Cnemidophorus ruatanus, Mccranie, James R. & Hedges, S. Blair, 2013

Mccranie, James R. & Hedges, S. Blair, 2013, A review of the Cnemidophorus lemniscatus group in Central America (Squamata: Teiidae), with comments on other species in the group, Zootaxa 3722 (3), pp. 301-316 : 304-308

Cnemidophorus ruatanus new combination

( Figs. 2 , 3 )

Cnemidophorus lemniscatus ruatanus Barbour 1928: 60 .

Cnemidophorus lemniscatus lemniscatus: Burt 1931: 30 (part). Cnemidophorus lemniscatus: Echternacht 1968: 152 (part). Cnemidophorus lemniscatus ruatanus: Wright 1993: 79 .

Cnemidophorus lemniscatus: Montgomery et al. 2007: 38 .

Holotype. MCZ 26759 from Coxen Hole, Isla de Roatán, Islas de la Bahía, Honduras.

Geographic distribution. Cnemidophorus ruatanus is known from Isla de Roatán, Isla de Utila, and Cayo Cochino Pequeño in the Islas de la Bahía, Honduras, and from the Río Motagua Valley and adjacent coastal plain of Guatemala eastward across northern Honduras to extreme northeastern Nicaragua. This species is largely a coastal form on the mainland, but does penetrate inland along some river valleys (i.e., Río Motagua in Guatemala into a tributary in adjacent Copán, Honduras; Río Viejo in Atlántida, Honduras; Fig. 4 ). The record of this species from southeastern Belize reported by Stafford & Meyer (1999) might represent a recent range expansion, or an introduction as thought by Stafford et al. (2010).

Diagnosis. Cnemidophorus ruatanus can be distinguished from other northern populations of the C. lemniscatus species group as follows: from C. l. espeuti of Islas de Providencia and San Andrés, Colombia, by having a single and distinct vertebral stripe, 27–32 subdigital lamellae on the fourth toe, and 14–17 subdigital lamellae on the fourth finger (versus vertebral stripe split, 33–35 subdigital lamellae on the fourth toe, and 17–19 subdigital lamellae on the fourth finger in C. l. espeuti ); from C. l. gaigei in having 11–13 in 31, 10 in 5, or 9 in 19 scales surrounding the three enlarged precloacal plates, and in those having only 9 scales the precloacal pair are completely separated by a single scale ( Fig. 5A ) in 15 or separated by distance of one-third to over one-half in 4 (versus 9–10 scales surrounding the three enlarged precloacal plates in all eight examined and the precloacal pair in broad contact, without a smaller interscale in 6 ( Fig. 5 B) and a small interscale not separating precloacal pair by more than one-third in 2 in C. l. gaigei [see Discussion below]). Burt (1931) synonymized C. ruatanus with C. l. lemniscatus , but the latter differs most notably from C. ruatanus in having a split or divided vertebral stripe (versus vertebral stripe single in C. ruatanus ). Cnemidophorus ruatanus also differs from C. l. lemniscatus in amount of molecular divergence ( Fig. 1 ).

Description. The following is based on 21 males (FMNH 283560–61, 283563–64, 283568; KU 192621, 203160, 220101; USNM 563589–90, 563593–94, 563596, 563598–99, 570397, 570399, 570401, 573206, 573209, 580929) and 14 females (FMNH 283562, 283566; KU 101334, 101340, 192622; USNM 563588, 563592, 563595, 563597, 573207–08, 573211–12, 580928). Cnemidophorus ruatanus is a moderately large lizard (maximum recorded SVL 113 mm [FMNH 283561]); dorsal head scales enlarged, smooth, platelike, with paired prenasal scales with short median sutures, a single frontonasal, paired prefrontals, a single frontal, paired frontoparietals, two pairs of parietal scales (occasionally one or two parietals can be partially divided), and a single interparietal scale; rostral not contacting frontonasal; postnasal not contacting prefrontal; prefrontal and first superciliary usually in contact; frontal ridge absent; posterior frontal suture usually aligned near midlength of supraocular 3; scales in frontoparietal region smooth, flat; interparietal varies from slightly narrower than, to slightly broader than, flanking parietals; 1–6 (2.8 ± 1.1) scales between fourth supraocular and inner-most parietal on each side; 1–6 (4.0 ± 1.0) scales between fourth supraocular and outer-most parietal on 34 sides; usually 4 supraoculars per side (occasionally a fifth smaller supraocular present posteriorly, rarely only 3 supraoculars present); median pair of occipital scales distinctly enlarged, much larger than first dorsal scale row; 17–28 (21.4 ± 2.5) occipitals (occipitals of Harvey et al. 2012 plus scales bordering frontoparietals); supratemporals moderately enlarged, separated from parietals; short rostral groove present or absent; nostril centered in nasal suture; nostril opening oval; first supraocular contacting second supraocular; 1.0 to 1.5 rows of lateral supraocular granules, 21–52 (29.0 ± 6.4) granules on each side; anterior extent of circumorbital semicircles in single row reaches between midlength and posterior third of supraocular 3, that of double row not reaching seam between supraoculars 3–4; 2–9 (4.2 ± 1.4) circumorbitals; 5–6 (usually 6) superciliaries per side, usually with second (occasionally third) elongated; 4 (rarely 5) suboculars per side, first on each side usually entire (occasionally divided), lower edge (when entire) or lower part (when divided) usually contacting supralabial 3 (rarely contacting both supralabials 3 and 4); subocular keel present; patch of enlarged scales located in front of auditory meatus; no auricular or preauricular flaps or folds; 1–1 loreals; usually 6 (rarely 5 or 7) supralabials and 5 (occasionally 6) infralabials per side; first supralabial straight ventrally, usually longer than second supralabial; lingual sheath absent; moveable eyelids present; pupil rounded; first pair of chinshields contacting infralabials, those chinshields separated only at posterior edge; intergular sulcus absent; 18–26 (22.2 ± 2.2) anterior gulars; 9–18 (14.0 ± 2.5) posterior gulars; gular patch of distinctly enlarged scales absent; intertympanic sulcus absent; sharp transition from anterior gulars to smaller posterior ones; mesoptychial scales moderately enlarged, bordered anteriorly by sharp transition to small scales; edge of gular fold not serrated; dorsal scales conical, 190–230 (212.0 ± 9.0) middorsals between occipitals and first enlarged caudal scale; 97–123 (106.9 ± 6.0) granules around midbody; middorsal scales subequal to lateral scales; chest scales large, flat; pectoral sulcus absent; ventral body scales large, platelike, squarish, juxtaposed, smooth, in 25–33 (28.9 ± 2.6) longitudinal (long axis of body) rows, in 8 transverse rows at midbody; scales immediately lateral to outside ventral plate on each side small and granular; paired enlarged terminal scales forming precloacal plate; 9–13 (10.6 ± 1.4) scales bordering all three enlarged plates in 55 (those listed above plus FMNH 283565, 283567; KU 101328 –30, 101332 –33, 101339, 101341 –42, 101344 –47, 101349 –50; USNM 69397–400); males usually with one smaller subtriangular scale flanking paired terminal plates and 1–2 other smaller rounded scales between precloacal plate and small precloacal spur; 4–6 (4.8 ± 0.6) precloacal scales; postcloacal buttons absent; pair of slightly enlarged postcloacal scales present in males, those enlarged scales absent in females; caudal annuli complete, tail lacking crests or dorsolateral row of serrated scales; proximal subcaudals keeled; preaxial and postaxial brachial scales separated by band of continuous moderately enlarged scales; large dorsal brachial and antebrachial scales 1.0–1.5 times as wide as long, both sets of enlarged scales smooth, extending well beyond centers of arm; postaxial brachial scales in continuous enlarged row with preaxial brachial scales; postaxial antebrachial scales slightly enlarged; 29–34 (31.4 ± 1.2) combined subdigital lamellae on fourth finger in 20; subarticular lamellae of Fingers III–IV homogeneous in size, entire; 54–64 (59.7 ± 2.3) subdigital lamellae on fourth toe in 19, distal ones smooth; subarticular lamellae of Toes III–IV divided, each scale smaller than other lamellae; no row of distinctly enlarged scales between Toes IV–V; small scales separating supradigital scales from subdigital lamellae continuous, or nearly continuous; denticulate fringe absent along postaxial edge of outer toes; fifth toe not reduced, claw extending beyond level of articulation of Toes III–IV; 5–11 (7.4 ± 1.4) prefemoral scales on left side; heels without expanded scales; tibiotarsal shields present; tibiotarsal spurs absent; 38–47 (42.4 ± 2.4) total femoral-abdominal pores in males, 33–46 (39.7 ± 3.5) in females; no gap between femoral and abdominal pores; 3–4 (most often 3) scales separating each femoral-abdominal pore series; SVL 61.7–113.4 (79.6 ± 12.2) mm in males, 60.3–75.7 (67.0 ± 4.1) mm in females; TAL/SVL 1.60–2.79 in 13 males, 1.70–2.36 in eight females; HL/ SVL 0.23–0.27 in males, 0.21–0.27 in females; HW/SVL 0.13–0.16 in males, 0.12–0.18 in females; HD/SVL 0.13–0.17 in males, 0.11–0.15 in females; HW/HD 0.93–1.13 in males, 1.04–1.23 in females; SL/SVL 0.09–0.11 in males, 0.09–0.12 in females; SHL/SVL 0.19–0.23 in males, 0.18–0.22 in females; foot length/SVL 0.29–0.37 in males, 0.30–0.46 in females; hand length/SVL 0.11–0.16 in males, 0.12–0.16 in females.

Color in life of an adult male (FMNH 283564): middorsal longitudinal band Ground Cinnamon (239) with two dark brown stripes in dorsolateral field of band; middorsal band bordered by Dark Brownish Olive (129) stripe (about six granules wide), that dark stripe bordered below by narrow Chartreuse (158) dorsolateral stripe; lateral surface of body greenish yellow with scattered brownish green mottling; top of head Emerald Green (163) along outer edges, becoming brown with green tinge medially; anterior surfaces of fore- and hind limbs Opaline Green (162D); ventral surfaces of head, throat, and chest Robin’s Egg Blue (93); belly white with pale blue tinge; outer two ventral plates on each side Emerald Green; ventral surfaces of limbs and subcaudal surface pale blue, except palms and soles Robin’s Egg Blue. Color in life of another adult male (KU 220101): dorsal surface of head tan; lateral surface of head yellow-green, grading to turquoise blue on lips; outer edge of supraoculars outlined by yellow-green line; middorsal band tan, flanked by dark brown stripe; lateral surfaces yellowish green on neck and near groin, pinkish tan between neck and groin with yellow to white spots; dorsal surfaces of forelimbs olive green; dorsal surfaces of hind limbs tan with white spots; dorsal surface of tail tan; chin turquoise blue; chest pale turquoise blue; belly green with bluish cast; ventral surface of forelimbs white; ventral surface of hind limbs turquoise blue. Color in life of an adult female (UF 150291): middorsum of body Drab-Gray (119D) with slightly paler paramiddorsal stripes; dorsolateral field Hair Brown (119A) bounded by pale tan stripes; lateral and ventrolateral fields Chartreuse (158) bounded by pale tan to white stripes; venter Light Sky Blue (116D) grading to pale bluish green laterally; dorsum of head Grayish Horn Color (91); limbs grayish brown with white spots dorsally, pale grayish blue ventrally; tail Drab-Gray dorsally, pale bluish green ventrally.

Color in alcohol: juveniles have 8 longitudinal white to dirty white stripes dorsally and laterally on a body that is dark brown laterally and slightly paler brown middorsally; adults and juveniles of both sexes have a complete, pale brown middorsal swath extending from posterior edge of head to base of tail; adult females have that middorsal swath bordered laterally by thin paler brown to cream stripe that is in turn bordered below by a broad dark brown stripe that extends to posterior end of body, last mentioned stripe also bordered below by a cream to pale brown, thin stripe, with area below that thin stripe pale brown with or without indistinct cream stripe or cream to pale yellow small spots in adult females; adult males have middorsal swath with or without evidence of paler brown, thin border stripe, but with a dark brown broader border stripe (as in females) extending to posterior end of body, with area below that dark brown border stripe with cream to pale yellow spots in males; ventral surfaces of head and body cream to white with pale bluish gray tinge, or entirely black, except throat region pale blue to pale gray in males. Adults of both sexes from the Bay Islands have pale blue ventral surfaces, whereas those from the mainland can have either pale blue or black ventral surfaces. Adults of both sexes from Isla de Roatán retain the pale stripes to a larger size than do those from the remaining Honduran populations.

Habitat. Cnemidophorus ruatanus occurs at low elevations (sea level to 400 m elevation) in the Lowland Moist Forest and Lowland Dry Forest formations of Holdridge (1967). This is a sun loving species that occurs in open areas, such as beach vegetation and along open, gravely or sandy riverbanks. Both sexes exhibit the habit of pausing while moving about to lift and shake one of its forelegs. It has been collected or seen active during every month of the year in Honduras. Montgomery et al. (2007, 2011) provided studies on the habitats and natural history of a population of C. ruatanus on Cayo Cochino Pequeño, Honduras.

Echternacht (1968) compared a series of Cnemidophorus ruatanus from the Honduran mainland to a series of Cnemidophorus lemniscatus group specimens from El Real, Panama. Echternacht (1968) noted that the El Real specimens frequently had ten ventral plates across the venter at the level of the fifteenth ventral plate. Examination of a series of 15 of the El Real population revealed that all 15 have ten transverse (across the venter) ventrals at the level of the fifteenth ventral or at one or two ventral rows posterior to that point. The Panamanian C. lemniscatus group populations are also the only populations of the genus Cnemidophorus (sensu Harvey et al. 2012) known from the Pacific versant. We herein describe the Panamanian population as a new species of Cnemidophorus .

Figures

FIGURE 1. Phylogenetic tree of lizards of the Cnemidophorus lemniscatus complex from a maximum-likelihood analysis of DNA sequences of three mitochondrial genes (12 S rRNA, 16 S rRNA, and cytochrome b). A scale bar is indicated. The numbers at nodes are Bayesian posterior probabilities (left) and bootstrap (right) support values. The tree is rooted with the teiid lizard Kentropyx borkiana (not shown).

FIGURE 2. Adult male (FMNH 283564) of Cnemidophorus ruatanus comb. nov. (SVL 76.6 mm).

FIGURE 3. Adult female (FMNH 283567) of Cnemidophorus ruatanus comb. nov. (SVL 62.4 mm).

FIGURE 4. Map showing the known localities (circles) for Cnemidophorus ruatanus com. nov. The squares represent localities from which tissue samples were also sequenced.

FIGURE 5 A. Cloacal plate region of Cnemidophorus ruatanus comb. nov. (KU 101329) showing the small scale separating the enlarged terminal precloacal plates; 5 B Cloacal plate region of Cnemidophorus gaigei comb. nov. (USNM 117471) showing the enlarged terminal precloacal plates in broad contact medially.