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Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)

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Old descriptions and lack of taxonomic revisions have caused confusion regarding diagnosis and identification of some species of Gonatodes. Gonatodes caudiscutatus (Günther, 1859) was described in the genus Gymnodactylus, based on four specimens (two males and two females) from the western slopes of the Andes, in Ecuador. Later this species was allocated to the genus Goniodactylus and more recently to Gonatodes, but it was never redescribed in a detailed way using the characters presently used to distinguish between species. The species was found in several herpetological collections misidentified as Gonatodes concinnatus and at least some of the records of G. caudiscutatus from Colombia were based on Gonatodes riveroi Sturaro and Avila-Pires, 2011. Herein we redescribe and diagnose Gonatodes caudiscutatus on the basis of 41 specimens from 11 localities in Ecuador, and photos of the syntypes of Gymnodactylus caudiscutatus and Gonatodes collaris, compare the species with its congeners, and present data on its distribution.
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Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859)
(Squamata: Sphaerodactylidae)
Author(s): Marcelo José Sturaro and Teresa C.S. Avila-Pires
Source: South American Journal of Herpetology, 8(2):132-145. 2013.
Published By: Brazilian Society of Herpetology
DOI: http://dx.doi.org/10.2994/SAJH-D-13-00002.1
URL: http://www.bioone.org/doi/full/10.2994/SAJH-D-13-00002.1
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Redescription of the Gecko Gonatodes caudiscutatus
(Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro1,2,* and Teresa C.S. Avila-Pires1,3
1 Museu Paraense Emílio Goeldi / CZO. Caixa Postal 399, CEP66017-970, Belém, Pará, Brazil.
2 Programa de Pós-Graduação em Zoologia, Universidade Federal do Pará - Museu Paraense Emílio Goeldi.
3 E-mail: avilapires@museu-goeldi.br
* Corresponding author, email: marcelosturaro@gmail.com
Abstract. Old descriptions and lack of taxonomic revisions have caused confusion regarding diagnosis and identification of some species
of Gonatodes. Gonatodes caudiscutatus (Günther, 1859) was described in the genus Gymnodactylus, based on four specimens (two males and
two females) from the western slopes of the Andes, in Ecuador. Later this species was allocated to the genus Goniodactylus and more recently
to Gonatodes, but it was never redescribed in a detailed way using the characters presently used to distinguish between species. The species
was found in several herpetological collections misidentified as Gonatodes concinnatus and at least some of the records of G.caudiscutatus
from Colombia were based on Gonatodes riveroi Sturaro and Avila-Pires, 2011. Herein we redescribe and diagnose Gonatodes caudiscutatus on
the basis of 41 specimens from 11 localities in Ecuador, and photos of the syntypes of Gymnodactylus caudiscutatus and Gonatodes collaris,
compare the species with its congeners, and present data on its distribution.
Keywords. South America; Ecuador; Taxonomy; Distribution; Gonatodes collaris.
Resumo. Antigas descrições e a ausência de revisões taxonômicas têm causado confusões na diagnose e identificação de algumas espécies de
Gonatodes. Gonatodes caudiscutatus (Günther, 1859) foi descrito no gênero Gymnodactylus, baseado em quatro espécies (dois machos e duas
fêmeas) provenientes da encosta oeste dos Andes, no Equador. Mais tarde essa espécie foi alocada no gênero Goniodactylus e posteriormente
em Gonatodes, mas nunca foi redescrita de forma detalhada e usando os caracteres atualmente utilizados para distinguir as espécies. Essa
espécie foi encontrada em diversas coleções herpetológicas identificadas erroneamente como Gonatodes concinnatus. Ademais, ao menos
alguns dos registros de G.caudiscutatus para a Colômbia foram baseados em Gonatodes riveroi Sturaro and Avila-Pires, 2011. Nesse trabalho,
nós redescrevemos e diagnosticamos Gonatodes caudiscutatus com base em 41 espécimes de 11 localidades do Equador, e fotos dos síntipos
de Gymnodactylus caudiscutatus e Gonatodes collaris, comparamos essa espécie com as demais do gênero, e apresentamos dados sobre sua
distribuição.
INTRODUCTION
The sphaerodactylid genus Gonatodes encompasses
at present 30 species distributed in Central and South
America and the Antilles (Peters and Donoso-Barros,
1970; Rivero-Blanco, 1979; Avila-Pires, 1995; Esqueda,
2004; Powel and Henderson, 2005; Cole and Kok, 2006;
Barrio-Amorós and Brewer-Carías, 2008; Rivas and
Schargel, 2008; Rojas-Runjaic etal., 2010; Schargel etal.,
2010; Sturaro and Avila-Pires, 2011; Kok, 2011; Rivero-
Blanco and Schargel, 2012, Rivas etal., 2013). At present
it is also found in Florida, United States, and the Gala-
pagos Archipelago, Ecuador, as a recent introduction
(Wright, 1983; Hoogmoed, 1989; Krysko and Daniels,
2005). A detailed revision of the genus, including the 15
species known at that time, was elaborated by Rivero-
Blanco (1979) as a PhD thesis, but it was never published.
Avila-Pires (1995) presented detailed descriptions of the
Brazilian species of Gonatodes, a trend that continued in
the description of new species since then (Esqueda, 2004;
Powel and Henderson, 2005; Cole and Kok, 2006; Barrio-
Amorós and Brewer-Carías, 2008; Rivas and Schargel,
2008; Schargel etal., 2010; Sturaro and Avila-Pires, 2011;
Kok, 2011; Rivero-Blanco and Schargel, 2012).
Günther (1859) described Gymnodactylus caudiscu-
tatus based on specimens from the western slopes of the
Andes, in Ecuador, collected by Mr. Fraser. O’Shaughnessy
(1875) mentioned the species as Goniodactylus caudiscuta-
tus. Boulenger (1885), in his catalogue of the lizards of the
British Museum, considered the species under the genus
Gonatodes, its present taxonomic position.
Gonatodes collaris Garman (1892) was described
based on specimens from Wreck Bay, Chatham (San Cris-
tobal) Island, in the Galapagos Archipelago, Ecuador, col-
lected by Dr. Georg Baur on 25 June 1891. Heller (1903)
mentioned that he did not find this species in his expedi-
tion to the type locality. Van Denburgh (1912) suggested
that Baur’s specimens were from Guayaquil (currently
Santiago de Guayaquil, Guayas, Ecuador), possibly mis-
labeled, because other collectors had not found this spe-
cies and Baur had also collected in Guayaquil. Barbour
and Loveridge (1929) also mentioned the type locality of
G.collaris as perhaps Guayaquil. Later on, however, Wood
(1939) and Mertens (1963) reported specimens from the
type locality.
Vanzolini (1965) proposed that Gonatodes collaris
was a junior synonym of G.caudiscutatus, which was cor-
roborated by Rivero-Blanco (1979) and Wright (1983).
South American Journal of Herpetology, 8(2), 2013, 132–145
© 2013 Brazilian Society of Herpetology
Submitted: 27 December 2012
Accepted: 02 August 2013
Handling Editor: Oscar Alberto Flores-Villela
doi: 10.2994/SAJH-D-13-00002.1
The last author studied 21 individuals from Progreso, a
small farming community approximately three miles in-
land from Wreck Bay, and could not find differences be-
tween G. collaris and G. caudiscutatus. Considering the
absence of endemic Gonatodes in Galapagos and the re-
striction of the species to man-modified habitats, Wright
(1983) suggested that it represented a recent, man-in-
duced introduction to the island, from mainland Guaya-
quil, an opinion followed by Hoogmoed (1989).
Shreve (1947) described Gonatodes caudiscutatus fal-
conensis based on 16 specimens, 14 of which were from
Pauji, Acosta District, Falcon State, Venezuela (holotype:
MCZ 48878, paratypes: MCZ 48718, MCZ 48879–85,
and NMB 13867–72 [NMB record numbers according to
Naturhistorisches Museum Basel website, NMB 2010],
and one from Ojo de Agua, Acosta District, Falcon State,
Venezuela (MCZ 49054), all collected by H.G. Kluger, be-
tween June 11 and August 30, 1945, except MCZ 48718,
collected in 1944. According to Shreve (1947), G.c.falco-
nensis differed from the nominal subspecies mainly by col-
or pattern and size. Vanzolini (1968) and Rivero-Blanco
(1968) elevated G.caudiscutatus falconensis to species, tak-
ing into account the geographic distance between the two
taxa, a position followed by Peters and Donoso-Barros
(1970). Rivero-Blanco (1979), in his unpublished revision
of the genus Gonatodes, showed that G.caudiscutatus and
G.falconensis differed in subcaudal scale pattern, corrobo-
rating their status as distinct species.
Gamble et al. (2008a) presented a molecular phy-
logeny of Gonatodes, where G.caudiscutatus and G.eladioi
Nascimento, Avila-Pires, and Cunha, 1987 formed a clade
sister to all other species they examined (11 in total). This
same relationship was found by Schargel etal. (2010), who
included 21 species in their molecular analysis. Carvajal-
Campo and Torres-Carvajal (2012) reported the distribu-
tion of this species in Ecuador and some notes on color
in life, sexual dimorphism, and morphological variations.
The first author, during a visit to the National Muse-
um of Natural History, Washington, D.C., United States,
while examining specimens of Gonatodes concinnatus from
several herpetological collections, observed that G. cau-
discutatus was frequently misidentified as G.concinnatus.
A similar misidentification had already been reported by
Vanzolini (1955) and the problem apparently persists.
The lack of a detailed description of G.caudiscutatus may
be one of the reasons for this confusion. Herein we pres-
ent a detailed description of G.caudiscutatus, compare it
with its congeners, and present data on its distribution.
MATERIALS AND METHODS
We analyzed a total of 362 specimens of Gonatodes,
including 41 specimens of G.caudiscutatus from 11 locali-
ties (Appendix). Specimens are deposited in the following
institutions: American Museum of Natural History, New
York, New York, USA (AMNH); California Academy of Sci-
ences, San Francisco, California, USA (CAS); University of
Kansas, Museum of Natural History, Lawrence, Kansas,
USA (KU); Museum of Comparative Zoology, Harvard
University, Cambridge, Massachusetts, USA (MCZ); Mu-
seu Nacional do Rio de Janeiro, Universidade Federal de
Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ); Coleção
Herpetológica Osvaldo Rodrigues da Cunha, Museu Par-
aense Emilio Goeldi, Belém, Pará, Brazil (MPEG); Museu
de Zoologia, Universidade de São Paulo, São Paulo, Bra-
zil (MZUSP); Sam Noble Oklahoma Museum of Natural
History, Norman, Oklahoma, USA (OMNH); Nationaal
Natuurhistorisch Museum, Leiden, The Netherlands
(RMNH); Texas Cooperative Wildlife Collection, Depart-
ment of Wildlife Science, Texas A&M University, Col-
lege Station, Texas, USA (TCWC); University of Michi-
gan, Museum of Zoology, Ann Arbor, Michigan, USA
(UMMZ); National Museum of Natural History, Wash-
ington, D.C., USA (USNM). Additionally we examined
photos of the syntypes of Gymnodactylus caudiscutatus
(BMNH 1946.9.7.6–9) and Gonatodes collaris (BMNH
1946.8.22.99, 1946.8.23.1–2; MCZ 9432). Comparisons
were based on the remaining specimens examined and
the literature (Roze, 1963; Donoso-Barros, 1966; Rivero-
Blanco, 1979; Mijares-Urrutia and Arends, 1996; Esque-
da, 2004; Powell and Henderson, 2005; Cole and Kok
2006; Barrio-Amorós and Brewer-Carías 2008; Rivas and
Schargel, 2008; Schargel etal., 2010, Rojas-Runjaic etal.,
2010, Kok, 2011; Rivas etal. 2013).
Measurements were taken with an electronic cali-
per (to the nearest 0.1mm) and when necessary under
a stereomicroscope. Species description follows Sturaro
and Avila-Pires (2011). Tail length was measured only in
specimens with intact, non-regenerated tail. Statistics are
presented as “minimum–maximum (mean ± standard de-
viation, N=sample size)”. Descriptions of color in life are
based on field notes and color photos.
RESULTS
Gonatodes caudiscutatus (Günther, 1859)
(Figs.1–3,5–6)
Gymnodactylus caudiscutatus Günther, 1859:410 (4 syn-
types, BMNH 1946.9.7.6–9, type locality “Andes of
western Ecuador,” collected by “Mr. Fraser” [Louis
Fraser, see Thomas 1906:9,33]).
Goniodactylus caudiscutatus; O’Shaughnessy, 1875:265.
Gonatodes caudiscutatus; Boulenger, 1885:61; Fowler,
1913:169; Burt and Burt, 1931:244 (part), 1933:2
(part); Vanzolini, 1955:124, 1965:18, 1968:18; Pe-
ters and Donoso-Barros, 1970:129; Rivero-Blanco,
1979:73; Wright, 1983:32; Hoogmoed, 1989:15;
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires 133
South American Journal of Herpetology, 8(2), 2013, 132–145
Kluge, 1991:11; Olmedo and Cayot, 1994:8; Rösler
2000:84; Kluge 2001:11; Torres-Carvajal, 2001:3;
Lehr 2002:115; Cole and Kok, 2006:4; Gamble
etal., 2008a:91; Gamble etal., 2008b:271; Rivas and
Schargel, 2008:41; Schargel et al., 2010:322; Stu-
raro and Avila-Pires, 2011:30; Carvajal-Campos and
Torres-Carvajal, 2012:525; Almendáriz and Carr,
2012:190.
Gonatodes caudiscutatus caudiscutatus Wermuth, 1965:43.
Gonatodes [caudiscutatus] caudiscutatus Shreve, 1947:520.
Gonatodes collaris Garman, 1892:83 (4 syntypes,
BMNH 1946.8.22.99, 1946.8.23.1–2 [formerly
1899.5.5.152–154], MCZ 9432, type locality Wreck
Bay, Chatham [San Cristobal] Island, Galapagos,
collected by Dr. Georg Baur, 25 June 1891); Heller,
1903:60; Van Denburgh, 1912:410; Barbour and
Loveridge, 1929:269; Wood, 1939:1; Mertens,
1963:21; Wermuth, 1965:44.
Diagnosis
A moderately large Gonatodes, with maximum
snout-vent length (SVL) of 45.4mm (USNM 285433). Pu-
pil round. No distinctly enlarged supraciliary spine. Scales
around midbody 89–101. Ventral scales in a longitudinal
row 43–51. Subdigital lamellae 17–21 under fourth fin-
ger, 21–27 under fourth toe; proximal ones as wide as dig-
its. Mostly three (in some cases two) lateral rows of scales
on distal portion of fingers and toes. Tail with midventral
scales distinctly wider than long, forming a repetitive se-
quence of one single midventral in contact laterodistally
with one scale per side, followed by a single midventral
in contact laterodistally with two scales per side (1’1”).
Adult males with elongate, orange-yellow spots on dor-
sal surface of head, including a longitudinal stripe along
each canthus rostralis and recurved, intermittent stripes
posteriorly; an antehumeral white vertical stripe, anteri-
orly bordered by black; a posthumeral ocellus, white with
black margin; and orange-yellow gular region with some
gray spots lateroventrally.
Gonatodes caudiscutatus presents a subcaudal se-
quence 1’1”, which differs from all other species except
G. eladioi, G. infernalis Rivas and Schargel, 2008, G. li-
chenosus Rojas-Runjaic, Infante-Rivero, Cabello and Ve-
loso, 2010, G.naufragus Rivas, Ugueto, Schargel, Barros,
Velozo & Sánchez, 2013, G. superciliaris Barrio-Amorós
and Brewer-Carías, 2008, and Gonatodes timidus Kok,
2011. Gonatodes seigliei Donoso-Barros, 1966 presents
a combined pattern (1’1” + 1’1’1”). Gonatodes caudiscu-
tatus differs from G.eladioi in reaching SVL of 45.4mm
(<40mm in G.eladioi), by the absence in males of a light
vertebral band (present), and by presenting usually three
lateral scales on fingers and toes (two). Gonatodes cau-
discutatus is smaller than G. infernalis (SVL > 55 mm),
presents a posthumeral ocellus on each side (no ocellus
in G.infernalis), and usually three lateral scales on fingers
and toes (usually four). It differs from G.lichenosus by pre-
senting male color pattern distinct from that of females,
with orange stripes on head (no sexual dichromatism and
no vivid colors on head in G.lichenosus), by the absence
of clusters of enlarged, conical to spinelike scales on body
(present), and by the presence of 17–21 lamellae under
fourth finger (14–16) and 21–27 under fourth toe (18–
19). Gonatodes caudiscutatus is larger than G. naufragus
(SVL<27.2mm), and it presents a posthumeral ocellus
at each side (no ocellus in G.naufragus). From G.seiglei
it differs in the absence of body vermiculations in males
(body bluish-grey and black vermiculated in G. seiglei).
It differs from G.superciliaris by the absence of an elon-
gate supraciliary spine and of clusters of enlarged, spine-
like scales on flanks (both present in G.superciliaris), and
by the presence of a posthumeral ocellus (absent). Go-
natodes caudiscutatus differs from G.timidus Kok, 2011,
by its smaller size (adult reaching to nearly 51.6 in SVL
in G.timidus), and the presence of a posthumeral ocellus
at each side (no ocellus in G.timidus). Apart from differ-
ences in caudal sequence, G. caudiscutatus differs from
G. albogularis (Duméril and Bibron, 1836), G. antillensis
(Lidth de Jeude, 1887), G. atricucullaris (Noble, 1921),
G.daudini Powell and Henderson, 2005, G.petersi Don-
oso-Barros, 1967, G.vittatus (Lichtenstein and von Mar-
tens, 1856) and G.humeralis (Guichenot, 1855) in num-
ber of lateral scales on fingers and toes distally (mostly
three versus two). G.caudiscutatus is easily differentiated
from G. alexandermendensi Cole and Kok, 2006, G.astra-
lis Schargel, Rivas, Makowsky, Señaris, Natera, Barros,
Molina and Barrio-Amorós, 2010, G. hasemani Griffin,
1917, and G.superciliaris by the presence in these species
of an elongate supraciliary spine; from G. antillensis by
the shape of pupil, respectively round versus vertical; and
from G.daudini by the distinctly larger dorsals in this spe-
cies (89–101 scales around midbody in G.caudiscutatus,
39–44 in G.daudini).
Description
Maximum SVL in males 44.3mm (USNM 285435),
in females 45.4mm (USNM 285433). Tail round in cross
section, tapering toward tip, 1.0–1.2 (1.11±0.06, N=13)
times SLV. Head length 0.22–0.26 (0.24±0.01, N=41)
times SLV, 1.3–1.6 (1.43±0.06, N=39) times as long as
wide, 1.3–1.6 (1.43±0.08, N=39) times as wide as high.
Snout round, moderately elongate (Fig.1A), gently slop-
ing toward top of head. Neck slightly narrower than head
and anterior portion of body. Body cylindrical. Limbs
well-developed, lower arms 0.09–0.14 (0.11 ± 0.01,
N =41) times SVL, lower legs 0.12–0.20 (0.14 ± 0.01,
N=41) times SVL.
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires
134
South American Journal of Herpetology, 8(2), 2013, 132–145
Figure1. Gonatodes caudiscutatus from Palenque, Rio Palenque Science Center, Los Rios, Ecuador. (A)Dorsal and (B)ventral views of head; ventral views
of (C)right hand and (D)right foot (RMNH 40149); (E)ventral view of tail (RMNH 40151). Scale bar=5mm.
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires 135
South American Journal of Herpetology, 8(2), 2013, 132–145
Rostral convex, 1.6–2.2 (1.88±0.12, N=41) times
as wide as high; mid-posteriorly with a shallow depres-
sion and a longitudinal cleft, and slightly indented by
0–2 medial postrostrals. Postrostrals 2 (9.8%), 3 (70.7%)
or 4 (19.5%), lateral ones (supranasals) distinctly larger
than medial one(s) (N=41) (Fig.1A). Nasal bordered by
rostral, first supralabial, 3 postnasals, and supranasal; in-
ternostril distance 0.20–0.28 (0.24±0.02, N=39) times
head width. Supranasal roughly oval or semicircular, 1.0–
2.1 (1.30±0.20, N=41) times as wide as long. Postnasals
similar in size to adjacent loreals. Scales on snout convex,
hexagonal to round, juxtaposed, relatively uniform in
size. Canthus rostralis rounded. Loreal region with scales
slightly more elongate than those on snout, largest on
row adjacent to supralabials, 8–12 (10.0±0.9, N=41) lo-
reals on a line between postnasals and anterior corner of
eye. Top and posterior portion of head, as well as supraor-
bital region, with granular scales. A short supraciliary flap
present, anteriorly with 7–11 (8.8±1.0, N=41) enlarged
and flattened scales, among which 1–5 (2.6±0.9, N=35)
small, conical spines. Pupil round, eye diameter 0.19–0.24
(0.21±0.01, N= 39) times head length. Scales on tem-
poral region similar to those on top of head. Ear-opening
much smaller than eye, oval, posterior to, and at same lev-
el of, commissure of mouth. Supralabials 6–8 (6.8±0.6,
N=41), distinctly enlarged anteriorly and decreasing in
size posteriorly, 4–6 (5.2±0.4, N=41) to below center of
eye; followed to corner of mouth by small scales.
Mental large, distinctly wider anteriorly than poste-
riorly, with posterior margin forming a wide angle, 0.9–1.4
(1.11±0.08, N=41) times as wide as long; two (90.2%)
or three (9.8%) postmentals (Fig.1B). Scales on chin flat,
smooth, polygonal, juxtaposted, largest anteriorly, de-
creasing in size posteriorly. Infralabials 5–7 (5.8± 0.5,
N=41), distinctly enlarged anteriorly and decreasing in
size posteriorly, 3–5 (3.9±0.6, N=41) to below center of
eye, followed to corner of mouth by small scales.
Scales on nape small and granular, becoming slightly
larger on sides of neck. Scales on throat anteriorly granu-
lar, posteriorly flat, smooth, hexagonal or round, increas-
ing in size toward the posterior region, with a short tran-
sitional zone between the anterior and posterior parts.
Dorsals granular, increasing in size toward flanks.
Ventrals larger than dorsals, roughly hexagonal, flat,
smooth, imbricate, in oblique rows; 43–51 (46.7 ± 2.2,
N=39) scales along the midventral line between anterior
margin of forelimbs and vent; 16–21 (18.7±1.2, N=40)
scales in a transverse line at midbody, with a short transi-
tional zone between ventrals and scales on flanks. Scales
around midbody 89–101 (94.8±3.2, N=39). Scales on
preanal plate similar to ventrals, except for those bor-
dering vent, which are very small. Escutcheon present in
males on posterior portion of belly and on three to five
(mostly four) rows (N=8) (body–knee direction) on ven-
tral surface of thighs.
Scales on anterodorsal surface of forelimbs flat,
smooth, roundish, imbricate, largest close to the wrist;
on posterodorsal and ventral surfaces convex, smooth,
rhomboid, juxtaposed, relatively small. Scales on an-
terodorsal and ventral surface of thighs and ventral sur-
face of hind limbs flat, smooth, rhomboid, imbricate and
relatively large; on posterodorsal surface of thighs and
dorsal surface of lower legs smaller, convex, smooth,
round, subimbricate.
Lamellae under second (II) through fourth (IV) fin-
gers (proximal lamellae between parentheses): II: 13–16
(5–7), III: 15–21(5–8), and IV: 17–21(6–9) (Fig.1C). La-
mellae under second (II) through fourth (IV) toes (proxi-
mal lamellae between parentheses): II: 13–15 (5–6), III:
17–22 (6–9), and IV: 21–27 (11–15) (Fig.1D). Claws ex-
posed, non-retractile, between two basal scales. Fingers
and toes with three, occasionally two, lateral scales be-
tween fourth (counted from the claw toward the hand or
foot) subdigital lamellae and the dorsal scale.
Scales on tail dorsally and laterally relatively small,
rhomboid, flat, smooth, imbricate. On ventral surface of
tail scales smooth, flat, imbricate, increasing in size to-
ward midventral line. Midventral scales, except close to
the base, distinctly wider than long, forming a repetitive
sequence of one single midventral in contact laterodistal-
ly with one scale per side, followed by a single midventral
in contact laterodistally with two scales per side – 1’1” in
the coding by Avila-Pires (1995) (Fig.1E).
Color in preservative
Males with dorsal surface of head dark brown, with
a longitudinal white stripe at each side along the canthus
rostralis, from the snout to the supraocular region, and
two recurved (approximately U-shaped) white stripes
formed by elongate, irregular spots, anterior one wider
and starting on posterior corner of eyes, posterior one
starting on supralabials. A third, partial U-stripe may be
present, as well as other irregular spots on top of head.
A white, vertical antehumeral stripe, bordered anteriorly
by a black band. Dorsum blackish-gray with a paler verte-
bral band which varies from almost inconspicuous to con-
spicuous, bordered by a series of black spots with white
posterior margin. Flanks blackish-gray with white, bor-
dered by black, small spots; a conspicuous white ocellus,
bordered by black, is present posterior to arm insertion.
Limbs grayish-brown. Ventral surface of head and gular
region cream color with brown stripes and spots; belly
and underside of limbs dark gray (except on escutcheon
area, which is light gray), with a sharp delimitation on an-
terior level of arms between the cream and dark gray ar-
eas. Tail blackish-gray dorsally, dark gray ventrally; origi-
nal tail distally with white bands that form complete rings
around the tail (Figs.2A–B). Some males present the head
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires
136
South American Journal of Herpetology, 8(2), 2013, 132–145
Figure2. Variation in color pattern of Gonatodes caudiscutatus (in preservative). (A)Dorsal and (B)ventral color pattern variation in males (left USNM
285431, right USNM 285426); (C)dorsal (USNM 285427, USNM 285432 and USNM 285428) and (D)ventral (USNM 285427 and USNM 285428) color
pattern variation in females and juvenile. Scale bar=10mm.
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires 137
South American Journal of Herpetology, 8(2), 2013, 132–145
pattern as described above, but otherwise the pattern is
similar to that of females and juveniles (see below). It
may represent a transitional pattern, of subadults, but we
found specimens up to 40mm SVL with such a pattern,
while other specimens of 36mm SVL presented already
the typical adult male pattern.
Females and juveniles dorsally predominantly red-
dish-brown or grayish-brown. Head with two or three
recurved beige stripes posteriorly, similar to, but less evi-
dent than, those in males. A beige or tan vertebral band
starts on nape, continues along the body and occupies all
dorsal surface of the tail; on the body it is bordered at each
side by black spots, most of them with a dirty white poste-
rior margin; along the tail the spots at each side fuse and
form transverse bands, the white band becoming longer
toward the tip of the tail. Flanks with dark-brown or black
blotches, and a beige or tan, black margined, posthumer-
al ocellus (absent in TCWC 24958 and USNM 285436).
Some specimens also with small dirty-white spots on
flanks. Ventral surface predominantly cream or beige,
with brown irregular stripes and spots under head and on
gular region, brown-peppered on belly, brown and cream
variegated under anterior portion of tail. Underside of
(original) tail distally with alternating brown and white
bands (Figs.2C–D).
Color in life
USNM 285431, a male 43.5 mm SVL (shown in
Fig. 2A–B after preservation), was described in life as
“dark, orange throat and pattern on head; few blue spots
on sides” (from USNM catalogue, collector Eugene W.
Schupp). USNM 285426, a male 40.2mm SVL (Fig. 3),
was described as having “orange-yellow chin with white
anterior-medial spot; posthumeral ocellus light blue with
Figure3. Gonatodes caudiscutatus from Santo Domingo de los Colorados, 47kmS of Centro Científico, Rio Palenque, Pichincha, Ecuador. (A)Dorsal
view of USNM 285426, a male with SVL=40.2mm. (B)Ventral view of same male. (C)Dorsal view of USNM 285427, a female with SVL=38.6mm. All
photos by Roy W. McDiarmid.
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires
138
South American Journal of Herpetology, 8(2), 2013, 132–145
black; streaks on head rich yellow-orange” (R. McDiar-
mid, pers.comm.). From Fig.3A–B we can also see that
the specimen has a general dorsal color (head, body, fore-
limbs and part of hind limbs) with different shades of
gray, while tail and part of hind limbs are light drab with
black and beige spots; ventrally belly is light gray, tail
gray with white spots anteriorly, white and black bands
posteriorly.
For females, the only available note of color in life
for the specimens examined is of USNM 285432 (SVL
41.8 mm), described as “brown and black; white bands
on tip of tail” (from USNM catalogue, collector Eugene
W. Schupp). This description basically agrees with the
photo in life of USNM 285427, another female with SVL
38.6mm (Fig.3C).
Garman (1892), in the description of Gonatodes col-
laris, and Carvajal-Campos and Torres-Carvajal (2012)
also described the color in life of G.caudiscutatus.
Distribution
Gonatodes caudiscutatus occurs west of the Andes
in Peru (Departamento Tumbes) and Ecuador (Bolívar,
El Oro, Guayas, Los Rios, Manabi, Santo Domingos de
los Tsáchilas, Pichincha provinces). East of the Andes
it has been recorded in Morona-Santiago, Pastaza,
Tungurahua and Zamora-Chinchipe provinces, in Ec-
uador (see Discussion). It is found at present also on
Chatham (San Cristóbal) Island, Galapagos Archipel-
ago, probably as a recent introduction (Fowler, 1913;
Rivero-Blanco, 1979; Wright, 1983; Hoogmoed, 1989;
Lehr, 2002; Gamble et al., 2008a,b; Carvajal-Campos
and Torres-Carvajal, 2012; examined specimens;
Fig.4).
Peters and Donoso-Barros (1970) reported the oc-
currence of Gonatodes caudiscutatus on the Pacific coast
of Colombia, but apparently no specimen is known
from this country. Werner (1900) reported G. caudiscu-
tatus from Orocué and Bogota, Colombia. Burt and Burt
(1931, 1933) and Burt (1932) also reported G. caudiscu-
tatus from Colombia (Villavicencio, Gragoa, Guaicaraino
and Medina). Sturaro and Avila-Pires (2011) identified
the specimens from Villavicencio (part of material exam-
ined by Burt and Burt, 1931; AMNH 35280–87, AMNH
35290–91) and other localities from central Colombia as
G.riveroi, a species of the G.concinnatus group that they
described as new.
Figure4. Distribution of Gonatodes caudiscutatus. Closed circles= material examined; open circles= data from literature (Fowler, 1900; Rivero-Blanco,
1979; Wright, 1983; Hoogmoed, 1989; Lehr, 2002; Gamble etal., 2008a,b; Carvajal-Campos and Torres-Carvajal, 2012); red triangle= type locality of
Gonatodes collaris; square= specimen from Chichirota, Pastaza province, Ecuador (USNM 234501).
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires 139
South American Journal of Herpetology, 8(2), 2013, 132–145
DISCUSSION
Günther (1859), in the original description of Go-
natodes caudiscutatus, did not mention the number of
specimens of the type series. Boulenger (1885), in the
catalogue of lizards in the British Museum of Natural
History, mentioned four syntypes of G.caudiscutatus, of
both sexes. Rivero-Blanco (1979), based on information
by Dr. Garth Underwood, reported two males and two
females. Regarding G. collaris, Barbour and Loveridge
(1929), in the catalogue of types of reptiles and amphib-
ians in the Museum of Comparative Zoology, mentioned
one syntype and that “the other cotype mentioned in the
original description cannot be found anywhere”. Rivero-
Blanco (1979) reported three syntypes in the collection
of BMNH, all males. Even though Garman (1891:84),
in the original description, did not say how many speci-
mens he had before him, while describing the condition
of scales under tail (p. 84) refers to four specimens, which
is in agreement with the three specimens in BMNH and
one in MCZ. BMNH specimens may have been part of the
material from the Galapagos Islands purchased by this
museum in 1899, “containing numerous types of species
described by Dr. Baur” (Boulenger, 1906). Figures 5 and 6
present both series of syntypes.
Gonatodes caudiscutatus has been misidentified in
collections as G.concinnatus, which occurs also in Ecua-
dor east of the Andes. They differ, among other things, in
subcaudal pattern (1’1” in G.caudiscutatus; 1’1’1”, in some
cases with a proximal segment 1’1’2”, in G.concinnatus)
Figure5. Syntypes of Gonatodes caudiscutatus (voucher numbers from left to right: BMNH 1946.9.7.6–9). (A) Dorsal and (B) ventral views. Scale
bar=10mm. Photos by Gabriela Bittencourt.
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires
140
South American Journal of Herpetology, 8(2), 2013, 132–145
Figure6. Syntypes of Gonatodes collaris (voucher numbers from left to right: BMNH 1946.8.22.99; 1946.8.23.01–02, and MCZ 9432), a synonym of
G.caudiscutatus. (A)Dorsal and (B)ventral views. Scale bar=10mm. Photos provided by Gabriela Bittencourt (BMNH specimens), Museum of Compara-
tive Zoology and Harvard University (MCZ specimen).
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires 141
South American Journal of Herpetology, 8(2), 2013, 132–145
and in color pattern: G. caudiscutatus presents a white
antehumeral stripe (bordered anteriorly by black) and
a posthumeral ocellus, while G. concinnatus presents a
white bar (with black margins) in suprahumeral position;
besides, G. concinnatus has vermiculations on body, not
present in G.caudiscutatus.
Records of Gonatodes caudiscutatus east of the An-
des, in Ecuador, are recent and, as few species are known
to occur on both sides of the Andes, they raise questions
about either their identity or how they crossed the Andes.
Gamble (2008a,b), in a molecular phylogenetic analysis,
included a single tissue sample of G.caudiscutatus from
Limón, Ecuador (KU 218359, in Morona-Santiago prov-
ince, according to KU Herpetological Collection Online
Catalogue). Carvajal-Campos and Torres-Carvajal (2012)
reported more specimens from Limón, and from two
additional localities east of the Andes (Tungurahua and
Zamora-Chinchipe provinces). They compared specimens
from both sides of the Andes and found only small but
statistically significant differences on the mean number
of loreals and supraciliary spines. However, as remarked
by the authors, the sample was small and may have bi-
ased the results. A fifth locality is that of USNM 234501,
from Chichirota, Pastaza province (collected by Ramon
Olalla and Pablo Mena on February 1954), which extends
the known distribution almost 200 km eastward, well
into Amazonian Ecuador. This specimen was examined
by MJS and it was similar to those from west of the An-
des. Since the specimens reported by Carvajal-Campos
and Torres-Carvajal (2012) were found only near human
settlements, these authors raised the possibility that the
species colonized the areas east of the Andes through the
accidental dispersal by human settlers. Its recent intro-
duction in Galapagos (as data seem to indicate) reinforces
this possibility. On the other hand, its present occurrence
in at least four different places rather far apart east of the
Andes makes it less likely. Phylogeographic studies would
be useful to give an idea about how populations west and
east of the Andes relate to each other and for how long
they have been separated.
ACKNOWLEDGMENTS
We thank David Kizirian (AMNH), Jens Vindum
(CAS), Jucivaldo D. Lima (IEPA), Linda Trueb (KU), Rick
Feeney (LACM), Jose Rosado and Jonathan Losos (MCZ),
José P. Pombal-Jr and Ulisses Caramaschi (MNRJ), Hus-
sam Zaher (MZUSP), Janalle Caldwell (OMNH), Ronald
de Ruite (RMNH), Toby J. Hibbtts and Lee A. Fitzgerald
(TCWC), Greg Schneider (UMMZ), Ron Heyer, Roy McDi-
armid, Traci Hartsell, Addison Wynn and Robert Wilson
(USNM), for kindly loaning or allowing access to speci-
mens under their care. Gabriela Bittencourt provided
photos and Colin McCarthy (BMNH) important data on
the syntypes of Gymnodactylus caudiscutatus and three
syntypes of Gonatodes collaris. Jonathan B. Losos, Jose
Rosado and Jon Woodward (MCZ and Harvard Univer-
sity) provided photos and information about the fourth
syntype of G.collaris. Roy McDiarmid provided photos of
Gonatodes caudiscutatus in life. We also thank Marinus S.
Hoogmoed and Mateus Ferreira for useful comments re-
garding a previous version of this manuscript, and Aaron
M. Bauer and an anonymous reviewer for their comments
and suggestions to improve the manuscript. MJS received
a Short Term Visit Award from the Smithsonian Institu-
tion, sponsored by Ron W. Heyer, and is presently sup-
ported by a scholarship from Conselho Nacional de De-
senvolvimento Científico e Tecnológico (CNPq; process
PROTAX 159253/2010-1). TCSAP receives a research fel-
lowship from CNPq (process 304199/2010-9).
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APPENDIX
Specimens examined
Gonatodes albogularis (6): CUBA: San Carlos: Guantanamo (USNM 63220). NICARAGUA (MNRJ 3164, 10670). PANAMÁ:
Bocas del Toro; Cayo Nancy (USNM 338485, USNM 338487, USNM 338498).
Gonatodes antillensis (2): VENEZUELA: Orchilla Island (USNM 79231), Netherlands Antilles, Curacao, Ft. Nassau, Willam-
stad (USNM 94980).
Gonatodes annularis (22): BRAZIL: Amapá: Mazagão, Cachoeira Inajá, Rio Camaipi (MPEG 2667), Oiapoque, km90 of BR-
156, Aldeia Tukai (MPEG 24439), Serra do Navio (MPEG 15080, MPEG 15087, MPEG 15100, MPEG 15148, MPEG 19592,
MPEG 19627–31), Serra do Navio, Área Urucum (MPEG 19213); Pará: Cachoeira Porteira, Trombetas River (MPEG 16264),
Oriximiná, Cruz Alta, 6km from Trombetas River (MPEG 15396), Oriximiná, Porto Trombetas (MPEG 24198). FRENCH
GUIANA: Petit Saut, River Sinnamary (MPEG 15826, MPEG 15830, MPEG 15838, MPEG 15841–42). SURINAME: (MPEG
21823).
Gonatodes caudiscutatus (49): ECUADOR: “Andes of western Ecuador” BMNH 1946.9.7.6–9 (photos of the syntypes of
Gymnodactylus caudiscutatus); El Oro: 6mi. S. of Santa Rosa (CAS 85423); Guayas: Guayaquil (MZUSP 9181–90, MZUSP
54370–72; TCWC 24958-59), Balzar (MZUSP 9191), Bucay, region about (USNM 166147), 21km SW El Empalme, on
Quevedo-Guayaquil Mwy (USNM 166148), 3mi, E Milagro, Hacienda San Miguel, ca. 200mi (USNM 166144-45); Mana-
bi: 2km E Puerto de Cayo, ca 20mi. (USNM 166149-50), Chone, Hotel Turin (USNM 234503); Los Rios: Palenque, Rio
Palenque Science Center (RMNH 40149–153); Pastaza: Chichirota (USNM 234501); Pichincha: Santo Domingo de los Colo-
rados, 47kmS of, Centro Cientifico Rio Palenque (USNM 285426–36). Galapagos: Chatham Island, Wreck Bay BMNH
1946.8.22.99 and 1946.8.23.1–2, MCZ 9432 (photos of the syntypes of Gonatodes collaris).
Gonatodes ceciliae (5): TRININDAD: Arima, Spring Hill Estate Arima Vallei (RMNH 10160), Maracas, River ly Maracas
(RMNH 14913). VENEZUELA: Sucre: San Juan de lãs Galdonas (RMNH 40134–35; MZUSP 53553).
Gonatodes concinnatus (98): ECUADOR: “Pallatanga” BMNH 1946.9.7.15 (photos of syntype of Goniodactylus buckleyi);
Canelos: BMNH 1046.9.7.10–12 (photos of syntypes of Goniodactylus concinnatus), BMNH 1046.9.7.13–14 (photos of syn-
types of Goniodactylus buckleyi); Sucumbíos: Shushufindi, Rio Aguarico (USNM 234497), Santa Cecilia (KU 105290–91, KU
109572–73, KU 146643, KU 146645, KU 147588–89, KU 152496), Lago Agrio (KU 126683, KU 126685–86, KU 126688,
KU 297992), Limoncocha (LACM 73293–300, MZUSP 54655, KU 144378, MCZ 86414), Reserva de Produccíon Faunística
de Cuyabeno (RPF Cuyabeno), Estación Biológica e la Pontificia Universidade Catolica del Ecuador (OMNH 36406–11,
OMNH 36424). PERU: Loreto: San Jacinto (KU 222138), Moropón (TCWC 36665–71, TCWC 36743–46, TCWC 38994,
TCWC 41213–20, TCWC 41902–04, TCWC 41906–08, TCWC 42706–13, TCWC 42828, TCWC 43332, TCWC 44230,
TCWC 44283, TCWC 41763, MZUSP 28248–49, MZUSP 28260–63, MZUSP 28273–79, MZUSP 28311–13, MZUSP 28319,
MZUSP 28375, MZUSP 39218).
Gonatodes eladioi (1): BRAZIL: Pará: Marabá, Salobo, Serra dos Carajás (MPEG 18020).
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires
144
South American Journal of Herpetology, 8(2), 2013, 132–145
Gonatodes hasemani (22): BRAZIL: Pará: Juruá, Rio Xingu (MZUSP 67240–57); Rondônia: Porto Velho, Área de inunda-
ção da UHE de Samuel (MPEG 15569–71), Porto Velho, Rio Yamary, Cachoeira do Samuel, Território do Guaporé (MNRJ
10678).
Gonatodes humeralis (16): BRAZIL: Pará: Piçarra (MNRJ 16107), Tucuruí, Lago da UHE Tucuruí (MPEG 21881–87). PERU:
Loreto: Moropón (MZUSP 28280–87).
Gonatodes ligiae (10): VENEZUELA: Barinas: 8kmSW Santa Barbara, on San Cristobal Highway (TCWC 60231), Barinitas
(TCWC 57236–44).
Gonatodes nascimentoi (holotype and 21 paratypes): Brazil: Almeirim, Monte Dourado, forest near Estação Ecológica do Jari
(MPEG 23822–27, MPEG 27719); Anapu, Fazenda Caracol, right margin of Rio Xingu (MPEG 25596 [holotype], MPEG
25164, MPEG 25595, MPEG 25597, MPEG 25598–25601), Portel, Fazenda Riacho Monte Verde (MPEG 24643, MPEG
24644–50).
Gonatodes riveroi (holotype and 62 paratypes): COLOMBIA: Boyacá: Garagoa (CAS 71238, USNM 84968, USNM 92490–92);
Boyacá-Cundinamarca-Meta border: Guaucáramo (USNM 84972); Meta: San Juan de Arama (KU 192635-36); Villavicencio
(MZUSP 49153 [holotype], MCZ 132796–97; AMNH 35280–87, AMNH 35290–91, KU 110579–80, KU 151933–35, MCZ
19213, MCZ 77388–89, MCZ 110009–14, MCZ 110741–42, MZUSP 2145–46, MZUSP 49152, MZUSP 49154, USNM
84969); Villavicencio, Finca “El Buque” (MZUSP 49155–62); Villavicencio, Instituto “Roberto Franco”(MCZ 77391–97,
MCZ 146137); 5kmNE of Villavicencio (UMMZ 1272206); Rio Negro, c.12km from Villavicencio, Finca “Las Orquideas”
(MCZ 150022); Güejar River, Finca Guadualito (MZUSP 44777); Puerto Lopez (AMNH 97350, AMNH 98406).
Gonatodes rozei (19 paratypes): VENEZUELA: Miranda: 32.9kmN of Altagracia de Orituco, Guatopo National Park (TCWC
59300–59304, MZUSP 57574 [formerly TCWC 59305]); 32.9-34.4kmN of Altagracia de Orituco, Guatopo National Park
(TCWC 59306). Guarico: 25.2kmN of Altagracia de Orituco, Guatopo National Park (TCWC 59318); 5kmE of «Puesto la
Colonia,» Guatopo National Park (TCWC 59319–59320); 7kmW of San Francisco Macaira, La Elvira (TCWC 59309-59317,
TCWC 59321).
Gonatodes taniae (7): VENEZUELA: Aragua: Rancho Grande (MZUSP 57575; USNM 165605, USNM 196337), Rancho
Grande, Biological Station (USNM 142390), Rancho Grande, Parque Nacional Henri Pittier (USNM 247762-63; USNM
259166).
Gonatodes tapajonicus (18): BRAZIL: Pará: Cachoeira do Limão, Rio Tapajós (MZUSP 53669, MZUSP 53671–74, MZUSP
53676–77), Itaituba: Comunidade Aldeia Nova (MPEG 27708–27718).
Gonatodes vittatus (2): TRINIDAD: St. Patrick, Chathan Beach, on Erin Bay (USNM 314695–96).
Redescription of the Gecko Gonatodes caudiscutatus (Günther, 1859) (Squamata: Sphaerodactylidae)
Marcelo José Sturaro and Teresa C. S. Avila-Pires 145
South American Journal of Herpetology, 8(2), 2013, 132–145
... Sturaro & Avila-Pires, 2011 (Fig. 1C) Especie descrita recientemente, se consideraba que el taxón pertenecía a Gonatodes concinnatus sensu lato. Así mismo, la especie fue confundida con G. caudiscutatus (Günther 1859), pero los ejemplares pertenecen a G. riveroi (Sturaro & Avila-Pires 2013). ...
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