Content uploaded by Marcelo José Sturaro
Author content
All content in this area was uploaded by Marcelo José Sturaro on Oct 23, 2014
Content may be subject to copyright.
Accepted by A. Bauer: 28 Mar. 2011; published: 6 May 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2869: 1–36 (2011)
www.mapress.com/zootaxa/Article
1
Taxonomic revision of the geckos of the Gonatodes concinnatus complex
(Squamata: Sphaerodactylidae), with description of two new species
MARCELO JOSÉ STURARO1,2 & TERESA C.S. AVILA-PIRES1
1Museu Paraense Emílio Goeldi / CZO, CP 399, 66017-970 Belém, Pará, Brasil.
E-mail: (MJS) marcelosturaro@gmail.com and (TCSAP) avilapires@museu-goeldi.br
2Programa de Pós-Graduação em Zoologia, Universidade Federal do Pará / Museu Paraense Emílio Goeldi.
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1
Resumo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Gonatodes concinnatus (O’Shaughnessy, 1881). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Gonatodes ligiae Donoso-Barros, 1967 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Gonatodes tapajonicus Rodrigues, 1980 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Gonatodes nascimentoi sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
Gonatodes riveroi sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Species comparisons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Key to the species of Gonatodes concinnatus complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32
Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33
Abstract
The Gonatodes concinnatus complex, as here considered, consists of Gonatodes species characterized by a white
suprahumeral spot with black margins; vermiculations or not on back; and transversely enlarged scales under the tail,
showing the sequence 1’1’1”, and in some specimens 1’1’2” on the anterior portion. Two species are presently recognized
in this Amazonian complex: G. concinnatus and G. tapajonicus. New material collected in eastern Amazonia (states of
Pará and Amapá, Brazil) made it necessary to review these species. We analyzed several populations within this complex,
from Colombia, Ecuador, Peru, Venezuela and Brazil, including these new records. Specimens were separated in groups
defined on basis of color pattern. Stepwise discriminant function analyses were then performed to compare the external
morphology (measurements and scale counts) in these groups. Results support recognition of five taxa, corresponding to
G. concinnatus from western Amazonia, in Ecuador and northern Peru; G. l ig ia e from northwestern Venezuela (east of the
Andes); G. tapajonicus, from the Tapajós river basin, in Pará, Brazil; and two new species, one from eastern Amazonia,
in the states of Pará (north and south of the Amazon river) and Amapá, Brazil, and another from central Colombia, east
of the Andes. Diagnoses and descriptions of all species are presented.
Key words: lizard, South America, Amazonian rainforest, taxonomy
Resumo
O complexo Gonatodes concinnatus, como aqui considerado, consiste nas espécies de Gonatodes caracterizadas por uma
mancha suprahumeral branca com margens pretas; vermiculações ou não no dorso; e escamas alargadas sob a cauda,
apresentando a seqüência 1’1’1”, e em alguns espécimes 1’1’2” na porção anterior. Duas espécies são atualmente
STURARO & AVILA-PIRES
2 · Zootaxa 2869 © 2011 Magnolia Press
reconhecidas neste grupo amazônico, G. concinnatus e G. tapajonicus. Novos espécimes coletados no leste da Amazônia
(nos estados do Pará e Amapá, Brasil) fizeram necessária a revisão dessas espécies. Analisamos diversas populações
dentro desse complexo, provenientes da Colômbia, Equador, Peru, Venezuela e Brasil, incluindo os novos registros. Os
espécimes foram separados em grupos definidos com base no padrão de coloração. Análises discriminantes utilizando o
método por passos (stepwise) foram realizadas para comparar a morfologia externa (caracteres morfométricos e
merísticos) nesses grupos. Os resultados apoiam o reconhecimento de cinco táxons, correspondendo a G. concinnatus, da
Amazônia Ocidental, no Equador e norte do Peru; G. ligiae, do noroeste da Venezuela (a leste dos Andes); G. tapajonicus,
da bacia do Rio Tapajós, no Pará, Brasil; e duas novas espécies, uma da Amazônia oriental, nos estados do Pará (ao norte
e ao sul do Rio Amazonas) e Amapá, Brasil, e outra da Colômbia central, a leste dos Andes. As diagnoses e descrições de
todas as espécies são apresentadas.
Palavras-chave: lagarto, América do Sul, Floresta Amazônica, taxonomia
Introduction
The genus Gonatodes currently comprises 24 species distributed in Central and South America, in the Antilles and,
as a recent introduction, in Florida, United States (Peters & Donoso-Barros 1970; Rivero-Blanco 1979; Avila-Pires
1995; Esqueda 2004a; Krysko & Daniels 2005; Powel & Henderson 2005; Cole & Kok 2006; Barrio-Amorós &
Brewer-Carías 2008; Rivas & Schargel 2008; Rojas-Runjaic et al. 2010; Schargel et al. 2010). It is characterized by
having dorsal scales granular and juxtaposed; ventral scales larger than dorsals, flat, smooth and imbricate; femoral
and precloacal pores absent; escutcheon present in males, on posterior surface of belly and ventral aspect of thighs;
and free claws (Peters & Donoso-Barros 1970; Hoogmoed 1973; Avila-Pires 1995; Kluge 1995). Sexual
dimorphism is evident in color pattern, usually with colorful males and cryptic females. Male color pattern is an
important character for recognizing species, since differences in scale counts between species are frequently small
(Vanzolini 1968; Rivero-Blanco 1979).
Gonatodes concinnatus (O’Shaughnessy, 1881) and Gonatodes tapajonicus Rodrigues, 1980 have in common
the presence of a white suprahumeral spot with black margins; vermiculations on back (except in G. c. ligi ae ); and
transversely enlarged scales under the tail, showing the sequence 1’1’1”, and in some specimens 1’1’2” (under the
anterior portion of the tail), as defined by Avila-Pires (1995). According to Rodrigues (1980), Gonatodes
tapajonicus differs from G. concinnatus by its color pattern and the form of the mental. Recent material from
eastern Amazonia presented the same characteristics common to both species mentioned above, but with
differences in color pattern. In order to identify this material and considering the variation reported in G.
concinnatus, it was necessary to undertake a revision of this group, which we are calling “G. c onci nnat us
complex”.
O’Shaughnessy (1881) described Goniodactylus concinnatus based on three specimens from Canelos, Ecuador
(BMNH 1946.9.7.10–12), and Goniodactylus buckleyi based on one specimen from Pallatanga (probably in error,
see Rivero-Blanco 1968) and two from Canelos, Ecuador. Boulenger (1885) examined the types of Goniodactylus
concinnatus and G. buckleyi and observed that they represented the same species, respectively male and female; the
species was considered under the combination Gonatodes concinnatus. Subsequent authors followed Boulenger
(1885).
Gonatodes ligiae was described by Donoso-Barros (1967), who provided only a short description, based on
two specimens from Bosque de la Carabela (holotype) and Parque de Moromoy (paratype), Barinitas, Vezenuela.
Rivero-Blanco (1968), based on specimens from the type-locality but without examining the type material of G.
ligiae, considered it a synonym of G. concinnatus. Rivero-Blanco (1979) recognized it as a subspecies of G.
concinnatus, but he never published this work. The shape of the suprahumeral spots (a vertical bar in G.
concinnatus concinnatus and an ocellus in G. concinnatus ligiae) and the presence (G. c. concinnatus) or absence
(G. c. ligiae) of white spots or vermiculations on the back and flanks were pointed out as diagnostic characters.
Gonatodes tapajonicus was described by Rodrigues (1980) based on nine specimens from Cachoeira do Limão
(04º41’S, 56º21’W), Tapajós River, Pará, Brazil.
In this paper we evaluate the validity of the currently recognized species of the Gonatodes concinnatus
complex, and verify the status of additional populations from eastern Amazonia (states of Pará and Amapá, Brazil),
by analyzing variation of external morphology.
Zootaxa 2869 © 2011 Magnolia Press · 3
GONATODES CONCINNATUS COMPLEX
Material and methods
Specimens examined of the Gonatodes concinnatus complex are listed under each species and comparative
material appears in Appendix I. They are deposited in the following institutions: American Museum of Natural
History, New York, New York, USA (AMNH); California Academy of Sciences, San Francisco, California, USA
(CAS); Stanford University collection, now maintained in CAS (CAS-SU); Instituto de Pesquisas Científicas e
Tecnológicas do Estado do Amapá, Macapá, Amapá, Brazil (IEPA); University of Kansas, Museum of Natural
History, Lawrence, Kansas, USA (KU); Natural History Museum of Los Angeles County, Los Angeles, California,
USA (LACM); Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (MCZ);
Museu Nacional do Rio de Janeiro, Universidade Federal de Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ); Museu
Paraense Emilio Goeldi, Belém, Pará, Brazil (MPEG); Museu de Zoologia, Universidade de São Paulo, São Paulo,
Brazil (MZUSP); Sam Noble Oklahoma Museum of Natural History, Norman, Oklahoma, USA (OMNH);
National Natuurhistorisch Museum, Leiden, The Netherlands (RMNH); Texas Cooperative Wildlife Collection,
Department of Wildlife Science, Texas A&M University, College Station, Texas, USA (TCWC); University of
Michigan, Museum of Zoology, Ann Arbor, Michigan, USA (UMMZ); National Museum of Natural History,
Washington, D.C., USA (USNM). We examined photos of all syntypes of Goniodactylus buckleyi and
Goniodactylus concinnatus, which are deposited in The Natural History Museum (former British Museum of
Natural History), London, United Kingdom (BMNH). We could not examine the type specimens of Gonatodes
ligiae, which are deposited in the Museo de Zoología de la Universidad de Concepción, Chile (former personal
collection of R. Donoso-Barros).
Measurements and scale counts. Measurements were taken with an electronic caliper (to the nearest 0.1
mm), when necessary under a stereomicroscope, as follows: SVL (snout-vent length, from tip of snout to cloacal
opening); TL (tail length, from cloacal opening to tip of tail); HL (head length, from anteriormost point of rostral
to anterior margin of ear-opening); HW (head width, on the widest part of head); HD (head depth, on the highest
part of head); ED (eye diameter; between anterior and posterior corner of the eye); IND (internostril distance,
between medial margins of nasal scales); SSL (supranasal scale length, between anterior and posterior corners of
scale); SSW (supranasal scale width, between lateral corners of scale); RSL (rostral scale length, between anterior
and posterior corners of scale); RSW (rostral scale width, between lateral corners of scale); MSL (mental scale
length, between anterior and posterior corners of scale); MSW (mental scale width, between lateral corners of
scale); LAL (lower arm length, from tip of elbow to wrist); LLL (lower leg length, from knee to ankle); HSL
(suprahumeral spot length, measured between anterior and posterior margins); HSH (suprahumeral spot height,
measured between lateral and medial ends).
Scale counts were taken under a stereomicroscope, as follows: SL (supralabials: distinctly enlarged scales
along the upper jaw); IL (infralabials: distinctly enlarged scales along the lower jaw); PR (postrostral scales: scales
in contact to rostral); SP (small postrostral scales: small scales in contact to rostral medially); PN (postnasal scales:
scales in contact with posterior portion of nasal); LS (loreal scales: in a line between postnasals and anterior corner
of orbit); PM (postmental scales: in contact to posterior portion of mental); SAM (scales around midbody, counted
midway between fore- and hind limbs); VLR (ventral scales in a longitudinal row, counted along a midventral line,
from anterior margin of forelimbs to anterior margin of hind limbs); VTR (ventral scales in a transversal row,
counted midway between fore- and hind limb); SCS (supraciliary scales: enlarged and flattened scales on anterior
portion of supraciliary flap); SSC (supraciliary spines: scales with conical shape on the anterior portion of
supraciliary flap); PL2F, PL3F, PL4F (proximal lamellae under respectively second, third and fourth fingers,
counted from base of finger to the sharp angle between first and second phalanges); PL2T, PL3T, PL4T (proximal
lamellae under respectively second, third and fourth toes, counted from base of toe to the sharp angle between first
and second phalanges); DL2F, DL3F, DL4F (distal lamellae under respectively second, third and fourth fingers,
counted between the sharp angle between first and second phalanges and ungual scale); DL2T, DL3T, DL4T
(distal lamellae under respectively second, third and fourth toes, counted between the sharp angle between first and
second phalanges and ungual scale); LS2F, LS3F, LS4F (lateral scales on distal part of respectively second, third
and fourth fingers, between fourth subdigital lamella counted from ungual basal scale toward the hand [distally to
proximally] and dorsal scale); LS2T, LS3T, LS4T (lateral scales on distal part of respectively second, third and
fourth toes, between fourth subdigital lamella counted from the ungual scale toward the foot [distally to
proximally] and dorsal scale); RSE (rows of escutcheon scales under thigh, counted along a line between the
anterior and the posterior aspects of the thigh).
STURARO & AVILA-PIRES
4 · Zootaxa 2869 © 2011 Magnolia Press
Measurements and scale counts were taken on the right side of the body, except when this side was damaged. In
this case data from the left side of the body were used.
Sex was determined by the presence of an escutcheon on the posterior portion of belly and ventral surface of
thighs in males; this feature is absent in females. Specimens under 35 mm SVL were defined as juveniles.
Statistical analysis. We used a discriminant function analysis for all scale counts and measurements (except tail
length) together, to test if groups defined a priori on basis of color pattern could be differentiated by a combination of
other morphological characters (Tabachnick & Fidell 2001). Prior to the analysis we confirmed the normality of the
data using the Shapiro-Wilk test. Measurements may be influenced by factors such as sexual dimorphism,
developmental stage and indeterminate growth, which makes it important to use size-free analysis for such data.
Strauss (1985) and Reis et al. (1990) proposed a model to eliminate the size effect on the discriminant analysis, which
consists of regressing each variable separately on the first principal component of a principal component analysis and
then applying the discriminant function analysis to the residuals obtained from these regressions. The nonparametric
resampling method of jackknifing was used to test the statistical significance of the canonical functions based on the
correct classification rate (McGarigal et al. 2000). Due to the high number of variables and low number of individuals
in most samples, we applied the stepwise method in the discriminant function analysis and adjusted the F-enter and F-
remove to select N-1 variables (where N is the number of specimens of the smallest group), thus increasing the
robustness of the analyses. The forward direction option was used, so that at each step all variables were reviewed and
evaluated to determine which one would contribute most to the discrimination between groups. The variables in each
step that showed best discrimination power between groups were then used in the subsequent step of the analysis.
Missing scale counts and morphometric values were estimated using the mean value for that character within each
group. Missing values never represented more than 1.6% of the total number of cases. For all analyses we used the
statistic software SYSTAT for Windows, version 12 (Systat Software, San Jose, California, USA).
Groups considered in the discriminant analysis were defined by color pattern of preserved male specimens, but
some observations were based on color in life. Females and juveniles were assigned to groups based on their locality
and proximity to certain male patterns, since few or no geographic differences in female pattern were found.
Species description and comparisons. Species description follows Avila-Pires (1995) with the addition of some
morphometric and scale count characters. Statistics are presented as “minimum–maximum (mean ± standard
deviation, N=sample size).” Tail length was measured only in specimens with intact, non-regenerated tails.
We compared the species of Gonatodes concinnatus complex with other congeners based on specimens examined
(Appendix I) and the literature (Donoso-Barros 1966; Rivero-Blanco 1979; Esqueda 2004a; Powell & Henderson
2005; Cole & Kok 2006; Barrio-Amorós & Brewer-Carías 2008; Rivas & Schargel 2008; Rojas-Runjaic et al. 2010;
Schargel et al. 2010). Within the G. concinnatus complex, the most informative characters selected by the
discriminant function analysis were graphically compared, either using boxplots (meristic characters) or linear
regressions (morphometric characters).
Results
We analyzed 376 specimens of Gonatodes, of which 255 were of the Gonatodes concinnatus complex. We
distinguished five different color patterns (see Table 1; Fig. 1), which were used to define the groups for the
discriminant analysis (herein named A, B, C, D and E, as for the color patterns). Stepwise discriminant function
analysis using both scale counts and measurements revealed highly significant differences among groups (Wilks's
Lambda=0.025, df=36; approx. F=42.4, df=907, p=0.00). The nine variables selected as the most powerful
discriminators among groups are listed in Table 2. Scales around midbody was selected first, classifying 61.7% of the
specimens, followed by suprahumeral spot height, mental scale width, rostral scale length, ventral scales in a
longitudinal row, lateral scales of second finger, distal lamellae under second toe, proximal lamellae under second toe,
and lateral scales of second toe, whose additions improved the classification criterion to respectively 77.3%, 84.0%,
87.5%, 90.2%, 91.0%, 91.4%, 92.2% and 93.4%. The jackknifed classification matrix correctly classified unknown
specimens 92.6% of the times (Table 3). The first, second and third discriminant functions explained, respectively,
45.4%, 36.8% and 11.0% of the total variance in the nine variables. The first two discriminant functions provided
separation between C+D (which overlap), A, B and E groups, even though with a small overlap between A and E (Fig.
2). The third discriminant function separates groups C and D almost completely. Characters with the largest loadings
on the first discriminant function were scales around midbody and ventral scales in a longitudinal row (Table 2).
Zootaxa 2869 © 2011 Magnolia Press · 5
GONATODES CONCINNATUS COMPLEX
Mental scale width and suprahumeral spot height had the highest loadings on the second function, and the
suprahumeral spot height and distal lamellae under second toe were the most important on the third function.
We have, therefore, five groups that are distinct from each other in terms of color pattern and a number of external
characters and that are separated geographically (Fig. 3), indicating that they constitute independent lineages that
deserve to be recognized as separate species. Considering the type-localities of the three names already available for
this complex, we conclude that pattern A refers to Gonatodes concinnatus; pattern B to G. ligiae, here resurrected as a
valid species; and pattern C to G. tapajonicus. No names are available that correspond to patterns D and E, which are
therefore described as new species. A description of each species is presented below.
TABLE 1. Comparison of groups A–E of Gonatodes concinnatus complex recognized on basis of color pattern.
TABLE 2. Results of size-free stepwise discriminant function analysis of scale counts and measurements comparing the five
groups identified on basis of color pattern. 1DF: loadings on the first discriminant function; 2DF loadings on the second
discriminant function; 3DF loadings on the third discriminant function. See Material and Methods for abreviation of characters.
TABLE 3. Classification matrix and Jackknifed classification matrix of size-free stepwise discriminant function analysis using
scale counts and measurements.
Patterns Throat Head dorsally Back and limbs
A Beige, immaculate Beige without vermiculations Brown and beige vermiculations
B Light brown, immaculate Light brown, immaculate Light brown, immaculate
C White with black oblique stripes
(in life, vivid yellow with red-
dish-brown oblique stripes)
Brown and beige vermiculations (in life,
vivid yellow and reddish-brown vermicu-
lations)
Brown and beige vermiculations (in
life, vivid yellow and reddish-
brown vermiculations
D White or dark-brown with black
and white oblique stripes (in life,
orange with orange-yellow
oblique stripes)
Dark-brown and beige vermiculations (in
life, reddish brown with drab or orange-
yellow vermiculations)
Dark-brown and beige vermicula-
tions (in life, reddish-brown with
drab or orange-yellow vermicula-
tions)
E Brown, immaculate Beige without vermiculations Brown and beige small vermicula-
tions
Scale count/
Measurement F(+ent,-rem) Wilks's Lambda Approx. F-ratio p-value 1DF 2DF 3DF
SAM 131.913 0.321 131.913 0 0.593 0.217 0.067
HSH 60.441 0.163 91.884 0 0.008 -0.502 0.842
MSW 42.871 0.096 77.662 0 0.019 -0.808 -0.264
RSL 25.867 0.068 67.174 0 0.031 -0.467 -0.135
VLR 19.927 0.051 59.084 0 0.466 -0.205 -0.347
LS2F 18.733 0.039 54.665 0 0.208 -0.19 0.114
DL2T 12.597 0.033 49.866 0 0.114 0.353 0.353
PL2T 9.249 0.028 46.274 0 0.144 -0.258 -0.077
LS2T 8.078 0.025 42.372 0 0.245 0.066 0.15
Classification Matrix Jackknifed Classification Matrix
Pattern A B C D E %correct A B C D E %correct
A 133001595.7% 133001595.7%
B 0 10000100.0% 0 900190.0%
C 0 01900100.0% 0 01900100.0%
D 0 0 4 20 0 83.3% 0 0 5 19 0 79.2%
E 3 3005790.5% 3 3005790.5%
Total 136 13 23 21 62 93.4% 136 12 24 20 63 92.6%
STURARO & AVILA-PIRES
6 · Zootaxa 2869 © 2011 Magnolia Press
FIGURE 1. Dorsal and ventral color patterns in preservative in the Gonatodes concinnatus complex. (A) pattern A (MZUSP
54655); (B) pattern B (TCWC 57237); (C) pattern C (MZUSP 53674, holotype); (D) pattern D (MPEG 25596, holotype); and
(E) pattern E (MZUSP 49153, holotype). Scale bar = 5 mm.
Zootaxa 2869 © 2011 Magnolia Press · 7
GONATODES CONCINNATUS COMPLEX
FIGURE 2. Bivariate scattergrams from stepwise discriminant function analysis using scale counts and measurements. Red =
Pattern A. Cyan-green = Pattern B. Lime-green = Pattern C. Yellow = Pattern D. Blue = Pattern E.
Gonatodes concinnatus (O’Shaughnessy, 1881)
(Figs.1A, 4, 5, 6)
Goniodactylus concinnatus O’Shaughnessy, 1881: 237 (syntypes BMNH 1046.9.7.10–12, formerly BMNH 80.12.8.29–31,
type-locality: Canelos, Ecuador, collected by Buckley).
Goniodactylus buckleyi O’Shaughnessy, 1881: 238 (syntypes BMNH 1046.9.7.13–14, formerly BMNH 80.12.8.32–33, type-
locality: Canelos; BMNH 1946.9.7.15, formerly BMNH 80.12.8.34, type-locality: Pallatanga [probably in error, see
Rivero-Blanco, 1968: 105, 1979: 92]; all in Ecuador, collected by Buckley).
Gonatodes concinnatus; Boulenger, 1885: 61–62; Burt & Burt, 1933: 2; Vanzolini, 1955: 123, 1968: 25 (part); Wermuth, 1965:
44; Mechler, 1968: 331 (part); Rivero-Blanco, 1968: 104 (part); Peters & Donoso-Barros, 1970: 132 (part); Dixon & Soini,
1975: 19, 1986: 23; Duellman, 1978: 195; Rodrigues, 1980: 313; Duellman & Mendelson III, 1995: 358; Moravec,
Tuanama & Burgos, 2001: 51; Bartlett & Bartlett, 2003: 158 (part); Powell & Henderson, 2005: 714; Cole & Kok, 2006: 4;
Gamble, Simon, Colli & Vitt, 2008: 271.
Gonatodes concinnatus concinnatus; Rivero-Blanco, 1979: 92 (part).
STURARO & AVILA-PIRES
8 · Zootaxa 2869 © 2011 Magnolia Press
FIGURE 3. (A) Geographic distribution of Gonatodes concinnatus complex. (B) Detailed geographic distribution of
Gonatodes concinnatus. Red = Gonatodes concinnatus (Pattern A). Cyan-green = Gonatodes ligiae (Pattern B). Lime-green =
Gonatodes tapajonicus (Pattern C). Yellow = Gonatodes nascimentoi sp. nov. (Pattern D). Blue = Gonatodes riveroi sp. nov.
(Pattern E). Open symbols = type-localities; half-black symbol = literature record; question mark = Pallatanga, Ecuador,
probably in error; triangle = locality referred to only as ‘Rio Napo’. Numbers refer to localities cited under each species. One
number may represent more than one locality.
Zootaxa 2869 © 2011 Magnolia Press · 9
GONATODES CONCINNATUS COMPLEX
Material examined (numbers in bold between parentheses refer to Fig. 3): ECUADOR. “Pallatanga” (1): Photos of
BMNH 1946.9.7.15 (formerly BMNH 80.12.8.34, syntype of Goniodactylus buckleyi), leg. Buckley. SUCUMBÍOS.
Shushufindi, Rio Aguarico (2): USNM 234497, female. Santa Cecilia (3): KU 105290–91, KU 109572–73, KU
146643, KU 146645, KU 147588–89, KU 152496, 4 males and 5 females. Lago Agrio (4): KU 126683, 126685–86,
126688, 2 males and 2 females; KU 297992, female, leg. J. Lee. Limoncocha (5): LACM 73293–300, a juvenile, 5
males and 2 females, leg. W.R. Heyer; MZUSP 54655, male, leg. K. Miyata; KU 144378, MCZ 86414, a juvenile and
a male. Reserva de Producción Faunística de Cuyabeno (RPF Cuyabeno), Estación Biológica de la Pontificia
Universidad Catolica del Ecuador (6): OMNH 36406–11, 36424, 3 males and 4 females, leg. L.J. Vitt. Reserva de
Producción Faunística de Cuyabeno (RPF Cuyabeno), Saladero de Dantas (6): OMNH 36412–14, a juvenile, a male
and a female, leg. L.J. Vitt. Corporación Estatal Petrolera Ecuatoriana (CEPE) oil field (near Tarapoa, near Lago
Agrio), Cuyabeno (7): MCZ 163233, male. SUCUMBIOS–ORELLANA. Rio Napo (undefined locality - 8): UMMZ
90813, female, leg. W.M. Clark. NAPO. George Kiederle Hacienda, southern bank Rio Napo, 2 km W Puerto Napo
(9): USNM 166135–37, 2 males and a female, leg. J.A. Peters. Rio Misahualli (10): USNM 234490, female, leg. M.
Olalla. 1 km E Jatun Sacha (11): OMNH 36377–78, a male and a female. San José de Sumaco (12): AMNH 28879,
female, leg. C. Ollalo. ORELLANA. Between Loreto and Concepcion (13): CAS-SU 16529–30, a male and a female,
leg. J. Olalla and C.A. Olalla. Loreto (14). MZUSP 3382–83, 2 females, leg. J. Olalla; USNM 234494, male, leg. J.
Ollala. Rio Punino, affluent of Rio Payamino above Coca (15): MCZ 131046, juvenile. Rio Payamino (16): USNM
234491–93, a male and 2 females, leg. M. Olalla. Mouth of Rio Coca (17): USNM 234495, male, leg. J. Olalla.
PASTAZA. Rio Pindo (18): USNM 234500, male, leg. R. Olalla. Canelos (19): Photos of BMNH 1046.9.7.10–12
(syntypes of Goniodactylus concinnatus, formerly BMNH 80.12.8.29–31), 3 males (Fig. 4), Photos of BMNH
1046.9.7.13–14 (syntypes of Goniodactylus buckleyi, formerly BMNH 80.12.8.32–33), 2 females, leg. Buckley.
CAS-SU 15814–15, CAS-SU 16531, a male and 2 females, leg. A. Proana. Sarayacu, (20): MCZ 37703–04, KU
121088–89, a juvenile and 3 females. Rio Villano (21), USNM 234499, female, leg. C. Estrella. Nuevo Golandrina,
on trail W toward Rio Curaray, 130 km S of Coca (22): USNM 321064, female, leg. R.P. Reynolds and W. Lamar.
Tiguino (UNOCAL Base Camp), 130 km S of Coca (22): USNM 321057, female, leg. R. P. Reynolds and W. Lamar.
Rio Conambo (23): USNM 234502, female, leg. R. Olalla. MORONA-SANTIAGO. Cusuime, Rio Cusuime, 60 km
airline SE Macas (24): AMNH 113666–67, 2 males, leg. B. Malkin. Taisha, Macas (25). USNM 234498, male, leg.
M. Ollala; USNM 283922, USNM 283924, a juvenile and a female, leg. C. M. Fugler. PERU: LORETO. San Jacinto
(26): KU 222138, female, leg. R. A. Leschen. Moropón (27): 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, a juvenile, 25 males and 12 females; leg. P. Soini; TCWC 41763, juvenile, leg. J.R. Dixon;
MZUSP 28248–49, MZUSP 28260–63, MZUSP 28273–79, MZUSP 28311–13, MZUSP 28319, MZUSP 28375,
MZUSP 39218, 3 juveniles, 12 males and 4 females, leg. P. Soini. Explorama Lodge, junction Rio Yanamono and Rio
Amazonas (28): KU 220370, juvenile, leg. W. E. Duellman. Iquitos (29): TCWC 41137–38, a juvenile and a female,
leg. P. Soini. ACEER on Quebrada Grande, close to junction Rio Sucusari and Rio Napo (30): KU 220371, male, leg.
W. E. Duellman. Yanamono (31): MZUSP 28354–55, a juvenile and a female, leg. P. Soini. Rio Orosa (32). MZUSP
56657, male, leg. P. Soini. Estirón, Rio Ampiyacu (33): MZUSP 13458, male, leg. B. Malkin.
Diagnosis. A relatively large Gonatodes, with maximum SLV of 52.6 mm. Scales around midbody 109–131.
Ventral scales in a longitudinal row 49–61. Proximal subdigital lamellae as wide as digits, in total 17–22 under fourth
finger, 21–27 under fourth toe. Three or four lateral rows of scales on distal portion of fingers and toes. Tail with
midventral scales distinctly wider than long, forming a repetitive sequence of two single midventrals (one after the
other), each in contact with one laterodistal scale per side, followed by a divided (only on proximal portion of tail,
when present) or single midventral in contact with two laterodistal scales per side (respectively 1’1’2” and 1’1’1”). A
white suprahumeral bar, bordered by black, present both in males and females, although more conspicuous in males;
it reaches dorsally between the dorsolateral and middorsal regions. Males with head dorsally without vermiculations;
back and limbs with a vermiculated pattern of dark and light spots; no dark streaks on gular region (at least in
preservative).
Description. Maximum SVL in males of 52.6 mm (TCWC 36669), in females of 49.2 mm (KU 146645). Tail
round in cross section, tapering toward tip, 1.0–1.4 (1.15 ± 0.09, N=37) times SLV. Head length 0.21–0.27 (0.24 ±
0.01, N=137) times SLV, 1.3–1.6 (1.45 ± 0.06, N=136) times as long as wide, 1.2–1.7 (1.42 ± 0.08, N=135) times as
wide as high. Snout round, moderately elongate (Fig. 5A), gently sloping toward top of head. Neck slightly narrower
than head and anterior portion of body. Body cylindrical. Limbs well developed, lower arm 0.11–0.15 (0.14 ± 0.01,
N=139) times SVL, lower leg 0.13–0.17 (0.15 ± 0.01, N=139) times SVL.
STURARO & AVILA-PIRES
10 · Zootaxa 2869 © 2011 Magnolia Press
Rostral convex, 1.6–2.9 (2.07 ± 0.16, N=139) times as wide as high; posterior margin with a shallow
depression medially and slightly indented by 1–3 medial postrostrals, with or without a median cleft extending
anteriorly. Postrostrals 3 (72.0%), 4 (19.4%), or 5 (8.6%), laterals ones (supranasals) distinctly larger than medial
one(s) (N=139) (Fig 5A). Nasal bordered by rostral, first supralabial (only a narrow contact in some specimens), 3
(92.1%) or 4 (7.9%) postnasals (N=139), and supranasal; internostril distance 0.20–0.28 (0.24 ± 0.02, N=136)
times head width. Supranasal roughly oval or semicircular, 0.8–1.5 (1.16 ± 0.13; N=139) times as wide as long.
Postnasals slightly larger than, or 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, 9–13 (10.4 ± 0.8, N=138) loreals on a line between
postnasals and anterior corner of eye. Top and posterior portion of head, as well as supraorbital region, with
granular scales. A short supraciliary flap present, anteriorly with 6–12 (8.4 ± 1.1, N=133) enlarged and flattened
scales, among which 0–7 (2.0 ± 1.3; N=133) small, conical spines. Pupil round, eye diameter 0.19–0.26 (0.21 ±
0.01, N=129) times head length. Scales on temporal region similar to those on top of head. Ear-opening much
smaller than eye, oval, posterior to, and at same level of, commissure of mouth. Supralabials 5–7 (6.2 ± 0.5;
N=139), distinctly enlarged anteriorly and decreasing in size posteriorly, one or two of them posterior to centre of
eye, followed to corner of mouth by small scales.
Mental large, distinctly wider anteriorly than posteriorly, with posterior margin forming a wide angle, 0.9–1.4
(1.1 ± 0.1; N=137) times as wide as long; 1 (1.4%), 2 (82.7%), 3 (12.9%) or 4 (2.9%) postmentals (Fig. 5B). Scales
on chin flat, smooth, polygonal, juxtaposed, larger anteriorly, decreasing in size posteriorly. Infralabials 4–7 (5.5 ±
0.7; N=138), distinctly enlarged anteriorly and decreasing in size posteriorly; 0–3 of them posterior to centre 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
granular, posteriorly flat, smooth, hexagonal or round, increasing in size toward the posterior region, with a short
transitional zone between the anterior and posterior parts.
Dorsals granular, increasing in size toward the flanks. Ventrals larger than dorsals, roughly hexagonal, flat,
smooth, imbricate, in oblique rows; 49–61 (56.1 ± 2.1; N=119) scales along the midventral line between anterior
margin of forelimbs and vent; 17–20 (18.5 ± 0.9; N=126) scales in a transverse line at midbody, with a short
transitional zone between ventrals and scales on flanks. Scales around midbody 109–131 (120.7 ± 4.6; N=110).
Scales on preanal plate similar to ventrals, except for those bordering vent, which are very small. Escutcheon
present in males on posterior portion of belly and on 3–5 (mostly four) rows (body-knee direction) on ventral
surface of thighs.
Scales on anterodorsal surface of forelimbs flat, smooth, roundish or rhomboidal, imbricate, largest close to the
wrist; on posterodorsal and ventral surface of forelimbs convex, smooth, rhomboidal, juxtaposed, relatively small.
Scales on anterodorsal surface of thighs and ventral surface of hind limbs flat, smooth, roundish or rhomboidal,
imbricate and relatively larger; on posterodorsal surface of thighs and dorsal surface of lower legs smaller, convex,
smooth, rhomboidal, juxtaposed.
Lamellae under second (II) through fourth (IV) fingers (proximal lamellae in parentheses): II: 14–18 (5–7),
III: 16–21 (5–7), and IV: 17–22 (5–9) (Fig. 5C). Lamellae under second through fourth toes (proximal lamellae in
parentheses): II: 14–18 (5–7), III: 16–22 (5–8), and IV: 21–27 (9–13) (Fig. 5D). Claws exposed, non-retractile,
between two basal scales. Fingers and toes with three, occasionally four or two (USNM 234497; USNM 234494
and TCWC 42706), lateral scales distally, between the fourth (counted from the claw towards the hand) subdigital
lamella and the dorsal scale.
Scales on tail dorsally and laterally relatively small, rhomboidal, flat, smooth, imbricate. On ventral surface of
tail scales smooth, flat, imbricate, increasing in size toward midventral line. Midventral scales, except close to the
base of the tail, distinctly wider than long, forming a repetitive sequence of two single midventrals (one after the
other), each in contact laterodistally with one scale per side, followed by a divided (only on proximal portion of
tail, when present) or single midventral scale in contact laterodistally with two scales per side – respectively 1’1’2”
and 1’1’1” in the codification of Avila-Pires (1995: Figure 2) (Fig. 5E).
Color in preservative. In males (Figs. 1A, 4), dorsal surface of head beige, without markings. Back and
flanks, base of tail and hind limbs with relatively large, beige and brown vermiculations. A large, conspicuous,
white suprahumeral bar, bordered by black, extending dorsally at least up to the dorsolateral region, in some cases
almost reaching the middorsal region; never in the form of an ocellus, but Vitt & Torre (1996) present the photo of
a specimen with the white bar divided into two spots by a transversal black band. Ventral surface of head, gular
Zootaxa 2869 © 2011 Magnolia Press · 11
GONATODES CONCINNATUS COMPLEX
region and chest beige or reddish-brown, without oblique streaks; belly gray; underside of limbs beige. Tail brown
and/or black dorsally, white and/or brown ventrally. Escutcheon area (belly and thighs) light gray.
In females, dorsal surface of head and limbs with brown and black irregular spots; back gray with dorsolateral
pairs of black spots and, in some specimens, pairs of beige spots; flanks gray with black and brown spots.
Suprahumeral bar conspicuous, white with black margins, similar in extension but thinner than that of males.
Ventral surface of head and gular region white with dark oblique streaks, in contact or not at midventral line; belly
and underside of limbs light gray. Tail brown and/or black dorsally, white and/or brown ventrally; original tail
distally with white bands that form complete rings around the tail.
FIGURE 4. Gonatodes concinnatus (syntypes, BMNH 1946.9.7.10–12). (A) Dorsal, (B) ventral views. Scale bar = 50 mm.
(Photo by Colin McCarthy).
STURARO & AVILA-PIRES
12 · Zootaxa 2869 © 2011 Magnolia Press
FIGURE 5. Gonatodes concinnatus (MZUSP 54655). (A) Dorsal and (B) ventral views of head; ventral views of (C) right
hand and (D) right foot; (E) ventral view of tail. Scale bar = 1 mm.
Zootaxa 2869 © 2011 Magnolia Press · 13
GONATODES CONCINNATUS COMPLEX
FIGURE 6. Gonatodes concinnatus from Cuyabeno, Sucumbios, Ecuador. (A) Adult male (LJV 5014) and (B) adult female
(OMNH 36409; LJV 5046) (Photos by Laurie Vitt).
STURARO & AVILA-PIRES
14 · Zootaxa 2869 © 2011 Magnolia Press
Color in life. Color in life has been described by Duellman (1978) and Vitt & de la Torre (1996). In males (Fig.
6A) head, neck and forelegs orange or reddish brown with cream spots; suprahumeral bar white bordered by black;
body olive-green or brown, with reddish brown or yellowish white vermiculations that may present a black margin.
Throat orange with cream streaks, belly yellowish gray, and tail black with white rings, or the entire belly and tail
black.
In females (Fig. 6B), head, body and tail grayish tan or drab gray with irregular crossbands, brown or black
anteriorly and white posteriorly; distally the tail becomes black and white banded. Throat cream with dark streaks;
venter creamy tan or yellow. Iris reddish brown with a cream circle around the pupil.
Distribution. Western Amazonia, in Ecuador and northern Peru (Fig. 3A–B).
Remarks. Gonatodes ligiae Donoso-Barros, 1967, from Venezuela, is considered in this paper as a valid species.
Material from Colombia previously identified as G. concinnatus is here described as a new species, G. riveroi.
Gonatodes ligiae Donoso-Barros, 1967
(Figs. 1B, 7, 8)
Gonatodes ligiae Donoso-Barros, 1967 (holotype V 126, according to Rivero-Blanco, 1979 in the Museo de Zoología de la
Universidad de Concepción, Chile; type-locality: Bosque de la Carabela, near Barinitas, Barinas, Venezuela), Donoso-
Barros, 1968: 108
Gonatodes concinnatus; Rivero-Blanco, 1968: 104 (part); Peters & Donoso-Barros, 1970: 132 (part).
Gonatodes concinnatus ligiae; Rivero-Blanco, 1979: 92; Rivero-Blanco & Barrio-Amorós, 2002: 67; Esqueda, 2004b: 162.
Material examined (numbers in bold between parentheses refer to Fig. 3): VENEZUELA. BARINAS. 8 km SW
Santa Barbara, on San Cristobal Highway (41): TCWC 60231, female, leg. R. H. Dean. Barinitas (44): TCWC
57236–44, 2 juveniles, 3 males and 4 females, leg. C. Rivero-Blanco.
Diagnosis. A medium-sized Gonatodes, with maximum SLV of 40.7 mm. Scales around midbody 109–128.
Ventral scales in a longitudinal row 45–52. Proximal subdigital lamellae as wide as digits, in total 15–17 under fourth
finger, 18–21 under fourth toe. Two, occasionally three, lateral rows of scales on distal portion of fingers and toes. Tail
with midventral scales distinctly wider than long, forming a repetitive sequence of two single midventrals (one after
the other), each in contact laterodistally with one scale per side, followed by a single midventral in contact
laterodistally with two scales per side (1’1’1”). A white suprahumeral bar, bordered by black, present both in males
and females, although more conspicuous in males; it extends dorsally to approximately the dorsolateral region. Males
with head and trunk dorsally without vermiculations; no dark streaks on gular region.
Description. Maximum SVL in males of 40.7 mm (TCWC 57242), in females of 40.2 mm (TCWC 57240). Tail
round in cross section, tapering toward tip, 0.8–1.2 (1.01 ± 0.18, N=4) times SLV. Head length 0.22–0.27 (0.24 ± 0.02,
N=10) times SLV, 1.4–1.6 (1.48 ± 0.07, N=136) times as long as wide, 1.2–1.4 (1.31 ± 0.06, N=10) times as wide as
high. Snout round, moderately elongate (Fig 7A), gently sloping toward top of head. Neck slightly narrower than head
and anterior portion of body. Body cylindrical. Limbs well developed, lower arm 0.11–0.14 (x=0.13 ± 0.01, N=10)
times SVL, lower leg 0.13–0.15 (0.14 ± 0.01, N=10) times SVL.
Rostral convex, 1.8–2.4 (2.00 ± 0.20, N=10) times as wide as high; posterior margin with a shallow depression
medially and slightly indented by zero or one medial postrostral, with a median cleft extending anteriorly. Postrostrals
2 (10%) or 3 (90%), laterals ones (supranasals) distinctly larger than medial one when this is present (N=10) (Fig 7A).
Nasal bordered by rostral, first supralabial, three postnasals (N=10), and supranasal; internostril distance 0.23–0.30
(0.25 ± 0.03, N=10) times head width. Supranasal roughly oval, 1.0–1.3 (1.14 ± 0.08; N=10) times as wide as long.
Postnasals slightly larger than, or 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, 7–8 (7.8 ± 0.42, N=10) loreals on a line between postnasals and
anterior corner of eye. Top and posterior portion of head, as well as supraorbital region, with granular scales. A short
supraciliary flap present, anteriorly with 6–8 (6.6 ± 0.84, N=10) enlarged and flattened scales, among which 0–2 (1.2
± 0.79; N=10) small, conical spines. Pupil round, eye diameter 0.21–0.27 (0.23 ± 0.02, N=10) times head length.
Scales on temporal region similar to those on top of head. Ear-opening much smaller than eye, oval, posterior to, and
at same level of, commissure of mouth. Supralabials 4–6 (5.5 ± 0.71; N=10), distinctly enlarged anteriorly and
decreasing in size posteriorly, one or two of them posterior to centre of eye, followed to corner of mouth by small
scales.
Zootaxa 2869 © 2011 Magnolia Press · 15
GONATODES CONCINNATUS COMPLEX
FIGURE 7. Gonatodes ligiae (TCWC 57237). (A) Dorsal and (B) ventral views of head; ventral views of (C) right hand and
(D) right foot; (E) ventral view of tail. Scale bar = 1 mm.
STURARO & AVILA-PIRES
16 · Zootaxa 2869 © 2011 Magnolia Press
FIGURE 8. Gonatodes ligiae from Parque Moromoy, Barinitas, Barinas, Venezuela. (A) Adult male and (B) adult female
(Photos by Carlos Rivero-Blanco).
Zootaxa 2869 © 2011 Magnolia Press · 17
GONATODES CONCINNATUS COMPLEX
Mental large, distinctly wider anteriorly than posteriorly, with posterior margin forming a wide angle, 1.2–1.4
(1.31 ± 0.06; N=10) times as wide as long; 2 (70%) or 3 (30%) postmentals (Fig. 7B). Scales on chin flat, smooth,
polygonal, juxtaposted, larger anteriorly, decreasing in size posteriorly. Infralabials 5–6 (5.1 ± 0.32; N=10),
distinctly enlarged anteriorly and decreasing in size posteriorly; 1–3 of them posterior to centre 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
granular, posteriorly flat, smooth, hexagonal or round, increasing in size toward the posterior region, with a short
transitional zone between the anterior and posterior parts.
Dorsals granular, increasing in size toward the flanks. Ventrals larger than dorsals, roughly hexagonal, flat,
smooth, imbricate, in oblique rows; 45–52 (48.6 ± 2.3; N=9) scales along the midventral line between anterior
margin of forelimbs and vent; 17–19 (17.8 ± 1.0; N=9) scales in a transverse line at midbody, with a short
transitional zone between ventrals and scales on flanks. Scales around midbody 92–107 (101.5 ± 5.5; N=9). Scales
on preanal plate similar to ventrals, except for those bordering vent, which are very small. Escutcheon present in
males on posterior portion of belly and on 3–4 (mostly three) rows (body-knee direction) on ventral surface of
thighs.
Scales on anterodorsal surface of forelimbs flat, smooth, roundish or rhomboidal, imbricate, largest close to the
wrist; on posterodorsal and ventral surface of forelimbs convex, smooth, rhomboidal, juxtaposed, relatively small.
Scales on anterodorsal surface of thighs and ventral surface of hind limbs flat, smooth, roundish or rhomboidal,
imbricate and relatively larger; on posterodorsal surface of thighs and dorsal surface of lower legs smaller, convex,
smooth, rhomboidal, juxtaposed.
Lamellae under second (II) through fourth (IV) fingers (proximal lamellae in parentheses): II: 10–14 (4–5),
III: 13–17 (4–6), and IV: 15–17 (5–6) (fig 7C). Lamellae under second through fourth toes (proximal lamellae in
parentheses): II: 12–16 (4–6), III: 15–18 (5–6), and IV: 18–21 (8–10) (Fig. 7D). Claws exposed, non-retractile,
between two basal scales. Fingers and toes with two, occasionally three, lateral scales distally, between the fourth
(counted from the claw towards the hand) subdigital lamella, and the dorsal scale.
Scales on tail dorsally and laterally relatively small, rhomboidal, flat, smooth, imbricate. On ventral surface of
tail scales smooth, flat, imbricate, increasing in size toward midventral line; midventral scales, except close to the
base of the tail, distinctly wider than long, forming a repetitive sequence of two single midventrals (one after the
other), each in contact laterodistally with one scale per side, followed by a slightly larger single midventral scale in
contact laterodistally with two scales per side – 1’1’1” in the codification of Avila-Pires (1995: Figure 2) (Fig. 7E).
Color in preservative. In males (Fig. 1B), dorsal surface of head light brown, spotless. Back and flanks, base
of tail and hind limbs light-brown. A large, conspicuous, white suprahumeral bar, bordered by black, extending
dorsally to approximately the dorsolateral region. Ventral surface of head, gular region and chest beige or light-
brown, without oblique streaks; belly gray; underside of limbs beige. Tail brown dorsally, beige or light-brown
ventrally. Escutcheon area (belly and thighs) light gray.
In females, dorsal surface of head and limbs with brown and black irregular spots; back gray with dorsolateral
pairs of black spots and, in one specimen (TCWC 57243), a beige middorsal line and pairs of dark-brown spots on
base of tail; flanks gray with black and brown spots. Suprahumeral bar conspicuous, white with black margins,
thinner than that of males, extending dorsally to approximately the dorsolateral region. Ventral surface of head and
gular region white with dark oblique streaks, in contact or not at midventral line; belly and underside of limbs light
gray. Tail brown with black spots dorsally, white or light-brown ventrally; original tail distally with white bands
that form complete rings around the tail.
Color in life. Descriptions were based on photos of specimens from Parque Moromoy, Barinitas, Barinas,
Venezuela, sent to us by Carlos Rivero-Blanco, and description of coloration in Rivero-Blanco (1979). In males
(Fig. 8A), head dorsally brownish yellow, laterally and neck greenish yellow. Back greenish yellow with a
brownish yellow vertebral band, flanks slate gray, occasionally with one or two minute white spot(s), with an
irregular black margin. Suprahumeral bar white with black margins (in one of the specimens photographed by C.
Rivero-Blanco, there is a small white spot in the black margin above the white bar). Head ventrally and gular
region dusty yellow, in some specimens with orange spots; remaining ventral region gray, lighter on escutcheon
areas. Tail dark brown dorsally, lighter below; original tail may present a banding pattern, seen also in juveniles.
In females (Fig. 8B), dorsal region mottled with several hues of gray and brownish-gray. At each side a
dorsolateral series of dark brown spots may be present. Suprahumeral bar white with dark brown, irregular
STURARO & AVILA-PIRES
18 · Zootaxa 2869 © 2011 Magnolia Press
margins. Ventral region light creamy gray with a pattern of brown chevrons on gular region; underside of tail
cream, occasionally with dark and light rings present.
Distribution. Northwestern Venezuela, on the region of Cordillera de Mérida (Fig. 3A).
Remarks. Rivero-Blanco (1968) recorded this species as G. concinnatus from Reserva Florestal Ticoporo,
Socopós, Barinas, Venezuela (number 42 in Fig. 3A). Rivero-Blanco and Barrio-Amorós (2002) reported G. c.
ligiae from distrito Páez, 7 km NE of La Victoria, Apure, Venezuela (number 40 in Fig. 3A). Esqueda (2004b)
recorded G. c. ligiae from Calderas, Barinas, Venezuela (number 43 in Fig. 3A), not far from the type-locality.
Gonatodes tapajonicus Rodrigues, 1980
(Figs. 1C, 9, 10)
Gonatodes tapajonicus Rodrigues, 1980: 309 (holotype MZUSP 53676, type-locality: Cachoeira do Limão, Rio Tapajós, Pará,
Brazil); Vanzolini, 1986: 10; Avila-Pires, 1995: 283.
Material examined (numbers in bold between parentheses refer to localities in Fig. 3): BRAZIL. PARÁ. Município
Itaituba. Comunidade Aldeia Nova ("Aldêia Akay Muuybu"), anteriorly known as Comunidade Boa Fé, right
margen of Rio Tapajós (04o 41' 58.2"S and 56o 22' 37.0"W) (45): MPEG 27708–27718, 2 juveniles, 5 males and 4
females, leg. M.J. Sturaro, P.L.V. Peloso and J.O. Gomes. Cachoeira do Limão, right margin of Rio Tapajós (~4o41’S
and 56o21’W) (46): MZUSP 53676 (holotype), male; MZUSP 53669, 53671–74, 53677 (paratypes), a male and 5
female, leg. M.T. Rodrigues. Floresta Nacional de Altamira, Distrito de Moraes de Almeida (06º 1’8”S and 55º
18’17”W) (47): MPEG 25095, female, leg. C. Lima and R.R. Silva.
Diagnosis. A relatively large Gonatodes, with maximum SVL of 55 mm. Scales around midbody 118–133.
Ventral scales in a longitudinal row 53–63. Proximal subdigital lamellae as wide as digits, in total 18–22 under fourth
finger, 22–27 under fourth toe. Three, occasionally four, lateral rows of scales on distal portion of fingers and toes.
Tail with midventral scales distinctly wider than long, forming a repetitive sequence of two single midventrals (one
after the other), each in contact laterodistally with one scale per side, followed by a divided (only on proximal portion
of tail, when present) or single midventral in contact laterodistally with two laterodistal scales per side (respectively
1’1’2” and 1’1’1”). A white suprahumeral ocellus, bordered by black, present both in males and females, although
more conspicuous in males. Males with a vermiculated pattern of light and dark (vivid yellow and reddish brown in
life) spots both on head and body dorsally; gular region with dark oblique streaks (yellow with reddish brown streaks
in life).
Description. Maximum SVL in males of 53 mm (MZUSP 53676), in females of 55 mm (MZUSP 53671)
(Rodrigues 1980). Tail round in cross section, tapering toward tip, 1.1–1.3 (1.21 ± 0.10, N=6) times SLV. Head length
0.22–0.27 (0.23 ± 0.01, N=19) times SLV, 1.4–1.6 (1.47 ± 0.06, N=19) times as long as wide, 1.3–1.5 (1.41 ± 0.06,
N=19) times as wide as high. Snout round, moderately elongate (Fig 9A), gently sloping toward top of head. Neck
slightly narrower than head and anterior portion of body. Body cylindrical. Limbs well developed, lower arm 0.13–
0.15 (0.14 ± 0.01, N=19) times SVL, lower leg 0.12–0.16 (0.15, ± 0.01, N=19) times SVL.
Rostral convex, 1.8–2.4 (2.11 ± 0.17, N=19) times as wide as high; posterior margin with a shallow depression
medially and slightly indented by 1–3 medial postrostrals, with or without a medial cleft extending anteriorly.
Postrostrals 3 (79%), 4 (10.5%), or 5 (10.5%), laterals ones (supranasals) distinctly larger than medial ones (N=19)
(Fig 9A). Nasal bordered by rostral, first supralabial (only a narrow contact in some specimens), 3 (68.4%) or 4
(31.6%) postnasals (N=19), and supranasal; internostril distance 0.2–0.3 (0.25 ± 0.02, N=19) times head width.
Supranasal roughly oval or circular, 1.0–1.5 (1.20 ± 0.14; N=19) times as wide as long. Postnasals slightly larger than,
or 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, 10–13 (11.1 ± 0.8, N=19) loreals on a line between postnasals and anterior corner of eye. Top
and posterior portion of head, as well as supraorbital region, with granular scales. A short supraciliary flap present,
anteriorly with 7–10 (8.6 ± 1.0, N=19) enlarged and flattened scales, among which 0–4 (2.6 ± 1.1; N=19) small,
conical spines. Pupil round, eye diameter 0.20–0.27 (0.23 ± 0.01, N=19) times head length. Scales on temporal region
similar to those on top of head. Ear-opening much smaller than eye, oval, posterior to, and at same level of,
commissure of mouth. Supralabials 5–8 (6.7 ± 0.7; N=19), distinctly enlarged anteriorly and decreasing in size
posteriorly, one or two of them posterior to centre of eye, followed to corner of mouth by small scales.
Zootaxa 2869 © 2011 Magnolia Press · 19
GONATODES CONCINNATUS COMPLEX
Mental large, distinctly wider anteriorly than posteriorly, with posterior margin forming a wide angle, 1.1–1.4
(1.25 ± 0.11; N=19) times as wide as long; 1 (10.5%), 2 (63.2%) or 3 (26.3%) postmentals (Fig. 9B). Scales on chin
flat, smooth, polygonal, juxtaposted, larger anteriorly, decreasing in size posteriorly. Infralabials 5–7 (5.8 ± 0.7;
N=19), distinctly enlarged anteriorly and decreasing in size posteriorly; 1–3 of them posterior to centre 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
granular, posteriorly flat, smooth, hexagonal or round, increasing in size toward the posterior region, with a short
transitional zone between the anterior and posterior parts.
Dorsals granular, increasing in size toward the flanks. Ventrals larger than dorsals, roughly hexagonal, flat,
smooth, imbricate, in oblique rows; 53–63 (57.4 ± 3.3; N=19) scales along the midventral line between anterior
margin of forelimbs and vent; 18–21 (19.2 ± 0.9; N=18) scales in a transverse line at midbody, with a short
transitional zone between ventrals and scales on flanks. Scales around midbody 118–133 (124.2 ± 3.2; N=18).
Scales on preanal plate similar to ventrals, except for those bordering vent, which are very small. Escutcheon
present in males on posterior portion of belly and on four rows (body-knee direction) on ventral surface of thighs.
Scales on anterodorsal surface of forelimbs flat, smooth, roundish or rhomboidal, imbricate, largest close to the
wrist; on posterodorsal and ventral surface of forelimbs convex, smooth, rhomboidal, juxtaposed, relatively small.
Scales on anterodorsal surface of thighs and ventral surface of hind limbs flat, smooth, roundish or rhomboidal,
imbricate and relatively larger; on posterodorsal surface of thighs and dorsal surface of lower legs smaller, convex,
smooth, rhomboidal, juxtaposed.
Lamellae under second (II) through fourth (IV) fingers (proximal lamellae in parentheses): II: 15–18 (5–6),
III: 18–20 (5–7), and IV: 18–22 (6–8) (fig 9C). Lamellae under second through fourth toes (proximal lamellae in
parentheses): II: 14–18 (5–7), III: 16–22 (5–8), and IV: 22–27 (10–12) (Fig. 9D). Claws exposed, non-retractile,
between two basal scales. Fingers and toes with three, occasionally four, lateral scales distally, between the fourth
(counted from the claw towards the hand) subdigital lamella, and the dorsal scale.
Scales on tail dorsally and laterally relatively small, rhomboidal, flat, smooth, imbricate. On ventral surface of
tail scales smooth, flat, imbricate, increasing in size toward midventral line; midventral scales, except close to the
base of the tail, distinctly wider than long, forming a repetitive sequence of two single midventrals (one after the
other), each in contact laterodistally with one scale per side, followed by a divided (only on proximal portion of
tail, when present) or single midventral scale in contact laterodistally with two scales per side – respectively 1’1’2”
and 1’1’1” in the codification of Avila-Pires (1995: Figure 2) (Fig. 9E).
Color in preservative. In males (Fig 1C), dorsal surface of head with relatively large reddish-brown and beige
vermiculations. Sides of head with oblique and/or longitudinal, beige and brown bands. Back and flanks, base of
tail and limbs with beige and brown vermiculations. A large, conspicuous, white suprahumeral ocellus with black
margin. Ventral surface of head and gular region beige with oblique black streaks, in contact or not at midventral
line; chest beige; belly dark-brown; underside of limbs and base of tail brown. Dorsal surface of tail brown, ventral
surface reddish-brown; distal part in one specimen with a beige ring around the tail. Escutcheon area (belly and
thighs) light gray.
In females, dorsal surface of head and limbs with brown and beige irregular spots. Sides of head with beige and
brown oblique and/or longitudinal bands. Back gray with dorsolateral pairs of black and white spots; flanks gray
with black and brown irregular spots. A moderately large, conspicuous, white suprahumeral ocellus, with black
margin, smaller than that of males. Ventral surface of head and gular region white with dark oblique streaks, in
contact or not at midventral line; belly and underside of limbs light gray. Tail brown and/or black dorsally, white
and/or brown ventrally; original tail distally with white bands that form complete rings around the tail.
Color in life. In males (Fig. 10A–B), head dorsally and laterally reddish-brown and vivid yellow.
Suprahumeral ocellus white with black margin. Vermiculations on back brown and yellow, smaller vermiculations
on flanks bluish-white and black. Head ventrally and gular region vivid yellow with reddish-brown streaks, chest
vivid yellow. Remaining ventral region dark gray, lighter on escutcheon areas (belly and thighs). Tail dark gray to
black all around, except near the base, where it is similar to the back.
In females (Fig. 10C–D), head dorsally and lateraly gray and brown. Suprahumeral ocellus white with black
margin. Back brown with pairs of black and gray spots. Head ventrally and gular region beige with light-yellow
and gray streaks, chest white. Remaining ventral region white. Tail anteriorly brown with black spots dorsally,
orange ventrally; posteriorly with black and white rings.
STURARO & AVILA-PIRES
20 · Zootaxa 2869 © 2011 Magnolia Press
FIGURE 9. Gonatodes tapajonicus (paratype, MZUSP 53674). (A) Dorsal and (B) ventral views of head; ventral views of (C)
right hand and (D) right foot; (E) ventral view of tail. Scale bar = 1 mm.
Zootaxa 2869 © 2011 Magnolia Press · 21
GONATODES CONCINNATUS COMPLEX
FIGURE 10. Gonatodes tapajonicus from Aldeia Nova, righ margin of Rio Tapajós, Pará, Brazil. Adult male (A) in dorsal
(MPEG 27714, SVL = 52.5 mm) and (B) ventral (MPEG 27718, SVL = 48.7 mm) views. Adult female (C) in dorsal (MPEG
27715, SVL = 39.5 mm) and (D) ventral (MPEG 27716, SVL = 51.1 mm) views (Photo C by Pedro L. Peloso).
Distribution. Interfluvium Tapajós–Xingu, state of Pará, Brazil (Fig. 3A).
Remarks. Rodrigues (1980) pointed out that the type-series was collected in an area with predominance of the
palm Euterpe oleracea (“açaizal”), along a trail known as ‘Boa Fé’, in front of Cachoeira do Limão, Tapajós river,
at approximately 04o41´S 45o21´W. It is possible that it is the same locality where MPEG 27708–27718 have been
collected, community ‘Boa Fé’ being the old name of this locality. Similar to the type-series, these specimens were
in an “açaizal” in a flooded depression within the forest; they were found mostly on the palm Euterpe oleracea, one
specimen on a tree trunk, between 30 and 200 cm above the ground. Contrary to what was observed by Rodrigues
(1980), we also found G. humeralis in the “açaizal”, in syntopy with G. tapajonicus. The Euterpe oleracea forest
was one of the few in the area still well-preserved, due to the exploitation in other similar environments of the palm
heart for the food market. The species was not found in the more disturbed ‘açaizais’, nor outside this habitat.
Gonatodes nascimentoi sp. nov.
(Fig. 1D, 11, 12)
Holotype (number in bold between parentheses refer to localities in Fig. 3): MPEG 25596 (field number BML
446), an adult male from Fazenda Caracol, right margin of Rio Xingu, Município Anapu, Pará, Brazil (3o27’30”S
and 51o40’33”W) (50), collected in pitfall trap inside primary forest, by A. A. Lima, M. J. Sturaro and R. A. T.
Rocha, 12 January 2008.
Paratypes (numbers in bold between parentheses refer to localities in Fig. 3): BRASIL. AMAPÁ. Município
Ferreira Gomes. Floresta Nacional do Amapá (01º06’37’’N and 51º53’37’’W) (48): IEPA/FL 361, male, leg. J.
Lima, June 2005. PARÁ. Município Almeirim. Monte Dourado, forest near Estação Ecológica do Jari (0o35’27’’S
and 52º44’09’’W) (49): MPEG 23822–27, 5 males and a female, leg. T. A. Gardner and M. A. Ribeiro-Jr, May–
June 2005; MPEG 27719, female, leg. M. J. Sturaro and J. O. Gomes, January 2009. Município Anapu. Type-
STURARO & AVILA-PIRES
22 · Zootaxa 2869 © 2011 Magnolia Press
locality (50): MPEG 25164, male, leg. A. A. Lima and R. A. T. Rocha, November 2007; MPEG 25595, 25597, a
juvenile, a female and a male, leg. A. A. Lima, M. J. Sturaro and R. A. T. Rocha, January 2008; MPEG 25598–
25601, a male and 3 females, leg. A. A. Lima, M. J. Sturaro, R. A. T. Rocha, P. L. V. Peloso and F. S. Rodrigues,
March 2008. Município Portel. Fazenda Riacho Monte Verde (~ 3o15’S 50o19’W) (51): MPEG 24643, female,
leg. T.C.S. Avila Pires and J.O. Gomes, February 2007; MPEG 24644–50, 6 males and a female, leg. J. O. Gomes,
March 2007.
Diagnosis. A relatively large Gonatodes, with maximum SLV of 55.7 mm. Scales around midbody 118–143.
Ventral scales in a longitudinal row 49–60. Proximal subdigital lamellae as wide as digits, in total 18–23 under
fourth finger, 23–28 under fourth toe. Three, occasionally four, lateral rows of scales on distal portion of fingers
and toes. Tail with midventral scales distinctly wider than long, forming a repetitive sequence of two single
midventrals (one after the other), each in contact laterodistally with one scale per side, followed by a divided (only
on proximal portion of tail, when present) or single midventral in contact laterodistally with two scales per side
(respectively 1’1’2” and 1’1’1”). A white suprahumeral bar, bordered by black, both in males and females (but
thinner in the latter); from very short (hardly higher than long) to almost reaching the middorsal region. Males with
a vermiculated pattern of light and dark (in life respectively drab or orange-yellow, and reddish-brown) spots on
head and body dorsally; gular region light with dark oblique streaks or redddish with white oblique streaks (orange
with orange-yellow streaks in life).
Description. Maximum SVL in males of 55.7 mm (MPEG 23822), in females of 54.0 mm (MJS 035). Tail
round in cross section, tapering toward tip, 1.1–1.3 (1.15 ± 0.09, N=8) times SLV. Head length 0.21–0.28 (0.24 ±
0.02, N=24) times SLV, 1.2–1.6 (1.40 ± 0.08, N=24) times as long as wide, 1.2–1.5 (1.40 ± 0.10, N=24) times as
wide as high. Snout round, moderately elongate (Fig 11A), gently sloping toward top of head. Neck slightly
narrower than head and anterior portion of body. Body cylindrical. Limbs well developed, lower arm 0.14–0.17
(0.15 ± 0.01, N=24) times SVL, lower leg 0.13–0.17 (0.15 ± 0.01, N=24) times SVL.
Rostral convex, 1.8–3.0 (2.01 ± 0.28, N=24) times as wide as high; posterior margin with a shallow depression
medially and slightly indented by 0–2 medial postrostrals, with or without a medial cleft extending anteriorly.
Postrostrals 2 (12.5%), 3 (83.3%), or 4 (4.2%), laterals ones (supranasals) distinctly larger than medial one(s),
when these are present (N=24) (Fig 11A). Nasal bordered by rostral, first supralabial (only a narrow contact in
some specimens), 3 postnasals (N=24), and supranasal; internostril distance 0.14.–0.21 (0.17 ± 0.01, N=24) times
head width. Supranasal roughly oval, circular or semicircular, 0.9–1.4 (1.13 ± 0.13; N=24) times as wide as long.
Postnasals slightly larger than, or 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, 10–13 (10.2 ± 0.8, N=24) loreals on a line between
postnasals and anterior corner of eye. Top and posterior portion of head, as well as supraorbital region, with
granular scales. A short supraciliary flap present, anteriorly with 7–12 (9.5 ± 1.4, N=24) enlarged and flattened
scales, among which 0–3 (1.2 ± 1.3; N=24) small, conical spines. Pupil round, eye diameter 0.20–0.27 (0.22 ± 0.02,
N=24) times head length. Scales on temporal region similar to those on top of head. Ear-opening much smaller than
eye, oval, posterior to, and at same level of, commissure of mouth. Supralabials 5–7 (6.2 ± 0.5; N=24), distinctly
enlarged anteriorly and decreasing in size posteriorly, one or two of them posterior to centre of eye, followed to
corner of mouth by small scales.
Mental large, distinctly wider anteriorly than posteriorly, with posterior margin forming a wide angle or a right
angle, 1.2–1.4 (1.28 ± 0.7; N=24) times as wide as long; 2 (95.8%), or 3 (4.2%) postmentals (Fig. 11B). Scales on
chin flat, smooth, polygonal, juxtaposted, larger anteriorly, decreasing in size posteriorly. Infralabials 5–8 (6.4 ±
0.9; N=24), distinctly enlarged anteriorly and decreasing in size posteriorly; 1–3 of them posterior to centre 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
granular, posteriorly flat, smooth, hexagonal or round, increasing in size toward the posterior region, with a short
transitional zone between the anterior and posterior parts.
Dorsals granular, increasing in size toward the flanks. Ventrals larger than dorsals, roughly hexagonal, flat,
smooth, imbricate, in oblique rows; 52–60 (56.3 ± 2.4; N=22) scales along the midventral line between anterior
margin of forelimbs and vent; 18–24 (21.2 ± 1.3; N=22) scales in a transverse line at midbody, with a short
transitional zone between ventrals and scales on flanks. Scales around midbody 118–143 (129.9 ± 6.4; N=22).
Scales on preanal plate similar to ventrals, except for those bordering vent, which are very small. Escutcheon
present in males on posterior portion of belly and on 4–5 rows (body-knee direction) on ventral surface of thighs.
Zootaxa 2869 © 2011 Magnolia Press · 23
GONATODES CONCINNATUS COMPLEX
Scales on anterodorsal surface of forelimbs flat, smooth, roundish or rhomboidal, imbricate, largest close to the
wrist; on posterodorsal and ventral surface of forelimbs convex, smooth, rhomboidal, juxtaposed, relatively small.
Scales on anterodorsal surface of thighs and ventral surface of hind limbs flat, smooth, roundish or rhomboidal,
imbricate and relatively larger; on posterodorsal surface of thighs and dorsal surface of lower legs smaller, convex,
smooth, rhomboidal, juxtaposed.
Lamellae under second (II) through fourth (IV) fingers (proximal lamellae in parentheses): II: 15–19 (4–6),
III: 18–22 (5–6), and IV: 18–23 (6–8) (Fig 11C). Lamellae under second through fourth toes (proximal lamellae in
parentheses): II: 15–19 (5–6), III: 18–23 (5–7), and IV: 23–28 (9–12) (Fig. 11D). Claws exposed, non-retractile,
between two basal scales. Fingers and toes with three, occasionally four, lateral scales distally, between the fourth
(counted from the claw towards the hand) subdigital lamella, and the dorsal scale.
Scales on tail dorsally and laterally relatively small, rhomboidal, flat, smooth, imbricate. On ventral surface of
tail scales smooth, flat, imbricate, increasing in size toward midventral line; midventral scales, except close to the
base of the tail, distinctly wider than long, forming a repetitive sequence of two single midventrals (one after the
other), each in contact laterodistally with one scale per side, followed by a divided (only on proximal portion of
tail, when present) or single midventral scale in contact laterodistally with two scales per side – respectively 1’1’2”
and 1’1’1” in the codification of Avila-Pires (1995: Figure 2) (Fig. 11E).
Measurements of holotype (in millimeters): SVL 48.7, TL 54.0, HL 11.7, HW 7.9, HD 6.8, ED 2.6, IND 2.0,
SSL 0.8, SSW 1.12, RSL 1.3, RSW 2.5, MSL 2.6, MSW 3.2, LAL 8.1, LLL 7.8, HSH 5.34, HSL 2.6.
Scale counts of holotype: SAM 127 scales, VLR 55, VTR 21, SL 6, IL 5, PR 3, SP 1, PN 3, LS 12, PM 2, SCS
8, SSC 3, PL2F 5, DL2F 12, LS2F 3, PL3F 4, DL3F 14, LS3F 4, PL4F 6, DL4F 13, LS4F 4, PL2T 5, DL2T 12,
LS2T 4, PL3T 5, DL3T 15, LS3T 3, PL4T 10, DL4T 13, LS4T 3, RSE 4.
Color in preservative. In males (Fig. 1D), dorsal surface of head with reddish-brown and beige
vermiculations. Sides of head, in some specimens, with oblique and/or longitudinal, beige and brown bands. Back
and flanks, base of tail and hind limbs with relatively large, beige color and dark-brown vermiculations. A
conspicuous, white suprahumeral vertical bar with black margin, which varies from only slightly to distinctly
higher than long (see under “Remarks” below). Ventral surface of head and gular region beige with black streaks,
in some specimens dark-brown with beige streaks, in contact or not at midvental line; chest beige; belly dark-
brown; underside of limbs and base of tail brown. Dorsal surface of tail brown, underside dark-brown or black; a
white band may be present distally, forming a complete ring around the tail. Escutcheon area (belly and thighs)
light gray.
In females, dorsal surface of head and limbs with brown and beige irregular spots. Back gray with dorsolateral
pairs of black and white spots; flanks gray with black and brown irregular spots. Suprahumeral bar conspicuous,
white with black margins, thinner than that of males; in some cases they almost reach the middorsal region. Ventral
surface of head and gular region white with black oblique streaks, in contact or not at midventral line; belly and
underside of limbs light-gray. Tail gray dorsally and white ventrally; original tail distally with white and black
bands that form complete rings around the tail.
Color in life (Photos of MPEG 23822, 23824–25, 25596–99; field notes of MPEG 24643): In males (Fig.
12A–B), head dorsally and laterally with orange and orange-yellow, or orange and drab, vermiculations. On back
vermiculations may be smaller than, or similar in size to, those on head and the lighter spots are frequently
bordered by black (occasionally also on head). Suprahumeral bar or ocellus white with black margins. Flanks with
smaller, bluish-white and black vermiculations. Head ventrally and gular region orange with orange-yellow
streaks, chest orange. Remaining ventral region plumbeous, lighter on escutcheon areas (belly and thighs). Tail
dark gray to black all around, except near the base, where it is similar to the back; distal portion with white spots.
Color in life of MPEG 24643, a male, was described based on the color guide of Smithe (1975): Head dorsally
“buff” (24) and “mahogany red” (132B); body “brick red” (132A) with “chamois” (123D) vermiculation; a white
bar with black margins; lower flanks “blackish neutral gray” (82) with “pearl gray” (81) spots. Ventrally, head and
chest (until forearm) “buff” (24) with convergent “scarlet” (14) streaks; otherwise ventral region and tail mainly
“blackish neutral gray” (82), midventrally medium (84) to light (85) neutral gray; tail dorsally with a darker and
lighter pattern (TCSAP, field notes).
Females (Fig. 12C–D) dorsally brown or grayish-brown, with a paired series of dark brown to black irregular,
transversely elongate, spots. Suprahumeral bar white with black margins, thinner than those in males. Head
ventrally and gular region white with black streaks, chest light-gray. Tail gray with black spots all around; posterior
portion with white spots.
STURARO & AVILA-PIRES
24 · Zootaxa 2869 © 2011 Magnolia Press
FIGURE 11. Gonatodes nascimentoi sp. nov. (holotype, MPEG 25596). (A) Dorsal and (B) ventral views of head; ventral
views of (C) right hand and (D) right foot; (E) ventral view of tail. Scale bar = 1 mm.
Zootaxa 2869 © 2011 Magnolia Press · 25
GONATODES CONCINNATUS COMPLEX
FIGURE 12. Gonatodes nascimentoi sp. nov. from Anapu, Rio Xingu, Pará, Brazil. Adult male (paratype, MPEG 25598, SVL
= 46.7 mm) in dorsal (A) and ventral (B) views; adult female (paratype, MPEG 25599, SVL = 39.7 mm) in dorsal (C) and
ventral (D) views. (Photos by Pedro L. Peloso).
Distribution. Eastern Brazilian Amazonia, in the states of Pará and Amapá. Until now it is only known south
of the Amazon river in the interfluvium Xingu–Tocantins, Pará state, and north of the Amazon river east of Paru
river, in the states of Pará and Amapá (Fig. 3A).
Etymology. The species is named after Francisco Paiva do Nascimento, who worked for many years in Museu
Paraense Emílio Goeldi and contributed to our knowledge of Amazonian reptiles, most of all about the snakes from
eastern Amazonia.
Remarks. The suprahumeral spot varies among populations and, to a smaller degree, within populations. In
the only specimen from Amapá examined, the spot has the shape of a short bar, almost an ocellus. Specimens from
Pará, north of the Amazon River, show a vertical bar reaching dorsally at least the dorsolateral region, in some
cases almost the middorsal region. South of the Amazon, all specimens from Monte Verde present a short vertical
bar (not reaching the dorsolateral region, but always with dorso-ventral axis longer than antero-posterior axis),
while in the specimens from Anapu the vertical bar is similar to those from northern Pará (although one specimen
presents a vertical bar and a small ocellus above the bar, on both sides).
Specimens were collected by active search or in pitfall traps, in primary or little-disturbed forest. Six
specimens were found on the buttresses or trunks of live trees, between 15–50 cm above ground; one was inside the
loose bark of a dead tree, 120 cm above ground; a male and a female were on a fallen trunk; and at the Xingu river
a pair was found on a rock, 50 cm above ground. At this locality specimens hid inside rock crevices or tree trunk
holes when disturbed. Weights vary between 0.2–5.1 g.
STURARO & AVILA-PIRES
26 · Zootaxa 2869 © 2011 Magnolia Press
Gonatodes riveroi sp. nov.
(Figs. 1E, 13, 14)
Gonatodes concinnatus: Vanzolini, 1955: 126 (part); 1968: 26 (part); Mechler, 1968: 331 (part); Peters & Donoso-Barros, 1970
(part); Rivero-Blanco, 1979: 94 (part); Sanchez, Castaño & Cardenas, 1995: 317; Bartlett & Bartlett, 2003: 158 (part).
Holotype (number in bold between parentheses refer to localities in Fig. 3): MZUSP 49153 (field number
FMEDEM 1465), adult male, from Villavicencio, State of Meta, Colombia (~4o09’S and 73o37’W) (37), collected
by W. W. Lamar and F. Medem, 22 October 1977.
Paratypes (numbers in bold between parentheses refer to localities in Fig. 3): COLOMBIA. BOYACÁ.
Garagoa (34): CAS 71238, USNM 84968, USNM 92490–92, a male and 4 females, leg. Nicéforo Maria, 1931.
BOYACÁ–CUNDINAMARCA–META border. Guaicáramo (according to Paynter Jr., 1997 in the hills near
confluence of Rio Guavio with Rio Upía, at eastern base of the eastern Andes) (35): USNM 84972, female, leg.
Nicéforo Maria, 1931. META. San Juan de Arama (36): KU 192635–36, a male and a female, leg. M. Chin, August
1967. Type-locality (37): MCZ 132796–97, a juvenile and a female, July 1972; AMNH 35280–87, AMNH 35290–
91, 5 males and 5 females, leg. Nicéforo Maria, January 1927; KU 110579–80, a juvenile and a male, collector not
specified, July 1967; KU 151933–35, 3 males, collector not specified, May 1967; MCZ 19213, male, January
1924; MCZ 77388–89, a male and a female, collector and date not specified; MCZ 110009–14, a juvenile, 3 males
and 2 females, June 1968; MCZ 110741–42, 2 males, leg. G. Gorman, June 1968; MZUSP 2145–46 (former
AMNH 35292–93), 2 females, leg. Nicéforo Maria, no date; MZUSP 49152, female, leg. M. Lugor and F. Medem,
October 1977; MZUSP 49154, female, leg. W. W. Lamar and F. Medem, October 1977; USNM 84969, male, leg.
Nicéforo Maria, 1931. Villavicencio, Finca “El Buque” (37): MZUSP 49155–62, 3 males and 5 females, leg.
E.Thierry and F. Medem, October 1977. Villavicencio, Instituto 'Roberto Franco' (37): MCZ 77391–97, 2
juveniles, 2 males and 3 females, collector not specified, July 1963; MCZ 146137, male, collector not specified,
October 1974. 5 km NE of Villavicencio (37): UMMZ 127206, male, leg. W. Mobely and K. Adler, July 1965. Rio
Negro, c. 12 km from Villavicencio, Finca 'Las Orquideas' (“below” [south of?] road Bogotá–Villavicencio) (37):
MCZ 150022, male, leg. F. Medem, 1975. Güejar River, Finca Guadualito (38): MZUSP 44777, female, leg. L.
Klein and F. Medem, September 1967. Puerto Lopez (39): AMNH 97350, juvenile, leg. A.V. Hutchison and
Mahoney, January 1966; AMNH 98406, female, leg. V.H. Hutchison, April 1966.
Diagnosis. A moderately large Gonatodes, with maximum SLV of 47.6 mm. Scales around midbody 96–123.
Ventral scales in a longitudinal row 44–55. Proximal subdigital lamellae as wide as digits, in total 14–21 under
fourth finger, 17–25 under fourth toe. Two or three lateral rows of scales on distal portion of fingers and toes. Tail
with midventral scales distinctly wider than long, forming a repetitive sequence of two single midventrals (one
after the other), each in contact with one laterodistal scale per side, followed by a divided (only on proximal portion
of tail, when present) or single midventral in contact with two laterodistal scales per side (respectively 1’1’2” and
1’1’1”). A white suprahumeral bar bordered by black, present both in males and females, although more
conspicuous in males; it reaches dorsally at least up to the dorsolateral region, in some females it almost reaches
the middorsal region; never in the form of an ocellus. Males with head dorsally without vermiculation; back and
limbs finely vermiculated; gular region without dark streaks.
Description. Maximum SVL in males of 47.6 mm (USNM 84968), in females of 47.1 mm (USNM 92490).
Tail round in cross section, tapering toward tip, 1.0–1.3 (1.17 ± 0.11, N=11) times SLV. Head length 0.22–0.28
(0.24 ± 0.01, N=61) times SLV, 1.3–1.6 (1.46 ± 0.05, N=57) times as long as wide, 1.2–1.8 (1.38 ± 0.10, N=56)
times as wide as high. Snout round, moderately elongate (Fig. 13A), gently sloping toward top of head. Neck
slightly narrower than head and anterior portion of body. Body cylindrical. Limbs well developed, lower arm 0.11–
0.16 (0.13 ± 0.01, N=62) times SVL, lower leg 0.12–0.16 (0.14 ± 0.01, N=62) times SVL.
Rostral convex, 1.6–2.3 (1.96 ± 0.13, N=61) times as wide as high; posterior margin with a shallow depression
medially and slightly indented by 0–3 medial postrostrals, with or without a median cleft extending anteriorly.
Postrostrals 2 (4.9%), 3 (82%), 4 (4.9%) or 5 (8.2%), lateral ones (supranasals) distinctly larger than medial one(s),
when these are present (N=61) (Fig 13A). Nasal bordered by rostral, first supralabial (only a narrow contact in
some specimens), 2 (96.8%) or 3 (3.2%) postnasals (N=63), and supranasal; internostril distance 0.20.–0.27 (0.24 ±
0.02, N=55) times head width. Supranasal roughly oval or circular, 0.9–1.6 (1.23 ± 0.15; N=62) times as wide as
long. Postnasals slightly larger than, or 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
Zootaxa 2869 © 2011 Magnolia Press · 27
GONATODES CONCINNATUS COMPLEX
elongate than those on snout, largest on row adjacent to supralabials, 9–11 (9.8 ± 0.7, N=61) loreals on a line
between postnasals and anterior corner of eye. Top and posterior portion of head, as well as supraorbital region,
with granular scales. A short supraciliary flap present, anteriorly with 6–12 (8.3 ± 1.0, N=60) enlarged and
flattened scales, among which 0–5 (1.7 ± 1.4; N=60) small, conical spines. Pupil round, eye diameter 0.19–0.26
(0.22 ± 0.01, N=56) times head length. Scales on temporal region similar to those on top of head. Ear-opening
much smaller than eye, oval, posterior to, and at same level of, commissure of mouth. Supralabials 5–8 (6.1 ± 0.6;
N=62), distinctly enlarged anteriorly and decreasing in size posteriorly, one or two of them posterior to centre of
eye, followed to corner of mouth by small scales.
Mental large, distinctly wider anteriorly than posteriorly, with posterior margin forming a moderate or wide
angle, 1.1–1.5 (1.27 ± 0.08; N=59) times as wide as long; 1 (1.6%), 2 (76.2%), 3 (19.0%), or 4 (3.2%) postmentals
(Fig. 13B). Scales on chin flat, smooth, polygonal, juxtaposted, larger anteriorly, decreasing in size posteriorly.
Infralabials 4–7 (5.4 ± 0.7; N=63), distinctly enlarged anteriorly and decreasing in size posteriorly; one or two of
them posterior to centre 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
granular, posteriorly flat, smooth, hexagonal or round, increasing in size toward the posterior region, with a short
transitional zone between the anterior and posterior parts.
Dorsals granular, increasing in size toward the flanks. Ventrals larger than dorsals, roughly hexagonal, flat,
smooth, imbricate, in oblique rows; 44–56 (50.8 ± 2.3; N=58) scales along the midventral line between anterior
margin of forelimbs and vent; 15–21 (18.0 ± 1.2; N=62) scales in a transverse line at midbody, with a short
transitional zone between ventrals and scales on flanks. Scales around midbody 96–123 (110.1 ± 5.6; N=60). Scales
on preanal plate similar to ventrals, except for those bordering vent, which are very small. Escutcheon present in
males on posterior portion of belly and on 3–5 rows (body-knee direction) on ventral surface of thighs.
Scales on anterodorsal surface of forelimbs flat, smooth, roundish or rhomboidal, imbricate, largest close to the
wrist; on posterodorsal and ventral surface of forelimbs convex, smooth, rhomboidal, juxtaposed, relatively small.
Scales on anterodorsal surface of thighs and ventral surface of hindlimbs flat, smooth, roundish or rhomboidal,
imbricate and relatively larger; on posterodorsal surface of thighs and dorsal surface of lower legs smaller, convex,
smooth, rhomboidal, juxtaposed.
Lamellae under second (II) through fourth (IV) fingers (proximal lamellae in parentheses): II: 13–17 (5–6),
III: 15–19 (4–6), and IV: 14–21 (5–8) (Fig 13C). Lamellae under second through fourth toes (proximal lamellae in
parentheses): II: 12–18 (5–7), III: 16–21 (5–8), and IV: 17–25 (8–12) (Fig. 13D). Claws exposed, non-retractile,
between two basal scales. Fingers and toes with three, occasionally two, lateral scales distally, between the fourth
(counted from the claw towards the hand) subdigital lamella, and the dorsal scale.
Scales on tail dorsally and laterally relatively small, rhomboidal, flat, smooth, imbricate. On ventral surface of
tail scales smooth, flat, imbricate, increasing in size toward midventral line; midventral scales, except close to the
base of the tail, distinctly wider than long, forming a repetitive sequence of two single midventrals (one after the
other), each in contact laterodistally with one scale per side, followed by a divided (on proximal portion of tail,
when present) or single midventral scale in contact laterodistally with two scales per side – respectively 1’1’2” and
1’1’1” in the codification of Avila-Pires (1995: Figure 2) (Fig. 13E).
Measurements of holotype (in millimeters): SVL 42.5, TL 46.0 (distally regenerated), HL 9.8, HW 6.7, HD
4.7, ED 2.0, IND 1.7, SSL 0.7, SSW 0.8, RSL 1.1, RSW 2.1, MSL 2.2, MSW 2.6, LAL 5.3, LLL 6.2, HSH 2.8,
HSL 1.57.
Scale counts of holotype: SAM 116 scales, VLR 51, VTR 17, SL 6, IL 6, PR 3, SP 1, PN 3, LS 9, PM 2, SCS
8, SSC 2, PL2F 6, DL2F 11, LS2F 3, PL3F 6, DL3F 13, LS3F 3, PL4F 7, DL4F 11, LS4F 3, PL2T 6, DL2T 11,
LS2T 3, PL3T 6, DL3T 13, LS3T 3, PL4T 11, DL4T 12, LS4T 3, RSE 4.
Color in preservative. In males (Fig. 1E), dorsal surface of head reddish-brown, spotless. Back and flanks,
base of tail and hind limbs with relatively small, beige and brown vermiculations. A large, conspicuous, white
suprahumeral bar with black margins, extending dorsally at least up to the dorsolateral region. Ventral surface of
head, gular region and chest brown, without oblique streaks; belly dark-brown or dark-gray; underside of limbs
beige. Tail dark brown dorsally and brown ventrally. Escutcheon area (belly and thighs) light gray.
In females, dorsal surface of head and limbs light-brown or gray, with black irregular spots; brown and black
oblique and/or longitudinal bands may be present on sides of head. Back gray with dorsolateral pairs of black
spots; flanks gray with black and brown irregular spots. A moderately large, conspicuous, white suprahumeral bar,
STURARO & AVILA-PIRES
28 · Zootaxa 2869 © 2011 Magnolia Press
FIGURE 13. Gonatodes riveroi sp. nov. (holotype, MZUSP 49153). (A) Dorsal and (B) ventral views of head; ventral views
of (C) right hand and (D) right foot; (E) ventral view of tail. Scale bar = 1 mm.
Zootaxa 2869 © 2011 Magnolia Press · 29
GONATODES CONCINNATUS COMPLEX
FIGURE 14. Gonatodes riveroi sp. nov. from Villavicencio, Meta, Colombia (UMMZ 127206, SVL = 41.9 mm). (Photo by
Kraig Adler).
with black margins, but less defined than that on males; in some specimens it almost reaches the middorsal region.
Ventral surface of head and gular region white with dark oblique streaks, in contact or not at midventral line; belly and
underside of limbs white or beige. Tail gray dorsally and white ventrally; original tail distally with white and black
bands that form complete rings around the tail.
Color in life (photo of UMMZ 127206, male): Head dorsally and laterally brown. Back with brown and yellow
vermiculations. Suprahumeral bar white with black margins. Tail dark-brown (Fig. 14).
Distribution. Central Colombia east of the Andes (Fig. 3A). Sanchez et al. (1995) reported G. concinnatus in
Colombia from “Amazonas, Casanare, Boyacá, Cundinamarca, Meta, Pasto, Orinoco, Guaviare.” It is necessary,
however, to verify whether all these records refer to the species here described.
Etymology. The species is named after Dr. Carlos Rivero-Blanco, who made an extensive study on Gonatodes, as
well as contributions to environmental conservation in Venezuela.
Remarks. Rivero-Blanco (1979) considered the specimens from Villavincencio and vicinity, in Meta, Colombia,
as intergrades between Gonatodes concinnatus concinnatus, from Peru and Ecuador, and G. c. ligiae, from Venezuela.
Species comparisons
The five species considered here under the term Gonatodes concinnatus complex are easily distinguishable from the
remaining 22 species of Gonatodes by the presence in males of a moderately large, white suprahumeral spot with
black margin, and vermiculations on back (the latter absent in Gonatodes ligiae). A white antehumeral bar is present
in G. ceciliae Donoso-Barros, 1966, G. humeralis (Guichenot, 1855), and G. ocellatus (Gray, 1831). G. seig liei is
described by Donoso-Barros (1966) as presenting a “prescapular” ocellus, while Rivero-Blanco (1979) indicates that
females present a “prehumeral” ocellus. This latter author refers to the spot in G. concinnatus as “scapular” or
“suprascapular”, indicating a more anterior position of the spot in G. seigliei in relation to the species of the G.
STURARO & AVILA-PIRES
30 · Zootaxa 2869 © 2011 Magnolia Press
concinnatus complex. None of the remaining species have a similar bar or spot on the humeral region or
vermiculations on back.
The species of the G. concinnatus complex differ moreover from G. alexandermendesi Cole & Kok, 2006, G.
hasemani Griffin, 1917, G. infernalis Rivas & Schargel, 2008, and G. astralis Schargel et al., 2010, by the absence of
a very elongate spine on the supraciliary flap. They differ from G. annularis Boulenger, 1887, G. caudiscutatus
(Günther, 1859), G. el adio i Nascimento, Avila-Pires & Cunha, 1987, G. lichenosus Rojas-Runjaic, Infante-Rivero,
Cabello & Velozo, 2010, G. infernalis, G. hasemani, G. astralis, and G. seigliei by the subcaudal sequence (1’1’2” in G.
annularis, 1’1” anteriorly and 1’1’1” posteriorly in G. seigliei, midventrals not enlarged in G. hasemani and G.
astralis, 1’2” anteriorly and 1’1” posteriorly in G. lichenosus, 1’1” in the remaining three species). From G. antillensis
(Lidth de Jeude, 1887) they differ by the shape of the pupil (complex and vertical in G. antillensis).
Gonatodes tapajonicus and G. nascimentoi reach a larger SVL than Gonatodes albogularis (Duméril & Bibron,
1836), G. antillensis, G. atricucullaris Noble, 1921, G. caudiscutatus, G. daudini Powell & Henderson, 2005, G.
eladioi, G. falconensis Shreve, 1947, G. humeralis, G. petersi Donoso-Barros, 1967, and G. vittatus (Lichtenstein,
1856), none of which exceed 42 mm in SVL.
Within the Gonatodes concinnatus complex, color pattern separates three groups – (1) vermiculation present on
head and body, gular region with dark streaks (G. tapajonicus and G. nascimentoi sp. nov.); (2) vermiculation only on
body, no dark streaks on gular region (G. concinnatus and G. riveroi sp. nov.); and (3) head and back without
vermiculation, no dark streaks on gular region (G. li giae ). There are also differences in SVL, with G. ligi ae being
smallest (maximum SVL 40.7 mm), G. riveroi sp. nov. intermediate (47.6 mm), and the three remaining species larger
(52.6 mm in G. concinnatus, 55 mm in G. tapajonicus, 55.7 mm in G. nascimentoi sp. nov.). Moreover, G. l ig ia e and G.
riveroi sp. nov. have 2–3 lateral scales on the distal part of the digits, whereas in the three largest species 3–4 scales
are present. G. tapajonicus and G. nascimentoi sp. nov. differ from each other mainly by their colors and the shape of
the suprahumeral spot – yellow and brown with a suprahumeral ocellus in G. tapajonicus, orange and orange-yellow
with a suprahumeral bar in G. nascimentoi sp. nov.
The main characters pointed out by the discriminant analysis were scales around midbody and ventral scales in a
longitudinal row, which separated G. concinnatus, G. ligiae and G. riveroi sp. nov. from each other, and G. lig ia e and
G. riveroi sp. nov. from G. tapajonicus and G. nascimentoi sp. nov. (Fig. 15A–B). G. concinnatus was more similar in
these characters to G. tapajonicus and G. nascimentoi sp. nov., but they differed in the relative width of the mental
scale and the height of the suprahumeral spot (Fig. 15C–D). Finally, G. tapajonicus and G. nascimentoi sp. nov. could
be distinguished by the height of the suprahumeral spot and by the number of distal lamellae under the second toe
(Fig. 15D–E). Since we found only one meristic character separating these two species, and there is some overlap
between these counts, we compared them with a t-test and the difference is highly significant (t = 5.272; df = 41; p <
0.00).
Even though the shape of suprahumeral spot distinguishes some of these species, we also found that they tend to
vary geographically within the same species. If we compare, for instance, the suprahumeral spot of G. tapajonicus and
G. nascimentoi sp. nov., the former species always has an ocellus, whereas in the latter it is distinctly a bar, even if
only slightly higher than long. However, when we compare their height and length, a few specimens of G. nas ci men to i
sp. nov. have suprahumeral spots similar to those of G. tapajonicus, and each population exhibits different shapes
(Fig. 16). This character, therefore, seems to be very plastic and should be considered with care.
Key to the species of Gonatodes concinnatus complex
1 Dark streaks on gular region and vermiculations on dorsal surface of head absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Dark streaks on gular region and vermiculations on dorsal surface of head present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Back and limbs immaculate; maximum SVL < 42 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonatodes ligiae
- Back and limbs vermiculated; maximum SVL > 47 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Dorsum with brown and beige small vermiculations; 2–3 lateral scales on the distal part of the digits; 96–123 scales around
midbody; 44–55 ventral scales in a longitudinal row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonatodes riveroi
- Dorsum with brown and beige large vermiculations; 3 (occasionally four or two) lateral scales on the distal part of the digits;
109–131 scales around midbody; 49–61 ventral scales in a longitudinal row . . . . . . . . . . . . . . . . . . . . . Gonatodes concinnatus
4 A white suprahumeral bar, bordered by black, from very short to almost reaching the middorsal region present in males and
females (but thinner in the latter); in life, throat orange with orange-yellow oblique stripes. . . . . . . . . . Gonatodes nascimentoi
- A white suprahumeral ocellus, bordered by black, present in males and females (although more conspicuous in males); throat
vivid yellow with reddish-brown oblique stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonatodes tapajonicus
Zootaxa 2869 © 2011 Magnolia Press · 31
GONATODES CONCINNATUS COMPLEX
FIGURE 15. Variation in main characters pointed out by the discriminant analysis in the five species of the G. concinnatus
complex. (A) Scales around midbody (SAM). (B) Ventral scales in a longitudinal row (VLR). (C) Correlation between width
(MSW) and length (MSL) of the mental scale. (D) Correlation of height (HSH) and length (HSL) of the suprahumeral spot. (E)
Number of distal lamellae under the second toe (DL2T). Boxplots (A B, E) indicate maximum, first quartile, median, third
quartile and maximum, and when present outliers (*).
STURARO & AVILA-PIRES
32 · Zootaxa 2869 © 2011 Magnolia Press
FIGURE 16. Comparison of the suprahumeral spot shapes between four populations of G. nascimentoi and the species G.
tapajonicus. HSH = height of suprahumeral spot. HSL = length of suprahumeral spot. Both scales in millimeters.
Discussion
Male color pattern is an important character to distinguish species in Gonatodes, some of which are very similar in
external morphology and scale counts (Vanzolini 1968; Rivero-Blanco 1979; Rodrigues 1990). It is important,
however, to take into consideration that polymorphism in color pattern is known in some species of Gonatodes –
e.g., in Gonatodes annularis (Rivero-Blanco 1968; 1979; Hoogmoed 1973), G. hasemani (Avila-Pires 1995) and G.
alexandermendesi (Schargel et al. 2010). In the case of the species here recognized, the groups initially identified
by color pattern showed also morphological differences, indicating that they constitute different taxa and not only
color morphs. The two most similar taxa are G. tapajonicus and G. nascimentoi, separated mainly by differences in
color, but even in this case they differ in one meristic character, which indicates that they represent independent,
distinct lineages. Following the General Lineage Species Concept (de Queiroz 1998), we interpret them as distinct
species.
Considering the similarities, we assume that the species here studied form a monophyletic group, but this needs
to be tested. G. concinnatus was recovered as the sister species of G. humeralis by Gamble et al. (2008), a
relationship partially corroborated by Schargel et al. (2010), who also included G. antillensis in their analysis,
concluding that G. humeralis was sister to the clade G. concinnatus–G. antillentis (none of the studies included G.
tapajonicus in the analyses). If our assumption of monophyly of the species here studied is correct, the G.
concinnatus complex + G. antillensis would have the same age (estimated as early Miocene by Gamble et al. 2008)
as G. humeralis. It is interesting to observe that the G. concinnatus complex and G. hu mera lis are the most
widespread Gonatodes species in Amazonia. It is possible, therefore, that they have a similar, parallel history, even
though, at least concerning external morphology, divergence within the G. concinnatus complex has been more
accentuated than in G. humeralis.
Zootaxa 2869 © 2011 Magnolia Press · 33
GONATODES CONCINNATUS COMPLEX
Gonatodes nascimentoi sp. nov. seems to be restricted to eastern Amazonia, on both sides of the Amazon
River. Even though no other lizard is known with exactly the same distribution, there are some examples of
lizards that are present on both sides of the lower Amazon, e.g. Arthrosaura kockii and Tretioscincus agilis
(Avila-Pires 1995). Ayres and Clutton-Brock (1992) have shown that for primates the lower Amazon acted less
as a barrier than the middle course of the river. The dynamics of island formation and river channel changes near
the Amazon mouth may have allowed such movements across the lower Amazon.
Gonatodes humeralis occurs in sympatry with many larger species of the genus, e.g. G. annularis, G.
concinnatus, G. hasemani, G. tapajonicus (Avila-Pires 1995; Dixon & Soini 1975; Rivero-Blanco 1979;
Moravec et al. 2001), and it was also found in sympatry with G. nascimentoi. Sympatry between two large
species has been reported only for Venezuela (Schargel et al., 2010). In our fieldwork in the Xingu River,
Gonatodes nascimentoi sp. nov. was found on the right margin of the river, while G. hasemani was found on the
left margin, thus these two species may be parapatric. G. tapajonicus and G. hasemani are both known from the
interfluvium Xingu–Tapajós (G. hasemani also west of the Tapajós), but until now they are not known from the
same or close-by localities. North of the Amazon river G. nascimentoi sp. nov. occurs partially in the same
general area as G. annularis, but again up to the present there is no record of syntopy. The exact distribution of
G. annularis and Gonatodes nascimentoi sp. nov., G. hasemani and G. tapajonicus, whether they occur together
or what separates them, and in case they occur in syntopy how they interact, are at the moment open questions.
Acknowledgments
David Kizirian (AMNH), Jens Vindum (CAS), Linda Trueb (KU), Jucivaldo D. Lima (IEPA), Rick Feeney
(LACM), Jose Rosado and Jonathan Losos (MCZ), José P. Pombal-Jr and Ulisses Caramaschi (MNRJ), Hussam
Zaher (MZUSP), Janalle Caldwell (OMNH), Ronald de Ruite (RMNH), Toby J. Hibbtts and Lee A. Fitzgerald
(TCWC), Greg Schneider (UMMZ), Ron Heyer, Roy McDiarmond, Traci Hartsell, Addison Wynn and Robert
Wilson (USNM), kindly loaned or allowed access to specimens under their care. Amanda Lima, Jerriane
Gomes, Jucivaldo Lima, Marco Antônio Ribeiro Jr. and Ulisses Galatti permitted examination of specimens
under study by them. Jerriane Gomes, Marco Antônio Ribeiro Júnior., Pedro L. V. Peloso and Toby A. Gardner
provided photos and/or data from G. nascimentoi sp. nov., Miguel T. Rodrigues (USP) photos of Gonatodes
tapajonicus, Kraig Adler (Cornell University) photos of Gonatodes riveroi sp. nov., Carlos Rivero-Blanco
photos of Gonatodes ligiae, and Colin McCarthy (BMNH) photos and important information on the syntypes of
Goniodactylus concinnatus and Goniodactylus buckleyi. Pedro Bernardes helped with photos of some color
patterns. Charles J. Cole (AMNH), Marinus Hoogmoed (MPEG), Paulo Passos (MNRJ) and Miguel T.
Rodrigues gave useful suggestions regarding a previous version of the manuscript. Part of the material here
studied was obtained during expeditions supported by the Brazilian Council for the Development of Science/
CNPq (process 473287/04-8), United Kingdom’s Darwin Initiative, Grupo Orsa, Precious Woods Pará, and
FIDESA/Belo Monte project. MJS thanks Amanda Lima, Francílio Rodrigues, Pedro L. Peloso, Mariana
Araguaia and Reginaldo Rocha for their help and companionship during fieldwork. The first author is especially
thankful to Ron W. Heyer and Smithsonian Institution for a Short Term Visit Award, and to all the USNM
Amphibian and Reptile division staff for their support and friendship during his stay at the division. MJS was
supported by a scholarship from CNPq.
Literature cited
Avila-Pires, T.C.S. (1995) Lizards of Brazilian Amazonia (Reptilia: Squamata). Zoologische Verhandelingen, 299, 1–706.
Ayres, J.M. & Clutton-Brock, T.H. (1992) River boundaries and species range size in Amazonian primates. The American
Naturalist, 140, 531–567.
Bartlett, R.D. & Bartlett, P. (2003) Reptiles and Amphibians of the Amazon: An Ecotourist’s Guide. University Press of
Florida, Gainesville, 291 pp.
Barrio-Amorós, C.L. & Brewer-Carías, C. (2008) Herpetological results of the 2002 expedition to Sarisariñama, a tepui in
Venezuelan Guayana, with the description of five new species. Zootaxa, 1942, 1–68.
STURARO & AVILA-PIRES
34 · Zootaxa 2869 © 2011 Magnolia Press
Boulenger, G.A. (1885) Catalogue of the Lizards in the British Museum (Natural History.) Vol. I. Geckonidae,
Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. Trustees of the British Museum, London, xii + 436 pp., Pls. 1–
32.
Boulenger, G.A. (1887) On a new geckoid lizard from British Guiana. Proceedings of the Zoological Society of London,
1887, 153–154.
Burt, C.E. & Burt, M.D. (1933) A preliminary check list of the lizards of South America. Transactions of the Academy of
Science of St. Louis, 28, 1–104.
Cole, C.J. & Kok, P.J.R. (2006) A new species of gekkonid lizard (Sphaerodactylinae: Gonatodes) from Guyana, South
American. American Museum Novitates, 3524, 1–13.
de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: a conceptual
unification and terminological recommendations. In: Howard, J.D. & Berlocher, S.J. (Eds.), Endless Forms, Species
and Speciation. Oxford University Press, New York. pp. 57–75.
Dixon, J.R. & Soini, P. (1975) The reptiles of the upper Amazon Basin, Iquitos Region, Peru: I. Lizards and amphisbaenians.
Milwaukee Public Museum, Contributions in Biology and Geology, 4, 1–58.
Dixon, J.R. & Soini, P. (1986) The reptiles of the upper Amazon Basin, Iquitos Region, Peru. Milwaukee Public Museum,
Milwaukee, 154 pp.
Donoso-Barros, R. (1966) Dos nuevos Gonatodes de Venezuela. Publicación Ocasional del Museo Nacional de Historia
Natural, Santiago de Chile, 11, 1–32.
Donoso-Barros, R. (1967) Diagnosis de dos nuevas especies del género Gonatodes de Venezuela. Noticiario Mensual Del
Museo Nacional de Historia Natural de Santiago, 129, no pagination.
Donoso-Barros, R. (1968) The lizards of Venezuela (Check List and Key). Caribbean Journal of Science, 8, 105–122.
Duellman, W.E. (1978) The biology of an Equatorial Herpetofauna in Amazonian Ecuador. Miscellaneous Publications,
Museum of Natural History, University of Kansas, 65, 1–352.
Duellman, W.E. & Mendelson III, J.R. (1995) Amphibians and reptiles from northern Departamento Loreto, Peru: taxonomy
and biogeography. The University of Kansas Science Bulletin, 55, 329–376.
Duméril, A.M.C. & Bibron, G. (1836) Erpetologie Générale ou Histoire Naturelle Complète des Reptiles. Vol.3. Librairie
Encyclopédique Roret, Paris, 517 pp.
Esqueda, L.F. (2004a) Una nueva especie de Gonatodes (Squamata: Gekkonidae) proveniente del piedemonte cisandino de
Venezuela. Herpetotropicos, 1, 32–39.
Esqueda, L.F. (2004b) Apéndice II. Registros Complementarios de la Sierra de Perijá, el Macizo de Tamá y de bajas
elevaciones. In: La Marca, E. & Soriano, P, Reptiles de Los Andes de Venezuela. Fundación Polar, Conservación
Internacional, CODEPRE-ULA, Fundacite Mérida, BIOGEOS, Mérida, Venezuela, pp. 161–165.
Gamble, T.; Simons, A. M.; Colli, G.R. & Vitt, L.J. (2008) Tertiary climate change and the diversification of the Amazonian
gecko genus Gonatodes (Sphaerodactylidae, Squamata). Molecular Phylogenetics and Evolution, 46, 269–277.
Gray, J.E. (1831) A synopsis of the species of the class Reptilia. In: Griffith, E. & Pidgeon, E. (Eds.) The Class Reptilia
Arranged by the Baron Cuvier, with Specific Descriptions. Whittaker, Treacher, and Co., London, pp. 1–110.
Griffin, L.E. (1917) A list of the South American lizards of the Carnegie Musuem, with descriptions of four new species.
Annals of the Carnegie Museum, 11, 304–320, Pls. 32–35.
Guichenot, A. (1855) Reptiles. In: Castelnau F. (Ed.), Animaux Nouveaux ou Rares Recueillis pendant l'Expédition dans les
Parties Centrales de l'Amérique du Sud, de Rio de Janeiro à Lima, et de Lima au Pará; Exécutée par Ordre du
Gouvernement Français pendant les Années 1843 a 1847, sous la Direction du Comte Francis de Castelnau. Reptiles.
Tome second. P. Bertrand, Paris, pp. 1–95, Pls 1–18.
Proceedings of the Zoological Society of London, 1859, 402–420, Pl. 20.
Hoogmoed, M.S. (1973) Notes on the herpetofauna of Surinam IV: The lizards and amphisbaenians of Surinam.
Biogeographica, 4, 1–419.
Krysko, K.L. & Daniels, K.J. (2005) A key to the geckos (Sauria:Gekkonidae) of Florida. Caribbean Journal of Science, 41,
28–36.
Kluge, A.G. (1995) Cladistic relationships of sphaerodactyl lizards. American Museum Novitates, 3199, 1–23.
Lichtenstein, H. (1856) Nomenclator Reptilium et Amphibiorum Musei Zoologici Berolinensis. Namenverzeichniss der in
der zoologischen Sammlung der Königlichen Universität zu Berlin aufgestellten Arten von Reptilien und Amphibien
nach ihren Ordnungen, Familien und Gattungen. Königlichen Akademie der Wissenschaften, Berlin, 48 pp.
Lidth de Jeude, T.W. van (1887) On a collection of reptiles and fishes from the West Indies. Notes from the Leyden Museum,
9, 129–139. Pls. 2.
McGarigal, K., Cushman, S. & Stafford, S.G. (2000) Multivariate Statistics for Wildlife and Ecology Research. Springer-
Verlag, New York, 283 pp.
Mechler, B. (1968) Les geckonidés de la Colombie. Revue Suisse de Zoologie 75, 305–371.
Moravec, J., Tuanama, I.A. & Burgos, A.M. (2001) Reptiles recently recorded from the surroundings of Iquitos
(Departamento Loreto, Peru). asopis Národního muzea, ada p írodov dná, 170, 47–68.
Nascimento, F.P., Avila-Pires, T.C.S. & Cunha, O.R. (1987) Os répteis da área de Carajás, Pará, Brasil (Squamata). II.
Boletim do Museu Paraense Emílio Goeldi, Série Zoologia, 3, 33–65.
Zootaxa 2869 © 2011 Magnolia Press · 35
GONATODES CONCINNATUS COMPLEX
Noble, G.K. (1921) Some new lizards from northwestern Peru. Annals of the New York Academy of Sciences, 29, 133–139.
O'Shaughnessy, A.W.E. (1881) An account of the Collection of Lizards made by Mr. Buckley in Ecuador, and now in the
British Museum, with descriptions of the new species. Proceedings of the Zoological Society of London, 1881, 227–
245, Pls. 22–25.
Paynter Jr., R.A. (1997) Ornithological gazetteer of Colombia. Museum of Comparative Zoology, Harvard University,
Cambridge, Massachusetts, 537 pp.
Peters, J.A. & Donoso-Barros, R. (1970) Catalogue of the Neotropical Squamata: part II. Lizards and amphisbaenians.
United States National Museum Bulletin, 297, 1–293.
Powell, R. & Henderson, R.W. (2005) A new species of Gonatodes (Squamata: Gekkonidae) from the West Indies.
Caribbean Journal of Science, 41, 709–715.
Reis, S.F.; Pessôa, L.M. & Strauss, R. (1990) Applicaion of size-free canonical discriminant analysis to studies of
geographic differentiation. Revista Brasileira de Genética, 13, 509–520.
Rivas, G.A. & Schargel, W.E. (2008) Gecko on the rocks: an enigmatic new species of Gonatodes (Sphaerodactylidae) from
Inselbergs of the Venezuelan Guayana. Zootaxa, 1925, 39–50.
Rivero-Blanco, C. (1968) Un genero y dos especies de tuqueques (Sauria: Sphaerodactylinae) citados por primera vez para
Venezuela, con notas sobre la distribucion de otras especies poco conocidas. Memoria de La Sociedad de Ciencias
Naturales la Salle, 27, 103–119.
Rivero-Blanco, C. (1979) The Neotropical lizard genus Gonatodes Fitzinger (Sauria: Sphaerodactylinae). PhD Thesis,
Texas A&M University. 233 pp.
Rivero-Blanco, C. & Barrio-Amorós, C.L. (2002) Geographic Distribution. Sauria: Gonatodes concinnatus ligiae.
Herpetological Review 33, 67.
Rodrigues, M.T. (1980) Descrição de uma nova espécie de Gonatodes da Amazonia (Sauria, Gekkonidae). Papéis Avulsos
de Zoologia, 33, 309–314.
Rojas-Runjaic, F.J.M.; Infante-Rivero, E.E.; Cabello, Velozo, P. (2010) A new non-sexually dichromatic species of the
genus Gonatodes (Sauria: Sphaerodactylidae) from Sierra de Perijá, Venezuela. Zootaxa, 2671, 1–16.
Sanchez, H.; Castaño, O. & Cardenas, G. (1995) Diversidad de los reptiles en Colombia. In: Rangel-Ch, J.O. (Ed.),
Colombia Diversidad biotica I. Guadalupe, Santafé de Bogota, Colombia, pp. 277–325.
Schargel, W.E.; Rivas, G.A.; Makowsky, R.; Señaris, J.C.; Natera, M.A.; Barros, T.R. & Barrio-Amorós, C.L. (2010)
Phylogenetic systematics of the genus Gonatodes (Squamata: Sphaerodactylidae) in the Guayana region, with
description of a new species from Venezuela. Systematics and Biodiversity, 8, 321–339.
Shreve, B. (1947) On Venezuelan reptiles and amphibians collected by Dr. H. G. Kugler. Bulletin of the Museum of
Comparative Zoology, 99, 519–537.
Smithe, F.B. (1975) Naturalist’s Color Guide. The American Museum of Natural History, New York. 8 pp.
Strauss, R.E. (1985) Evolutionary allometry and variation in body form in the South American catfish genus Corydoras
(Callichthyidae). Systematic Zoology, 34, 381–396.
Tabachnick, B.G. & Fidell, L.S. (2001) Using Multivariate Statistics. Allyn & Bacon, Boston, Massachusetts, 966 pp.
Vanzonili, P.E. (1955) Sôbre Gonatodes varius (Auguste Duméril) com notas sôbre outras espécies do gênero (Sauria,
Gekkonidae). Papéis Avulsos de Zoologia, 12, 119–132.
Vanzolini, P.E. (1968) Lagartos brasileiros da família Gekkonidae (Sauria). Arquivos de Zoologia, 17, 1–84.
Vanzolini, P.E. (1986) Addenda and corrigenda to the Catalogue of Neotropical Squamata. Smithsonian Herpetological
Information Service, 70, 1–25.
Vitt, L.J. & de la Torre, S. 1996. Guia para la Investigacion de las Lagartijas de Cuyabeno - A Research Guide to the
Lizards of Cuyabeno. Museo de Zoologia (QCAZ), Centro de Biodiversidad y Ambiente, Pontificia Universidad
Católica del Ecuador, Monografia 1, Quito, 165p.
Wermuth, H. (1965) Liste der rezenten Amphibien und Reptilien: Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich,
89, i–xxii, 1–246.
STURARO & AVILA-PIRES
36 · Zootaxa 2869 © 2011 Magnolia Press
APPENDIX I. Additional material examined. Total number of specimens per species indicated in parentheses.
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,
Willamstad (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 (40): ECUADOR: 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); Manabi: 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); Pichincha: Santo Domingo de los
Colorados, 47 km S of, Centro Cientifico Rio Palenque (USNM 285426–36)
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 eladioi (1): BRAZIL: Pará: Marabá, Salobo, Serra dos Carajás (MPEG 18020).
Gonatodes hasemani (22): BRAZIL: Pará: Juruá, Rio Xingu (MZUSP 67240–57); Rondônia: Porto Velho, Área de
inundação 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 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 vittatus (2): TRINIDAD: St. Patrick, Chathan Beach, on Erin Bay (USNM 314695–96).
