Enyalioides heterolepis ( Bocourt 1874 )

Torres-Carvajal, Omar, Etheridge, Richard & Queiroz, Kevin De, 2011, A systematic revision of Neotropical lizards in the clade Hoplocercinae (Squamata: Iguania), Zootaxa 2752, pp. 1-44 : 13-19

Enyalioides heterolepis ( Bocourt 1874)

Proposed standard English name: spiny woodlizards

Proposed standard Spanish name: lagartijas de palo espinosas

Enyalus heterolepis Bocourt (1874:1) . Holotype: MHNP 4067, from “Veragua [ Panama; ( Guibé 1954)].”

Enyalius heterolepis Günther (1885-1902:56).

Enyalioides heterolepis Boulenger (1885:114) ; Burt & Burt (1930:9; 1931:265; 1933:23); Peters & Donoso-Barros (1970:114).

Enyalioides insulae Barbour in Bangs et al. (1905:100). Syntypes: MCZ 6983, 167581, from “Gorgona Island [Departamento Cauca], Colombia,” synonymy fide Parker (1926:550).

Enyalioides Mocquardi Despax (1911:10) . Syntypes: MHNP 06-226–228, from “l’Équateur,” synonymy fide Burt and Burt (1931:265).

Diagnosis. This species can be distinguished from other species of Enyalioides by having scattered, projecting, tetrahedral large scales on dorsum, flanks, and hind limbs, which are conspicuous in juveniles and adults of both sexes. The only other species of Enyalioides with scattered, projecting dorsal scales is E. cofanorum , which differs from E. heterolepis in lacking projecting scales on the hind limbs.

Description. (1) dorsal head scales conical or multicarinate, strongly projecting dorsally; (2) posterior superciliaries not enlarged relative to adjacent scales; (3) scales on lateral edge of skull roof just posterior to superciliaries sometimes (55.6%) slightly more projecting than adjacent scales; (4) all pretympanic scales similar in size; (5) gular scales conical or multicarinate, strongly projecting ventrally; (6) dorsal neck scales projecting, heterogeneous in size, largest scales tetrahedral; lateral neck scales similar in size to smallest dorsal neck scales; (7) vertebrals larger than adjacent dorsals, forming distinct raised middorsal crest that extends onto tail as a pair of crests; (8) nuchal region with continuous and sometimes (16.7%) paired middorsal crest; (9) dorsals distinctly keeled and heterogeneous in size, with largest scales similar in size to vertebrals; (10) continuous longitudinal row of raised, enlarged scales between dorsals and flank scales present; (11) scales on flanks keeled and usually (83.3%) heterogeneous in size, with largest scales similar in size and shape to vertebrals; (12) ventrals usually (66.7%) smooth; (13) fore limb scales keeled dorsally, smooth or slightly keeled ventrally; (14) hind limb scales keeled dorsally, smooth or slightly keeled ventrally; scattered enlarged scales present dorsally; dorsal scales of pes heterogeneous in size; (15) caudals heterogeneous, increasing in size posteriorly on each segment (6–8 scales in lateral view), not modified as conspicuous spines ( Fig. 3 ); (16) tail compressed laterally. Meristic and morphometric characters are presented in Table 1 .

Coloration in life ( Fig. 4 ). Juveniles and adults of both sexes of Enyalioides heterolepis have a distinct black or dark brown triangular mark or anterodorsally oriented stripe between commisure of mouth and ventral margin of eye.

Adult females (KU 76052, 113494, 169853): dorsal background reddish brown or olive green, sometimes with olive grey or green mottling laterally, or reddish-brown suffusion; cream spot ventral or posterior to ear sometimes present; venter creamy tan with reddish-brown suffusion laterally, or reddish brown with cream flecks; throat grey with brown streaks or black spot; iris reddish brown with yellow ring around pupil (C.W. Myers [21 July 1963] and W.E. Duellman [10 June 1975] field notes).

Adult males (KU 76048, 76051, 113490, QCAZ 8654): dorsal background brown, reddish brown, or yellowish brown, with darker lines forming a reticulate pattern; scattered pale blue spots on body sometimes present; head yellowish green, reddish brown, or yellowish brown; labials golden yellow or greenish yellow, sometimes followed by an orange streak posterior to mouth commisure; orange or white spot behind ear sometimes present; flanks similar to dorsum, or with greenish background; chin yellow or cream; light or dark brown vertical stripe from fore limb insertion to scapular region (sometimes continuous middorsally) sometimes present; green or blue flecks on dorsal aspect of limbs sometimes present; gular region yellow (sometimes streaked with brown), with black medial mark posteriorly; venter dull yellow or brown laterally and blackish medially, or creamy tan with pinkish brown mottling; ventral surface of limbs greenish; tail similar to dorsum, or bluish grey dorsally and pale brown ventrally, sometimes with blue flecks; iris brown with yellow ring around pupil, sometimes with grayish-white area dorsally; tongue flesh white with dark grey tip (C.W. Myers [21 July 1963, 17–19 April 1967] and color photographs).

Juveniles (KU 76053, 96688, 146657, 164166): dorsal background brown or reddish brown, with brown or reddish brown transverse bars; light brown V-shaped mark behind head sometimes present; flanks olive tan or light green, with brown spots; yellow or yellowish-green vertical bar from fore limb insertion to vertebral crest; head olive brown; cream spot behind ear sometimes present; gular region and belly cream, pale green, or dull yellow, with brown or red streaks and light brown or reddish flecks, respectively; black gular patch; iris brown or reddish brown with yellow ring around pupil (C.W. Myers [10 September 1964] and W.E. Duellman [10 April 1972, 28 March 1975] field notes and photographs).

Natural history. Most adult specimens have been found sleeping on tree trunks at night 20–170 cm above ground, heads-up if in vertical position, sometimes next to streams. Some juveniles and adults were collected on forest floor or under logs by day. A female (SVL = 89 mm; QCAZ 2025) collected in March 1990 in Provincia Pichincha, Ecuador, had two enlarged vitellogenic follicles on each side (4.43–8.75 mm X 5.23–9.25 mm). This species can be found in primary forest, secondary forest, or plantations.

Distribution. Enyalioides heterolepis occurs at elevations between 0–1000 m in Panama, the western slopes of the Andes and adjacent Pacific lowlands of Colombia and Ecuador, and Gorgona island, Colombia ( Fig. 6 ). This species is known to occur in sympatry with E. oshaughnessyi in northwestern Ecuador and Morunasaurus groi in northwestern Colombia. E. heterolepis also occurs close to E. oshaughnessyi in southwestern Colombia and Morunasaurus groi in Panama.

Remarks. Torres-Carvajal & de Queiroz (2009) found strong support (bootstrap value = 100) for the monophyly of E. heterolepis in a maximum likelihood analysis of mitochondrial DNA sequence data that included two specimens from Panama and one specimen from Ecuador. Sequences of the two individuals from Panama were more similar to each other (maximum-likelihood corrected distance = 0.025) than they were to the individual from Ecuador (0.063 and 0.068).

Figures

FIGURE 3. Caudal segments from the anterior third of the tail of ten species of Hoplocercinae in lateral view. (A) Enyalioides cofanorum, (B) E. heterolepis, (C) E. laticeps, (D) E. microlepis, (E) E. oshaughnessyi, (F) E. palpebralis, (G) E. praestabilis, (H) E. rubrigularis, (I) E. touzeti, and (J) Morunasaurus annularis. Scale bars = 5 mm. Illustration by D. Paucar.

FIGURE 4. Photographs of live lizards representing twelve species of Hoplocercinae.

FIGURE 6. Distribution of Enyalioides heterolepis (triangles) and E. laticeps (dots).