 1830
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Ϯ
ϯ
ϰ
ϱ
ϲ
ϳ
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Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
x
x
x
x
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
V
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
?
x
x
x
x
x
x
x
x
x
x
x
21
ŝŽƌĞŐŝŽŶƐ
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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Ϯ
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ϱ
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x
ϲ
ϳ
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ϵ
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Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
x
x
x
x
x
x
x
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
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x
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x
x
x
x
x
x
x
x
x
x
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x
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x
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x
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x
x
x
x
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
ϭ
Ϯ
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ϰ
ϱ
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ϳ
ϴ
ϵ
ϭ
Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
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)
x
x
x
x
x
x
x
x
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
x
x
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
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x
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x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
23
ŝŽƌĞŐŝŽŶƐ
Genus Plesiodipsas
Harvey, Rivas, Caicedo
and Rueda-Almonacid
2008
Plesiodipsas perijanensis
(Alemán 1953)
Genus Pseudoboa
Schneider 1801
Pseudoboa coronata
Schneider 1801
Pseudoboa neuwiedii
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
ϭ
Ϯ
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Ϭ
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
x
x
x
x
x
x
x
x
x
x
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
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x
x
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x
x
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x
x
x
x
x
x
x
x
x
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
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
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 1854)
Micrurus nattereri
(Schmidt 1952)
ϭ
Ϯ
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ϰ
ϱ
ϲ
ϳ
ϴ
ϵ
ϭ
Ϭ
x
x
ϭ ϭ
ϭ Ϯ
ϭ
ϯ
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
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x
25
ŝŽƌĞŐŝŽŶƐ
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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ϭ Ϯ
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ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
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x
x
x
x
x
x
x
x
x
x
x
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x
.......continued on the next page
RIVAS ET AL.
ŝŽƌĞŐŝŽŶƐ
Genus Typhlophis
Fitzinger 1843
Typhlophis squamosus
(Schlegel 1839)
FAMILY
LEPTOTYPHLOPIDA
E Stejneger 1891
Genus Epictia Gray
1845
Epictia albifrons
(Wagler 1824)
Epictia goudotti
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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ϵ
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ϭ Ϯ
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ϯ
ϭ
ϰ
ŶĚĞŵŝĐ
ƐƉĞĐŝĞƐ
/ŶƚƌŽĚƵĐĞĚ
ƐƉĞĐŝĞƐ
ƐƉĞĐŝĞƐ
dŚƌĞĂƚĞŶĞĚ
ƐƉĞĐŝĞƐ
x
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x
x
x
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-
27
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
3
2
2
3
2
2
3
Family
30
3
16
25
20
19
23
Genera
121
6
35
55
60
53
62
Species
366
6
49
67
108
69
80
% of total
of species
100
1.6
13.4
18.3
29.5
18.9
21.9
Endemics
121
0
9
3
31
5
13
% endemics
33.1
0
18.4
4.5
28.7
7.3
16.3
Threatened 22
6
0
3
1
3
1
% threatened
100
0
4.5
0.9
4.4
1.3
6.0
29
TABLE 2 (continued)
Coastal
area
Central Coastal
Range
Eastern Coastal
Range
Orinoco
Delta
Llanos
Guayana
Amazonia
Pantepui
Order
2
3
3
3
3
3
3
1
Family
11
24
20
24
23
26
22
6
Genera
15
69
56
52
56
86
76
20
Species
18
102
81
69
70
146
131
37
% of total of
species
4.9
27.9
22.1
18.9
19.1
39.9
35.8
10.1
Endemics
0
24
13
3
1
9
9
25
% endemics
0
23.5
16.1
4.4
1.4
6.2
6.9
67.6
Threatened
0
1
6
3
4
3
4
0
1.0
7.4
4.4
5.7
2.1
3.1
0
% 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.
31
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.
33
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.
RIVAS ET AL.
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
35
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).
RIVAS ET AL.
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.
37
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.
RIVAS ET AL.
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.
39
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
RIVAS ET AL.
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
41
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.
RIVAS ET AL.
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
43
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.
RIVAS ET AL.
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.
45
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-
RIVAS ET AL.
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
47
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-
RIVAS ET AL.
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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63
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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.
RIVAS ET AL.

Reptiles of Venezuela: an updated and commented checklist

Transcripción

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