Cnemidophorus nigrigula, Arias, Federico, Carvalho, Celso Morato De, Rodrigues, Miguel Trefaut & Zaher, Hussam, 2011

Arias, Federico, Carvalho, Celso Morato De, Rodrigues, Miguel Trefaut & Zaher, Hussam, 2011, Two new species of Cnemidophorus (Squamata: Teiidae) of the C. ocellifer group, from Bahia, Brazil, Zootaxa 3022, pp. 1-21 : 3-9

Cnemidophorus nigrigula sp. nov.

( Fig. 1 , 2 )

Holotype. MZUSP 93721 (field number MRT 3313), adult male, from Santo Inácio (10°39’S, 42°37’W), state of Bahia, Brazil, elevation 520 m, collected by Miguel T. Rodrigues et al., on 5 May 2000.

Paratypes. MZUSP 93735 to 93750 (field number, respectively MRT 3378 to 3393), collected by Miguel Trefaut Rodrigues from 4–7 May 2000, and MZUSP 93807, 93819 (field number, respectively MRT 3105, 3274), collected by Miguel T. Rodrigues, on 5 May 2000; same locality data as for the holotype. Male adults: MZUSP 93737, 93740, 93741, 93742, 93743, 93744, 93748, 93749, 93807 and 93819. Female adults: MZUSP 93735, 93736, 93738, 93739, 93745, 93746, 93747 and 93750.

Diagnosis. Cnemidophorus nigrigula is a member of the C. ocellifer group with which it shares the presence of granules in the supraorbital semicircles and the absence of anal spurs in males. The species differs from all other members of the group by having a gular and occasionally the sublabial region dark black, ontogenetic changes in male color pattern, sexual dimorphism in size, and a pronounced sexual dichromatism. Furthermore, Cnemidophorus nigrigula differs from C. jalapensis by having 15–20 (x= 17) femoral pores (11– 16, x= 13.3 in C. jalapensis ), 27–32 (x= 30) scales around tail (19–26, x= 22.7), 190–220 (x= 208.6) dorsal scales (200–250, x= 225.9), lateral spots present (lateral spots absent), and a larger body size (smaller, SVL= 53.45 mm). Cnemidophorus nigrigula differs from C. mumbuca by having 27–32 (x= 30) scales around tail (19–27, x= 22.6 in C. mumbuca ), dorsal stripes absent in adult males (present), row of bright turquoise ocelli on lower lateral fields (bluish white ocelli, only males). Cnemidophorus nigrigula differs from C. ocellifer by having 15–20 (x= 17) femoral pores (14–17, x= 16 in C. ocellifer ), 190–220 (x= 208.6) dorsal scales (172–188, x= 181), and dorsal stripes absent in adult males (present). Cnemidophorus nigrigula differs from C. confusionibus by having 27–32 (x= 30) scales around tail (22– 28, x= 24.9 in C. confusionibus ), 190–220 (x= 208.6) dorsal scales (188–211, x= 201.6), and lateral spots turquoise (yellow). Cnemidophorus nigrigula differs from all members of the littoralis subgroup ( C. littoralis , C. abaetensis and C. venetacaudus ) by having enlarged scales in temporal region, posterior to third subocular (absent), 2–3 rows of enlarged scales in dorsal part of arm (one row), 5 supraciliaries (6–7), 8 longitudinal rows of ventral scales (8– 10), 26–29 (x= 27) transverse ventral scale rows (29–38), 15–20 (x= 17.5) femoral pores (21–45), and a brown tail (bluish green). Furthermore, it differs from C. littoralis and C. abaetensis by the absence of a mid-dorsal stripe (present in both species), and absence of a stripe on the tail (present in both species). Cnemidophorus nigrigula differs from C. venetacaudus by having lateral ocelli on flanks (absent), and from C. cyanurus sp. nov. (see below) by having 15–20 (x= 17.5) femoral pores (31–38, x= 34.5 in C. cyanurus ), 8 longitudinal rows of ventrals (8–10), 87– 105 (x= 95) scales around midbody (110–126, x= 118.2), 5 superciliaries (6–7), mid-dorsal stripe absent (present), lateral spots on flanks (absent), and a brown tail (bright blue-green).

Description of holotype. Measurements: snout-vent-length (SVL) 81.11 mm, trunk length 34.19 mm, head length 20.71 mm, head width 12.99 mm, head height 11.49 mm, tail length 174.97 mm (2.16 times longer than SVL), humerus length 7.43 mm, femur length 14.76 mm, tibia length 11.1mm, foot length 25.92 mm, and arm length 51.78 mm.

Snout moderately pointed. Rostral large, wider than high, visible from above, separated from frontonasal by the midline contact between nasals. Anterior and posterior nasal in broad contact by an oblique suture. Nostril rounded, in lower part of suture. Frontonasal sub-hexagonal, contacting nasals and prefrontals. Prefrontals roughly trapezoidal, pentagonal, in broad contact along midline; contacting laterally nasal, loreal and first supraocular. Frontal approximately pentagonal, longer than wide, and wide anteriorly, separated from second, third and fourth supraoculars by a row of granules. Two frontoparietals, approximately pentagonal, separated from supraoculars by a row of granules. Five parietals, external ones smaller; interparietal sub-pentagonal, longer and wider than others, bordered at each side by parietals. Occipital scales irregular and variable in size. Four supraoculars on each side, second and third largest, first in contact with loreal, prefrontal, frontal and first supraciliary. Five supraciliaries on each side, first and second largest, others small, sub-equal, only first supraciliary in contact with first supraocular, all others separated from supraoculars by a row of granules. Loreal single, large, in contact with nasal, prefrontal, first supraocular, first supraciliary, preocular, first subocular, and third and fourth supralabials. Preocular narrow, higher than wide, in contact with first subocular, loreal, and small scales in ocular region. Three suboculars on each side, anteriormost subocular keeled, approximately pentagonal, in contact with fourth and fifth supralabial; second subocular keeled, longer than anterior, approximately rectangular, in contact with fifth, sixth and seventh supralabials; third subocular smooth, approximately round. A continuous keel runs from preocular to second subocular. Seven supralabials on each side. Temporal region with irregular scales, granular centrally, enlarged ventrally. A supratemporal row with moderately large scales, decreasing in size posteriorly. Ear opening large, round, higher than wide with smooth margins. All dorsal and lateral head scales juxtaposed, smooth (except for keeled preoculars and suboculars). Symphysal anteriorly ellipsoid, posteriorly in contact with first infralabials and postsymphysal, forming two wide angles. Postsymphysal single, pentagonal, in contact with first and second infralabials; followed by five pairs of enlarged chinshields. First pair of chinshields in broad contact along midline, in contact with second and third infralabials; longer than others. The second, third and fourth pairs of chinshields separated from infralabials by a row of small granules. Seven infralabials on each side; followed posteriorly by a series of small scales extending to commissure of mouth; first infralabials the smallest. Gular region divided in two areas: anterior region with enlarged, round scales, in transverse rows, delimited posteriorly by a line uniting lower margin of ear openings; posterior region covered with granules, in transverse rows, bordered posteriorly by the antegular fold. Gular and antegular folds marked by granules; three enlarged scale rows between the two folds. Scales on nape and sides of neck similar to dorsals. Dorsals and scales on flanks granular, rounded, smooth, sub-imbricate; 200 scales along a middorsal line from nape to base of tail; 99 scales around mid-body (excluding ventrals), 32 scales around tail. Ventral scales large, smooth, rectangular (wider than long), imbricate, in 29 longitudinal rows; 8 ventral scales in transverse rows across mid-body. Ventral scales separated from scales on flanks by row of moderately large scales. Preanal plate with three enlarged scales, one central surrounded laterally by small scales and two on posterior region. Femoral pores in a continuous row along each thigh, medially with a short gap; 9 pores on right side, 8 pores on left side. Scales on base of tail rectangular, smaller than ventrals, in transverse rows; keeled and slightly mucronate dorsal and laterally, smooth ventrally, imbricate. Tail scales becoming gradually longer and narrower from the base to tip; subcaudal scales becoming keeled distally. Limbs with large, smooth, imbricate scales on dorsal aspect of upper arms, antero-dorsal aspect of forearms, antero-ventral aspect of thighs, and ventral aspect of lower legs; elsewhere scales small, granular. Larger scales on upper arms in longitudinal rows. Forearms with one row of enlarged scales, wider than long. Anterior scales on thigh decreasing in size proximally, with three rows of enlarged scales. Lower legs with two rows of enlarged, hexagonal scales. Patch of scales modified in form on shin small on the heel. Ventral aspect of hands and feet granular; one enlarged tubercle at base of pollex. Sub-digital lamellae single; lamellae under left and right fourth finger 16; under left and right fourth toe 30.

Color in life. ( Fig. 1 ) Dorsal and dorso-lateral surfaces of the head, body, limbs and tail light brown, Lateral surface of the neck and flanks light gray. Dorso-lateral surface of the body ornamented with a row of regularly spaced bright turquoise ocelli that extends from the level of the forelimbs to the anterior third of the tail. Light gray area of the flanks between limbs with irregular bright turquoise ocelli. Throat and, occasionally labial region, black. Belly and ventral surface of limbs and tail white. Lateral surface of tail greenish white.

Variation in females and juveniles. Sexual dichromatism ( Fig. 3 ) and size dimorphism are evident (Table 1). In females, the dorsum of head, limbs and tail are brown. The anterior half of body is light brown, while the posterior half is bright green. There are three dorsolateral diffuse stripes on each side of the body. The paravertebral stripe extends from the neck to the base of the tail, while the dorsolateral stripe runs from the supraciliaries to the first third of the tail, and the ventrolateral originate on the subocular region, cross the ear, the body and extend through the hindlimbs to reach the feet. Irregular dark spots are disposed between the paravertebral and dorsolateral stripes, while a row of regularly spaced turquoise ocelli is present between the dorsolateral and ventrolateral stripes, from the forelimbs to the hindlimbs. Below this series of ocelli, there are irregular bluish spots that reach the margin of the ventrolateral scales. The lateral surface of the head is light gray, while the throat is dark black. The belly and the ventral surface of the limbs and tail are light white.

group. Supraocular scales (SO); supraciliary scales (SC); fourth finger lamellae (FFL): fourth toe lamellae (FTL); number of

granules around midbody (SAM); number of scales around tail ( SAT); number of dorsal granules (D); longitudinal rows of

enlarged scales in dorsal part of arm (RH); longitudinal rows of ventral scales (VL); transverse rows of ventral scales (VT);

total number of femoral pores (FP); enlarged scales in temporal region posterior to third subocular (EST).

Males exhibit ontogenetic changes in color pattern ( Fig. 4 ). In juveniles ( Fig. 5 A), the coloration is similar to that of females, with the distinction that stripes are bright white and dorsolateral bands are dark. In subadult males ( Fig. 5 B), the anterior half of the body is brown, and stripes are lost, while the posterior half of the body is bright green and stripes becoming diffuse. On the flanks, large regular turquoise-blue ocelli extend from the forelimbs to the first third of the tail. Bluish spots that reach the venter are irregularly disposed below the larger ocelli. Adult males ( Fig. 5 C) show the characteristic coloration of the holotype.

Color in preservative. Dorsal parts of head, body, limbs, and tail dark. Lateral region of the body with a series of regularly spaced turquoise-blue ocelli, from the forelimbs to first third of the tail. The lateral surface of the head and flanks (between the limbs) is light grey, with irregular turquoise-blue ocelli. Throat dark black. Belly and ventral region of limbs and tail white.

Variation. Variation in meristic characters is summarized in Table 1, while variation in measurements is given in Table 2 .

Etymology. The specific epithet, nigri, is a Latin adjective meaning ‘‘black,’’ and gula, a Latin noun in apposition, meaning throat, in allusion to the characteristic black coloration of the throat in this species.

Figures

FIGURE 1. Cnemidophorus nigrigula from Santo Inácio, Bahia state, Brazil. A) Dorsal view of an adult male. B) Ventral view of an adult male. Photo: Miguel Rodrigues.

FIGURE 2. Cnemidophorus nigrigula, holotype, MZUSP 93721, adult male. A) Dorsolateral view of the head; B) lateral view of head; C) ventral view of the head; D) preanal plate and femoral pores. Photo: J. C. Arredondo.

FIGURE 3. Sexual dichromatism in Cnemidophorus nigrigula. A) Dorsal view of an adult female. B) Dorsal view of an adult male. Both specimens from Santo Inácio, Bahia state, Brazil.

FIGURE 4. Ontogenetic change in color pattern in males of Cnemidophorus nigrigula. A) Juvenile. B) Sub adult. C) Adult.

FIGURE 5. Photo in life of Cnemidophorus nigrigula, showing color variation in males. A) Juvenile. B) Sub adult. C) Adult.

Tables

TABLE 2. Character variation (given as means, range in parentheses) for five members of the Cnemidophorus ocellifer sub-

Character	C. nigrigula sp nov. (n= 83)	C. jalapensis (n= 2)	C. confusionibus (n= 19)	C. mumbuca (n= 11)	C. ocellifer (n= 52)
SO	4 (4–4)	4 (4)	4 (4-4)	3.9 (3–5)	4 (4-4)
SC	5 (5)	5 (5)	5 (5)	5 (5)	5 (5)
TV	27.4 (26–29)	25.5 (25–26)	28.2 (27–29)	27 (24–29)	27.2 (26–28)
LV	8 (8)	7 (6–8)	8 (8)	8 (6–8)	8 (8)
FP	17 (15–20)	13 (12–14)	17.5 (16–21)	16.7 (14–20)	16 (14–17)
FFL	17 (16–18)	15.5 (15–16)	15.6 (15–17)	16.3 (13–19)	16 (15–17)
FTL	31.3 (30–32)	27 (26–28)	30.4 (29–35)	27.4 (24–32)	30 (28–31)
RH	2 (2)	3 (3)	3 (3)	3 (3)	3 (3)
SAM	99.8 (97–105)	96.5 (91–102)	95 (87–105)	101.0 (91–117)	94.6 (92–100)
SAT	30 (27–32)	24 (22–26)	24.9 (22–28)	22.6 (19–27)	28 (25–30)
D	208.6 (190–220)	206 (200–212)	201.6 (188–211)	229.6 (194–271)	181 (172–188)
EST	present	present	present	present	present