Holotype. QCAZ 8073 (Fig. 1), an adult male from Alto Tambo, 5 km on road to Placer, Bosque Integral Otokiki, 0.90600°N; -78.60600°W (DD), 620 m, Provincia Esmeraldas, Ecuador, collected on 2 May 2010 by I.G. Tapia, D. Almeida-Reinoso, J.M. Guayasamin and L.A. Coloma.

Figure 1. 

Holotype (QCAZ 8073, adult male, SVL = 119 mm) of Enyalioides altotambo in dorsal (top) and ventral (bottom) views. Photographs by Luis A. Coloma.

Paratype. ECUADOR: Provincia Esmeraldas: QCAZ 6671, adult female, Alto Tambo, Balthazar river, 0.90000°N; -78.61667°W, 645 m, collected on 5 November 2005 by F. Ayala-Varela and I.G. Tapia.

Enyalioides altotambo differs from other species of Enyalioides, except for E. oshaughnessyi, in having dorsal scales that are both smooth and homogeneous in size. It can be distinguished from E. oshaughnessyi (character states in parentheses) by the following characters: iris brown in both sexes (iris bright red in both sexes); scales on lateral edge of skull roof just posterior to superciliaries strongly projected (moderately projected); adults of both sexes with light green spots on dorsum (if present, spots turquoise or blue); adult males with a black medial patch on gular region not extending dorsally to form an antehumeral bar (black patch under gular fold extending dorsally to form a short antehumeral bar); scales on flanks almost homogenous in size (flank scales heterogeneous in size, with a few enlarged, circular, keeled scales); pale postympanic stripe on lateral aspect of neck in both sexes (pale postympanic spot in both sexes), posterior surface of thighs without enlarged scales (scattered enlarged scales), tail length/total length 0.57–0.60 (0.59–0.62).

Male (Fig. 1); SVL = 119 mm; TL = 160 mm; maximum head width = 21.9 mm; head length = 29.8 mm; head height = 20.3 mm; dorsal head scales keeled or multicarinate, projected dorsally; parietal eye present; eight scales immediately posterior to superciliaries conical, dorsolaterally projected, and conspicuously larger than adjacent scales; temporal scales small, pyramidal, low; one large conical pretympanic scale; superciliaries 17; canthals five; postrostrals three; supralabials 13 if counted to a point below middle of eye; rostral divided into three small scales, similar in size to adjacent supralabials; one longitudinal row of lorilabials between suboculars and supralabials at level of middle of eye, longitudinal rows of lorilabials anterior to this point two; loreal region with small, keeled, and juxtaposed scales; nasal at level of supralabials V–VI; infralabials 11 if counted to a point below middle of eye; mental (1.68 mm wide × 1.98 mm high) slightly wider and 1.5 times higher than adjacent infralabials; postmentals three; gulars ventrally projected and separated from each other by skin covered with tiny granular scales; gular fold complete midventrally, extending dorsally and posteriorly to form an antehumeral fold; neck with some oblique folds, and a dorsolateral row of enlarged scales; distal aspect of oblique fold immediately anterior to antehumeral fold with approximately six enlarged scales similar in size to gulars, but more than three or four times the size of adjacent fold scales.

Vertebral crest strongly projected and decreasing in size posteriorly, with vertebrals on neck at least four times higher than those between hind limbs; crest bifurcates at a point approximately 10 mm posterior to the cloaca, and extends onto tail about 1/3 its length; body flanks between fore and hind limbs with slight dorsolateral fold; scales on dorsolateral fold slightly larger than adjacent scales; dorsal and flank scales small, smooth, imbricate, more or less homogeneous in size; ventral scales imbricate, keeled, rectangular or rhomboid, with a posterolateral mucron; ventrals more than twice the size (area) of dorsals.

Limb scales keeled dorsally and keeled or feebly keeled ventrally; scales on dorsal and posterior aspects of thighs heterogeneous in size, with most scales less than half the size of those scales on anterior and ventral aspects, separated from each other by skin covered with tiny granular scales; subdigitals on finger IV 25; subdigitals on toe IV 29; femoral pores on each side one; tail laterally compressed and gradually tapering posteriorly; caudal scales smooth at the base of tail, becoming keeled and imbricate towards tip, gradually 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 seven scales long dorsally.

(Fig. 1). Head light green with a few black and dark brown scales; superciliaries, canthals and labials yellow; bluish cream blotch, wider than high, behind tympanum; pretympanic scales bluish cream; dorsal body background light green with a fine dark brown reticulation and scattered bluish cream scales; vertebrals yellowish green; tail green with incomplete dark brown rings; black irregular marks on limbs, covering most of hands and feet; chin white; gular region bluish cream anterolaterally grading into yellowish green and then bluish green posteriorly, with a posteromedial black patch; ventral aspect of body, limbs and tail dirty cream; flank color pattern extending onto ventrolateral aspect of body; iris brown with golden ring around pupil.

Variation in meristic and morphometric characters of Enyalioides altotambo are presented in Table 2. The single female paratype (QCAZ 6671; Fig. 2) is similar in lepidosis and color patterns to the holotype. It differs from the holotype in lacking a black gular patch, and in having a longer pale postympanic stripe, a yellow chin, and a yellow gular region. Furthermore, the scales on the lateral edge of the skull roof and those forming the vertebral crest are more projected in the female (Fig. 2); however, this variation could be ontogenetic rather than sexual because the female is larger (SVL = 134 mm) than the male (SVL = 119).

Figure 2. 

Paratype (QCAZ 6671, adult female, SVL = 132 mm) of Enyalioides altotambo. Photograph by Luis A. Coloma.

Enyalioides altotambo is only known from two adjacent localities at 620–645 m in the Chocoan rainforests of northwestern Ecuador (Fig. 3). Female paratype QCAZ 6671 was found at 5:30 pm with its head facing up on a tree trunk.

Figure 3. 

Distribution of Enyalioides altotambo (circles), E. anisolepis (triangles) and E. sophiarothschildae (squares).

The specific epithet is a noun in apposition and refers to Alto Tambo, Provincia Esmeraldas, Ecuador, a village on the Ibarra-San Lorenzo road where Enyalioides altotambo was discovered.

Although previously referred to Enyalioides oshaughnessyi, the possibility that the specimens here named Enyalioides altotambo represented a distinct species was recognized in previous studies. In a phylogenetic analysis of hoplocercine lizards, Torres-Carvajal and de Queiroz (2009) found “E. oshaughnessyi” to be paraphyletic relative to E. touzeti based on three samples of “E. oshaughnessyi”. One of them corresponded to the paratype of Enyalioides altotambo, and was sister to a clade containing the sister taxa E. touzeti and E. oshaughnessyi. Torres-Carvajal et al. (2011) noted that the color of the iris in live specimens of “E. oshaughnessyi” from Alto Tambo was not bright red as in live specimens of “E. oshaughnessyi” from other localities and suggested that the two forms represented separate species. Nonetheless, these authors found no other differences between the two potential species and refrained from associating the name E. oshaughnessyi with one versus the other because the type locality data of E. oshaughnessyi is vague (“Ecuador”), and the color of the iris was not recorded in its original description (Boulenger 1881). Here we recognize known populations other than that at Alto Tambo as E. oshaughnessyi based on the enlarged, circular and keeled scales scattered on the flanks of E. oshaughnessyi (absent in E. altotambo), as described and illustrated in its original description (Fig. 4; Boulenger 1881).

Figure 4. 

Holotype of Enyalioides oshaughnessyi (MRHN [Museum Royal d’Histoire Naturelle, Belgium] 2009, adult male). Illustration taken from original description (Boulenger 1881).

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.

Figure 5. 

Holotype of Enyalioides anisolepis (QCAZ 12537, adult male, SVL = 130 mm). Top: dorsolateral view; middle: ventral view; bottom: lateral view of head. Photographs by Omar Torres-Carvajal.

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.

Figure 6. 

Paratypes of Enyalioides anisolepis. Lateral (left) and ventral (right) views of an adult male (top, QCAZ 12527, SVL = 111 mm), an adult female (middle, QCAZ 12552, SVL = 101 mm), and a juvenile (bottom, QCAZ 12535, SVL = 59 mm). Photographs by Omar Torres-Carvajal.

Enyalioides anisolepis can be distinguished from other species of Enyalioides, except for E. heterolepis, by having conical dorsal head scales (only in E. anisolepis and E. heterolepis) and scattered, projecting, large scales on the dorsum, flanks, and hind limbs (also in E./Morunasaurus annularis and E./M. groi), which are conspicuous in adults of both sexes (Fig. 7). Besides occurring on opposite sides of the Andes, E. anisolepis differs from E. 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).

Figure 7. 

Close-up of left dorsum of Enyalioides anisolepis (QCAZ 12537, holotype) showing scattered enlarged scales. Scale bar = 5 mm. Photograph by Omar Torres-Carvajal.

The only other species of Enyalioides with scattered, projecting dorsal scales is E. cofanorum, which differs from E. anisolepis in lacking projecting scales on the hind limbs, and in being smaller in size (maximum SVL in males and females of E. cofanorum 107 mm and 109 mm, respectively; 130 mm and 119 mm in E. anisolepis). Additionally, adults of both sexes of E. cofanorum have a brownish background (marked sexual dichromatism in E. anisolepis).

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.

(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).

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.

Holotype. CORBIDI 647 (Fig. 8), an adult male from Río Lejía in the trail La Cueva-Añazco Pueblo, -6.83655°S; -77.48603°W (DD), 1700 m, Provincia Mariscal Cáceres, Región San Martín, Perú, collected on 2 February 2008 by P.J. Venegas.

Paratypes (2). PERU: Región San Martín: Provincia Mariscal Cáceres: MUSM 21883-84, adult males, El Dorado, -6.76666°S; -77.54500°W, 1600m, collected on 5 December 2003 by P.J. Venegas.

Figure 8. 

Holotype of Enyalioides sophiarothschildae sp. n. (CORBIDI 647, adult male, SVL = 135 mm). Top: dorsolateral view; middle: lateral view of head; bottom: ventral view. Photograph by Pablo J. Venegas.

Enyalioides sophiarothschildae can be distinguished from other species of Enyalioides, except for E. laticeps, by having caudal scales that are relatively homogeneous in size on each caudal segment; in all other species of Enyalioides, the dorsal and lateral caudals increase in size posteriorly on each caudal segment, and the largest (posteriormost) caudals on each segment are mucronate or have some kind of projection (Torres-Carvajal et al. 2011). Enyalioides sophiarothschildae differs from E. laticeps (character states in parentheses) in color patterns: gular region in males white with a black medial patch scattered with turquoise scales (orange or dirty cream with longitudinal brown, reddish-brown, bluish, or orange streaks, and a large brown or black medial blotch at the level of the gular fold); chest in males grayish white with a turquoise tone (usually an orange tone); labials and chin immaculate white (cream or green in many tones, but never immaculate white).

Male (Fig. 8); SVL = 135 mm; TL = 223 mm; maximum head width = 28.4 mm; head length = 34.7 mm; head height = 24.3 mm; dorsal head scales uni- or multicarinate, projected dorsally; parietal eye present; 3–4 scales immediately posterior to superciliaries conical, dorsolaterally projected, and slightly larger than adjacent scales; temporal scales small, multicarinate, separated from each other by tiny granular scales; no distinctly enlarged pretympanic scales; superciliaries 13; canthals six; postrostrals three; supralabials 10 if counted to a point below middle of eye; rostral (3.14 mm wide × 1.47 mm high) slightly wider than adjacent supralabials; single longitudinal row of lorilabials between suboculars and supralabials at level of middle of eye, two longitudinal rows of lorilabials anterior to this point; loreal region composed of small, smooth, and juxtaposed scales, some of which are separated from each other by tiny granular scales; nasal at level of supralabials III–IV; left and right infralabials nine if counted to a point below middle of eye; mental (2.77 mm wide × 2.60 mm high) slightly wider and 1.5 times higher than adjacent infralabials; postmentals two; gulars ventrally projected and separated from each other by skin covered with tiny granular scales; gular fold complete midventrally, extending dorsally and posteriorly to form antehumeral fold; neck with several oblique folds, and a dorsolateral row of enlarged scales; distal part of oblique fold immediately anterior to antehumeral fold with approximately 10 enlarged scales similar in size to gulars, but more than twice the size of adjacent fold scales.

Vertebral crest strongly projected and decreasing in size posteriorly, with vertebrals on neck at least four times higher than those between hind limbs; crest bifurcates posteriorly and extends onto tail less than ¼ its length; body flanks between fore and hind limbs with slight dorsolateral and ventrolateral longitudinal folds; scales on dorsolateral folds similar in size to adjacent scales; dorsal and flank scales small, keeled, imbricate, more or less homogeneous in size, and separated from each other by skin covered with tiny granular scales; ventral scales imbricate, smooth or slightly keeled, rectangular or rhomboid, with a posterolateral mucron; ventrals more than twice the area of dorsals.

Limb scales keeled dorsally and smooth or slightly keeled ventrally; scales on dorsal and posterior surfaces of thighs heterogeneous in size, with most scales less than half the size of those on anterior and ventral surfaces, separated from each other by skin covered with tiny granular scales; subdigitals on finger IV 17; subdigitals on toe IV 25; femoral pores on each side four; tail laterally compressed and gradually tapering posteriorly; caudal scales strongly keeled and imbricate, not gradually 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.

(Fig. 8). Head dark green with large black blotch between the eye and the tympanum; loreal region, nasal scale, labials and chin white; white blotch on posterior end of mandible; neck greenish brown dorsally and dark brown laterally, with a white rhomboidal blotch extending longitudinally from tympanum to scapular region; dorsal body background dark brown with scattered green scales and pale spots; limbs dark brown with green transverse bands; tail dark green with scattered dark brown marks; vertebral crest with intermixed green and dark brown scales; gular region white with a black posteromedial patch bearing scattered turquoise scales; chest grayish white with a turquoise tone anteriorly; belly grayish white with scattered, faint, pale brown blotches; ventral surface of limbs grayish white, with a longitudinal faint turquoise stripe along the thighs; tail grayish white; iris silver peripherally and dark brown centrally, with dark brown reticulations; silver ring around pupil.

Variation in meristic and morphometric characters of Enyalioides sophiarothschildae are presented in Table 2. One male paratype (MUSM 21883) differs from the holotype in having some scattered dark brown blotches on the throat.

Enyalioides sophiarothschildae is known from the northeastern slopes of the Cordillera Central in Peru between 1600–1700 m (Fig. 3). This species is only known from two adjacent localities, the trail to La Cueva-Añasco Pueblo in the drainage of the Lejía river and El Dorado in the drainage of the Blanco river, both tributaries of the Huallabamba river in the northern part of the Huallaga river basin. This area corresponds to the Selva Alta (400–1000 m) and Yungas (300–2300 m) ecoregions (Brack 1986; Peñaherrera del Aguila 1989).

Individuals of Enyalioides sophiarothschildae were found active by day in primary forest. The holotype was found crossing a trail and tried to hide between the roots of a big tree when approached for capture. One of the paratypes climbed up a tree three meters above the ground when approached. The other paratype was found sitting on a big root.

The specific epithet is a noun in the genitive case and is a patronym honoring Sophia Rothschild in recognition of her financial support for the improvement of the herpetological collection of CORBIDI through the BIOPAT Program.