ZOOSYSTEMA • 2005 • 27 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com
825
Taxonomy of the genus Mabuya
(Reptilia, Squamata, Scincidae) in Venezuela
Aurélien MIRALLES
Muséum national d’Histoire naturelle,
Département Écologie et Gestion de la Biodiversité,
FRE 2696 – Adaptation et Évolution des Systèmes ostéomusculaires,
case postale 55, 55 rue Buffon, F-75231 Paris cedex 05 (France)
amiral@mnhn.fr
Gilson RIVAS FUENMAYOR
Museo de Historia Natural La Salle,
Apartado postal 1930, 1010-A, Caracas (Venezuela)
anolis30@hotmail.com
César L. BARRIO-AMORÓS
Fundación AndígenA,
Apartado postal 210, 5101-A, Mérida (Venezuela)
cesarlba@yahoo.com
Miralles A., Rivas Fuenmayor G. & Barrio-Amorós C. L. 2005. — Taxonomy of the genus
Mabuya (Reptilia, Squamata, Scincidae) in Venezuela. Zoosystema 27 (4) : 825-837.
ABSTRACT
Venezuela is a key location in terms of the distribution of the genus Mabuya
Fitzinger, 1826, given both its intermediate position between the Amazonian
region, Middle America and the Caribbean Islands, and its remarkable
diversity of bioregions. Based on material from different international major
collections, a synthesis of the knowledge on Mabuya species from Venezuela
and the neighbouring islands of Margarita, Trinidad and Tobago is given.
New biogeographical data are also added, extending considerably the
distribution of two species, M. falconensis Mijares-Urrutia & Arends, 1997
and M. nigropunctata (Spix, 1825). An identification key and drawings of
Venezuelan species are given for the first time. This study constitutes a
preliminary step for a revision of this genus which has been controversial and
confusing both taxonomically and nomenclaturally. Moreover, it reveals the
existence of many different populations in Venezuelan Andes and the coastal
range mountains which cannot be at present identified to any known species.
KEY WORDS
Reptilia,
Squamata,
Scincidae,
Mabuya,
Neotropical species,
Trinidad,
Venezuela,
Andes.

INTRODUCTION
For a long time the definition of the genus
Mabuya Fitzinger, 1826 included a large number
of species occurring in tropical areas of Africa,
Asia and the New World. More than 110 species
were placed in Mabuya according to its former
definition (Greer & Broadley 2000). However,
this genus has been recently divided into four dif-
ferent genera (Mausfeld et al. 2002): 1) Chinonia
Gray, 1845 (Cape Verdian clade); 2) Euprepis
Wagler, 1830 (Afromalagasy clade), later repla-
ced by Bauer (2003) for Trachylepis Fitzinger,
1843, the oldest valid designation for Euprepis
placing this genus in the synonymy of
Mabuya s.s.; 3) Eutropis Fitzinger, 1843 (Asian
clade); and 4) the name Mabuya being reserved to
the American clade. Because all Neotropical spe-
cies of Mabuya, with the exception of M. atlanti-
ca from Fernando de Noronha Island, Brazil,
appear to form a monophyletic group (Greer et
al. 1999; Mausfeld et al. 2002; Carranza &
Arnold 2003), the genus Mabuya will be consid-
ered in the present paper as an exclusively neotro-
pical genus, encompassing Central America,
South America, and Caribbean Is lands, as
suggested by Mausfeld et al. (2002).
Systematics of this American genus is proble-
matic. Mainly because no general revision of
this group has been undertaken since Dunn
(1936) and the more recent contributions are
mostly limited to descriptions of new taxa
(Mijares-Urrutia & Arends 1997; Rodrigues
2000) or restricted to the Brazilian region
(Ávila-Pires 1995; Rodrigues 2000; Pinto &
Ávila-Pires 2004). Additionally, Mabuya is see-
mingly a morphologically conservative genus,
and because it lacks a comprehensive taxonomic
study there is a poor understanding of the varia-
tion of morphological characters used for dia-
gnos ing spec ies . Consequent ly , spec ies
boundaries in this genus have remained contro-
versial for more than a century. The situation is
complicated by the fact that the types of
M. mabouya, M. nigropunctata and M. unimar-
ginata have been lost.
In this paper we review the taxonomic status of
species of Mabuya in Venezuela as well as the
Miralles A. et al.
826
ZOOSYSTEMA • 2005 • 27 (4)
RÉSUMÉ
Taxinomie du genre Mabuya (Reptilia, Squamata, Scincidae) au Venezuela.
Le Venezuela est une zone clé en terme de répartition pour le genre Mabuya
Fitzinger, 1826, tant pour sa position intermédiaire entre la région
amazonienne, l’Amérique centrale et les îles caraïbes, que pour sa remarquable
diversité de biorégions. Suite à l’examen de spécimens provenant de
différentes collections internationales de référence, une synthèse des
connaissances portant sur les espèces de Mabuya du Venezuela et des îles
voisines de Margarita, Trinidad et Tobago, est proposée. De nouvelles
données biogéographiques sont éga lement inc luses, élargissant
considérablement les aires de répartitions de deux espèces, M. falconensis
Mijares-Urrutia & Arends, 1997 et M. nigropunctata (Spix, 1825). Une clé de
détermination des espèces vénézuéliennes ainsi que des illustrations détaillées
sont proposées pour la première fois. Ce travail constitue une approche
préliminaire indispensable au projet de révision de ce genre dont la
systématique est si confuse et controversée, tant d’un point de vue
taxinomique que nomenclatural. De plus, il souligne l’existence de plusieurs
populations présentes dans les Andes vénézuéliennes et la chaîne côtière,
identifiables à aucune espèce déjà décrite.
MOTS CLÉS
Reptilia,
Squamata,
Scincidae,
Mabuya,
espèces néotropicales,
Trinidad,
Venezuela,
Andes.

neighbouring islands of Trinidad and Tobago.
Venezuela is in a key location in terms of the
distribution of Mabuya, given its intermediate
position between the Amazonian region, Middle
America and the Caribbean Islands. Additionally,
Venezuela is remarkable for its diversity of
bioregions which contain different assemblies of
species of Mabuya. We have listed five known
species in Venezuela, three of them being consi-
dered until now as endemic to the country.
Moreover, we had the opportunity to study many
different populations of Mabuya, which appa-
rently does not belong to any of the known
species. Phylogenetic and morphological studies
to establish the taxonomic status of those popula-
tions are in preparation.
MATERIALS AND METHODS
The scale nomenclature used to describe speci-
mens was based on Ávila-Pires (1995).
Diagnoses, based both on the literature and the
examined material, are intended to distinguish
between taxa known from Venezuela and the
Guianan region. Drawings were made with a
stereomicroscope Leica MS5 equipped with a
camera lucida. The specimens, preserved in 70%
ethanol, are housed in the American Museum of
Natural History, New York (AMNH), the
Carnegie Museum, Pittsburgh (CM), the Field
Museum, Chicago (FMNH), the Los Angeles
County Museum (LACM), the Museo de
Historia Natural La Salle, Caracas (MHNLS),
the Muséum national d’Histoire naturelle, Paris
(MNHN), the Museu Paraense Emilio Goeldi,
Belém (MPEG), the Nationaal Natuurhistorisch
Museum Naturalis, Leiden (RMNH), the Sam
Noble Oklahoma Museum of Natural History,
Norman (OMNH), the Colección de Anfibios y
Reptiles del Laboratorio de Biogeografía,
Universidad de Los Andes, Mérida (ULABG)
and the University of Michigan Museum of
Zoology, Ann Arbor (UMMZ). Other cited
acronyms refer to the Australian Museum,
Sydney (AM), the Estación Biológica de Rancho
Grande of Maracay (EBRG), the Museum of
Comparative Zoology, Harvard (MCZ), the
Museu de Zoologia da Universidade de São
Paulo (MZUSP) and the University of Texas,
Arlington (UTA).
RESULTS AND DISCUSSION
STATE OF KNOWLEDGE ON THE NOMENCLATURE
OF SOUTH AMERICAN MABUYA
The nomenclatural history of Mabuya is proble-
matic and controversial because some names
available have been repeatedly applied incorrectly
to different species. In this section we present a
brief summary of the nomenclatural problems of
South American Mabuya.
The binomen Mabuya mabouya, firstly created by
Lacepède (1788) to refer to an Antillean species,
was later used to refer to a complex of species
encompassing a wide range, from Mexico and the
Caribbean islands, to Amazonian region (Dunn
1936; Burger 1952; Peters & Donoso-Barros
1970; Hoogmoed 1973; Schwartz & Henderson
1991; Ramirez-Pinilla et al. 2002). Vanzolini
(1981) considered it very improbable that an
Antillean species, M. mabouya, would be so
widely distributed and present in Brazil. This
supposition was recently reinforced by Miralles
(2005), who designated a neotype for this species
restricted to Southern Lesser Antilles. Rebouças-
Spieker (1981b) described M. ficta for the
Brazilian Amazonia, but this species was later
considered a synonym of M. bistriata (Spix,
1825) by Ávila-Pires (1995), based on the exami-
nation of the lectotype of M. bistriata, designated
by Hoogmoed & Gruber (1983). Consequently,
Ávila-Pires (1995) used for the other Amazonian
species (erroneously called M. bistriata by
Vanzolini [1981]) the name M. nigropunctata
(Spix, 1825) and designated a neotype for this
species. In conclusion, M. nigropunctata is the
correct name for the most widely distributed
species in the northern part of South America.
Mabuya bistriata is recognized as a distinct
species, distributed along large rivers of Brazilian
Amazonia (Ávila-Pires 1995) and French Guiana
(Massary et al. 2001). Rodrigues (2000) does not
Taxinomie des Mabuya vénézueliens (Reptilia, Scincidae)
827
ZOOSYSTEMA • 2005 • 27 (4)

share this point of view, considering that the
lectotype of M. bistriata and the neotype of
M. nigropunctata do not agree with the figures
drawn by Spix in 1825, in their original descrip-
tions. As a consequence, he advocates using the
names M. bistriata and M. ficta (sensu Rebouças-
Spieker [1981a, b]). However, given the lack of
significant details in Spix’s descriptions and
drawings, and the absolute necessity to respect
and stabilize the zoological nomenclature, we
prefer following the definition of M. bistriata and
M. nigropunctata given by Ávila-Pires (1995).
In conclusion, we recognize 17 distinct species of
Mabuya on the South American continent:
M. agilis (Raddi, 1823); M. agmosticha
Rodrigues, 2000; M. arajara Rebouças-Spieker,
1981; M. bistriata (Spix, 1825) (sensu Ávila-Pires
[1995]); M. caissara Rebouças-Spieker, 1974;
M. carvalhoi Rebouças-Spieker & Vanzolini,
1990; M. cochabambae Dunn, 1936; M. croizati
Horton, 1973; M. dorsivittata Cope, 1862;
M. falconensis Mijares-Urrutia & Arends, 1997;
M. frenata (Cope, 1862); M. guaporicola Dunn,
1936; M. heathi Schmidt & Inger, 1951;
M. macrorhyncha Hoge, 1946; M. meridensis
Miralles, Rivas & Schargel, 2005; M. nigropalma-
ta Andersson, 1918 and M. nigropunctata (Spix,
1825) (sensu Ávila-Pires [1995]) (Ávila-Pires
[1995]; Mijares-Urrutia & Arends 1997;
Rodrigues 2000; Mausfeld & Lötters 2001). Five
of them, M. carvalhoi, M. croizati, M. falconensis,
M. meridensis and M. nigropunctata, are known
from Venezuela (Horton 1973; Mijares-Urrutia &
Arends 1997; Molina 1998; Miralles et al. 2005).
SYSTEMATICS OF VENEZUELAN SPECIES
Although Ávila-Pires (1995) had clarified the sta-
tus of Amazonian species of Mabuya, the taxono-
my of northern South American species remains
poorly understood. In Venezuela, Gorzula &
Señaris (1999) adopted the name M. bistriata
(= M. nigropunctata sensu Ávila-Pires [1995]) for
populations formerly referred to M. mabouya.
Our observations indicate that M. bistriata is not
known from Venezuela and that population
referred to this species are either M. nigropunctata
or populations that cannot be assigned with
confidence to any known species. However,
because M. bistriata is likely to occur in southern
Venezuela we have decided to include this species
in the identification key presented at the end of
this paper.
The five known Venezuelan species belonging to
the genus Mabuya can be separated into two very
distinct phenotypic groups. The first one is for-
med by M. carvalhoi and M. croizati which are
presumably sister species united by the following
synapomorphies: frontoparietals fused, snout
acute and a high number of nuchal scales (most
often three to four pairs). The second group for-
med by M. fa lconensis, M. meridensis and
M. nigropunctata, does not seem to form a clade
and share only plesiomorphic characteristics: pai-
red frontoparietals, a more rounded snout and a
single pair of nuchals.
SYSTEMATICS
Family SCINCIDA E Gray, 1825
Genus Mabuya Fitzinger, 1826
Mabuya carvalhoi
Rebouças-Spieker & Vanzolini, 1990
(Fig. 1A, B)
Mabuya sp. nov. – O’Shea 1989: 68.
Mabuya carvalhoi Rebouças-Spieker & Vanzolini,
1990: 377. — Ávila-Pires 1995: 573. — Molina 1998:
149. — Rodrigues 2000: 314.
TYPE MATERIAL. — Holotype: Ilha de Maracá, state of
Roraima, Brazil (MZUSP 66679). Paratypes: same
locality (MZUSP 66671 to 66678, 66680, 69648,
69649, AMNH 137372 [= MZUSP 64958 in the
original description]).
MATERIAL EXAMINED. — Brazil. Roraima state, Ilha
de Maracá, 1 paratype (AMNH 137372).
Venezuela. Amazonas state, Coyowa-Terri, 1 spec.
(MHNLS 14471).
DISTRIBUTION (Fig. 5). — Mabuya carvalhoi has been
reported from the southeastern border of the
Amazonas state in Venezuela (Molina 1998) and from
Roraima state in Brazil (Rebouças-Spieker &
Vanzolini 1990).
DIAGNOSIS. — Mabuya with fused prefrontals and fron-
toparietals, parietal plates in contact with each other
Miralles A. et al.
828
ZOOSYSTEMA • 2005 • 27 (4)

behind the interparietal plate, three to four pairs of
nuchals. Four supraciliaries, the first usually the longest.
Palms and soles light coloured, but scales on these areas
often with a darker margin. Body with six longitudinal
dark stripes; two dorsal stripes running from supraocu-
lars scales to the base of the tail where they fuse, two
dorsolateral stripes and two lateral stripes.
Mabuya croizati Horton, 1973
(Fig. 1C, D)
Mabuya croizati Horton, 1973: 75. — Molina 1998:
149. — Rebouças-Spieker & Vanzolini 1990: 377. —
Rodrigues 2000: 314.
Taxinomie des Mabuya vénézueliens (Reptilia, Scincidae)
829
ZOOSYSTEMA • 2005 • 27 (4)
A C
B D
A-C
D
FIG. 1. — Lateral view of the head and dorsal view of the anterior part of the body of Mabuya carvalhoi Rebouças-Spieker &
Vanzolini, 1990 (AMNH 137372) (A, B) and M. croizati Horton, 1973 (CM 7982) (C, D). Scale bars: 4 mm.

Mabuya nigropalmata – Burt & Burt 1933: 86 (part). —
Dunn 1936: 554 (part).
TYPE MATERIAL. — Holotype: Mt Turimiquire,
Sucre state, Venezuela (AMNH 29314). Paratypes:
Mt Turimiquire (AMNH 29215); Elvecia (MCZ
39735, 39736).
MATERIAL EXAMINED. — Venezuela. Anzoategui state,
Turimiquire massif, summit of Cerro La laguna,
2200 m, 1 spec. (EBRG 3797). — Sucre state,
Turumiquire massif, holotype (AMNH 29314);
Elvecia, Turumiquire massif, 3 specs (CM 7978,
7982, 7988); Turumiquire massif, 10 specs (FMNH
17796, 17797, 17798-428 to 17798-430, 17799-434
to 17799-438).
DISTRIBUTION (Fig. 5). — Mabuya croizati seems to be
endemic to the Turimiquire massif in northeastern
Venezuela. This species is reported for the first time in
the Anzoategui state, in the western part of the
Turimiquire massif (EBRG 3797).
DIAGNOSIS. — Mabuya with paired prefrontals and
fused frontoparietals, parietal plates in contact
with each other behind the interparietal plate, two to
four pairs of nuchals. Five, rarely six, supraciliaries
subequal in size. Palms and soles dark coloured. Body
with six longitudinal dark stripes; two dorsal stripes
running from supranasals or frontonasal, to the base of
the tail where they fuse, two dorsolateral stripes and
two lateral stripes.
REMARKS
Mabuya croizati is geographically separated from
its presumably sister species, M. carvalhoi, by
lowlands and the Orinoco River. Steyermark
(1966) recognized that the flora of the
Turimiquire massif shares many species with the
Venezuelan Guyana. Such correlated observa-
tions suggest that those two bioregions could
have been in contact in the past.
Mabuya falconensis
Mijares-Urrutia & Arends, 1997
(Fig. 2A, B)
Mabuya falconensis Mijares-Urrutia & Arends, 1997:
595. — Molina 1998: 149.
TYPE MATERIAL. — Holotype: Cerro Santa Ana,
Península de Paraguaná, Falcón state, Venezuela
(EBRG 1727). Paratypes: Miranda, Falcón state
(ULABG 2115); Colina, Falcón state (ULABG 2116).
MATERIAL EXAMINED. — Colombia. Guajira state, Rio
Barbacoa Arroyo de Arenas, 1 spec. (UMMZ 54793).
Venezuela. Carabobo state, Lago de Valencia, 2 specs
(MHNLS 5511, 5512); Palma Solar, Morón, 1 spec.
(MHNLS 5852); Bahia de Patanemo, 2 specs
(MHNLS 6087, 6302). — Falcón state, Sierra San
Luis, Curimagua, 1 spec. (MHNLS 9526); Península
de Paraguaná, Reserva Biologíca de Monte Cano,
200 m, 1 spec. (MHNLS 17095); Tucacas, 2 specs
(UMMZ 55927, 55932). — Miranda state,
Higuerote, 1 spec. (MHNLS 16654). — Sucre state,
Chacopata, 1 spec. (MHNLS 9040). — Yaracuy state,
Boquerón, 1 spec. (UMMZ 55924).
DISTRIBUTION (Fig. 6). — Until now, M. falconensis
was considered as an endemic xerophilous species of
the Falcón state, only known from a small area around
its type locality (Mijares-Urrutia & Arends 1997).
During our revision, we have discovered many other
localities for this species. East of the Falcón state, spec-
imens have been found in Yaracuy state, at Boqueron
(UMMZ 55924), in Carabobo state (MHNLS 5511,
5512, 5852, 6087, 6302) and in Sucre state, in the
Península de Araya (MHNLS 9040). West of Falcón
state, a specimen from the northeastern Colombia,
Guajira, has been also clearly identified as belonging to
this species (UMMZ 54793). Those observations
extend markedly its distribution along the northern
coast. The same pattern of distribution is found in
Cnemidophorus arenivagus Markezich, Cole &
Dessauer, 1997, a species endemic to the semiarid
coast of northern South America (Markezich et al.
1997).
DIAGNOSIS. — Mabuya with paired prefrontals and
frontoparietals, parietal plates in contact with each
other behind the interparietal plate, a single pair of
nuchals. Four supraciliaries, the second longest. Palms
and soles light coloured. On each side a wide, dark lat-
eral ill-defined band. Dorsolateral light stripes absent.
REMARKS
Color pattern of M. falconensis can be very
variable in a same population. Some specimens
have the dorsum nearly lacking all dark spots
(MHNLS 9526) whereas others have a high
density of spots aligned in eight narrow longitu-
dinal stripes (MHNLS 17095, UMMZ 55927).
Mabuya falconensis shares superficial characters with
M. mabouya, a geographically neighbouring species
from Southern Lesser Antilles (Miralles 2005): these
two possibly closely related species inhabit xerophi-
lous environment (Mijares-Urrutia & Arends 1997;
Breuil 2002) and have in common an accute snout
and a similar colour pattern. Mabuya falconensis can
be distinguished from M. mabouya in having four
supraocular (versus three) and a smaller size.
Miralles A. et al.
830
ZOOSYSTEMA • 2005 • 27 (4)

Mabuya meridensis
Miralles, Rivas & Schargel, 2005
(Fig. 3A, B)
Mabuya meridensis Miralles, Rivas & Schargel, 2005: 3.
TYPE MATERIAL. — Holotype: Mérida, Mérida state,
Venezuela (ULABG 1570). Paratypes: region of
Mérida (AMNH 13405 to 13408, 13526; ULABG
4153, 4281; UMMZ 57435).
MATERIAL EXAMINED. — Venezuela. Mérida state,
holotype (ULABG 1570); vicinity of Mérida,
8 paratypes (AMNH 13405 to 13408, 13526;
ULABG 4153, 4281; UMMZ 57435); 3 specs
(MHNLS 923, 1393, 17081).
D I STRIBUTION (F ig . 5) . — Mabuya mer iden-
s i s seem s to be endemic to the v ic in i t y of
Mérida, between 1300 m and 2200 m above sea
level.
Taxinomie des Mabuya vénézueliens (Reptilia, Scincidae)
831
ZOOSYSTEMA • 2005 • 27 (4)
A C
B
D
A, B
C, D
FIG. 2. — Lateral view of the head and dorsal view of the anterior part of the body of Mabuya falconensis Mijares-Urrutia & Arends,
1997 (MHNLS 5512) (A, B) and M. nigropunctata (Spix, 1825) (MHNLS 16203) (C, D). Scale bars: 4 mm.

DIAGNOSIS. — Mabuya with paired prefrontals and
frontoparietals, parietal plates in contact with each
other behind the interparietal plate, a single pair of
nuchals. Four supraciliaries, the second longest. Palms
and soles dark coloured. Seven dark band along body:
one very thin vertebral stripe; two dorsolateral stripes
ill-defined, formed by a succession of aligned dots; two
upper lateral and two lower lateral stripes.
Mabuya nigropunctata (Spix, 1825)
(Fig. 2C, D)
Scincus nigropunctatus Spix, 1825: 24.
Euprepis surinamensis Hallowell, 1856: 154.
Mabuia surinamensis – Cope 1862: 186.
Mabuia agilis var. nigropunctata Boulenger, 1887: 192
(part).
Mabuia aurata – Boulenger 1887: 189 (part). —
Goeldi 1902: 535 (part).
Mabuia agilis – Goeldi 1902: 534 (part).
Mabuya agilis – Burt & Burt 1933: 86 (part). — Burt
& Myers 1942: 49 (part).
Mabuya mabouya mabouya – Dunn 1936: 540 (part). —
Amaral 1937a: 203 (part). — Peters &Donoso-Barros
1970: 199 (part). — Hoogmoed 1973: 209 (part).
Mabuya agilis agilis – Amaral 1937b: 1743 (part).
Mabuya mabouya – Hoogmoed 1979 : 278. — Rivero-
Blanco & Dixon 1979: 296.
Mabuya bistriata – Rebouças-Spieker 1981a: 123;
1981b: 162. — Vanzolini 1981: 196. — Gorzula &
Señaris 1999: 147. — Rodrigues 2000: 315. —
Murphy 1997: 150. — Carranza & Arnold 2003: 279
(part).
Mabuya nigropunctata – Ávila-Pires 1995: 584. —
Massary et al. 2001: 19.
TYPE MATERIAL. — Holotype lost, from the original
type locality of “Ecgá”, correct original spelling being
“Ega”, at present Tefé, state of Amazonas, Brazil
(Ávila-Pires 1995). Neotype: Santa Rita, Município de
Maraã, left margin of Rio Japurá (Lago Paricá), state of
Amazonas, Brazil (MPEG 15248) (designated by
Ávila-Pires 1995).
MATERIAL EXAMINED. — Brazil. Acre state, 5 km N
Porto Walter, W of Rio Juruá, 4 specs (OMNH
37048 to 37051). — Amazonas state, Município de
Castanho, 40 km S Manaus at km 12 on road
to Autazes, 18 specs (OMNH 37681 to 37698). —
Maranhão state, Nova Vida, 25 km E of Rio
Gurupi, BR 316, 5 specs (MPEG 10690, 10691,
10693, 10695, 10698). — Para state, CEMEX,
Agropecuaria Treviso LTDA, 101 km S and 18 km E
Santarém, 15 specs (OMNH 36828 to 36842). —
Rondonia state, Rio Formoso, Parque Estadual
Guajará-Mirim, 90 km N Nova Mamoré, 7 specs
(OMNH 37411 to 37417). — Roraima state, 7 km E
Rio Ajaraní, BR 210, 10 specs (OMNH 36313 to
36322).
Colombia. Amazonas state, Leticia, 2 specs
(CM 55601, 55602).
Ecuador. Napo province, San Francisco, Rio Napo
200 m, 1 spec. (UMMZ 84742). — Sucumbios
Miralles A. et al.
832
ZOOSYSTEMA • 2005 • 27 (4)
A
B
FIG. 3. — Lateral view of the head (A) and dorsal view of the
anterior part of the body (B) of holotype of Mabuya meridensis
Miralles, Rivas & Schargel, 2005 (ULABG 1570). Scale bar:
4 mm.

province, Reserva Faunistica Cuyabeno (RPF-
Cuyabeno), 1 spec. (OMNH 36514).
French Guiana. St Eugene, 11 specs (MNHN 1996-
4570 to 1996-4572, 1997-2206 to 1997-2213).
Guyana. Cuyuni-Mazaruni region, Kartabu, on the
Mazaruni-Potaro, 5 specs (AMNH 15120 to 15122,
18183, 21326).
Peru. Loreto department, Rio Yuvineto, right bank of
Rio Putumayo, 2 specs (MNHN 1978-2412, 1978-
2413).
Trinidad and Tobago. Trinidad island, 1 spec.
(AMNH 64528); Trinidad island, San Rafael, 8 specs
(FMNH 49901 to 49908); Trinidad island, 1 spec.
(UMMZ 79919).
Venezuela. Amazonas state, Parima B, 1 spec.
(MHNLS 16389). — Aragua state, Quebrada, right
bank of the Río San Miguel, national park Henri
Pittier, 1 spec. (MHNLS 17080). — Bolívar state,
foot of the Roraima Mountain, La Gran Sabana,
5°10’N, 60°47’W, 2 specs (MHNLS 11544, 11545);
Serranía del Supamo, Cerro Santa Rosa, 1 spec.
(MHNLS 15532). — Delta Amacuro state, Managas,
2 specs (LACM 31469, 31470); Burojoida, 1 spec.
(MHNLS 4543). — Nueva Esparta state, Margarita
Island, Cerro Copey, 2 specs (MHNLS 3401, 3402). —
Miranda state, SW Araira, Hacienda La Ceiba, 1 spec.
(MHNLS 16652); Guatire, 3 specs (MHNLS 16655,
16651, 16658). — Sucre state, Península de Paria, Las
Melenas, 1 spec. (MHNLS 15533); Península de
Paria, Macuro, 1 spec. (MHNLS 16203).
Suriname. Marowijne, 10 km N Wanekreek, 1 spec.
(RMNH 28080). — Airstrip Paloemeu, 1 spec.
(RMNH 15629). — Afobaka, 1 spec. (RMNH
15593). — Awarra savannah, Maratakka river, 1 spec.
(RMNH 16453). — Paramaribo, 1 spec. (RMNH
15633). — Sipaliwini, 1 spec. (RMNH 15648). —
Nickerie, km 117 on road to Amotopo, Kabalebo area,
1 spec. (RMNH 28580); Blanche Marie, 2 specs
(RMNH 16468, 16469).
DISTRIBUTION (Fig. 6). — Mabuya nigropunctata
appears to be by far the most common species of
Mabuya in the Guyano-Amazonian region: this species
is known in Brazil from the Amazonian region, the
Mato Grosso state, a portion of the Atlantic forests
and the cerrado region (Ávila-Pires 1995); in
Amazonian part of Ecuador from the province of
Sucumbios (Vitt & de la Torre 1996) and the
province of Napo (UMMZ 84742); and everywhere in
French Guiana (Massary et al. 2001) and Suriname
(Hoogmoed 1973). We have also identified some
specimens from Colombia (CM 55601, 55602) as
belonging to this species.
DIAGNOSIS. — Mabuya with paired prefrontals and
frontoparietals, parietal plates generally separated by
the interparietal plate, a single pair of nuchals. Four to
six, mostly five, usually subequal supraciliaries. Palms
and soles dark, covered by small tubercules. On each
side a wide, dark lateral band, bordered or not by dor-
sal and ventral light stripes, of which the dorsal one,
when present, usually ill-defined.
REMARKS
In mainland Venezuela, specimens that can
clearly be assigned to M. nigropunctata, following
Ávila-Pires (1995), have been found in six
different states: Amazonas, from Parima B
(MHNLS 16389); Aragua, from the Henri
Pittier national park (MNHLS 17080); Bolívar,
from the foothills of Cerro Santa Rosa, a tepui in
serranía del Supamo (MHNLS 15532), and from
the foothills of Roraima tepui (MHNLS 11544,
11545); Delta Amacuro, from Burojoida
(MHNLS 4543) and Managas (LACM 31469,
31470); Miranda from Araira (MHNLS 16652)
and Guatire (MHNLS 16655) and Sucre, in the
península de Paria, from the path from Las
Melenas to Cerro el Humo (MHNLS 15533)
and from Macuro (MHNLS 16203). All those
localities, except the two situated in the small
coastal plain of Higuerote, Miranda state, and the
one in the Henri Pittier national park, Aragua
state, are in what Barrio-Amorós (1998) called
“arco amazónico oriental” (eastern Amazonian
arch) and what Gorzula & Señaris (1999) called
“Venezuelan Guayana”. We can infer that
M. nigropunctata is present in most of the eastern
part of the country. Moreover, two specimens
Taxinomie des Mabuya vénézueliens (Reptilia, Scincidae)
833
ZOOSYSTEMA • 2005 • 27 (4)
FIG. 4. — Map of Venezuela showing the major bioregions.

from Margarita Island (state of Nueva Esparta),
collected in 1953 by Felipe Martín in Cerro
Copey (MHNLS 3401, 3402), are also M. nigro-
punctata.
Populations have been registered on the islands of
Trinidad and Tobago. They have been consid-
ered by many authors as M. bistriata (Murphy
1997; Carranza & Arnold 2003). Their assigna-
tion to this species appears improbable to us,
given that this species is otherwise restricted to
the Guyano-Amazonian region and is apparently
absent from Venezuela. We studied specimens
from Trinidad island, confirming them to be
M. nigropunctata (AMNH 64528, FMNH
49901 to 49908, UMMZ 79919). By extension,
we consider highly probable that the specimen
from Tobago, Bacolet state (AMNH 73087),
identified by Murphy (1997), was not M. bistriata
but M. nigropunctata.
ANDEAN MABUYA:
STRONG POLYMORPHISM
OR A SPECIES COMPLEX?
MATERIAL EXAMINED. — Mabuya sp. A: Venezuela.
Aragua state, Colonia Tovar (MHNLS 17088, 17090-
17092). — Distristo capital, Caracas, Parque Nacional
El Avila, Sector Mariperez (MHNLS 13262). —
Miranda state, Pico Naiguata (MHNLS 16656,
16657); Sector El Amarillo, San Antonio de Los Altos
(MHNLS 13734, 13735). — Trujillo state, near
Bocono, front of the Laguna Negra (MHNLS 16648,
16649). — Vargas state, Club Casa de Campo Tovar
(MHNLS 17089, 17093).
Mabuya sp. B: Venezuela. Zulia state, Rio Escalante,
SW lake Maracaibo, sector el Cañon (MHNLS
16647); Mara Municipality, Borders of the riecito
Maché, river basin of the Cachirí river, sector La
Orchila, Sierra de Perijá (MHNLS 16648); Finca
Onia, 10 km del Vigia, via San Cristobal, gravid female
(MHNLS 16671); extracted from the uterus of the
female MHNLS 16671 (MHNLS 17035 to 17040).
During our review of Venezuelan Mabuya, we
studied many specimens from all over the conti-
nent. It clearly appeared that more than every-
where else in South America, a very high diversity
of populations occurs collectively in the coastal
range and the Andes of Venezuela (Fig. 5). These
populations all have a single pair of enlarged
nuchals and two separated frontoparietals, but are
readily distinguished between them and from
known species of Mabuya in the cephalic scala-
tion, color pattern or size. For the time being we
prefer to include those populations in an “Andean
species complex”. In addition to the recently des-
cribed M. meridensis, two putative species have
Miralles A. et al.
834
ZOOSYSTEMA • 2005 • 27 (4)
FIG. 5. — Occurence of Mabuya carvalhoi (●), M. croizati (●) and
M. meridensis (■) in Venezuela. Occurences of Andean species
complex have been also added: Mabuya sp. A from the coastal
range and the state of Trujillo (■) and Mabuya sp. B from the
Maracaibo lake bassin (■).
F IG. 6 . — Occurence of Mabuya fa lconens is (● ) and
M. nigropunctata (● ) in Venezuela, and of M. falconensis in
neighbouring Colombia.

835
ZOOSYSTEMA • 2005 • 27 (4)
been identified in the Andean region (Mabuya sp.
A from the coastal range and Trujillo state, and
Mabuya sp. B from the Maracaibo lake basin). A
study based on molecular data and detailed mor-
phological analysis is in preparation to critically
test the validity of these putative species.
CONCLUSIONS
This preliminary work shows that the genus
Mabuya is widely distributed through Venezuela,
with the exception of los Llanos of theOrinoco, dry
lowland open formations (Rivero-Blanco & Dixon
1979) between the Andes and the Guiana shield
(Fig. 4). But above all, it reveals that many Mabuya
from the highlands cannot be identified to any
known species, allowing us to focus now on those
undetermined populations. Indeed, taxonomic stu-
dies on mountainous Mabuya are virtually non-exis-
tent despite their ubiquitous representation in those
highlands by many apparently different species. It
seems highly probable that the complex geography
of those altitudinal regions involved numerous para-
patric speciation phenomena in the genus.
Descriptions of new species and molecular phyloge-
netic analysis in preparation should permit us to cla-
rify this very confuse situation.
Taxinomie des Mabuya vénézueliens (Reptilia, Scincidae)
KEY TO THE SPECIES OF VENEZUELAN SKINKS OF THE GENUS MABUYA
In Venezuela, Mabuya can be easily distinguished from all other lizard genera by the presence of a
translucent disk (or window) in the lower eyelid, two supranasals, and uniform cycloid body scales.
Mabuya of the Andean species complex are included in the following key. Mabuya bistriata is also
included in the key, even if it has not yet been reported in Venezuela. Indeed, given the distribution
and ecology of this species, its presence in the southern part of the country seems possible.
1. A single pair of nuchal scales, two separated frontoparietal scales .......................... 2
— More than a single pair of nuchal scales, frontoparietals fused into a single scale .. 3
2. Presence of a vertebral dark stripe .................................... M. meridensis (Fig. 3A, B)
— Absence of vertebral dark stripes .......................................................................... 4
3. Prefrontals plates fused into a single scale, two dark dorsals stripes start up to the
eyes, four supraciliaries ...................................................... M. carvalhoi (Fig. 1A, B)
— Two separated prefrontals plates, the two dark dorsals stripes start up to the nostrils,
five supraciliaries ................................................................ M. croizati (Fig. 1C, D)
4. Four supraciliaries with second longest, parietals scales always in contact with each
other behind interparietal scale .............................................................................. 5
— Five or six supraciliaries subequal in size, parietal scales generally separated by inter-
parietal scale ............................................................................................................
................................................................................ M. nigropunctata (Fig. 2C, D)
5. Palms and soles light coloured (same colour than belly) ........................................ 6
— Palms and soles dark coloured (darker than belly) ................................................ 7
6. Long snout, ill-defined dark lateral stripe at each side composed of small fused dark
dots and not bordered dorsally by a light stripe, no dorsolateral dark stripes ............
........................................................................................ M. falconensis (Fig. 2A, B)
— Short snout, well defined dark lateral stripes at each side and bordered dorsally by a
light stripe, two short dorsolateral dark stripes ...................................... M. bistriata
7. Two dorsolateral dark stripes, well defined white lateral stripes ................................
........................................................ Mabuya sp. A (coastal range and Trujillo state)
— No dorsolateral dark stripes, ill defined white lateral stripes Mabuya sp. B (Zulia state)

Acknowledgements
We are grateful to Teresa C. S. de Ávila-Pires
(MPEG) for sharing with us her interest on the
nomenclature of this genus, and to Jean-Pierre
Gasc and Annemarie Ohler (MNHN), Allen E.
Greer (AM), César R. Molina (MARN) and
Walter Schargel (UTA) for reading and criticizing
the manuscript. All curators, Linda S. Ford
(AMNH), Stephen P. Rogers (CM), Alan Resetar
(FMNH), James Hanken and José P. Rosado
(MCZ), Ivan Ineich (MNHN), Teresa C. S. de
Ávila-Pires (MPEG), Laurie J. Vitt (OMNH),
Enrique La Marca (ULABG) and Ronald A.
Nussbaum and Gregory E. Schneider (UMMZ)
are also greatly acknowledged. We also thank
Annemarie Ohler for welcoming one of us (A.M.)
at the UMS 602 Taxinomie et Collections
(Reptiles et Amphibiens), Département
Systématique et Évolution (MNHN).
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Taxinomie des Mabuya vénézueliens (Reptilia, Scincidae)
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