Content uploaded by Miralles Aurélien
Author content
All content in this area was uploaded by Miralles Aurélien on Jan 03, 2016
Content may be subject to copyright.
HERPETOLOGICAL JOURNAL, Vol. 16, pp. 1-7 (2006)
A NEW SPECIES OF MABUYA (REPTILIA, SQUAMATA, SCINCIDAE) FROM THE
CARIBBEAN ISLAND OF SAN ANDRÉS, WITH A NEW INTERPRETATION OF
NUCHAL SCALES: A CHARACTER OF TAXONOMIC IMPORTANCE
AURÉLIEN MIRALLES
Département d’Ecologie et de Gestion de la Biodiversité, FRE 2696 – Adaptation et évolution des systèmes
ostéomusculaires, 55 rue Buffon, Muséum National d’Histoire Naturelle, Paris, France
A new species of Mabuya lizard from the isolated Caribbean island of San Andrés is described.
This species is closely related to Mabuya pergravis Barbour, 1921, another poorly known species
from Providencia Island, 87 km NNE of San Andrés. Unfortunately this new species, known from
a single specimen, is now probably extinct. It differs from M. pergravis in many morphological
characters such as a smaller size and very different patterns of coloration, but most importantly
in the presence of a very high number of nuchal scales. A new definition of this last character,
which is of systematic importance in the genus Mabuya, is also given and discussed.
Key words: insular endemism, Mabuya berengerae, San Andrés Archipelago, skink
species, then later regarded as a subspecies of M.
mabouya (Dunn & Saxe, 1950; Valdivieso & Tamsitt,
1963). However, I consider M. pergravis as a species in
its own right, even though it has some characteristics in
common with the two other, widely distributed Carib-
bean species, M. mabouya and M. sloanii (considered
by Dunn & Saxe [1950] as two populations of the same
species M. mabouya, the first one being native from the
southern Lesser Antilles and the second one from both
the northern Lesser Antilles and Greater Antilles).
Mabuya pergravis differs from its geographical closest
neighbour, M. sloanii, in having a longer snout, a larger
size and in the absence of two dorsal stripes. It differs
also from M. mabouya in having four supraoculars
(versus three) and a longer snout (Miralles, 2005), this
last point being highly supported by the very wide gap
of about 2200 km which separates their respective geo-
graphical distributions.
During a systematic revision of the genus Mabuya, a
specimen (UMMZ 127884, firstly identified as a M.
pergravis in the collection catalogue of the UMMZ)
collected from the Island of San Andrés, was found.
The presence of the genus on the island was never men-
tioned until now. After a comparison with two
specimens of M. pergravis, USNM 13875 (holotype)
and USNM 76947, it became clear the specimen from
San Andrés does not conform to the species from the
adjacent island of Providencia, nor with any other New
World Mabuya. Even though M. pergravis and the
specimen from San Andrés share some characteristics,
such as a very long and pointed snout, some major char-
acteristics distinguish the two forms easily, such as
differences in scalation, size and colour patterns (Fig.
2). Given that (1) San Andrés island was never in con-
tact with Providencia (Parsons, 1956), and is 87 km
away from it; and (2) the specimen from San Andrés
differs in many morphological characteristics from
Mabuya pergravis, I hypothesize that the specimen
UMMZ 127884 belongs to a species unknown to sci-
ence. This paper describes this new species.
Correspondence: A. Miralles, Département d’Ecologie et de
Gestion de la Biodiversité, FRE 2696 – Adaptation et
évolution des systèmes ostéomusculaires, 55 rue Buffon,
Muséum National d’Histoire Naturelle, 75005 Paris, France.
E-mail: amiral@mnhn.fr
INTRODUCTION
The San Andrés Archipelago is made up of two major
isolated islands in the south-west Caribbean Sea, San
Andrés (St. Andrew) and Providencia (Old Providence).
These small Colombian islands are situated 197 km and
235 km respectively from the Nicaraguan coasts.
Providencia lies at 87 km north–north-east of San
Andrés. The latter is situated at 365 km in the north off
the coast of Panamá, while Providencia is at 626 km from
Jamaica, its nearest neighbour among the Antillean Is-
lands (Fig. 1). Despite their proximity to each other and
relationship to the continental landmass, San Andrés and
Providencia are thought to have been uplifted independ-
ently in the early Tertiary and to have never had a land
bridge either between them, or between the mainland
(Parsons, 1956). San Andrés is made of uplifted Tertiary
and Pleistocene rocks, and Holocene terrestrial and
beach deposits (Geister, 1975).
Mabuya pergravis is an endemic species from the Is-
land of Providencia. This species, described in 1921 by
Barbour, was recognized by Dunn (1936, 1945) as a full
FIG. 1. Localisation of the two Islands of Providencia and San
Andrés, with map of San Andrés Island.
MATERIALS AND METHODS
According to its former definition, Mabuya Fitzinger,
1826 was considered as a worldwide distributed genus
of tropical Scincidae (with the exclusion of Australia
and Pacific islands). However, a recent phylogenetical
study (Mausfeld et al., 2002) demonstrated that all
Neotropical Mabuya species clearly form a mono-
phyletic lineage (except M. atlantica, an endemic
species from Fernando de Noronha Archipelago off Bra-
zil, belonging to the African lineage, firstly classified by
Mausfeld et al. (2002) as Euprepis Wagler, 1830, then
replaced by Bauer (2003) by Trachylepis Fitzinger,
1843, in accordance with the rules of the Code of Zoo-
logical Nomenclature). Following these authors, I
consider Mabuya in the present paper as an exclusively
Neotropical genus, encompassing Central America,
South America, and Caribbean islands.
The specimens examined for this study were pre-
served in 70% ethanol and deposited at the American
Museum of Natural History, New York (AMNH), the
Carnegie Museum, Pittsburgh (CM), the Field Museum
of Natural History, Chicago (FMNH), the Museo de
Historia Natural La Salle of Caracas (MHNLS), the Mu-
seum National d’Histoire Naturelle, Paris (MNHN), the
Sam Noble Oklahoma Museum of Natural History, Nor-
man (OMNH), the University of Michigan Museum of
Zoology, Ann Arbor (UMMZ) and the National Mu-
seum of Natural History, Smithsonian Institution,
Washington (USNM).
Drawings were made with a stereomicroscope
LEICA MS5 equipped with a camera lucida. The scale
nomenclature, scale counts and measurements used in
the description of specimen are based after Avila-Pires
(1995). New characters of first rank in systematics for
the genus Mabuya, proposed by Greer & Broadley
(2000), and Greer & Nussbaum (2000) were also added
to the description. Moreover, a new definition of the
nuchal scales is given in the discussion of the present
paper.
MABUYA BERENGERAE SP. NOV.
Holotype. UMMZ 127884. An unsexed probably
subadult or adult specimen from San Andrés Island (be-
tween Morgans Cave and Punta Sur), Departamento de
Archipielago de San Andrés, Colombia [located be-
tween 12° 28' and 12° 36'N, and 81° 40' and 81° 44'W],
collected on 6 August 1967 by C. F. Walker. The col-
lector identified this specimen as a M. pergravis, and
wrote accompanying field notes “basking on leaf three
feet above the ground when first seen” (G. Schneider,
pers. comm.).
Diagnosis. Mabuya berengerae is a particularly un-
common species which can be easily distinguished from
all other Mabuya species, including its closest relative
M. pergravis, by the following combination of charac-
ters: an extremely long, slender and pointed snout, the
highest number of enlarged nuchal scales of any known
species of Mabuya (Table 1), a high number of
subdigital lamellae under the fourth toes (19) and a con-
trasting reticulated pattern on the upper side of the head,
the neck and the back, also unique in the genus (Fig. 2B,
3).
Description of the holotype. UMMZ 127884 (Fig
2A-B, 3). Very good state of preservation. Snout-vent
length 52.8 mm, tail (regenerated) length 28.8 mm,
Head length 11.7 mm. Rostral wider than high. Fore and
hind limbs easily touching each other when adpressed
against body. A pair of internasals in median contact. A
frontonasal, approximately hexagonal, as wide as long,
laterally in contact with anterior loreal scale. A pair of
quadrilateral prefrontals, separated medially, and in
contact with frontonasal, both anterior and posterior
loreals, first supraciliary, first and second supraoculars,
and frontal. Frontal rhomboid (almost triangular), in
contact with all the length of the second supraocular. A
pair of frontoparietals, each in contact with frontal, the
supraoculars except the first, parietal and interparietal.
Interparietal triangular, separated from nuchals by
parietals; parietal eye distinct. Parietal overlapping the
upper temporal scale. Four supraoculars, the second one
being the longest and largest. The posteriormost
supraocular in contact with the frontal is the second one.
Four supraciliaries, the second the longest. Nostril in
posterior part of the nasal. A small postnasal, bordered
by supranasal, anterior loreal and first supralabial. An
anterior and a posterior loreal, both elongate. A
frenocular and two pre-suboculars. Lower eyelid undi-
vided, with a transparent disk. A single row of small
FIG. 2. Drawings of the head of the holotype of Mabuya
berengerae (UMMZ 127884). A, lateral view of the head; B,
dorsal view of the anterior part of the body; and of the
holotype of Mabuya pergravis (USNM 13875); C, lateral
view of the head; D, dorsal view of the anterior part of the
body. Scale bar = 3 mm.
A. MIRALLES2
scales across the dorsal edge of the eyelid window.
Eight supralabials, the sixth being the widest and form-
ing the lower border of the eyelid. Two pretemporal
scales, a single primary temporal, two secondary
temporals in contact and three tertiary temporals. Tem-
poral scales imbricate, smooth, cycloid, not distinctly
delimited from the scales on the nape and the sides of the
neck. Ear-opening relatively small, oval, with undulat-
ing anterior margin and smooth posterior margin.
Mental scale wider than long, posterior margin straight.
Postmental scales and a pair of chin shields in contact
with infralabials. First pair of chin shields in contact me-
dially, second pair separated by a smaller cycloid scale.
Gulars similar to ventrals. Eight infralabials. One pair of
primary nuchal scales, the left one being laterally fused
with the left upper tertiary temporal; number of second-
ary nuchal scales asymmetric: five on the right side, four
on the left one (see Discussion). Other scales on nape
similar to dorsals. On lateral sides of neck, scales
slightly smaller. Dorsal scales cycloid, imbricate,
smooth; 56 in a longitudinal row. Ventrals similar to
dorsals; 37 in a longitudinal row. No distinct boundaries
between dorsals, laterals and ventrals; 28 scales around
midbody. Preanal plate with scales similar to ventral.
Scales on tail similar to dorsals, except for its posterior
part which is regenerated. Palms and soles light col-
oured, covered with small tubercles, subequal in size.
Both regions delimited by a row of larger and flatter
scales. Subdigital lamellae smooth, single, under fourth
finger 14 (right) and 15 (left), under fourth toe 19 (on
each side). Finger and toes clawed; toes length in the
following order: I < II < III = V < IV.
The colour in life is unknown. In preservative, the
background colour of the flanks and the upper side of
the head, neck, back, legs and tail is bronze, with six
very narrow and slightly darker dorsal lines running
from the middle of the nape to the base of the tail.
Supraciliaries, supranasals and lips are very light cream
A NEW MABUYA FROM SAN ANDRÉS ISLAND
TABLE 1. Number of primary and secondary pairs of enlarged nuchals (nuchals I and II) in some species of Mabuya. The table is
divided in two parts: species without pair of secondary enlarged nuchal (upper part), and species with them (lower part). When
specimens had an asymmetric number of nuchals on each side of the nape, the highest number of the two was scored. Data for
species followed by an asterisk (*) are entirely or partly based on literature (Dunn & Saxe, 1950; Rebouças-Spieker & Vanzolini,
1990; Avila-Pires, 1995 and Hoge, 1946).
Species Presence of nuchals I Number of nuchals II
none 1 none 1 2 3 >4
M. bistriata (n=23) - 23 23 - - - -
M. falconensis (n=5) - 5 5 - - - -
M. frenata (n=6) - 6 6 - - - -
M. heathi (n=4) - 4 4 - - - -
M. nigropunctata (n=91) 2 89 90 1 - - -
M. mabouya (n=11) - 11 11 - - - -
M. pergravis (n=20)* - 20 15 5 - - -
M. unimarginata (n=18) - 18 18 - - - -
M. berengerae (n=1) - 1 - - - - 1
M. carvalhoi (n=4)* - 4 - - 3 1 -
M. croizati (n=14) - 14 - 1 8 5 -
M. nigropalmata (n=5)* - 5 - - 5 - -
M. macrorhyncha (n=3)* - 3 - 3 - - -
M. sloanii (n=19) - 19 - 13 4 1 -
FIG. 3. Photograph of the holotype of Mabuya berengerae (UMMZ 127884). Scale bar = 1 cm.
3
coloured. A brown lateral strip, itself spotted with
darker dots, extends each side from the nostril (loreals,
upper third part of the supralabials, around eyes and
temporals), along the upper half part of the ear-opening,
and continues in a lighter reticulated gradation on the
neck to above the arms (width on neck about three scale
rows). Three nearly imperceptible light lines are sup-
posed to have been more discernible in the living
animal: a vertebral line, running from nape to the tail,
and two dorsolateral ones, on each side along the lateral
brown strips, from the lateral part of the parietals to
above the arms. Many dark brown symmetric spots ac-
centuate posterior sutures of nearly all supracephalic
scales, others elongate spots run on the neck. The back,
sides and upper side of the legs are spotted with small
fused dark dots, forming many transverse narrow
stripes. The ventrum, lower side of head, neck, lower
side of the limbs and tail are cream-coloured with some
grey marks due to the transparency of the ventral skin.
Etymology. The name has been given in honour of my
wife Bérengère, for her invaluable support during all my
studies.
Remark. It appears quite probable that this species,
known from a single specimen, is now extremely rare.
For this reason, I avoided the dissection of the holotype
that would be necessary to determine its sex.
DISCUSSION
CONSERVATION, ENDEMISM AND BIOGEOGRAPHY
During the seventeenth and eighteenth centuries, San
Andrés apparently supported extensive natural stands of
“Cedar” (possibly Cedrela odorata), which were deci-
mated by early colonists (Emmel, 1975). Most of the
island is presently devoted to the cultivation of coconut
palms, Cocos nucifera (covering the southern half of the
island) and Colombian tourism. The human population
of San Andrés increased from 17 000 in 1967 (Emmel,
1975) to over 50 000 in 1984 (Johnson, 1988) and is
likely growing still. Gonzalez et al. (1995) comment on
the state of conservation of the original biotopes on San
Andrés and Providencia islands “It has been an unequal
struggle: so many hectares of Primary forest mainly
from San Andrés, have been sacrificed to give space for
hundreds of cement tons”. Unfortunately, Mabuya
berengerae is only known from a single specimen
caught in 1967. Given the small size of the island, its
very damaged ecosystem (Emmel, 1975; Barriga et al.,
1985; Johnson, 1988; Gonzalez et al., 1995) and the fact
that this specimen is the only one known, it appears
highly probable that this species is now endangered, if
not extinct. For those reasons, a faunistic survey on this
island seems to be urgently needed, in particular for a re-
assessment of the current conservation status of M.
berengerae, but also in a more general perspective, to
have an accurate assessment of the degree of endemism
on this island and to elaborate preservation strategies to
protect it.
Indeed, the San Andrés Archipelago appears to have
a unique evolutionary history. Despite the fact that few
studies were published on the fauna of San Andrés Is-
land, five endemic taxa of terrestrial vertebrates have
been described in addition to Mabuya berengerae, from
it: three reptiles out of thirteen known from the Archi-
pelago (Valdivieso & Tamsitt, 1963), including a
dipsadine snake species, Coniophanes andresensis Bai-
ley, 1937, a species of polychrotid (also present on
Providencia), Norops concolor (Cope, 1862), and a sub-
species of gekkonid lizard, Sphaerodactylus argus
andresensis Dunn & Saxe, 1950; but also two passerines
birds including a species of mimid, Mimus magnirostris
Cory, 1887 and a species of vireonid, Vireo caribaeus
Bond & Meyer de Schauensee, 1942. On the other hand,
no amphibian or mammal species appears to be endemic
to the island. So high a degree of endemism in so small
an area (about 44 km2) strongly suggests a unique his-
tory and a prolonged isolation of this island.
Given the large distance that separates the San
Andrés Archipelago from the other Caribbean islands,
its biogeographical affinities are not clear. Its original
fauna and flora seem to be a mix of both Mesoamerican
and Caribbean elements (Riley, 1975; Smith et al.,
1989; Gonzalez et al., 1995). Although it seems certain
that M. berengerae and M. pergravis are sibling species
(their extremely elongate and pointed snout being the
most remarkable synapomorphy of their monophyletic
lineage), the origin of this clade on the archipelago is
still unresolved. On the basis of their scale characters,
Dunn & Saxe (1950) conclude that the Mabuya from
Providencia seemed to be more closely related to the
Lesser Antillean populations than to those from the ad-
jacent mainland. Indeed, M. pergravis and M.
berengerae share with M. mabouya (present in the
southern Lesser Antilles from Dominica, Guadeloupe,
Martinique and Saint Lucia) a slender muzzle and a
short lateral strip from snout to above arms. They share
also with M. sloanii (present in northern Lesser Antilles
and in the Great Antilles) the presence of secondary
nuchal scales. Contrary to the biogeographical trends
based on botanical or entomological studies, no affini-
ties seem to be shared by Mesoamerican species of
Mabuya and the San Andrés Archipelago’s clade.
However, species belonging to the genus Mabuya are
often very similar and external morphological charac-
ters are usually used for studying their taxonomy.
Unfortunately, most of these characters could easily re-
sult from convergence. For this reason, only a real
phylogenetical approach would precisely determine the
relationships of this small clade.
WHAT IS A NUCHAL SCALE?
The organisation of the nuchal scales, transversally
enlarged dorsal scales of the nape, is a major character-
istic in the systematics of the genus Mabuya, and more
generaly in all the lygosomines skinks. Notably, it repre-
sents one of the most important diagnostic characters for
A. MIRALLES4
the new species described in the present paper. Until
now, no real definition was given to explain what a
nuchal scale is, this point being somewhat unclear in the
literature. Most of the authors who worked on the sys-
tematic of the genus Mabuya considered more or less
implicitly nuchal scales as any scales present on the
nape, in contact with parietals, and larger than other cy-
cloid scales (= small scales covering the body) present
on the back, (Dunn, 1936; Hoogmoed, 1973; Avila-
Pires, 1995; Rodrigues, 2000). Recently, Mayer &
Lazell (2000: 872) proposed that the pair of scales on
the nape as a pair of nuchal scales if their combined
width is more than 75% the width of the parietals.
However, the width of dorsal scales can vary gradu-
ally in some specimens, from the small cycloid scales of
the back to the wide scales of the nape. Consequently, it
becomes impossible to distinguish the limit between
these two categories of scales, the count becoming sub-
jective. Therefore, I propose to present here a more
accurate definition for the nuchal scales. This one is
based on a hypothesis of homology. A nuchal scale is
regarded as an enlarged scale of the nape, occupying
transversally the place of two or more rows of dorsal
cycloid scale. Thus, the antero-posterior transition from
nape to back, between nuchal and dorsal scales is sharp,
one longitudinal row of nuchals giving rise to two rows
of cycloïds. In the case of a gradual variation of width
between a single row of small dorsal scale and a single
row of large scale on the nape, all those scales will be
considered as “enlarged dorsal cycloid ones”, not as
nuchals. Additionally, I have distinguished two catego-
ries of nuchals (Fig. 4). The “primary pair of enlarged
nuchals, or nuchals I” (Fig. 4A-B) is in contact with the
posterior border of the parietal scales. Each scale of this
always singular pair occupies the equivalent of three
transverse rows of dorsal cycloid scales (sometimes
two). The nuchals I are present in all species belonging
to the genus Mabuya sensu lato. The “secondary pairs of
enlarged nuchals, or nuchals II” (Fig. 4B) are aligned
along the vertebral axis, posterior to the “nuchals I”.
Their number could vary from one to four/five pairs
(holotype of M. berengerae) depending on the species.
Each “nuchal II” occupies the equivalent of two trans-
verse rows of dorsal cycloid scales. They are always
present in six species, M. berengerae, M. carvalhoi, M.
croizati, M. nigropalmata, M. macrorhyncha and
Mabuya sloanii. They are nearly always absent in M.
arajara, M. bistriata (sensu Avila-Pires, 1995), M.
caissara, M. cochabambae, M. dorsivittata, M.
falconensis, M. frenata, M. guaporicola, M. heathi, M.
pergravis, M. mabouya (sensu Miralles, 2005), M.
nigropunctata (sensu Avila-Pires, 1995) and M.
unimarginata (sensu Savage, 2002), (Avila-Pires, 1995;
Dunn, 1936; Mausfeld & Lötters, 2001; Mijares-Urrutia
& Arends, 1997; Rebouças-Spieker, 1981; Rodrigues,
2000). On the other hand, they seem to be occasionally
present in only four species, M. agmosticha, M.
lineolata, M. macleani and M. pergravis (Cochran,
1941; Dunn & Saxe, 1950; Mayer & Lazell, 2000;
Rodrigues, 2000). In conclusion, the characteristic
“presence or absence of secondary nuchals scales” con-
stitutes a new useful diagnostic character given its very
low intraspecific variability for the majority of Mabuya
species.
ACKNOWLEDGEMENTS
I am very grateful to J.-P. Gasc (MNHN), J. P.
Michels (ZMA) and an anonymous reviewer for useful
comments and rereading the manuscript. Many thanks
also to L. S. Ford (AMNH), S. P. Rogers (CM), A.
Resetar (FMNH), G. Rivas Fuenmayor and J. C. Señaris
(MHNLS), I. Ineich (MNHN), A. Estep and L. J. Vitt
(OMHN), R. A. Nussbaum and G. E. Schneider
(UMMZ), and to R. W. Heyer (USNM) for providing
access to specimens, but also to A. Ohler for welcoming
me in the UMS 602 Taxinomie et Collections – Reptiles
et Amphibiens, of the Departement Systematique et
Evolution (MNHN).
REFERENCES
Avila-Pires, T. C. S. (1995). Lizards of Brazilian
Amazonia (Reptilia: Squamata). Zoologische
Verhandelingen (Leiden) 599, 1-706.
Barbour, T. (1921). Some reptiles from Old Providence
Island. Proceedings of the New England Zoölogical
Club 7, 81-85.
Barriga, T., Hernández, C. & Jaramillo, T. (1985). La Isla
de San Andrés. Contribuciones al conocimiento de su
ecología, flora, fauna y pesca. Bogotá, Colombia:
Universidad Nacional de Colombia.
Bauer, A. M. (2003). On the identity of Lacerta punctata
Linnaeus, 1758, the type species of the genus Euprepis
Wagler, 1830, and the generic assignment of Afro-
Malagasy skinks. African Journal of Herpetology 52,
1-7.
Cochran, D. M. (1941). The Herpetology of Hispaniola.
Bulletin of the United States National Museum 177, I-
VII + 1-398.
FIG. 4. Details of scales on the nape of two specimens of
Mabuya: (A) specimen with only a pair of primary nuchals
(M. nigropunctata, OMNH 36834) and (B) specimen with a
pair of primary nuchals and three pairs of secondary nuchals
(M. carvalhoi, AMNH 13732). P: parietal scale; 1: primary
nuchal scale; 2: secondary nuchal scale; *: rows of cycloid
scales homologue to nuchals ones (only showed for the left
side).
A NEW MABUYA FROM SAN ANDRÉS ISLAND 5
Dunn, E. R. (1936). Notes on American mabuyas.
Proceedings of the Academy of Natural Sciences,
Philadelphia 87, 533-557.
Dunn, E. R. (1945). The Amphibians and Reptiles of the
Colombian Caribbean Islands San Andrés and
Providencia. Reprint from Caldasia 3, 363-365. In:
Contribution to the Herpetology of Colombia 1943-
1946. Reprinted from the Revista de la Academia
Colombiana de Ciencias Exactas, Físicas y Naturales
and from Caldasia, Boletín del Instituto de Ciencias
Naturales de la Universidad Nacional de Colombia-
Bogotá. Appendix: Herpetological publications of
Emmett Reid Dunn, 1915-1957. Privately printed M.
T. D. by Dunn, E. R. 1957, i-ix + 1-296.
Dunn, E. R. & Saxe, L. H. (1950). Results of the
Catherwood-Chaplin West Indies expedition, 1948.
Part V. Amphibians and Reptiles of San Andrés and
Providencia. Proceedings of the Academy of Natural
Sciences, Philadelphia 102, 141-165.
Emmel, T. C. (1975). The Butterfly faunas of San Andrés
and Providencia Islands in the Western Caribbean.
Journal of Research on the Lepidoptera 14, 49-56.
Geister, J. (1975). Riffbau und geologische Entwicklungs-
geschichte des Insel San Andrés. Stuttgarter Beiträge
zur Naturkunde Serie B (Geologie und Paläontologie)
15, 1-203.
Gonzalez, F., Diaz, J. N. & Lowy, P. (1995). An ilustrated
Flora of San Andrés and Providencia, with spezial
reference to useful plants / Flora illustrada de San
Andrés y Providencia, con énfasis en las plantas
útiles. Colombia: Convenio Sena, Universidad
Nacional de Colombia, Instituto de Ciencias Naturales.
Greer, A. E. & Broadley, D. (2000). Six characters of
systematic importance in the scincid lizard genus
Mabuya. Hamadryad 25,1-12.
Greer, A. E. & Nussbaum, R. A. (2000). New character
useful in the Systematics of the Scincid Lizard Genus
Mabuya. Copeia 2000, 615-618.
Hoge, A. R. (1946). Um novo lagarto da ilha da Queimada
Grande. Memoria do Insituto Butantan 19, 241-248.
Hoogmoed, M. S. (1973). Notes on the herpetofauna of
Surinam IV. The Lizards and Amphisbaenians of
Surinam. Biogeographica 4, i-ix + 1-419.
Johnson, T. H. (1988). Biodiversity and conservation in
the Caribbean: profiles of selected islands. ICBP
Monograph No. 1. Cambridge: International Council
for Bird Preservation.
Mausfeld, P. & Lötters, S. (2001). Systematics of Mabuya
frenata cochabambae Dunn, 1935 (Reptilia, Sauria,
Scincidae) from Inner-Andean Dry Valleys of Bolivia.
Studies on Neotropical Fauna and Environment 36,
49-55.
Mausfeld, P., Schmitz, A., Böhme, W., Misof, B.,
Vrcibradic, D. & Rocha, C. F. D. (2002). Phylogenetic
affinities of Mabuya atlantica Schmidt, 1945, endemic
to the Atlantic Ocean Archipelago of Fernando de
Noronha (Brazil): necessity of partitioning the genus
Mabuya Fitzinger, 1826 (Scincidae: Lygosominae).
Zoologischer Anzeiger 241, 281-293
Mayer, G. C., Lazell, J. (2000). A new species of Mabuya
(Sauria: Scincidae) from the British Virgin Islands.
Proceedings of the Biological Society of Washington
113, 871-886.
Mijares-Urrutia, A. & Arends, A. (1997). Un nuevo
Mabuya (Squamata: Scincidae) de la costa semiárida
del noreste de Venezuela. Revista Brasilieira de
Biologia 57, 595-601.
Miralles, A. (2005). The identity of Lacertus mabouya
Lacepède, 1788, with description of a neotype: an
approach toward the taxonomy of New World Mabuya.
Herpetologica 61, 46-53.
Parsons, J. J. (1956). San Andrés and Providencia, English
speaking islands in the western Caribbean. University
of California Publications in Geography 12, 1-84.
Rebouças-Spieker, R. (1981). Sobre uma nova espécie de
Mabuya do nordeste do Brazil (Sauria, Scincidae).
Papéis Avulsos de Zoologia 34, 121-123.
Rebouças-Spieker, R. & Vanzolini, P. E. (1990). Mabuya
carvalhoi, espécie do estado de Roraima, Brasil
(Sauria, Scincidae). Revista Brasileira de Biologia 50,
377-386.
Riley, N. D. (1975). A field guide to the butterflies of the
West Indies. London: Collins.
Rodrigues, M. T. (2000). A new species of Mabuya
(Sauria, Scincidae) from the semiarid caatingas of
northern Brazil. Papéis Avulsos de Zoologia 41, 313-
328.
Savage, J. M. (2002). The Amphibians and Reptiles of
Costa Rica: A Herpetofauna between Two Continents,
between Two Seas. Chicago: University of Chicago
Press.
Smith, D. S., Munroe, E. & Ramos, S. J. (1989).
Biogeographical Affinities of the Butterflies of the
Southwest Caribbean Islands, San Andrés and
Providencia. Journal of the Lepidopterists’ Society 43,
274-288.
Valdivieso, D. & Tamsitt, J. R. (1963). A check list and
key to the Amphibian and Reptiles of Providencia an
San Andrés. Caribbean Journal Science 3, 77-79
A. MIRALLES
Accepted: 1.12.04
6
APPENDIX
SPECIMENS EXAMINED AND LOCALITIES (N=159):
Mabuya berengerae sp. nov. (n=1): COLOMBIA:
San Andrés Island: UMMZ 127884 (holotype), be-
tween Morgans Cave and Punta Sur ( = South End).
Mabuya carvalhoi (n=1): BRAZIL: Roraima:
AMNH R-137372 (paratype), Ilha de Maraca.
Mabuya croizati (n=14): VENEZUELA: Sucre:
AMNH R-29314 (holotype), Turumiquire massif; CM
7978, 7982, 7988, Elvecia, Turumiquire massif; FMNH
17796, 17797, 17798-428 to 17798-430, 17799-434 to
17799-438, Turumiquire massif.
Mabuya falconensis (n=5): VENEZUELA:
Carabobo: MHNLS 5511, 5512, Lago de Valencia;
FALCÓN STATE: 5526, Sierra San Luis, Curimagua;
UMMZ 55927, 55932, Tucacas.
Mabuya mabouya (n=11): LESSER ANTILLES:
Dominica: MNHN 2003-0838, UMMZ 83323.
Guadeloupe: MNHN 2902, 1889-0664. Martinique:
MNHN 738, 739, 1785, 5110, 5421, 5424. Unspecified
island: MNHN 2903.
Mabuya nigropunctata (n=98): BRAZIL: Acre:
OMNH 37048 to 37051, 5 Km N Porto Walter, inland
from the Rio Juruá. Amazonas: OMNH 37681 to
37698, Município de Castanho, 40 Km S Manaus at Km
12 on road to Autazes. Maranhão: MPEG 10690,
10691, 10693, 10695, 10698, Nova Vida, 25 Km dist.
of Rio Gurupi, BR 316. Para: MPEG 8605, 8608, 8611,
8616, 8642, Bela Vista, Viseu ; MPEG 12194, 12196,
12199, 12200, 12218 between the rivers Tocantins and
Mojú, 12 miles of the barrage Tucurui; OMNH 36828
to 36842, CEMEX, Agropecuaria Treviso LTDA, 101
Km S and 18 Km E Santarém. Rondonia: OMNH 37411
to 37417, Rio Formoso, Parque Estadual Guajará-
Mirim, 90 Km N Nova Mamoré. Roraima: OMNH
36313 to 36322, 7 km E Rio Ajaraní, BR 210. FRENCH
GUIANA: MNHN 1996-4570 to 1996-4572, 1997-
2206 to 1997-2213, St Eugene; MNHN 2002-0612,
2002-0613, St Marcel. TRINIDAD & TOBAGO:
AMNH R-64528, Trinidad island FMNH 49901 to
49908, Trinidad island, San Rafael, UMMZ 79919,
A NEW MABUYA FROM SAN ANDRÉS ISLAND
Trinidad island. VENEZUELA: Amazonas: MHNLS
16389, Parima B, Bolívar: MHNLS 15532, Serranía del
Supamo, Cerro Santa Rosa; MHNLS 11545, foot of the
Roraima Moutain. Delta Amacuro: MHNLS 4543,
Burojoida. Sucre: MHNLS 15533, Peninsula de Paria,
Las Melenas; MHNLS 16203, Peninsula de Paria,
Macuro.
Mabuya lineolata (n=1): GREATER ANTILLES:
Dominican Republic: AMNH R-42145 (Holotype),
Monte Cristy province, not far from Monte Cristy, near
the bank of the Rio Yaque del Norte.
Mabuya pergravis (n=2): COLOMBIA: Providencia
island: USNM 13875 (Holotype), USNM 76947
Mabuya sloanii (n=19): BAHAMAS: Turks and
Caicos islands: UMMZ117393, West Caicos, on W
side. GREATER ANTILLES: Jamaica: UMMZ 85861,
Portland Point. Puerto Rico: CM 23775, 23776, FMNH
215, UMMZ 124819, Mona Island; OMNH 32878, un-
specified locality. LESSER ANTILLES: Anguilla: CM
115480, Brimegin, W of Foutain Hill, CM 115481,
unlocalised locality. British Virgin Islands: CM 17357,
17358, Anegada; UMMZ 80582 Virgin Gorda. Saint-
Barthélemy: MNHN 1997-6064, 2003-0843,
2003-0844, unlocalised locality. US Virgin Islands:
MNHN 554 (holotype), 1088, St Thomas; UMMZ
73821, St Thomas, Buck Island; UMMZ 80586, St
Thomas, Capella Island.
Mabuya unimarginata complex (n=7): COSTA
RICA: Puntarenas: UMMZ 117575, 1 Mile E of Volcan
de Buenos Aires cone finca; UMMZ 145875, 16 Km S
San Vito, on Highway 16 Las Cruces field station,
1250m. MEXICO: Yucatan: AMNH R-38863 to R-
38865, Chichen Itza; UMMZ 113548, Calcehtok.
NICARAGUA: Leon: OMNH 36173 to 36175, Vulcan
Momotombo. Rio San Juan: OMNH 36177 to 36179, by
Isla de Diamante on Rio San Juan. Rivas: OMNH
36176, Chococente. PANAMA: Bocas del Toro:
UMMZ 142636, Bocas del Toro. Los Santos: CM
43593 to 43594, Los Santos ; CM 43595, Santa Maria.
Panama: UMMZ 135354, Gorgas Memorial Laboratory
Field Station on the Rio Bayano, 250 m alt.
7
8
