Ameiva reticulata, Landauro, Caroll Z., García-Bravo, Antonio & Venegas, Pablo J., 2015

Landauro, Caroll Z., García-Bravo, Antonio & Venegas, Pablo J., 2015, An endemic new species of Ameiva (Squamata: Teiidae) from an isolated dry forest in southern Peru, Zootaxa 3946 (3), pp. 387-400 : 388-397

Ameiva reticulata sp. nov.

Figs. 1–4 , 5 (A–D)

Holotype. CORBIDI 0 8816 ( Figs. 1–2 ), an adult male collected on 31 July 2010 by A. García in the Valle Seco del Mantaro (12°5'26.916''S, 74°41'55.968''W, WGS 84) at 1411 m.a.s.l., District of Surcubamba, Province of Tayacaja, Region of Huancavelica, Peru.

Paratypes. A total of 35 specimens, all from the Province of Tayacaja, Region of Huancavelica, Peru: one adult male ( CORBIDI 1095) from Barropata (12°17'34.812''S, 74°40'58.826''W, WGS 84), at 1528 m.a.s.l., District of Surcubamba, collected on 28 April 2008 by C.Z. Landauro; one adult male ( CORBIDI 9919) from Intivilca (12°19' 51.4''S, 74° 37'53.1''W, WGS 84) at 2238 m.a.s.l., District of Colcabamba, collected on 0 4 April 2011 by D. Amaya; one adult male ( CORBIDI 9917) and three adult females ( CORBIDI 9918, 9920, 9921) from Jatuspata (12°15'1.2''S, 74°41'33.6''W, WGS 84) at 2609 m.a.s.l., District of Surcubamba, collected on 0 7 April 2011 by D. Amaya; four adult males ( CORBIDI 10084, 10086, 10090, 10092), four juvenile males ( CORBIDI 10081, 10082, 10087, 10094), seven adult females ( CORBIDI 10075, 10076, 10078, 10085, 10088, 10089, 10093) and five juvenile females ( CORBIDI 10077, 10079, 10080, 10083, 10091), all from Valle Seco del Mantaro (12°5'43.073''S, 74°42'45.002'' W, WGS 84) at 1180 m.a.s.l., District of Surcubamba, collected on 0 1 and 0 2 November 2011 by A. Urbay and C.Z. Landauro; one adult female ( CORBIDI 13623) from Campamento Limonal (12°14'5.184'' S, 74°41'52.100'' W, WGS 84) at 1438 m.a.s.l., District of Surcubamba, collected on 31 August 2013 by A. Escóbar; eight specimens from Pichiu District of Colcabamba: one juvenile ( CORBIDI 13622) at 1974 m.a.s.l. (12°20'9.890'' S, 74°39'26.755'' W, WGS 84), collected on 10 September 2013 by C.Z. Landauro; two juveniles ( CORBIDI 13620, 13621) at 2003 m.a.s.l. (12°19'53.815'' S, 74°39'6.943'' W, WGS 84), collected on 11 September 2013 by C.Z. Landauro; one adult male ( CORBIDI 13625) and one adult female ( CORBIDI 13628) at 2169 m.a.s.l. (12°19'15.902'' S, 74°38'51.498'' W, WGS 84), collected on 14 September 2013 by A. Escóbar; one adult female ( CORBIDI 13629) at 2169 m.a.s.l. (12°19'32.913'' S, 74°39'5.017'' W, WGS 84), collected on 15 September 2013 by A. Escóbar; one adult male ( CORBIDI 13626) and one adult female ( CORBIDI 13624) at 1868 m.a.s.l (12°21'48.634'' S, 74°37'27.249'' W, WGS 84), collected on 18 September 2013 by A. Escóbar.

Diagnosis. A medium-sized Ameiva distinguished from all congeners by the following combination of characters: (1) maximum SVL in males 149 mm; (2) dorsal head scales smooth; (3) frontal single; (4) frontoparietal and parietal plates in contact with interparietals; (5) 17–24 (both sides) scales, usually in single row between supraoculars and supraciliaries; (6) 8–17 occipitals, usually subequal to first dorsal row; (7) 19–25 anterior gulars; (8) middle anterior gulars polygonal and usually distinctly enlarged; (9) patch of distinctly enlarged posterior gulars usually present; (10) 8–14 posterior gulars between antegular and gular folds; (11) enlarged mesoptychial scales subequal or larger than largest gulars; (12) postbrachials moderately to distinctly enlarged; (13) 181–237 scales between interparietals and base of tail; (14) 102–137 dorsal scales across midbody; (15) ventrals in 28–32 transverse rows, and in eight longitudinal rows; (16) in life, adult individuals with head, arms, dorsum and flanks pale brown or grayish brown, legs and tail bright green or turquoise, flanks with light green or turquoise ocelli; (17) throat in adults white; (18) dorsum finely reticulated without a pale vertebral stripe along dorsum; (19) in life, dorsum and arms of juveniles brown with an olive green tint, legs and tail turquoise, and flanks with pale yellow ocelli; (20) associated with interandean dry forest.

Comparison with other species. Ameiva reticulata shares the characters described by Harvey et al. (2012) for the A. ameiva complex (e.g. A. ameiva , A. atrigularis , A. pantherina , and A. praesignis ): frontal entire, its posterior suture usually contacting second supraocular or aligned with suture between second and third supraoculars; frontal ridge absent; interparietal entire; parietal series (including interparietal) composed of five relatively long plates; narial suture passes through nostril; first supralabial usually curved; first subocular usually separated from supralabials by anterior expansion of the second subocular; intertympanic crease present; pectoral sulcus absent; ventral scales in 10–12 longitudinal rows; plate-like antebrachials continuous with or narrowly separated from brachial scales; combined femoral and abdominal pores number 28–45; fifth toe reduced; complete row of scales separate supradigital and subdigital lamellae along postaxial edge of each toe; tip of snout of adult males never reddish; adults without dorsolateral and vertebral stripes, or vertebral stripe only apparent on posterior dorsum; flanks with pale ocelli.

Within the Ameiva ameiva complex, A. reticulata most resembles A. ameiva , a species widely distributed throughout Amazonia ( Ugueto & Harvey 2011), and differs from it as follows (characteristics of A. ameiva in parentheses): maximum SVL in males 149 mm (163 mm) and in females 129 mm (146 mm); frontal usually pentagonal (hexagonal or roughly pentagonal in some specimens); fourth supraocular separated from parietals by one or two rows of circumorbital scales (up to four rows of circumorbital scales); 17–24 scales laterally between supraoculars and supraciliaries laterally (19–31scales); ventrals in eight longitudinal rows (ten longitudinal rows); 181–237, mean= 199.9 ± 36.52, dorsal scales along the middorsal line (209–287, mean = 252.6 ± 16.65); 102–137, mean = 123.4 ± 8.24 dorsal scales at midbody in a transverse row (135–175, mean = 154.3 ± 8.8); 19–25, mean = 22.27 ± 1.56 anterior gulars (18–32, mean = 25.6 ± 2.64); preanal shield with four rows of enlarged scales (5–7); in juveniles, dorsal surface brown or olive green with black reticulation and white pale ocelli on flanks (first third or half of dorsum green and rest brown, flanks dark brown without ocelli; Fig. 5 E); in adults, dorsal surface of body brown or greenish-brown with a soft black reticulation, flanks completely brown or brown fading into greenish or turquoise towards hind limbs and with ill-defined light ocelli, tail turquoise (first third or half of dorsum brown with dark brown flecks, dots or blotches and remaining part of dorsum and tail green; Fig. 5 F).

From the other species of the Ameiva ameiva complex such as A. atrigularis , A. pantherina (both from Venezuela) and A. praesignis (from Costa Rica, Panama and Colombia) ( Ugueto & Harvey 2011), A. reticulata can be separated by having: a lower scale count with 181–237, mean = 252.6 dorsals along the middorsal line and 102– 137, mean = 123.4 dorsals across the midbody in a transverse row (263–361, mean = 309 and 134–179, mean = 151.3 in A. atrigularis ; 291–343, mean = 311.3 and 137–163, mean = 147.4 in A. pantherina ; and 237–348, mean = 289.3 and 111–157, mean = 133 in A. praesignis ). Ameiva atrigularis and A. praesignis are longer than A. reticulata , with a maximum SVL in males of 186 mm and 243 mm, respectively (149 mm in A. reticulata ). Further, throat coloration in adults is dark gray or black in A. atrigularis , dark gray in A. pantherina , cream or blue in A. praesignis (white in A. reticulata ).

Other species of the genus Ameiva that occur in Peru such as A. aggerescusans , A. concolor and A. nodam , are all restricted to the upper Marañón basin in northern Peru ( Koch et al. 2013), and share a transversely divided frontal plate, while it is entire in A. reticulata .

Description. In males maximum SVL is 149.1 mm (CORBIDI 9919) and maximum total length is 481.1 mm (CORBIDI 9919); in females maximum SVL is 129.1 mm (CORBIDI 10089) and maximum total length is 357.1 mm (CORBIDI 10089). Head pyramidal, 0.25–0.29 (0.27 ± 0.01, n = 16) times SVL in males; 0.23–0.30 (0.26 ± 0.016, n = 20) times SVL in females. Snout elongate, bluntly pointed; canthus rostralis distinct. Neck narrower than head and body. Body cylindrical. Limbs well developed; tibia 0.11–0.18 (0.15 ± 0.02, n = 16) times SVL in males; 0.12–0.16 (0.14 ± 0.01, n = 20) times SVL in females; foot 0.30–0.42 (0.37 ± 0.03, n = 16) times SVL in males; 0.28–0.41 (0.36 ± 0.03, n = 20) times SVL in females. Tail round in cross section, tapers toward tip, 1.93– 2.50 (2.26 ± 0.19, n = 11) times SVL in males; 1.76–2.40 (2.14 ± 0.24, n = 13) times SVL in females.

Rostral pentagonal, slightly wider than high, visible from above, bordered posteriorly by nasals, which form a short medial suture. Each nasal divided by an oblique suture. Nostril in lower part of suture and anterior to it, directed lateroposteriorly, slightly taller than long. Frontonasal octagonal, in contact with nasals, loreal and prefrontals. Prefrontals paired, roughly pentagonal, with a very long medial suture about twice as long as that between nasals, laterally in contact with loreal, first supraocular and first supraciliar. Frontal pentagonal (rectangular in CORBIDI 10082, CORBIDI 10092), longer than wide and wider anteriorly, its sutures with prefrontals form a wide angle, slightly straight, its sutures with frontoparietals almost forming a straight line; frontal laterally in contact with first and second supraocular. Pair of trapezoidal frontoparietals (divided into two posteriorly in CORBIDI 9918, 1007677, 10080, 10084, 10087, 10094 and 10095 and with a small scale accessory between frontal and frontoparietals in CORBIDI 0 9919, 10090, 10082, 10087, 10089 and 10094), wider than long and with a long medial suture; frontoparietals laterally in contact with second and third supraocular, and small circumorbital scales bordering fourth up to middle portion of third supraocular or beginning of second supraocular. Interparietal irregularly hexagonal, higher than wide, often narrower (but sometimes slightly wider) than adjacent parietals; interparietal longitudinally divided; sutures with parietals slightly oblique and curved; interparietal bordered at each side by two large, irregular parietals divided by an oblique suture (parietals divided into two in CORBIDI 10078); outermost parietal oval in shape and larger than inner parietal. Parietal series composed of five scales including interparietal. Occipitals 8–17 (13.61 ± 2.25, n = 36), irregular and heterogeneous in size; occipitals usually subequal in size to first dorsal row, less often moderately to distinctly larger. Supraoculars four at each side. Circumorbital semicircles formed by 4–8 scales at each side, 9–15 (11.25 ± 1.46, n = 36) combining both sides, reaching middle, posterior and anterior portion of third and second supraocular (extending completely around supraoculars except first supraocular); only one scale row between third supraocular and prefrontals; fourth supraocular separated from parietals by one or two rows of circumorbital scales. Laterally, all supraoculars except first separated from supraciliaries by a single row of small rectangular scales; 17–24 (20.72 ± 1.59, n = 36) combining both sides. Supraciliaries five or seven (on one side), first highest, second longest. Loreal very large, trapezoidal and single, in contact with nasal, frontonasal, first supraciliary, first and second subocular, and third and fourth supralabial. Suboculars four; first subocular in preocular position, irregularly trapezoidal, longer than wide and slightly wider than second subocular; in contact with loreal, first supraciliary, small scales in ocular region, and second subocular; margin with loreal slightly curved or straight. Third subocular longest, all, except first, in contact with supralabials. Continuous keel from first subocular through entire length of third subocular. Postoculars small, approximately in one row of four or five irregularly trapezoidal scales. Palpebral disc opaque with enlarged scales on lower eyelid. Supralabials seven, sixth below center of eye. Row of enlarged supratemporals decreasing in size posteriad. Temporal region with polygonal scales, slightly smaller centrally than peripherally. External auditory meatus large, vertically oval, bordered by granular scales; anterior margin semicircular, posterior one straight. Tympanum recessed. All dorsal and lateral head scales juxtaposed and smooth.

Symphysal anteriorly ellipsoid, its posterior sutures forming wide angles with infralabials and postsymphysal. Postsymphysal single and pentagonal; in contact with first and second infralabial, followed by five enlarged chinshields. First pair in ample medial contact, only the first in contact with infralabials. Remaining chinshields separated from infralabials by one row of sublabials. Medial chin scales moderately small, elongate, convex, smooth, juxtaposed, subhexagonal, in slightly oblique rows, all subequal in size. Infralabials six, fifth below center of eye followed to commissure of mouth by smaller scales. Gular region divided into two areas: anterior region with polygonal and flat scales in slightly oblique rows that usually remain subequal or rarely grade to larger scales medially, delimited posteriorly by a line uniting the lower margins of the ear openings; middle anterior gular scales usually small, rarely moderately enlarged, 19–25 (22.27 ± 1.56, n = 36) scales along midline of chin from anteriormost to posteriormost anterior gular. Posterior gular region covered by smaller polygonal scales in transverse rows; posteriormost median gulars usually forming medial patch of moderately enlarged scales, 8–14 (10.72 ± 1.64, n = 36) scales midventrally from anteriormost posterior gular to antegular fold. Mesoptychial scales moderately enlarged (larger or, less often, subequal to largest anterior gulars), in about one or two rows, hexagonal, flat, smooth, and juxtaposed.

Scales on nape and sides of neck similar to dorsals. Dorsals and scales on flanks granular (slightly larger on dorsum than laterally), round, smooth, subimbricate, in approximately transverse rows; 181–237 (199.9 ± 36.52, n = 36) scales between occipitals and base of tail. Scales around midbody (excluding ventrals) 102–137 (123.47 ± 8.24, n = 36). Ventrals large, smooth, rectangular (wider than long), imbricate, in eight longitudinal (at midbody) and 28–32 (30.08 ± 0.93, n = 36) transverse rows; transition between ventrals and scales on flanks sharp. Preanal shield with four rows of enlarged scales; pattern of terminal scales of the preanal plate somewhat variable; often large median scale followed posteriorly by two subequal scales. Preanal plate surrounded anteriorly and laterally by smaller scales; posteriorly by much smaller scales. Femoral pores in continuous row along each thigh, 16–20 pores on each leg, 32–40 (35.88 ± 2.13, n = 36) pores combining both legs; each pore surrounded by four scales.

Scales on base of tail trapezoidal, smaller than ventrals, longer than wide, mostly keeled (smooth on base of tail on ventral surface). Caudal scales imbricate, in transverse rows, continuous around tail. Distal caudals longer and narrower, distinctly keeled, in transverse and approximately longitudinal rows. Arms with rows of very large, smooth, slightly imbricate, trapezoidal antebrachial scales on anterodorsal aspect of forearms and similar but smaller brachial scales on upper arms that extend almost to shoulder. Brachial row distinctly enlarged with smaller accessory row bordering upper margin and grading into granules posteriorly. Antebrachials and brachials in contact or separated by smaller scales at elbow. Dorsoposterior, posterior, and ventral aspect of arms with scales similar to dorsals, but slightly larger, except for a group (approximately one or two rows) of postbrachial scales on posterior aspect of upper arms, which are slightly more irregular. Legs with large, smooth, imbricate scales on anterior and ventral aspects of thighs, and ventral aspects of shanks. One row of large, trapezoidal scales anteriorly on thigh, gradually becoming smaller and irregular toward pores. On ventral aspect of shanks, four rows of large scales, anterior two trapezoidal, posterior one rhomboidal, decreasing in size from anterior toward posterior row. Elsewhere on legs scales similar to dorsals.

Subdigital lamellae transversely enlarged and single, moderately to distinctly tuberculate towards base. Tubercles most prominent under base of third toe, lamellae of outer toe continuing to heel. On palms, lamellae of inner finger continuing to wrist as large scales, increasing in size toward it; single large scale on postaxial side, following line of fifth finger, at short distance from wrist. Lamellae under fourth finger 14–17 (15.83 ± 0.69, n = 36), under fourth toe 27–36 (31.41 ± 1.79, n = 36).

Data of holotype. SVL 136.5 mm; head length 36.9 mm; tibia length 19.7 mm; foot length 45.5 mm; hand length 13.35 mm; tail length 285 mm; supralabials 7/7; infralabials 6/6; supraoculars 4/4; circumorbitals 5/5 extending to mid third supraocular; 10/11 scales between supraorbital and supraciliary scales; parietals 5; occipitals 16; occipitals larger than first dorsal row; 22 anterior gulars; mid anterior gulars small; 14 posterior gulars, midposterior gulars moderately enlarged; mesoptychial scales smaller than largest anterior gulars; dorsals between occipitals and base of tail 185; 120 scales around midbody (excluding ventrals); 8 longitudinal ventral rows; 30 transverse ventral rows; femoral pores 18/18; four scales forming preanal plate; 16 lamellae under fourth finger on right hand; 31 lamellae under fourth toe on right foot.

Color in preservative. Young specimens with dorsal surface of head and dorsal part of body uniformly brownish with medium-sized black spots at each side of vertebral region (most conspicuous towards sacrum). Sides of head uniformly brownish. Some specimens with almost half of vertebral region skyblue or turquoise. Dorsolateral surface black with regular turquoise spots, extending to base of tail. Upper limbs brownish with some small black scales. Hind limbs black with turquoise spots or turquoise with black spots. Throat pale, chest, ventral aspect of arms and most of abdomen semi pale (with a bluish tint). Ventral aspect of legs, posteriormost portion of abdomen, anal and subcaudal regions whitish with some dark scales.

Adult males have a turquoise brown dorsum with black reticulation or flecks ( Fig. 3 ; more visible in some females: Fig. 4 ); these markings remain conspicuous along dorsum from nape to base of tail. Flanks with small, regular, slightly scattered turquoise ocelli with distinct black margins more or less arranged in vertical rows; ocelli never extend onto dorsum. Top and sides of head uniformly light brown with black spots. Upper limbs light brown with black spots (smaller and less conspicuous than those on dorsum). Tail lighter than body with numerous scattered light and dark scales. Chin, throat and ventral aspect of arms pale, chest and most of abdomen pale or yellowish with a bluish tint; posteriormost portion of abdomen, anal area, and subcaudal region pale (a few dark scales present on subcaudal region). Ventral aspect of thighs pale-bluish without black spots. Ventral aspect of shanks uniformly pale-bluish.

Color in life. Dorsal surface of head light-brown with irregular black spots or flecks. Dorsal field of body predominantly greenish or turquoise covered by a thin black reticulation ( Fig. 5 A–B, D) which extends from nape to base of tail, becoming broader in last half of body in some specimens. Anterior to base of tail the dorsal surface is yellowish-brown with black reticulation. Flanks light brown fading into greenish or turquoise with light ocelli or bold reticulation from middle of flank to groin ( Fig. 5 D). One female (CORBIDI 10075) has a light longitudinal stripe with black borders from middle of flanks to groin. Ventral surface pale and tail pale bluish. Forelimbs light brown with black flecks. Hind limbs greenish or turquoise (in some juveniles brownish) covered by light spots with black margins or a black reticulation. Tail greenish or turquoise with a thin black reticulation similar to that on dorsum. Dorsum in juveniles brown with an olive green tint and thin black reticulation and white pale ocelli on flanks ( Fig. 5 C). Forelimbs light brown with black flecks as in adults and hind limbs gray or brown covered by pale spots and black reticulation.

Etimology. The specific epithet is an adjective derived from the latin word “ reticulatus” meaning “net-like” and refers to the soft net-like dorsal pattern of this species.

Distribution and natural history observations. Ameiva reticulata is known from four localities, all in the valley of the Mantaro River, Region of Huancavelica, in southern Peru, at elevations between 1113 m and 2609 m a.s.l. ( Fig. 6 ). The new species inhabits the seasonal dry forest of the Valle Seco del Mantaro ( Fig. 7 ) at the localities Barropata, Jatuspata, Intivilca, and Pichiu. Probably this species also occurs in the deep valley of the Las Pampas River in the Region of Ayacucho. The habitat at the aforementioned localities is seasonal dry forest with scattered croplands with plantations of corn Zea mays , avocado pear Persea sp., citrics and several species of fruit trees. The agriculture is more intensive in the surrounding areas of Pichiu village. Most individuals of A. reticulata were found at midday foraging actively at the base of cacti, shrubs (mainly thorn shrubs such as Acacia macracantha ) and stone walls. When threatened, individuals quickly hide under rocks or inside little self-dug burrows. The climate of the Mantaro River valley is usually dry and sunny.

Figures

FIGURE 1. Holotype of Ameiva reticulata sp. nov. (CORBIDI 8816, adult male, SVL = 136.5 mm). (A) dorsal view and (B) ventral view. Photographs by P. J. Venegas.

FIGURE 2. Holotype of Ameiva reticulata sp. nov. (CORBIDI 8816): dorsal (A), lateral (B) and ventral (C) views of head. Drawings by C. Koch.

FIGURE 3. Coloration and pattern of adult male paratypes of Ameiva reticulata sp. nov. in preservative: (A) Dorsal and (B) lateral views of CORBIDI 9917, (C) dorsal and (D) lateral views of CORBIDI 10086, and (E) dorsal and (F) lateral views of CORBIDI 10084. Photographs by P. J. Venegas.

FIGURE 4. Coloration and pattern of adult female paratypes of Ameiva reticulata sp. nov. in preservative: (A) dorsal and (B) lateral views of CORBIDI 10085, (C) dorsal and (D) lateral views of CORBIDI 10089, and (E) dorsal and (F) lateral views of CORBIDI 10075. Photographs by P. J. Venegas.

FIGURE 5. Ameiva reticulata sp. nov. (A – D) and A. ameiva (E-F) in life: (A) adult female of A. reticulata (CORBIDI 10076); (B) adult male of A. reticulata; (C) juvenile paratype (CORBIDI 13621); (D) adult female (CORBIDI 10088); (E) juvenile of A. ameiva from Madre de Dios; and (F) adult male of A. ameiva (CORBIDI 1713) from Yurimaguas. Photographs by C. Z. Landauro (A – D) and P. J. Venegas (E – F).

FIGURE 6. Map showing the distribution of Ameiva reticulata sp. nov. and A. ameiva in Peru. Triangles represent A. reticulata (red triangle is the type locality) and circles represent A. ameiva.

FIGURE 7. General landscape of the habitat of Ameiva reticulata sp. nov. at Valle Seco del Mantaro on November 2011. Photograph by C. Z. Landauro.