Enyalioides anisolepis , Torres-Carvajal, Omar, Venegas, Pablo J. & de Queiroz, Kevin, 2015
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Taxon classification Animalia Squamata Hoplocercidae
Enyalioides anisolepis sp. n.
Enyalioides anisolepis Proposed standard English name: rough-scaled woodlizards
Enyalioides anisolepis Proposed standard Spanish name: lagartijas de palo de escamas ásperas
Holotype. QCAZ 12537 (Fig. 5), an adult male from the eastern bank of the Mayo river, 4.5 km ESE Zumba, - 4.88605°S, - 79.08738°W (DD), 765 m, Provincia Zamora-Chinchipe, Ecuador, collected on 11 April 2014 by D.A. Paucar, D. Almeida-Reinoso, G. Galarza and D. Pareja.
Paratypes (14). ECUADOR: Provincia Zamora-Chinchipe: QCAZ 12521, juvenile with the same collection data as the holotype except - 4.88673°S, - 79.08744°W, 738 m; QCAZ 12527, adult male (Fig. 6) with the same collection data as the holotype except - 4.87147°S, - 79.08542°W, 738 m; QCAZ 12528, juvenile with the same collection data as the holotype except - 4.87136°S, - 79.08534°W, 738 m; QCAZ 12531, female with the same collection data as the holotype except - 4.87808°S, - 79.08956°W, 738 m; QCAZ 12535, juvenile (Fig. 6) with the same collection data as the holotype except - 4.88658°S, - 79.08747°W, 731 m; QCAZ 12536, juvenile with the same collection data as the holotype except - 4.88622°S, - 79.08737°W, 744 m; QCAZ 12552, female (Fig. 6) with the same collection data as the holotype except - 4.87589°S, - 79.08995°W, 741 m; QCAZ 12551, juvenile with the same collection data as the holotype except - 4.87521°S, - 79.08965°W, 724 m, collected on 12 April 2014; QCAZ 12517, adult male from Nuevo Paraíso, 700 m NW on road to Las Tres Aguas, - 4,87109°S, - 78,97579°W, 1742 m, collected on 10 April 2014 by the same collectors as the holotype; QCAZ 8395, female from Chito, sector Los Planes, - 4.89814°S, - 78.98095°W, collected on 16 February 2008 by S. Aldás-Alarcón; QCAZ 8515, female from Chito, sector Los Planes, - 4.89726°S, - 78.98191°W, collected on 18 February 2008 by S. Aldás-Alarcón; QCAZ 8428, female from Chito, 4.82037°S, - 78.96247°W, 1724 m, collected on 14 February 2008 by S. Aldás-Alarcón. PERU: Provincia San Ignacio: Región Cajamarca: CORBIDI 870, female from Alto Ihuamaca-Namballe, - 5.19448°S, - 79.08048°W, 1616 m, collected on 26 August 2008 by M. Dobiey; MUSM 20675, adult female from El Sauce, Tabaconas Namballe National Sanctuary, - 5.17897°S, - 79.16347°W, 1600 m, collected in April 2003 by C. Aguilar.
Enyalioides anisolepis can be distinguished from other species of Enyalioides , except for Enyalioides heterolepis , by having conical dorsal head scales (only in Enyalioides anisolepis and Enyalioides heterolepis ) and scattered, projecting, large scales on the dorsum, flanks, and hind limbs (also in Enyalioides / Morunasaurus annularis and Enyalioides / Morunasaurus groi ), which are conspicuous in adults of both sexes (Fig. 7). Besides occurring on opposite sides of the Andes, Enyalioides anisolepis differs from Enyalioides heterolepis (character states from Torres-Carvajal et al. 2011 in parentheses) in having fewer vertebral scales, 43-62, 50.87 ± 6.27 (52-98, 74.61 ± 10.39), a higher vertebral crest with the vertebrals on neck at least three times higher than those between the hind limbs (vertebrals on neck maximum twice as high as those between hind limbs), scattered dark spots on belly in juveniles and adults of both sexes (belly without scattered dark spots, blackish medially in some adult males), tail in adult males moderately compressed laterally (strongly compressed), and a marked sexual dichromatism (Fig. 6), with the dorsal background color greenish in males and brownish in females (both sexes with a brownish background).
The only other species of Enyalioides with scattered, projecting dorsal scales is Enyalioides cofanorum , which differs from Enyalioides anisolepis in lacking projecting scales on the hind limbs, and in being smaller in size (maximum SVL in males and females of Enyalioides cofanorum 107 mm and 109 mm, respectively; 130 mm and 119 mm in Enyalioides anisolepis ). Additionally, adults of both sexes of Enyalioides cofanorum have a brownish background (marked sexual dichromatism in Enyalioides anisolepis ).
Description of holotype.
Male (Fig. 5); SVL = 130 mm; TL = 220 mm; maximum head width = 28.7 mm; head length = 35.3 mm; head height = 24.6 mm; dorsal head scales keeled or multicarinate, those in the parietal region strongly projected dorsally; scales immediately posterior to superciliaries conical and dorsally projected, forming longitudinal row of ten (left) or nine (right) scales that extends posteriorly over supratemporal region; temporal scales small, tuberculate or keeled, juxtaposed; one enlarged pretympanic scale; superciliaries 17; canthals six; postrostrals three; supralabials 11 if counted to a point right below middle of eye, and 16 if counted to commisure of mouth; rostral (2.6 mm wide × 1.5 mm high) about twice as wide as adjacent supralabials; single longitudinal row of lorilabials between suboculars and supralabials at level of middle of eye, longitudinal rows of lorilabials anterior to this point 2-4; loreal region with small, smooth and keeled, juxtaposed scales; nasal at level of supralabials III and IV; infralabials nine (left) or eight (right) if counted to a point right below middle of eye, and 13 (left) and 11 (right) if counted to commisure of mouth; mental (2.8 mm wide × 1.5 mm high) wider and higher than adjacent infralabials; postmentals two; gulars projected, low; gular fold complete midventrally; small dewlap present; neck with several longitudinal and oblique folds.
Vertebral crest strongly projected and decreasing in size posteriorly, with vertebrals on neck at least three times higher than those between hind limbs; crest bifurcates at a point approximately 10 mm posterior to the cloaca, and extends onto tail about ¼ its length; flanks between fore and hind limbs with dorsolateral and ventrolateral longitudinal folds, as well as several oblique folds; axillary region with three vertical folds; scales on dorsolateral folds slightly larger than adjacent scales giving the fold the appearance of a low crest; scales between dorsolateral folds and vertebral crest heterogeneous in size, prominently keeled, and imbricate, with largest scales twice as large as smallest ones; neck and scapular region with scattered, large conical scales; flank scales ventral to dorsolateral folds similar to those dorsal to folds, with largest scales four times as large as smallest ones (Fig. 7); axillary region with conical scales forming two short vertical folds; ventral scales imbricate, keeled, rhomboidal, with a posterior mucron; ventrals more than three times the area of smallest dorsals.
Limb scales keeled dorsally and ventrally, homogeneous in size on fore limbs; scales on dorsal and posterior aspect of thighs heterogeneous in size, with most scales less than half the size of those on anterior and ventral aspects; scales on dorsal surface of shanks heterogeneous in size, with granular scales between large keeled scales; subdigitals on finger IV 17; subdigitals on toe IV 25; three femoral pores on left leg, two on right leg; tail laterally compressed and gradually tapering posteriorly; caudal scales strongly keeled and imbricate, increasing in size posteriorly on lateral and dorsal aspects of each caudal segment; caudals larger ventrally than dorsally; individual caudal segments three scales long ventrally and six scales long dorsally.
Color in life of holotype
(Fig. 5). Dorsal and lateral aspects of head with scattered black, brown, and pale green scales; labials greenish cream; dorsal background of body, limbs and tail brownish green with scattered pale green scales; vertebral crest pale green, the base and posterior surface of each vertebral scale dark brown; gular scales cream, the skin between them gray; orange patch on medial aspect of throat; chest and belly cream with a pale orange tint; ventral surface of limbs dirty cream with scattered brown spots; ventral surface of tail dirty cream proximally and brown distally; iris pale brown peripherally with dark brown reticulations, dark brown centrally; thin golden ring around pupil.
Variation in meristic and morphometric characters of Enyalioides anisolepis are presented in Table 2. Enlarged pretympanic scales are absent in more than half of the specimens; when present, they vary between 1-3. A few specimens have smooth scales intermixed with the keeled dorsals. Ventrals are keeled except for an adult female specimen (QCAZ 8428) that has smooth ventrals, and an adult male specimen (QCAZ 12517) that has smooth ventrals anteriorly and feebly keeled ventrals posteriorly. Caudal segments are 6-8 scales long laterally.
This species has a marked sexual dichromatism in background colors (green in males, brown in females; see Fig. 6). Adult male paratypes are very similar in color patterns to the holotype, except for having dark spots on the belly. A subadult male (QCAZ 12517) has scattered black flecks on the gular region.
Adult females share similar color patterns with juveniles (Fig. 6): dorsal background of head, body, limbs and tail dark or pale brown; flanks dark or pale brown with scattered dark spots, blotches, or transverse bands (cream flecks in QCAZ 8428); diagonal subocular dark stripe extending from subocular region to commisure of mouth; faint cream stripe extending longitudinally from tympanum to scapular region, except in specimen QCAZ 8428, which has instead a cream blotch posterior to tympanum; limbs with faint brown transverse bands; throat, chest, belly and ventral surface of limbs and tail pale brown or cream, with scattered dark spots on belly and thighs (dark spots absent in QCAZ 8428). In addition, juveniles generally have dark brown transverse bands on dorsum, dark flecks on head, and transverse rows of dark brown blotches on flanks. The neck and sides of head have a bright orange tint in one specimen (QCAZ 12535).
Distribution and ecology.
Enyalioides anisolepis is known to occur between 724-1742 m on the Amazonian slopes of the Andes in southern Ecuador and northern Peru (Fig. 3). It is known from Provincia Zamora-Chinchipe in extreme southern Ecuador and Región Cajamarca in northern Peru. Most specimens were found sleeping at night (7:00 pm– 1:00 am) between 0.2-1.5 m above ground on stems, leaves, and tree roots in primary and secondary forests. Nine of the 15 known specimens were found within 5 m of small streams.
The specific epithet anisolepis is a noun (in apposition) in the nominative singular and derives from the Greek words anisos (= unequal) and lepis (= scale). It refers to the heterogeneous scales on the dorsum, flanks and hind limbs of lizards of this species.
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