South American Journal of Herpetology, 8(1), 2013, 5–18
© 2013 Brazilian Society of Herpetology
A Distinctive New Species of Gonatodes
(Squamata: Sphaerodactylidae) from Isla La Blanquilla,
Venezuela, with Remarks on the Distribution of Some Other
Caribbean Sphaerodactylid Lizards
Gilson A. Rivas1,*, Gabriel N. Ugueto2, Walter E. Schargel3, Tito R. Barros1, Pablo Velozo4, Luz Esther Sánchez4
1
Museo de Biología, Facultad Experimental de Ciencias, La Universidad del Zulia. Apartado Postal 526, Maracaibo 4011, Estado Zulia, Venezuela.
11111 Biscayne Boulevard, #556, Miami, Florida 33181, USA.
3 Department of Biology, The University of Texas at Arlington. Arlington, Texas 76019, USA.
4 Laboratorio de protección y manejo de cuencas, Instituto Venezolano de Investigaciones Cientificas (IVIC). Maracaibo, Estado Zulia, Venezuela.
* Corresponding author. Email: anolis30@hotmail.com
2
Abstract. A new sphaerodactylid gecko of the genus Gonatodes is described from La Blanquilla Island, located 170 km north of the Venezuelan mainland. This new species exhibits the following suite of characters that immediately separate it from other taxa in the genus: vertically elliptic pupil, small size, uniformly reddish brown, non-sexually dimorphic coloration, and a subcaudal scale pattern type C (1’1”). The
new species is the fourth Gonatodes endemic to Caribbean islands, the other three being G. antillensis (Bonaire, Curaçao, the archipelagos of
Las Aves and Los Roques), G. daudini (Union Island, The Grenadines), and G. ocellatus (Tobago). Finally, we discuss the close phylogenetic
relationship (based on nuclear genes c-mos and NT3) between the new species and Gonatodes daudini, as well as its zoogeographical implications, showing interesting parallels with that of other Caribbean lizards.
Keywords. Gekkota; South America; Systematics; Taxonomy.
Resumen. Se describe un nuevo lagarto sphaerodactílido del género Gonatodes proveniente de la Isla de La Blanquilla, localizada 170 km al
norte de la tierra firme de Venezuela. La nueva especie se diferencia fácilmente de otros congéneres por la siguiente combinación de caracteres morfológicos: pupila verticalmente elíptica, diminuto tamaño, coloración uniforme, pardo rojiza en ambos sexos y un patron subcaudal
tipo C (1’1”). El nuevo taxón es el cuarto miembro del género Gonatodes endémico de una isla del mar caribe, las otras son G. antillensis (Bonaire, Curaçao, y los archipiélagos de Las Aves y Los Roques), G. daudini (Isla Unión, Las Granadinas) y G. ocellatus (Tobago). Finalmente, se
discute la cercana relación evolutiva entre la nueva especie y G. daudini, así como sus implicaciones zoogeográficas, mostrando interesantes
paralelismos con otras especies de lagartos del Caribe.
The taxonomy of reptiles from continental and oceanic islands of the Venezuelan Caribbean coast has, until
recently, remained unchanged and overlooked for many
years. Herpetological collecting efforts and research on
these islands began towards the end of the 1800s and
early 1900s by researchers and travelers who sporadically explored the area. The most important of these
early works are those by Peters (1873), Meek (1910), and
Hummelinck (1940), in which several species (e.g., Anolis blanquillanus Hummelinck 1940, Cnemidophorus nigricolor Peters 1873, and Phyllodactylus rutteni Hummelinck
1940) were described. However, in recent years, new research has brought to light the fact that the real diversity of herpetofauna from these islands has been greatly
underestimated (Ugueto et al., 2009; Ugueto and Harvey,
2010; Ugueto and Rivas, 2010; unpublished data for the
authors).
The Venezuelan Antilles (La Blanquilla, La Orchila,
and the archipelagoes of Las Aves, Los Hermanos, and
Los Roques) are separated from one another and from the
mainland by deep-sea trenches probably since their emergence. It is very likely that these have never been connected to the continent, for sure not during the last glacial
maxima (Neill et al., 2011), resulting in each possessing
a unique but depauparate herpetofauna. In this study,
we describe a new diminutive and distinctive sphaerodactylid lizard of the genus Gonatodes from one of these
islands. The new taxon was collected during recent fieldwork on Isla La Blanquilla (also known as Isla Blanca),
Venezuela. This species was apparently first collected at
the end of the 1930s by Hummelinck (1940) on the nearby Los Hermanos Archipelago but was never formally described. Hummelinck (1940) reported the specimens he
collected as “Gonatodes spec. (? Gymnodactylus aff.)” and
deposited them in the Nationaal Natuurhistorisch Museum (Leiden) and in the Zoölogische Museum (Amsterdam). The collection of the Zoölogisch Museum has been
incorporated into the collection of the National Natuurhistorisch Museum. Currently, they are now all housed
in the former, and will be integrated into a new center for
Submitted on: 18 February 2012
Accepted on: 25 January 2013
Handling Editor: Ana Prudente
doi: 10.2994/SAJH-D-12-18212.1
INTRODUCTION
South American Journal of Herpetology, 8(1), 2013, 5–18
biodiversity. For this reason we were unable to obtain a
loan of these specimens (Ronald de Ruiter “in litt”). Here
we describe this enigmatic species based on eight newly
collected specimens possessing a suite of unique features.
Additionally, we examine the phylogenetic position of
this new species based on molecular data.
MATERIALS AND METHODS
Descriptive methods
Specimens examined are deposited in the following
collections (Appendix I): at Museo de Biología, Universidad del Zulia, Maracaibo, Zulia, Venezuela (MBLUZ);
Museo de la Estación Biológica de Rancho Grande, Aragua, Venezuela (EBRG); Museo de Historia Natural La
Salle, Caracas, Venezuela (MHNLS); Milwaukee Public
Museum, Milwaukee, Milwaukee, USA (MPM); and the
Amphibian and Reptile Diversity Research Center, The
University of Texas at Arlington, Arlington, Texas, USA
(UTA).
Measurements of specimens were recorded with a
digital caliper (to the nearest 0.1 mm) and include snout–
vent length (SVL, from tip of snout to cloacal opening),
tail length (TL, from cloacal opening to tip of tail), head
length (HL, from the tip of the snout to the anterior margin of the ear opening), head width at widest section (HW,
at widest section of head), axilla-groin distance (AXG,
from posterior margins of arm insertion to anterior margin of hind limb insertion). Loreal scales were counted in
straight line between postnasals and anterior border of
orbit. Scale counts include ventral scales (VS, along a midventral line from anterior level of arm insertion to vent,
excluding the small granules at its edge) and scales around
midbody (SAMB). Terminology and scale counts follow
Avila-Pires (1995) and Rivero-Blanco and Schargel (2012).
Phylogenetic analysis
In order to examine the phylogenetic position of
the new species we conducted an analysis based on DNA
sequences from two nuclear genes (C-mos and NT3) that
have been previously used to investigate the systematics
of the genus (Gamble et al., 2008; Schargel et al., 2010).
The protocols for DNA isolation and PCR follow Schargel
et al. (2010). We included sequences of two individuals of
the new species (MBLUZ 1011 and 1013), as well as several Genbank (Benson et al., 2005) sequences from other
species in the genus and other sphaerodactylid genera to
be used as outgroups. The sequences used are listed in the
Appendix II with the corresponding Genbank accession
numbers. Sequences were imported into MEGA 5 (Tamura et al., 2011) and aligned using the ClustalW algorithm
6
(Thompson et al., 1994). MEGA 5 was also used for calculating genetic distances. The sequences from both genes
were concatenated in TREEFINDER Version of March
2011 (Jobb, 2011), and two partition filters (gene and
codon position) were added manually. We used the “propose model” option with default settings in TREEFINDER
to select, under the Corrected Akaike Information Criterion (AICc), the best-fit model of nucleotide substitution
for each partition. A maximum likelihood (ML) bootstrap analysis (1000 replications) was also conducted in
TREEFINDER loading the best-fit models for the different partitions and the partition rates set to optimum.
Results of the bootstrap analysis were summarized in a
consensus tree collapsing all clades that received a support value level less than 50.
Species description and results
Gonatodes naufragus sp. nov.
(Figs. 1–10, Table 1)
Gonatodes spec. (? Gymnodactylus aff.): Hummelinck
1940: 74 [four specimens collected in Morro Pando,
Archipelago Los Hermanos, Dependencias Federales, Venezuela; also observed in Morro Fondeadero].
Holotype. EBRG 5224 (Figs. 1, 4E), an adult female, collected on Playa Juan Gerardo (11°52’34.92”N–
64°37’36.68”W), 3 m above sea level (asl), La Blanquilla,
Dependencias Federales, Venezuela, 20 October 2010, by
Gilson A. Rivas, José J. Rodríguez and Ronnis Guevara.
Paratypes. MBLUZ 1010, an adult male, with
same data as holotype. MBLUZ 1011, female, between
Playa Juan Gerardo and Playa El Barco, La Blanquilla
(11°53’22.84”N–64°37’10.62”W) by Gilson A. Rivas, José
J. Rodríguez and Ronnis Guevara. MBLUZ 1012, female,
and MBLUZ 1013, hatchling, from Playa Piedra Ahogada,
La Blanquilla (11°49’23.87”N–64°38’09.15”W), 10 m asl,
obtained on 21 October 2010 by Gilson A. Rivas, José J.
Rodríguez and Ronnis Guevara. MBLUZ 1147, male, on
1 February 2012 by Gilson A. Rivas, Angel Fernández,
Jose J. Rodríguez and Jackeline Reid. MBLUZ 1146, juvenile, lomas de granito, La Blanquilla (11°51’53.42”N–
64°37’25.41”W), 15 m asl on 25 January 2012, 5 m asl.
by Gilson A. Rivas, Angel Fernandez, Jose J. Rodriguez
and Jackeline Reid. MBLUZ 1148, female, La Blanquilla
(11°51’32.24”N–64°37’27.17”W), 20 m asl. on 2 February
2012, by Gilson A. Rivas, Angel Fernandez, Jose J. Rodriguez and Jackeline Reid.
Diagnosis. The new species can be distinguished
from all congeners by the following unique combination
of characters: (1) small body size, with adults ranging
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
South American Journal of Herpetology, 8(1), 2013, 5–18
Table 1. Selected morphometric measurements (in mm) and scale counts of the type series of Gonatodes naufragus sp. nov. M = male, F = female,
J = juvenile, H = hatchling. EYN = eye-nostril distance. Other abbreviations are defined in the text. TL = corresponds to the complete tail of MBLUZ 1010
only; all other tails are incomplete or regenerated.
Specimen number
EBRG 5224
MBLUZ 1010
MBLUZ 1011
MBLUZ 1012
MBLUZ 1013
MBLUZ 1146
MBLUZ 1147
MBLUZ 1148
Sex
F
M
F
F
H
J
M
F
SVL
27.0
23.3
27.2
24.7
14.8
21.6
24.8
27.2
TL
25.2
26.2
11.5
–
13
7
23.4
12
from 23.3–27.2 mm SVL, (2) rostral distinctly acuminate and elongated, (3) pupil vertically elliptical, (4) absence of a clearly differentiated elongate supraciliary
spine, (5) dorsal scales bulky and relatively small but not
granular, (6) absence of clusters of distinctly enlarged
conical scales on sides, (7) 52–53 scales around midbody, (8) 35–38 ventral scales counted in a longitudinal
row, (9) escutcheon scales on posterior belly and ventral
HL
6.8
5.6
6.7
5.7
4
5.5
6.1
6.1
HW
4.9
3.9
4.5
4.3
2.7
4
4.6
4.2
EYN
1.7
1.5
1.7
1.6
0.9
1.4
1.6
1.7
AXG
12.2
10.3
11.1
10.5
5.6
8.3
9.8
12.3
VS
36
35
38
37
–
33
35
38
SAMB
63
60
67
–
65
66
60
surfaces of thighs in males evident, (10) one to two lateral
rows of scales on distal part of finger and toes (Fig. 2),
(11) midventral scales distinctly wider than long, forming
a repetitive sequence (only in the original tail) of a single
midventral scale in contact laterodistally with one scale
per side followed by a single midventral scale in contact
with two scales per side (1’1”sensu Avila-Pires, 1995; subcaudal pattern type C sensu Rivero-Blanco and Schargel,
2012, Fig. 3), (12) adults (both sexes) with uniform reddish brown dorsal coloration and hatchlings with striped
pattern, conspicuously different from that of adults.
Figure 2. Ventral aspect of right hand (left) and foot (right) of Gonatodes naufragus sp. nov. (paratype MBLUZ 1012, adult female).
Figure 1. Dorsal (upper), lateral (middle) and ventral (below) views of
head in Gonatodes naufragus sp. nov. (holotype EBRG 5224, adult female,
SVL 27.2 mm).
Figure 3. Ventral aspect of tail in Gonatodes naufragus sp. nov.: left,
paratype MBLUZ 1010, adult male with original tail; right, holotype
EBRG 5224, adult female with regenerated tail.
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
7
South American Journal of Herpetology, 8(1), 2013, 5–18
Description of holotype. An adult female, with
SVL 27.0 mm. Head approximately 1.3 times longer than
wide (HL: 6.8 mm; HW: 4.9 mm), 0.25 times SVL. Snout
4.1 mm long (0.60 times HL), acuminate in dorsal view,
45º sloping toward top of head. Neck barely narrower
than head and body; more elongated and thick than that
of other Gonatodes species, which gives the lizard an appearance similar to that of Sphaerodactylus. Body nearly
cylindrical but wider than high; AXG 11.5 mm. Limbs
short, well developed with short digits, fourth toe length
2.3 mm, 2.7 times shank length (6.3 mm). Tail thick,
round in cross section, tapering toward tip. TL 25.2 mm
in length. Tongue elongate, immaculate, with the tip
rounded, without medial cleft. Teeth small, conical and
subequal in length.
Rostral large, V-shaped, visible from above and
pointed, very sharp, contrary to that observed in other
Gonatodes in which it is less acuminate (e.g., G. antillensis and G. daudini) or rounded (most species); posterior margin slightly indented by median postrostral and
semidivided by a moderately long medial cleft. Three
postrostral scales; lateral ones (supranasals) distinctly
larger than median, which is slightly smaller than adjacent posterior scale. Nasal bordered by rostral, lateral
postrostral (supranasal), two postnasals, and first supralabial; nostril located relatively far from tip of the snout
(more distantly than in any other species of Gonatodes).
Postnasals roughly subequal in size to each other and
lower one both sides of the head; postnasals subequal
in size to scales on loreal region. Scales on top of snout
quadrangular, flat and juxtaposed, gradually becoming
slightly imbricate towards loreal region. Loreal scales
number about eight (right side) and nine (left side) on
a line between postnasals and anterior margin of orbit.
Scales decrease noticeably in size from the postrostrals
to the interocular area, posteriorly increasing slightly in
size toward the occipital region. Supraciliary flap well
developed, with a few conical supraciliaries slightly enlarged, but none forming a distinctly elongate spine. Pupil vertically oval. Supralabials 4 (both sides) to center
of eye, first two the largest (first slightly longest than
second), third slightly smaller and fourth much smaller,
followed by 3 (both sides) much smaller scales along lip
to rictus of mouth, all similar in size to scales on temporal region. Scales on temporal region similar in size
and shape to those on top of head. Ear opening roughly
rounded, about 1/5 the size of the orbit; a deep auditory
meatus.
Mental large, with angular posterior margin. Postmentals two, polygonal distinctly larger than adjacent
posterior scales and polygonal. Scales on chin directly
behind postmentals similar in shape to them, posteriorly rounded and much smaller; a few larger, polygonal,
juxtaposed scales adjacent to infralabials. Infralabials four (both sides) to center of eye, decreasing in size
8
posteriorly, the anterior two very large (first much longer
than second); anterior three infralabials projecting onto
the ventral plane, although the third barely visible from
below.
Scales on nape and sides of neck continuous with
those on posterior part of head (small, rounded, slightly
bulky and subimbricate) but slightly larger. Scales on the
gular region smooth and imbricate, with round posterior
margin, and with an anterior segment with much smaller
scales, and a posterior segment with larger scales. Dorsal
scales not granular, but round, bulky and subimbricate,
distinctly larger than those on top of head and neck; dorsolaterally and on flanks slightly larger than those middorsally. Limit between subimbricate scales on flanks and
imbricate ventrals not clearly demarcated. Ventral region
with scales distinctly larger than dorsals, similar on chest
and on belly, smooth, with round margin; ventrals in
oblique rows, with 36 scales. SAMB about 63, of which
about 12 are ventrals. There are diminutive scales around
the vent arranged irregularly.
Scales on dorsal part of tail larger than on body,
slightly bulky, imbricate, with round posterior margin;
limit between body and caudal scales not clearly demarcated. Subcaudal scales larger than those dorsally, smooth,
flat, imbricate, with round posterior margin; midventralscale single, conspicuously enlarged transversally, laterally
in contact with either two or three smaller adjacent caudal scales. The tail has been regenerated twice; midventral
scales of regenerated portions much shorter and wider
that those on original portion (see Fig. 3).
Scales on limbs rounded and juxtaposed, except
ventrally and on anterior surface of forearms and thighs
where they are smooth, flat and imbricate. Scales under
thighs and anterior part of belly, close to the vent, less
pigmented. Lamellae under first (I) through fifth (V) finger (right/left side): I: 5/6, II: 9/9, III: 10/10, IV: 11/12,
V: 8/7 (the fifth finger on right side is partially broken on
third lamellae). Lamellae under first (I) through fifth (V)
toe (right/left side): I: 5/5, II: 9/7, III: 10/11, IV: 14/14
(the fourth finger on left side is partially broken on seventh lamellae), V: 12/12. Fingers and toes with a single
or two lateral rows of scales distally. Claws exposed, nonretractile, between two basal scales (one dorsal and one
ventral).
Color of holotype in life. Dorsum uniformly dark
reddish brown with the tips of some scales on trunk pale
yellow (Fig. 4E). Head and neck scales are uniform without yellow color on the tips of the scales. Venter light
brown, with the abdominal region slightly darker. The
non-regenerated portion of the tail is similar in coloration
to that of dorsum; the remainder of the tail has been regenerated twice, the middle portion has light brown longitudinal stripes. Iris reddish brown with an ochraceousgolden ring around the pupil.
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
South American Journal of Herpetology, 8(1), 2013, 5–18
Variation in paratypes. Paratypes consist of three
adult females (SVL 24.7–27.2 mm), two adult males (SVL
23.3–24.8 mm), a juvenile (SVL 21.6 mm) and a hatchling
(SVL 14.8 mm). Table 1 provides scale counts and measurements of the type series. Supralabials 4–6, followed
by 2–3 much smaller scales along the lip to rictus of
Figure 4. Top: adult male (A) and adult female (B) Gonatodes antillensis in life. Gran Roque, archipelago Los Roques, Venezuela (both specimens uncollected). Center: dorsal view of the adult female paratype of G. naufragus sp. nov. in life (MBLUZ 1148) (C), and ventral view of the same specimen (D).
Bottom: Illustration of juvenile (above) and adult (below) Gonatodes naufragus sp. nov. in life. Drawings based on MBLUZ 1013 and EBRG 5224, respectively (E). Photographs: Luis Alejandro Rodríguez J.; Illustration: Gabriel N. Ugueto.
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
9
South American Journal of Herpetology, 8(1), 2013, 5–18
mouth, infralabials four, followed by 2–3 much smaller
scales along the lip to rictus of mouth. Loreal scales range
from seven to eight. Scales around midbody range from
57 to 67, of which 12–14 are ventrals. There are 33–38
scales on an imaginary line between the anterior part of
the insertion of the arm to the vent. Both males have an
evident escutcheon (Fig. 5) formed by a cluster of approximately 37 scales on the posterior portion of the abdomen,
distinctly different from other ventrals, unpingmented
except slightly on their outer margins. The same unpigmented scales are present under the thighs where they are
arranged in 3 (left leg)–4 (right) transverse rows of about
29 (right)–26 (left) scales each. Variation in the number of
lamellae under first (I) through fifth (V) finger is as follow:
I: 5–6, II: 8–9, III: 9–10, IV: 10–11, V: 7–9. Variation in the
number of lamellae under first (I) through fifth (V) toe is
as follow: I: 4–6, II: 8–9, III: 9–11, IV: 12–14, V: 10–12.
The coloration of the adult (Fig. 4C–D) is essentially
the same as in the holotype, except for MBLUZ 1147, a
presumably young male which has pale yellow reticulations overlaid on the head. The coloration of the single
Figure 5. Ventral view showing escutcheon areas (shaded in gray) on
posterior abdomen and ventral aspect of thighs in Gonatodes naufragus
sp. nov. (paratype MBLUZ 1010, adult male).
Figure 6. Map showing localities where Gonatodes naufragus sp. nov. was collected on La Blanquilla Island (black dots). Records of “Gonatodes spec.” by
Hummelinck (1940) on the neighboring archipelago of Los Hermanos (black triangles) that likely represent other populations of the new species.
10
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
South American Journal of Herpetology, 8(1), 2013, 5–18
hatchling (MBLUZ 1013) is the following: anterior flanks
and scapular region reddish with small yellow spots; dorsum reddish with a dark brown dorsolateral stripe with
narrow yellow borders at each side of the body, extending
from the nostril to the sides of anterior third of tail, with
interrupted on scapular region. Dorsum and tail brown
peppered with yellow, but less distinctly than on flanks
(see Fig. 4E). The subadult specimen (MBLUZ 1146) has a
color pattern intermediate between the hatchling and the
adults. It is reddish grey with tiny white dots on the side
of the body; the head is grey overlaid with red reticulations; the venter is grey.
Hummelinck (1940) briefly described the coloration
of adults and juveniles from the specimens he collected on
Los Hermanos Archipelago as follows:
Upperparts greyish (in life reddish), without
any well marked design, underparts much
lighter. Juveniles with a well marked laterodorsal, dark-brown stripe, abruptly beginning in
occipital region, narrowing and becoming more
vague on tail; each stripe with an irregular series of several spots in groundcolor medially,
with a little yellow dot in each centre anteriorly; two narrow yellow stripes below and above
the eye, disappearing in occipital region.
in a vertical terrace a few meters from the shore line in a
shaded, very humid, area. MBLUZ 1148 was found on the
ground inside the roots of a big tree. This area was covered
with abundant leaf litter (Fig. 7B). All localities where this
species has been observed come from the granitic area,
except the specimen from the patch from Playa Juan Gerardo and Playa El Barco, which come from an area where
the marine terraces are in contact with the granitic basement. It is particularly important, because the granite
basement is older than the terraces and offers more suitable and humid conditions with shaded areas than those
found on the terraces. We did not observe G. naufragus at
night; however, the vertically oval pupil suggests possible
crepuscular or nocturnal habits. The condition of the iris
is similar to that of G. antillensis, the only known species
of the genus with nocturnal habits.
On October 2010 we had the opportunity to observe
a feral cat around the camp. On the trip of 2012, several
cats were observed, including on the most distant parts of
the island. Several rats and mice were also observed. All
He also provided a brief morphological description
(e.g., snout acutely pointed, 12 lamellae beneath fourth
finger, 14 beneath fourth toe, 40–45 scales between anterior border of arm and vent). It is clear by the color
and morphological descriptions provide by Hummelinck
(1940) that Los Hermanos specimens are conspecific with
the species described here from Isla La Blanquilla. This is
not surprising since La Blanquilla is located with Los Hermanos Archipelago on a shallow marine platform (Maloney 1971).
Distribution and natural history. Gonatodes naufragus is known only from La Blanquilla, Dependencias
Federales and from at least two islets or “morros” from
Los Hermanos Archipelago, Venezuela (Fig. 6). The holotype EBRG 5224 and MBLUZ 1010 were found under
the bark of a large dead tree a few meters from the beach.
On the same tree, but about 1.5-2 m aboveground, two
Phyllodactylus rutteni were collected. MBLUZ 1011 was
collected on a rainy morning while removing some fallen
tree trunks on a coral substrate with scarce vegetation
matter. The juvenile (MBLUZ 1013) and an adult female
(MBLUZ 1012) were found around noon, under two decaying logs that maintained a higher level of moisture
than in the areas around it. We found shells of at least five
eggs and an unhatched egg inside a crevice of one of these
logs. MBLUZ 1146 was found under a rock just in the base
of a small three in storm scrub. MBLUZ 1147 was found
Figure 7. Typical landscape of hills in La Blanquilla Island (A). The hills
are formed from exposed granitic lithology sparsely covered with grasses. Sediments from erosion and detritus accumulate at the bottom of the
hills, which creates a matrix that retains moisture and allows for shrubs
and small trees to grow. Microhabitat of Gonatodes naufragus under a
Ficus tree (B) on La Blanquilla Island. Photographs by Lenin Parra (A)
and Gilson A. Rivas (B).
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
11
South American Journal of Herpetology, 8(1), 2013, 5–18
these three invasive species might affect the survival of
the native reptile fauna of the island.
La Blanquilla, is located approximately 100 km
northwest of Isla de Margarita. The island has an area
of 52.5 km2 and the highest altitude over 30 m above
sea level (Fig. 7A). On the isla La Blanquilla, a sequence
of reef limestone, called Blanquilla formation, crops out
on the island. This formation consists of three marine
terraces. The Blanquilla formation lies unconformably
above the basement of the island, which consists of the
Garantón Trodhjemite, of Paleocene age, while the age of
the Blanquilla formation is Pleistocene (Schubert, 1976).
Other species endemic to both Isla La Blanquilla and Los
Hermanos Archipelago are Anolis blanquillanus Hummelinck, 1940, Phyllodactylus rutteni Hummelinck, 1940,
and Cnemidophorus leucopsammus Ugueto and Harvey,
2010. Except for the Green Iguana, all lizards described
from these islands are endemic. Populations of P. rutteni
reported from other islands of the Venezuelan Antilles
represent undescribed species to be described by the authors elsewhere.
Etymology. The specific epithet naufragus is a Latin
masculine adjective, meaning “shipwrecked” or “castaway”, alluding to how a putative ancestor of this species
could have colonized Isla La Blanquilla, remaining stranded on this small island and evolving in isolation from its
congeners.
Comparisons. With a maximum SVL 31.1 mm (Rivera Rodríguez et al., 2011), Gonatodes daudini Powell and
Henderson 2005, from the tiny Union island, Grenadines,
was until now considered the smallest species of its genus
(Powell and Henderson, 2005). However, that title can
now be assigned to G. naufragus, with adult females (the
largest sex) reaching a maximum SVL of just 27.2 mm.
The diminutive size of both taxa readily separates them
from all the other 28 known species of Gonatodes (Rivero–
Blanco and Schargel, 2012), all of which reach or exceed
SVL 35 mm. Not only do G. naufragus and G. daudini share
a small size, but they exhibit a suite of other characters
unique to these two species. Both taxa have a conspicuously elongated and sharply pointed snout, with a larger
Figure 8. Schematic drawing of the shape of the pupil and the snout in three species of Gonatodes. (A) Gonatodes naufragus sp. nov. (paratype MBLUZ
1012, adult female). (B) Gonatodes daudini (MPM 33975, adult male). (C) Gonatodes antillensis (MBLUZ 1001, adult female). (D) Gonatodes aff. albogularis (MBLUZ 1036, adult male).
12
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
South American Journal of Herpetology, 8(1), 2013, 5–18
Figure 9. Dorsal squamation in three species of Gonatodes from Venezuela. (A) Gonatodes naufragus sp. nov. (holotype EBRG 5224, adult female). (B) Gonatodes antillensis (MBLUZ 1002, adult male). (C) Gonatodes aff. albogularis (MBLUZ 1037, adult male).
distance between the nostril and the tip of the rostral
scale compared to that in all other species of Gonatodes
(Fig. 8). The pointed snout in G. naufragus and G. daudini
is reminiscent of that in Sphaerodactylus.
Both Gonatodes naufragus and G. daudini are unique
in having relatively (G. naufragus) to conspicuously
(G. daudini) enlarged and bulky dorsals instead of the typical granules that are only slightly larger than the scales on
the top of the head found in all other Gonatodes (Fig. 9).
The diagnostic low counts of scales around the midbody
exhibited by both species (60–67 in G. naufragus and 33–
44 in G. daudini) reflect that. However, this character is
much more pronounced in G. daudini than in the new species (compare Fig. 9A-C with Powell and Henderson, 2005:
fig. 3), with much larger, bulkier and juxtaposed dorsals in
the former, noticeably smaller and subimbricate in G. naufragus. In some aspects, the condition exhibited by G. naufragus would appear to represent an intermediate stage
between the small granules typical of most Gonatodes and
the large scales of G. daudini. Additionally, both taxa are
atypical in that neither taxon is sexually dichromatic (see
Rivera Rodriguez et al., 2011 for data on G. daudini). In
most of Gonatodes sexual dichromatism is evident, with
males often being brightly ornamented whereas females
are cryptically colored. The only other Gonatodes reported to lack sexual dichromatism is the recently described
G. lichenosus from Sierra de Perijá in northwestern Venezuela (Rojas-Runjaic et al., 2010), which can be easily
distinguished by numerous other characters (i.e., larger
SVL, granular dorsal scales, rounded snout, clusters of
enlarged spine-like scales along flanks).
The new species differs from Gonatodes daudini in
other characteristics (data of the latter taxon in parentheses): dorsal coloration uniformly reddish brown (yellowish
to grayish-brown with one or more pairs of conspicuous
lateral ocelli), reddish brown iris (bright red iris), and a
vertically elliptical pupil (round). In this latter characteristic, G. naufragus differs from all other species of the genus except G. antillensis, which is currently believed to be
the only consistently nocturnal Gonatodes (Schargel et al.,
2010). Currently, it is not known whether G. naufragus is
also nocturnal in habits but the presence of a vertically elliptical pupil appears to indicate this possibility. The new
species can be easily separated from G. antillensis (data
of the latter taxon in parentheses) in having much larger
dorsals, with approximately 60 scales around midboby
(granular dorsals, with approximately 80 scales around
midbody), males with distinct escutcheon on abdomen
(escutcheon absent), and no sexual dimorphism in coloration (sexual dimorphism conspicuous; see Fig. 4A–B).
Sequence characteristics and phylogenetic
relationships
The selected model for all six partitions (all nucleotide sites separated by gene and codon position) under
AICc was HKY with or without gamma parameter depending on the partition. Because the nucleotide substitution
model was the same (HKY+G) for the first and second
codon positions for both genes, and because of the low
number of substitution for these codon positions, we
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
13
South American Journal of Herpetology, 8(1), 2013, 5–18
Figure 10. Consensus phylogenetic tree (consensus level: 50) of a maximum likelihood bootstrap analysis (bootstrap values shown on nodes) of two
nuclear genes (C-mos and NT3) for the genus Gonatodes.
14
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
South American Journal of Herpetology, 8(1), 2013, 5–18
decided to combine them into a single partition while
keeping them separated by gene. The “propose model” option in TREEFINDER as described above was conducted
once again with the new partition scheme yielding the
HKY+G model for all partitions except for the third codon position of C-mos, which was simply HKY. The ML
bootstrap consensus tree obtained is shown in Fig. 10.
The phylogenetic analysis suggests a strongly supported
(bootstrap value: 98) sister species relationship between
Gonatodes daudini and G. naufragus, corroborating with
our observations on the morphological similarities between both taxa. The relationship of this clade to other
Gonatodes remains uncertain, as the basal relationships
within the genus are poorly resolved.
of Cnemidophorus present on these islands. At least six
species of Cnemidophorus are distributed throughout
non-continental landmasses in the southern Caribbean
(Dutch and Venezuelan Antilles), whereas one closely
related taxon (C. vanzoi) is present on the tiny Maria
Major and Minor islands, off the southeastern coast of
St. Lucia (Ugueto and Harvey, 2010). The latter species
share numerous morphological characteristics with the
southern Caribbean Cnemidophorus (Harvey et al., 2012),
once again strengthening a probable zoogeographical link
between these islands. Considering the similarities, the
common ancestor of Gonatodes naufragus and G. daudini
could also have a South American origin. The putative ancestor must have been able to colonize these islands and
establish populations that gave rise to the two presently
known taxa.
DISCUSSION
The fact that Gonatodes naufragus and G. daudini
share unique morphological features within the genus,
along with the strongly supported sister relationship inferred, and low genetic divergence (uncorrected p is 0.007
and 0.008 for C-mos and NT3, respectively) leaves little
doubt of a close relationship and, perhaps, even relatively
recent divergence between both taxa. The relationship of
this clade to other Gonatodes remains uncertain, which is
consistent with the unresolved placement of G. daudini in
both the maximum parsimony and Bayesian analyses conducted by Schargel et al. (2010). However, we note that in
the Bayesian analysis conducted by Gamble et al. (2008),
G. daudini was recovered as the sister species to a clade
containing G. albogularis and G. vittatus. It is possible that
the higher resolution obtained by Gamble et al. (2008), as
far as basal relationships of Gonatodes, results from the
larger character dataset used by these authors.
The close phylogenetic relationship between Gonatodes naufragus (from La Blanquilla Island) and Gonatodes daudini (from Union Island, The Grenadines)
shows interesting parallels with that of other Caribbean
lizards. For example, species in the Anolis roquet series are
restricted to the southern Lesser Antilles and the islands
of La Blanquilla and Bonaire. Gorman and Stamm (1975)
examined the evolutionary relationships within the Anolis roquet series and concluded that A. blanquillanus from
La Blanquilla and A. bonairensis from Bonaire are closely
related to A. luciae from St. Lucia, north of the Grenada
bank. Yang et al. (1974) postulated two possible events for
how these anoles could have colonized the Lesser Antilles and the islands of the southern Caribbean. A primary
event of stepwise colonization of the Lesser Antillean island banks closest to the South American mainland followed by a secondary event where lizards from St. Lucia colonized La Blanquilla and then dispersed from the
latter westward towards Bonaire. An ancestor of South
American origin can also be inferred for whiptail lizards
Notes on the distribution of some other Caribbean
Gonatodes
The islands of Curaçao, Bonaire and La Orchila, as
well as the archipelagoes of Los Roques and Las Aves, are
all inhabited by at least one native species of Gonatodes.
The most widespread taxon is G. antillensis, found in Bonaire, Curaçao, Las Aves and Los Roques (although the
conspecificity of these populations has not been rigorously tested). Los Roques is also inhabited by G. aff. vittatus, whereas the small neighboring island of La Orchila
is populated by a taxon associated with G. albogularis, currently being described by the authors.
The situation in Aruba is less clear. Three species
of Gonatodes have been reported to occur on the island:
G. albogularis, G. antillensis and G. vittatus (van Buurt,
2001). Lidth de Jeude (1887) recorded G. antillensis from
Aruba and stated: “Many specimens were captured in Curaçao and Aruba”. Hummelinck (1940) did not find this
species on the latter island, in spite of extensively collecting there, and considered that Aruba could be safely
excluded as part of the distribution of G. antillensis.
However, van Buurt (2005) once again included G. antillensis as part of the Aruban herpetofauna and considered
that the species was probably introduced from Curaçao.
Cope (1885) reported G. albogularis from Aruba. Neither Ruthven (1923) nor Hummelinck (1940) recorded
new specimens from this island but van Buurt (2005)
still considered it part of the Aruban herpetofauna. According to van Buurt (pers. com.), he has never observed
either G. antillensis or G. albogularis on the island but reported them as present based on a checklist of the herpetofauna of Aruba by Odum (1992). Odum himself had
never observed G. albogularis or G. antillensis on Aruba
but preferred not to exclude them from his checklist
since his fieldwork on the island had been fairly short,
thus not being able to confidently demonstrate that the
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
15
South American Journal of Herpetology, 8(1), 2013, 5–18
lizards were not present on Aruba (van Buurt, pers. com.).
Whether G. albogularis and G. antillensis were or are still
present on Aruba cannot be assured at the present moment, but it is very likely that if they were ever found on
the island, this was due to human introductions. In the
past, building material (where these small lizards or their
eggs could have been inadvertently transported) would
ship from places such as Coro and Punto Fijo on nearby
mainland Venezuela (van Buurt, pers. com.), where G. albogularis is common. This could probably explain the possible historic occurrence of the latter species on Aruba.
The origins of G. antillensis on the island are more difficult to ascertain but they could also be attributed to a
mislabeled specimen.
Contrary to the situation of Gonatodes albogularis
and G. antillensis on Aruba, G. vittatus is quite common
and has been repeatedly recorded on the island (e.g., Ruthven, 1923; Hummelinck, 1940; van Buurt, 2005). However, G. vittatus also appears to have been introduced on
Aruba. Ruthven (1923) found it only in and about the
town of Oranjestad, although searches for this lizard were
carefully carried out in other localities of the island. Photographs of specimens from Aruba (e.g., photo 26 and 27,
p. 53 in van Buurt, 2005) show animals identical to those
found on the Venezuelan mainland. Purported absence
of native Gonatodes on Aruba is surprising because of its
proximity to the Venezuelan mainland and because the
island was probably connected to the mainland through
a land bridge at some point during the Pleistocene (van
Buurt, 2005). It would be the only southern Caribbean island lacking a native Gonatodes species. In contrast, Aruba
is home to other endemic lizards (Cnemidophorus arubensis and Phyllodactylus julieni), which belong to genera that
also have endemic taxa on other islands of the Dutch and
Venezuelan Antilles.
ACKNOWLEDGMENTS
The visit and discovery of the new species described
here was thanks to the development of the project: Recuperación de áreas del Caribe y/o de países miembros del ALBA,
under the direction of L.E. Sánchez. We greatfully thank
Luis Velasquez (Chipi) for the aquatic logistical support in
La Blanquilla and Los Hermanos archipelago. We thank E.
Briceño, A. Fernández, P. Granado, R. Guevara, L. Parra, J.
Reid, J.J. Rodríguez and A. Ruíz for their invaluable help
in the field. We also would like to thank G. van Buurt for
his helpful comments about the species present in Aruba
and Curaçao as well as to P.J.R. Kok and P. Granado for
their help with the figs. 4, 7 and map respectively. The
necessary permits were provided to Gilson A. Rivas by the
Ministerio del Poder Popular para el Ambiente, Venezuela
through the Facultad Experimental de Ciencias, Universidad del Zulia (collecting permit, Oficio No. 667, project
16
inventario nacional de Fauna), Maracaibo, and Fundación
La Salle de Ciencias Naturales (use of tissue samples, Oficio No. 0053, project sistemática, filogenía y taxonomía
de las lagartijas del género Gonatodes (Reptilia: Gekkonidae), Caracas. For help with molecular lab work we are indebted to C.L. Cox. Finally we would like to thank T.C.S.
Avila-Pires, R.C. Jadin P.J.R. Kok and an anonymous reviewer for their comments and suggestions to improve a
preliminary version of this manuscript.
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A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez
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South American Journal of Herpetology, 8(1), 2013, 5–18
APPENDIX I
List of comparative material examined
Gonatodes aff. albogularis (7). VENEZUELA: Dependencias Federales: La Orchila, MBLUZ 1035–1041.
Gonatodes antillensis (7). VENEZUELA: Dependencias Federales: Archipiélago de Las Aves (Barlovento), MBLUZ 999–1004.
Los Roques: Gran Roque, MHNLS 1031 [holotype of Gonatodes vitattus roquensis].
Gonatodes astralis (1). VENEZUELA: Bolívar: Serranía de los Pijiguaos, Distrito Cedeño, 600 m, MBLUZ 931 [paratype].
Gonatodes daudini (1). ST. VINCENT AND THE GRENADINES: Union Island, Water Rock Reserve on the northern slope of
Mt. Taboi above Chatham Bay (12°35’N, 61°25’W), MPM 33975 [paratype]
Gonatodes infernalis (2). VENEZUELA: Amazonas: sector El Infierno, on road Puerto Ayacucho-Gavilán, ca. 100 m, UTA
R-55378-79 [paratypes].
Gonatodes aff. vittatus (3). Los Roques: Gran Roque, MHNLS 1033, 1037, 1039 [paratypes of Gonatodes vitattus roquensis].
APPENDIX II
Genbank accession numbers for sequences (c-mos, nt3) used in the phylogenetic analysis
Gonatodes albogularis (GU139825, GU139877), Gonatodes albogularis (GU139826, GU139878), Gonatodes alexandermendesi
(GU139803, GU139856), Gonatodes antillensis (GU139827, GU139879), Gonatodes annularis (GU139807, GU139860),
Gonatodes astralis (GU139820, GU139873), Gonatodes caudiscutatus (EF534920, no sequence), Gonatodes concinnatus
(EF564070, no sequence), Gonatodes daudini (GU139832, GU139885), Gonatodes eladioi (EF564081, no sequence), Gonatodes falconensis (GU139829, GU139882), Gonatodes humeralis (GU139836, GU139889), Gonatodes infernalis (GU139816,
GU139869), Gonatodes ligiae (GU139831, GU139884), Gonatodes naufragus (JQ039940, JQ039941), Gonatodes ocellatus
(GU139839, GU139892), Gonatodes petersi (GU139841, GU139894), Gonatodes purpurogularis (GU139830, GU139883),
Gonatodes seiglei (GU139837, GU139890), Gonatodes taniae (GU139838, GU139891), Gonatodes timidus (GU139824, no
sequence), Gonatodes vittatus (GU139840, GU139893), Lepidoblepharis xanthostigma (GU139842, GU139895), Pseudogonatodes lunulatus (GU139844, GU139897), Pseudogonatodes manessi (GU139845, GU139898), Saurodactylus mauritanicus
(GU139846, GU139899), Sphaerodactylus molei (GU139843, GU139896).
18
A Distinctive New Species of Gonatodes (Squamata: Sphaerodactylidae) from Isla La Blanquilla, Venezuela,
with Remarks on the Distribution of Some Other Caribbean Sphaerodactylid Lizards
Gilson A. Rivas, Gabriel N. Ugueto, Walter E. Schargel, Tito R. Barros, Pablo Velozo, Luz Esther Sánchez