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Colobosaura modesta (REINHARDT & LÜTKEN, 1862)

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Higher TaxaGymnophthalmidae (Gymnophthalminae), Iphisini, Sauria, Gymnophthalmoidea, Squamata (lizards)
Subspecies 
Common NamesE: Bahia Colobosaura
Portuguese: Briba, Calango-cobra, Calanguinho, Lagartinho, Lagartinho-do-Folhiço, Lagarto, Lagarto-do-Folhiço 
SynonymPerodactylus modestus REINHARDT & LÜTKEN 1862: 218
Colobosaura modesta — BOULENGER 1887: 508
Colobosaura modesta — PETERS et al. 1970: 99
Colobosaura landii CUNHA 1977: 3
Colobosaura modesta— DO NASCIMENTO et al. 1987
Colobosaura modesta — WHITING et al. 2003
Colobosaura modesta — CACCIALI et al. 2016 
DistributionBrazil (Minas Gerais, Bahia, Sao Paulo, Goias, Mato Grosso do Sul, Mato Grosso, Maranhao, Para, Distrito Federal, Ceará, Piauí), Paraguay

Type locality: Morro da Garcza, north of Curvelo, Minas Gerais.  
Reproductionoviparous 
TypesHolotype: ZMUC (M. Andersen, pers. comm., 11 Aug 2011) 
DiagnosisDiagnosis. As generic diagnosis, and in addition the following features: three pairs of chinshields, of which anterior two very large, third pair distinctly smaller and short; 5-8 pairs of nuchals. Dorsal surface of head and back light brown, back with small, irregular, dark brown spots; flanks dark brown, well demarcated from back. N o light vertebral band (Avila-Pires 1995: 370 ff).

Description. Gymnophthalmid with maximum SVL in males of 55 mm (MPEG 9795), in females of 49 mm (MPEG 13992). Head 0.17-0.24 times SVL, relatively larger in juveniles, and in adult males (0.20-0.22) in relation to adult females (0.17-0.19); 1.4-1.7 (1.54 ± 0.08, n= 19) times as long as wide, 1.2-1.6 (1.31 ± 0.10, n= 16) times as wide as high. Neck only slightly narrower than, or as wide as, head and body. Body cylindrical to slightly depressed. Limbs short, forelimbs 0.16-0.25 times SVL, relatively shorter in larger specimens, hind limbs 0.28-0.38 (0.34 ± 0.03, n= 18) times (no allometric growth observed). Tail round incross section, tapering toward tip, 1.6 times SVL in the smallest specimen studied (SVL= 22mm), 1.8-2.4 (2.13 ± 0.23, n= 9) times SVL in the remaining ones (SVL between 27 mm and 55 mm, variation not correlated with size).
Tongue lanceolate, covered with imbricate, scale-like papillae, with a smooth, bifid tip. Anterior teeth conical, posterior teeth bicuspid or tricuspid.
Rostral trapezoidal in dorsal view, about two and a half times as wide as high, in broad contact with frontonasal. Frontonasal irregularly pentagonal or heptagonal, in contact with rostral, nasals, loreals (shortly) and prefrontals. Prefrontals form a short medial border (just in touch in MPEG 8513). Each prefrontal in contact with frontonasal, loreal, first and second supraoculars, and frontal. Frontal hexagonal, slightly longer than wide, incontact with prefrontals, second supraoculars, and frontoparietals. A pair of frontoparietals which form a medial suture longer than that between prefrontals. Each one in contact also with frontal, second (in most cases) and third supraoculars, parietal, and interparietal. Interparietal more than twice as long as wide, with sides approximately parallel and posterior margin angulate, truncate or roundish. Parietals wider and slightly shorter than interparietal. Three supraoculars, first smallest, second about as large as, or slightly larger than third. Two elongate supraciliaries, followed by a short scale which in some specimens is in a supraciliary position, in other specimens in a postocular position. First supraciliary widens anteriorly but does not expand dorsally. Nasal undivided, nostril in its inferior margin, medially. One roughly rectangular loreal, in contact with nasal, frontonasal, prefrontal, first supraocular, first supraciliary, preocular, frenocular, and first and second supralabials. Frenocular mostly present, relatively small, in some specimens fused with first subocular (holotype, ZMUC 4377, with frenocular on one side divided into one anterior and one posterior scale). One preocular, 2-4 narrow suboculars, and a wider postsubocular. One or two anterior, and one large, posterior, postoculars present. Large postocular dorsally in contact with third supraocular and parietal, ventrally either in contact with fifth supralabial, or separated from it by one of the other (smaller) postoculars. Lower eyelid with a semitransparent, undivided disc. Pupil round, lower margin may show one or a few small invaginations. Five supralabials, fifth largest, fourth below centre of eye. Supralabials followed by two postsupralabials, which decrease in size and almost reach ear-opening. Temporals 6-10, irregularly polygonal, smooth, juxtaposed, in oblique rows, slightly increasing in size posteriad. Posterior row, between parietal and ear-opening, formed by three relatively large scales. Ear-opening surrounded by small scales, with indented anterior margin. Tympanum recessed in an auditory meatus.
Mental wide, trapezoidal, rounded anteriorly. Postmental irregularly heptagonal, wider than long. Two large pairs of chinshields, in contact medially and with infralabials, followed by a third, much shorter, pair. Third pair usually in medial contact, and separated from infralabials by one wide and one narrow, elongate, scale. In some specimens third pair separated medially by a small scale, and so much reduced as not to be recognisable as chinshields. Five (rarely three or four) infralabials, fourth, or suture between third and fourth, below centre of eye. Infralabials followed by one or two postinfralabials, which almost reach ear-opening. All head scales juxtaposed, smooth.
Parietals followed by 5-8 pairs of trapezoidal, smooth, imbricate nuchals. Anterior pair of nuchals more than twice as wide as long, scales of posterior pair slightly longer than wide, other pairs intermediate. Between first pair of nuchals, more rarely also between second pair, may occur a medial, single scale. Scales on sides of neck flat, imbricate, smooth or multicarinate, with ellipsoid posterior margin; they increase slightly in size posteriad. Gulars with enlarged medial pair of scales (except, in some specimens, in the anterior row), slightly increasing in size posteriad, in 6-8, mostly seven, transverse rows (including collar). Collar not pronounced, formed by four scales.
Dorsals form transverse rows of narrow, imbricate scales, with truncate (hidden) anterior and sharply pointed posterior margins (causing a hexagonal impression), and with a wide, low keel; 28-32 (30.4 ± 1.2, n= 20) rows from first pair of nuchals to posterior level of hind limbs. Scales on flanks slightly wider, lanceolate; toward ventral region they gradually become smooth, and their posterior margins more rounded. Ventrals smooth, imbricate, in 18-21 (19.8 ± 1.1, n= 20) transverse rows, and in four longitudinal rows. Scales of medial pair of longitudinal rows larger, trapezoid; outer ventral scales roughly squarish, but with rounded posterior margins. Scales around midbody 26-29 (27.4 ± 1.1, n= 19). Most specimens examined with five pre-anal scales, of which a triangular one medially, two large paramedials that contact each other posterior to the medial triangular scale, and two small lateral ones. Para-medial scales completely separated in ZMUC 4377 (holotype), MPEG 8515, MPEG 14312, and MPEG 8513, in the former three specimens by an extra scale posterior to the medial one, in the latter specimen by the extension of the medial scale until the cloacal slit. In CEPB 498, MPEG 12178, and on one side of MPEG 13810 the lateral scales (which are variably developed i n other specimens) completely disappear. Pores well developed in adult males, between two scales, of which one much smaller than the other; 16-21 (18.7 ± 1.3, n= 12) in total. Preanal and femoral pores form a continuous row. Females without or with poorly developed pores (MPEG 13992 with two small femoral pores and one almost inconspicuous preanal pore on each side; MPEG 12907 with six pairs of scales similar to those carrying a pore, on each thigh, in at least three of which per side a small pore can be discerned; MPEG 8512 with 6-7 pairs of scales that might carry very inconspicuous pores).
Scales on tail similar to dorsals, but smaller. They form transverse rows of similar scales all around the tail.
Scales on forelimbs large, imbricate, smooth, except on ventral aspect ofupper arms, where they are much smaller. Hind limbs with very large, smooth, imbricate scales on anterior aspect of thighs; relatively large, smooth, imbricate scales on ventral aspect, bordered in males by the row of scales with pores; and granular scales on posterior and ventral aspects of thighs. Lower legs with large, smooth, imbricate scales o n their ventral aspect; slightly smaller scales o n anterior and posterior aspects, those on anterior aspect keeled; on dorsal aspect similar to the latter, but slightly decreasing in size. Fingers and toes well developed, with claws, except for first finger, which is reduced and clawless (according to Vanzolini & Ramos, 1977, this finger varies from tubercular, clawless, to relatively well developed). Lamellae under fingers single, not or slightly tuberculate, 9-11 (exceptionally eight or twelve) under fourth finger. Lamellae under toes mostly moderately tuberculate, under first, second and fifth toes single, under third and fourth toes mostly double; 14-17 (exceptionally 13 or 18) under fourth toe (counted along inner side) (Avila-Pires 1995: 370 ff).

Colour in life: MPEG 13992 (female), natal-brown (219A) on dorsal surface of head and most of back, changing on posterior part of back to chestnut (32); throughout back with small black spots. Flanks sepia (219). A narrow, dorsolateral, buff (124) stripe o n anterior part of body. Ventral region cream, translucent. According to Cunha (1977), the holotype of C. landii (MPEG 8513), when recently captured, had belly, underside of tail and of hind limbs light red (Avila-Pires 1995: 370 ff).

Color in preservative: dorsal region light brown, back scarcely to densely peppered with very small dark dots, and with small, irregular, dark-brown spots present in some specimens. A pale dorsolateral stripe, bordered dorsally by a dark-brown line, from nape, along body, and continuing along tail, although more conspicuous o n anterior part of body. Flanks dark-brown, with one more-or-less regular longitudinal row of small light dots, or with more numerous, widespread, white dots. Labials with alternate light and dark bars. Ventral region cream. Colour of tail similar to that of body, distally the dark lateral band may disappear. A dorsolateral row of light dots continues to tip of tail. Moreover, anterior part of tail with a middorsal dark-brown stripe, variably evident in different specimens. Colour pattern similar in males and females, but colours tend to be more contrasting in males (Avila-Pires 1995: 370 ff). 
CommentSynonymy: Perodactylus kraepelini has been removed from the synonymy of C. modesta and revalidated by Cacciali & Köhler 2017.

Distribution: Probably also in Bolivia (fide LANGSTROTH 2005). Not listed for Argentina by Avila et al. 2013, Zootaxa 3616 (3): 201–238, although there may have been questionable reports from Argentina.

Type species: Perodactylus modestus REINHARDT & LÜTKEN 1862 is the type species of the genus Colobosaura BOULENGER 1887. Note that the genus name Perodactylus is pre-occupied. 
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