A new species of Tropidurus (Squamata, Tropiduridae) of the semitaeniatus group from a semiarid area in Northeastern Brazil

Abstract

A new species of Tropidurus of the semitaeniatus group is described from northeastern Brazil. Tropidurus sp. nov. is morphologically similar to other species of the group in the marked dorsoventral flattening of the body, and the presence of a series of sublabial shields that are larger than the adjacent scales. It is distinguished by a single middorsal longitudinal light stripe that extends from the snout to the scapular region. The new species appears to have the largest body size of the group. The new species inhabits the Vale do Jaguaribe, in the eastern part of the state of Ceará, where there are many fis- sured rocky outcrops surrounded by hypoxerophytic caatinga vegetation.

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60 Accepted by S. Carranza: 30 May 2011; published: 27 Jun. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
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Copyright © 2011 · Magnolia Press
Zootaxa 2930: 60–68 (2011)
www.mapress.com/zootaxa/Article
A new species of Tropidurus (Squamata, Tropiduridae) of the semitaeniatus group
from a semiarid area in Northeastern Brazil
DANIEL CUNHA PASSOS1,4, DANIEL CASSIANO LIMA1,2,3 & DIVA MARIA BORGES-NOJOSA1
1Núcleo Regional de Ofiologia da Universidade Federal do Ceará – NUROF-UFC. Campus do Pici, Centro de Ciências, Bloco 905,
60.455-760, Fortaleza, Ceará, Brazil
2Programa de Pós-Graduação em Biodiversidade Animal, Universidade Federal de Santa Maria. Centro de Ciências Naturais e Exa-
tas, Prédio 17, sala 1140-D. Camobi, Km 9, 97105-900. Santa Maria, Rio Grande do Sul, Brazil
3Universidade Estadual do Ceará, Faculdade de Educação de Itapipoca. Av Mons. Tabosa s/n, 62.500-000, Itapipoca, Ceará, Brazil
4Corresponding author. E-mail: biologodanielpassos@gmail.com
Abstract
A new species of Tropidurus of the semitaeniatus group is described from northeastern Brazil. Tropidurus sp. nov. is mor-
phologically similar to other species of the group in the marked dorsoventral flattening of the body, and the presence of a
series of sublabial shields that are larger than the adjacent scales. It is distinguished by a single middorsal longitudinal
light stripe that extends from the snout to the scapular region. The new species appears to have the largest body size of the
group. The new species inhabits the Vale do Jaguaribe, in the eastern part of the state of Ceará, where there are many fis-
sured rocky outcrops surrounded by hypoxerophytic caatinga vegetation.
Key words: Tropidurus jaguaribanus sp. nov., Lizard, Caatinga
Resumo
Uma nova espécie de Tropidurus do grupo semitaeniatus é descrita do Nordeste do Brasil. Tropidurus sp. nov. é morfo-
logicamente semelhante às espécies do grupo semitaeniatus quanto ao evidente achatamento dorsoventral do corpo e pela
presença de uma série de escudos sublabiais aumentados em relação às escamas adjacentes. Entretanto, pode ser diferen-
ciada por apresentar no meio do dorso uma única faixa clara longitudinal que se prolonga do rostro até a região escapular.
Dentre as espécies do grupo, esta parece apresentar o maior porte corpóreo. A nova espécie habita localidades do Vale do
Jaguaribe, região leste do Estado do Ceará, onde predominam afloramentos rochosos graníticos fraturados, circundados
por caatinga hipoxerófita.
Palavras chave: Tropidurus jaguaribanus sp. nov., Lagartos, Caatinga
Introduction
Brazil has one of the most diverse assemblages of reptiles on the planet, with about 721 species including at least
241 lizards (Bérnils 2010). This great richness, one of the largest on Earth, results not only from the great extent of
the country, but also the diversity of ecosystems and the historical events of climatic and geographical changes dur-
ing the Pleistocene in South America (Rocha 1994).
Tropiduridae is one of families which contains the largest number of known species among the neotropical liz-
ards (Torres-Carvajal 2004). In recent decades, several phylogenetic studies have examined the relationships
among these species (Harvey & Gutberlet 2000; Frost et al. 2001), although some groups remain poorly studied. In
Brazil, there are 35 species of tropidurids, 17 of which belong to the genus Tropidurus (Bérnils 2010).
The genus Tropidurus Wied, 1820 consists of lizards whose distribution includes the South American mainland
east and west of the Andes, and the Galapagos Islands, occurring in open landscapes such as Caatinga, Cerrado,

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NEW SPECIES OF TROPIDURUS FROM NORTHEASTERN BRAZIL
and Chaco (Rodrigues 1987). This genus is widely diversified morphologically and ecologically. Its members
range from the small arenicolous T. hygomi, 15 cm in total length, to the generalist T. hispidus that can reach 32 cm
in length. Currently, the genus is subdivided into four species groups: T. spinulosus, T. torquatus, T. bogerti, and T.
semitaeniatus (Frost et al. 2001).
The semitaeniatus group contains three species: T. semitaeniatus (Spix 1825), T. pinima (Rodrigues 1984), and
T. helenae (Manzani & Abe 1990). They are saxicolous, inhabiting rocky outcrops, and occur exclusively in the
Brazilian Caatinga (Tropical Thorny Deciduous Savanna). Their characteristic marked dorsoventral flattening of
the body allows them to efficiently exploit crevices in rocks, where they adroitly find refuge from predators (Van-
zolini et al. 1980).
Tropidurus semitaeniatus is widely distributed in the Caatinga, and shows clear geographical variations (Rodri-
gues 2003). Studies in recent decades have contributed to understanding its ecology, especially on feeding and ther-
moregulatory behaviour (Vitt 1995; Ribeiro et al. 2008; Kolodiuk et al. 2009; Kolodiuk et al. 2010; Ribeiro &
Freire 2010; Ribeiro et al. 2010). Subsequent to the publication of a study on the reproductive biology of T. semi-
taeniatus by Vitt & Goldberg (1983), Rodrigues (1984) described T. pinima, warning that under the name T. semi-
taeniatus there could be many species still to be described. Although phylogenetic studies have been performed
with several species of the genus Tropidurus (Harvey & Gutberlet 2000; Frost et al. 2001), Rodrigues (2003)
emphasized the importance of further studies involving the systematics of the semitaeniatus group, arguing the
possibility of elucidation of the history of contacts between several areas of rocky outcrops in northeastern Brazil,
as well as the necessity for description of already-recognized new species.
Here we present the description of a new species of Tropidurus of the semitaeniatus group, from rocky out-
crops of the Jaguaribe Valley, a Caatinga region in northeastern Brazil.
Material and methods
The lizards were collected at the Fazenda Veneza (5º19’21’’ S and 38º11´58’’ W; Datum WGS 84), located in the
municipality of São João do Jaguaribe, eastern State of Ceará (Jaguaribe Valley) in northeastern Brazil, from
March 2008 through February 2009. The location is part of the Sertão, a region with semi-arid climate and vegeta-
tion, where trees reach about 12 m tall with a shallow, multi-branched canopy. The predominant soils feature a
basement with abundant displays of granite outcrops, locally called “lajeiros”. The lajeiros are as high as 2.0 m and
their many crevices serve as a hiding place for the lizards (Figure 1). This environment is ideal for saxicolous liz-
ards such as those in the semitaeniatus group, and the new species is abundant in the area.
The meristic and morphometric data used in the description were obtained from the right side of the individuals
examined. The variations of paratypes are presented in the format: range (mean ± one standard deviation), and are
given in proportion to snout-vent length (Table 1). The morphometric measurements were made with a digital cali-
per of 0.01 mm precision. The terminology used in describing the characters is based on taxonomic and systematic
literature for the genus Tropidurus (Rodrigues 1984; Rodrigues 1987; Manzani & Abe 1990; Frost et al. 2001).
The type series is deposited in the following scientific collections: Coleção Herpetológica da Universidade Federal
do Ceará (CHUFC), Museu de Zoologia da Universidade de São Paulo (MZUSP), Coleção Herpetológica da Uni-
versidade Federal de Santa Maria (ZUFSM), and Museu Paraense Emilio Goeldi (MPEG). Specimens of T. semi-
taeniatus, T. pinima, and T. helenae used for comparisons are listed in Appendix 1.
Results
Tropidurus jaguaribanus sp. nov.
Holotype. Adult male (CHUFC L 3860) collected in São João do Jaguaribe Municipality (5º19’21’’ S and
38º11´58’’ W), Ceará State, northeastern Brazil, 21 February 2009, by D. C. Lima and D. M. Borges-Nojosa (Fig-
ure 2).
Paratypes. All from the type-locality, collected from March 2008 through February 2009 by the same collec-
tors. Adult males (CHUFC L 3650, CHUFC L 3858), adult females (CHUFC L 3857, CHUFC L 3859, CHUFC L
3918), and unsexed juveniles (CHUFC L 3658, MPEG 28900, MZUSP 100739, ZUFSM L 0728).

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FIGURE 1. View of the type locality of Tropidurus jaguaribanus sp. nov.. Note the fissures in rocky outcrops.
Diagnosis. The color pattern of T. jaguaribanus sp. nov. allows it to be easily distinguished from the other spe-
cies of the semitaeniatus group (Figures 3 e 4). Tropidurus jaguaribanus sp. nov. differs from T. helenae and T.
pinima in having only one middorsal, longitudinal light stripe rather than three. This single stripe, which extends
dorsally from the snout to the scapular region, also allows it to be distinguished from T. semitaeniatus, because in
this latter, the single stripe always extends from the snout to the base of the tail. The stripe of T. jaguaribanus sp.
nov. may sometimes be absent or not evident in adult individuals, which also distinguishes it from the other species
of the group. Tropidurus jaguaribanus sp. nov. also has 5–6 supralabials, rather than 7–8 as in T. pinima and T.
semitaeniatus. Tropidurus jaguaribanus sp. nov. has a single subocular, in contact with the first canthal, distin-
guishing it from T. pinima, which has one scale between the subocular and the first canthal. Finally, the new species
differs from all the others by the prominently spined and keeled lateral scales on the trunk (Figure 5), which are vis-
ible to the naked eye in adult individuals and are inconspicuous in the other species and by the tarsal scales with
higher keels than in the other species of the group.
Description of the holotype. Measurements. Snout-vent length (SVL) 103.41 mm; tail length 138.18 mm;
body height at midline 12.37 mm; distance from snout to anterior margin of tympanum 24.12 mm; head height
10.14 mm; head width 21.47 mm; distance between limbs, from axillar region to inguinal region 43.73 mm; tail
base width 16.54 mm; femur length 25.10 mm; fourth finger length 10.57 mm; fourth toe length 16.38 mm.
Cephalic scales (Figure 2). Rostral scale smooth and broad. Nostril rounded, exposed laterally and inserted
into single nasal scale. Eight supraciliars elongated, narrow, oblique, and imbricate, forming small longitudinal
ridge. Supraocular scales smooth, flat, asymmetrical, with different dimensions, forming five rows on each side,
bounded by supraorbital semicircle. Two rows of small irregular supraoculars near eye, another two intermediate
rows of median scales, and one row of seven large supraocular scales, wider than long, transversely elongated.
Temporal scales small, irregular, decreasing in size posteriorly. Ear scales reduced, deep ear canal, and preauricular

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fringe with seven pyramidal scales. Occipital scale conspicuous, as wide as long, with irregular margins. Single
subocular, broad and long, in close contact with supralabials in posterior region. Six long supralabials, counting to
posterior margin of subocular. Seven long infralabials, counting to corresponding position of last supralabial. Men-
tal smooth, broad, and triangular, with posterior apex. A row of sublabial shields present between post-mentals and
infralabials, increasing in size posteriorly in relation to these. Small gulars, irregular, laterally imbricate, decreasing
in size posteriorly.
FIGURE 2. Tropidurus jaguaribanus sp. nov. (CHUFC L 3860, holotype). (A) lateral, (B) dorsal and (C) ventral views of the
head.
Trunk scales. Dorsal scales small, smooth, irregular, juxtaposed, almost indistinct, extending from temporal
scales to dorsal tail. Dorsal scales 139 in number, counting on dorsal midline from occipital scale to posterior mar-
gin of insertion of thigh. Lateral scales acicular, arranged in transverse rows, taller than wide, acutely keeled, per-
pendicular to trunk, revealing spiny aspect of flanks. Scales around middle of body (strap) 139 in number, counting
around middle of body between axilla and groin. Ventral scales smooth, juxtaposed, square and rectangular,
extending from gulars to vent. Ventral scales 68 in number, counting on ventral midline from level of anterior root
of arm to anterior insertion of thigh.
Limb scales. Arm scales large, imbricate and keeled, with scales in anterior region more strongly keeled than in
posterior region. Forearm scales large, imbricate and keeled, but with scales in posterior region more strongly
keeled than in anterior region. Carpus scales imbricate and slightly keeled in comparison to other parts of body.

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Femoral scales imbricate anteriorly, and imbricate and keeled posteriorly. Tibial scales large and imbricate, with
higher keels than femoral scales. Tarsum scales imbricate and strongly keeled. Infradigital lamellae keeled, 22
lamellae on fourth finger and 28 lamellae on fourth toe.
Tail scales. Dorsal scales of tail imbricate, slightly keeled. Lateral scales imbricate, strongly keeled. Ventral
scales smooth and imbricate.
Body folds. Three lateral neck folds, irregular and conspicuous, but without forming mite-pockets. One well-
marked ante-humeral fold. Two small axillary folds and four evident inguinal folds, lined with granular scales,
smaller than adjacent scales.
FIGURE 3. Dorsal and ventral views, in life, of male, female, adult and juvenile of Tropidurus jaguaribanus sp. nov. from the
type locality, illustrating the color pattern typical of the species.
FIGURE 4. Schematic illustration of the dorsal color pattern of the semitaeniatus group. (A) Tropidurus jaguaribanus sp. nov.,
(B) Tropidurus semitaeniatus, (C) Tropidurus helenae and (D) Tropidurus pinima.
FIGURE 5. Detail of the lateral scales and the spiny aspect of the flanks of Tropidurus jaguaribanus sp. nov.. Note the pres-
ence of mites.

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Sensory organs. Sensory organs present in cephalic scales, one or more from rostral to temporal scales, includ-
ing dorsal and lateral scales of head. Fingers, toes, and carpus also with sensory organs in their scales.
Color pattern in life (Figure 3). Trunk dark gray with small scattered light dots. One light stripe, longitudinal,
rostro-scapular, indistinct. Transverse trunk stripes, asymmetrical, extending from middorsal region to flanks, 9 on
each side. These transverse stripes have dark regions, immediately succeeded by irregular series of light spots.
Limbs with coloration similar to trunk. Ventral surface light-colored. Ventral surface of thighs and periphery of
vent with well-defined black pigmentation. Cloacal margin intense yellow.
Va ri at io n . The paratypes (N=9) show meristic characteristics similar to the holotype, with few variations (see
Table 1).
TABLE 1. Meristic and morphometric data of the type series of Tropidurus jaguaribanus sp. nov.. The variations of paratypes
are presented in the format: range (mean ± one standard deviation), and morphometric data are given in proportion to snout-
vent length (Table 1).
After the specimens have been placed in preservative fluid, the longitudinal stripes become lighter and the
tracks across the trunk acquire a staining pattern similar to the trunk.
Etymology. The specific epithet derives from the region of the state of Ceará where the species occurs, the
Jaguaribe Valley.
Distribution and natural history. Tropidurus jaguaribanus sp. nov. also occurs in other municipalities in the
Jaguaribe Valley in the state of Ceará. Another population was found on the Fazenda Amendoim (5º18’09’’ S and
38º54’05’’ W), Banabuiu Municipality, about 78 km east of the type locality in an area with many rocky outcrops
and climate conditions similar to the type locality.
CHARACTERS HOLOTYPE PARATYPES (N=9)
MERISTIC CHARACTERS
Supraciliary scales 8 7–9 (7.78 ± 0.67)
Widest supraorbital scales 7 7–8 (7.22 ± 0.44)
Preauricular fringe of scales 7 5–7 (6.44 ± 0.73)
Supra-labials scales 6 5–6 (5.56 ± 0.53)
Infra-labials scales 7 5–6 (5.89 ± 0.33)
Dorsal scales 139 129–149 (137.67 ± 6.69)
Strap scales 139 121–150 (139.11 ± 10.15)
Ventral scales 68 65–73 (69.11 ± 3.26)
Axillary folds 2 2–3 (2.44 ± 0.53)
Inguinal folds 4 2–4 (2.56 ± 0.73)
Lamellae in 4th finger 22 21–23 (21.78 ± 0.83)
Lamellae in 4th toe 28 24–28 (26.33 ± 1.12)
MORPHOMETRIC CHARACTERS (mm)
Snout-vent length 103.41 59.99–98.12 (75.58 ± 12.85)
Tail length/SVL 138.18 0.69–1.63 (1.28 ± 0.29)
Body height/SVL 12.37 0.10–0.15 (0.14 ± 0.02)
Snout-tympanum length/SVL 24.12 0.22–0.25 (0.24 ± 0.01)
Head height/SVL 10.14 0.09–0.11 (0.10 ± 0.01)
Head width/SVL 21.47 0.19–0.21 (0.20 ± 0.01)
Distance between limbs/SVL 43.73 0.41–0.50 (0.45 ± 0.03)
Tail base width/SVL 16.54 0.14–0.16 (0.15 ± 0.01)
Femur length/SVL 25.10 0.21–0.25 (0.23 ± 0.01)
Fourth finger length/SVL 10.57 0.09–0.14 (0.12 ± 0.02)
Fourth toe length/SVL 16.38 0.15–0.20 (0.17 ± 0.01)

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Tropidurus jaguaribanus sp. nov. is diurnal, and is apparently most abundant on rocky outcrops in these local-
ities. The new species shares this environment, and even the rock crevices with the similarly diurnal T. hispidus
(Spix, 1825) and with the phyllodactylids Phyllopezus periosus Rodrigues, 1986, P. pollicaris (Spix, 1825), and
Gymnodactylus geckoides Spix, 1825.
FIGURE 6. Specimen of Tropidurus jaguaribanus sp. nov., in life, illustrating the marked dorsoventral flattening of the body.
Discussion
The distribution of the semitaeniatus group coincides with nearly the entire area of the Caatinga morphoclimate
domain in northeastern Brazil, occurring in transitional regions with other biomes, and also particularly in open
areas above 1,000 m altitude (Rodrigues 1984). Vanzolini (1976) also reported the occurrence of these animals in
areas with vegetation typical of the Cerrado. Tropidurus helenae seems to be endemic to the Serra da Capivara in
southern Piauí state, while T. pinima occurs in the Serra do Assuruá on the right bank of the São Francisco River,
and in the Chapada Diamantina, in Rio das Contas Municipality, both in the state of Bahia.
Although these species occur in various vegetation types, their preferred microhabitat is basically groups of
rocks, their surfaces and crevices (Vanzolini et al. 1980). The exclusive occurrence in saxicolous environments can
be explained by their efficient exploitation of the environment, especially associated with specific morphological
adaptations such as the prominent dorsoventral body flattening (Figure 6) and strongly keeled tarsal scales, which
may impede capture by predators (Vanzolini et al. 1980; Vitt 1981; Rodrigues 1984).

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The dorsoventral flattening of the body and the presence of a row of sublabial shields that are larger than the
adjacent scales are common to the entire semitaeniatus group (Rodrigues 1984). Although the differences in folido-
sis among the four species in general are not diagnostic, the color pattern, in particular, allows accurate identifica-
tion of species.
The ecology of the semitaeniatus group is still poorly understood. Although some ecological aspects of T.
semitaeniatus have been studied in recent decades (e.g., Kolodiuk et al. 2010; Ribeiro & Freire 2010), studies
involving the other species of the group are scarce. Among the different species, T. jaguaribanus sp. nov. appears
to have the largest body, suggesting possible differences in ecological exploitation of the environment.
Since the different species of the semitaeniatus group are endemic in the Caatinga domain, this underscores the
need not only to clarify their taxonomy and systematics, but also to understand the ecological strategies of each
species and the evolutionary processes occurring in this group.
Key to the species of Tropidurus of the semitaeniatus group from northeastern Brazil
1 Single middorsal light stripe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Three dorsal light stripes, one middorsal, and two dorsalateral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
2 Middorsal light stripe extending from snout to the tail base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .T. semitaeniatus
- Middorsal light stripe extending from snout to the scapular region, may be absent or not evident in adults individuals . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. jaguaribanus
3 Middorsal light stripe extending from snout to the tail base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. pinima
- Middorsal light stripe extending from snout to the scapular region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .T. helenae
Acknowledgments
We thank the Museu de Zoologia da Universidade de São Paulo (MZUSP) for provide specimens of Tropidurus
pinima and T. helenae for comparative morphologic analysis, and two anonymous reviewers for suggestions and
comments; DCL thanks the Fundação Cearense de Apoio ao Desenvolvimento Científico e Tecnológico (FUN-
CAP) for doctorate fellowship (proc. no BDS-0017-00003.01.18/10); ICMBio for grant the collecting permits (per-
manent license 10893-1, reg. 472138) that supported this search; and L. G. Sales-Junior for logistic support in the
field works.
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Vitt, L.J. & Goldberg, S.R. (1983) Reproductive ecology of two tropical iguanid lizards: Tropidurus torquatus and Platynotus
semitaeniatus. Copeia, 1, 131–141.
APPENDIX 1. Specimens examined.
The list of examined tropidurids in comparative morphologic analysis follows below.
Tropidurus helenae: MZUSP 75053.
Tropidurus pinima: MZUSP 93517, MZUSP 93521, MZUSP 93523, MZUSP 93540, MZUSP 93545, MZUSP 93548, MZUSP
93549, MZUSP 93552, MZUSP 93557, MZUSP 93565.
Tropidurus semitaeniatus: CHUFC L 3737, CHUFC L 3767, CHUFC L3824, CHUFC L 3875, CHUFC L 3876, CHUFC L
1526, CHUFC L 1529, CHUFC L 1530, CHUFC L 1531, CHUFC L 1534.