Lepidoblepharis rufigularis, Batista, Abel, Ponce, Marcos, Vesely, Milan, Mebert, Konrad, Hertz, Andreas, Köhler, Gunther, Carrizo, Arcadio & Lotzkat, Sebastian, 2015

Batista, Abel, Ponce, Marcos, Vesely, Milan, Mebert, Konrad, Hertz, Andreas, Köhler, Gunther, Carrizo, Arcadio & Lotzkat, Sebastian, 2015, Revision of the genus Lepidoblepharis (Reptilia: Squamata: Sphaerodactylidae) in Central America, with the description of three new species, Zootaxa 3994 (2), pp. 187-221 : 197-201

Lepidoblepharis rufigularis sp. nov.

Figs. 3–5 , 11 .

Lepidoblepharis xanthostigma : all in part. (referring to populations in extreme southeastern Panama): Auth (1994); Young et al. (1999); Ibáñez et al. (2001); Köhler (2008); Jaramillo et al. (2010).

Holotype. Adult male SMF 50659, original field number AB 527 ( Figs. 3–5 , 11 ), collected on a hill 1 km north of Río Púcuro (8.057501°N, 77.370217°W, 1043 m elev.), Pinogana, Darién, Panama, on 0 8 July 2012 at 22:40 hrs by Abel Batista.

Diagnosis. Lepidoblepharis rufigularis (our sp. nov. 2) is characterized by the following combination of characters: (1) dorsal scales small, granular, and juxtaposed, ventral scales large, cycloid, flat, and imbricate; (2) scales on head small and granular; (3) four postrostral scales; (4) two short, barely discernible paramedian clefts in the more or less U-shaped posterior mental border; (5) five postmentals, the two median ones larger than the posteriorly adjacent chin scales; (6) 13 lamellae under fourth toe, 11 lamellae under fourth finger; (7) median subcaudals conspicuously wider than long, almost twice as wide as the laterally adjacent scales, with straight posterior margins arranged in a regular tail sequence of 1'1''; (8) ventral escutcheon consisting of 62 scales, almost twice as wide (13 scales) as long (7 scales); (9) subfemoral escutcheon consisting of 3–4 scales per thigh; (10) 17 longitudinal rows of ventral scales at midbody; (11) bilobate hemipenis; (12) SVL 25 mm.

Comparison with other species of the genus. Lepidoblepharis rufigularis can be differentiated from all species in the genus by its small size, number of lamellae under the fourth toe and finger, the reddish throat in males ( Fig. 11 ), and the configuration of the ventral escutcheon. In the following, we present comparisons to all other species within the genus, with the characteristics for L. rufigularis in parentheses. Lepidoblepharis rufigularis can be distinguished from the Panamanian species L. xanthostigma , L. sanctaemartae , L. emberawoundule , and Lepidoblepharis sp. nov. 3 (described below) by a genetic p-distance of 14–23% between individuals in 16S mtDNA. Lepidoblepharis xanthostigma is the most similar species, but has a different chin and throat coloration with dark reticulations on a pale background (orange background), and greatly enlarged median subcaudal scales which are more than two times as wide as the laterally adjacent subcaudal scales (enlarged but less than two times the width of laterally adjacent subcaudals, Fig. 3 ), usually 21 or more, very rarely 18, scales across snout (19), usually 18 or more, rarely 16 or 17, ventral scales a midbody (17), and an escutcheon long/wide ratio of 67–120% (54%). Lepidoblepharis emberawoundule , L. sanctaemartae , and Lepidoblepharis sp. nov. 3 (described below) have fewer than 10 lamellae under the fourth toe (13) and under the fourth finger (11). Additionally, L. sanctaemartae has large, flat, imbricate dorsal body scales (small, granular, and juxtaposed). To date, seven species of the genus Lepidoblepharis have been reported to possess ten or fewer lamellae under the fourth toe, i.e., to be short-toed ( L. miyatai , L. sanctaemartae , L. buchwaldi , L. montecanoensis , L. williamsi , L. peraccae , and L. microlepis ), and are therefore readily differentiable from the long-toed L. rufigularis (13 lamellae under the fourth toe). Of the remaining members of the genus, L. colombianus , L. conolepis , L. duolepis , L. festae , L. grandis , L. heyerorum , L. hoogmoedi , L. intermedius , and L. ruthveni are relatively to very large lizards for this genus with adult SVLs between 33 and 56 mm (25 mm). Additionally, L. conolepis and L. grandis have 14–20 lamellae under the fourth toe (13). The dorsal ground color in males of L. heyerorum is black with yellow dorsal markings (no yellow dorsal markings). The two long-toed specimens with granular dorsals reported as L. xanthostigma from Colombia by Ayala & Castro (1983) are similar to L. rufigularis in the number of ventral scales, but they have 22–25 scales across snout (19), an escutcheon with only 25 scales (62), the gular region with blotches (gular region with longitudinal bars), and an occipital pale W-shaped mark (no occipital mark at all).

Description of the holotype. Adult male as judged by everted hemipenes; SVL 25 mm, TL 33.0 mm (measured while the now broken tail was still intact), HL 5.6 mm, HW 4.1 mm, forelimbs 5.0 mm, hind limbs 9.3, shank 3.6 mm; rostral large, clearly visible from above, with a shallow, horseshoe-shaped posterior depression and a long median cleft; postrostrals four, including supranasals, one median postrostral slightly larger than posteriorly adjacent scales on snout and indenting the rostral; postnasals two, both about the same size as posteriorly adjacent loreal scales; scales on snout rounded and pointed backward; loreal scales subimbricate, elevated toward posterior and dorsal directions, six loreal scales on a longitudinal line between postnasals and orbit; 19 scales across snout between anterior sutures of second SPLs; scales on top of head small, granular, juxtaposed, generally pointing upward, about half the size of those on the middle area of snout; superciliary flap with two enlarged scales on anterior border, of which the first is slightly longer than the second, followed by 2–5 small and globular scales; supralabials three, posteriormost one below center of eye; ear-opening small, oval, in oblique orientation; mental large, posterior margin somewhat U-shaped, with two barely discernible, very short paramedian clefts; five enlarged postmentals, slightly larger than the posteriorly adjacent chin scales ( Fig. 3 B); scales on chin small, flat, rounded, and juxtaposed, on posterior region flat to flat-granular, slightly inclined backward; scales near posterior infralabials flat, subimbricate, and larger than scales in median area of chin; infralabials four, first largest and almost reaching anterior level of orbit, fourth below center of eye; throat with small granular scales, pointed and directed upward; dorsal scales on neck and body small, granular, and juxtaposed, at midbody and on posterior trunk pointed, granular, or, in frontal view, triangular, mostly directed posteriorly, some scales on flanks and in lumbar region pointed; dorsal skin on body too damaged to conduct longitudinal counts of middorsal scales; ventral scales flat, smooth, imbricate, with an ovoid posterior margin, increasing moderately in size from gular region to belly, posterior ventral scales longer than wide; 13 midventral scales in one HL, 23 between levels of axilla and groin, 28 to border of cloaca; ventral escutcheon patch with 62 scales, 7 long and 13 wide, escutcheon long/wide ratio 54%; subfemoral escutcheon with 3 scales in a single row under each thigh, and a barely discernible fourth scale in a second row under the right thigh; transition between ventrals and lateral scales abrupt; 17 longitudinal rows of ventrals at midbody; scales on precloacal plate similar to ventrals, except for those on border of cloaca, which are smaller; tail dorsally and laterally with flattened, smooth, and imbricate scales that are less elongate than ventrals (with a transitional zone at base of tail); underside of tail with a median row of moderately enlarged scales (usually no more than two times wider than the laterally adjacent subcaudal scales; Fig. 3 B), mostly with a repeated series of one median scale bordered laterodistally by one scale, followed by a slightly larger median scale in contact laterodistally with two scales, constituting a regular tail sequence of 1'1''; dorsal scales on forelimbs granular to conical; scales on hind limbs flat, smooth, and imbricate on anteroventral femoral and shank surfaces, granular elsewhere; fingers, from longest to shortest, IV-III-II-V-I; toes IV-III-II-V-I, fourth and third toes about the same length; 11 lamellae under fourth finger, 13 under fourth toe; claws enclosed by an ungual sheath composed of six scales, as typical for the genus.

Hemipenis morphology. The partially everted hemipenis of SMF 50659 ( Fig. 5 B) is a small organ; sulcus spermaticus bordered by well-developed and smooth sulcal lips; asulcate area of the truncus covered by papillate calyces; proximal portion of apex covered by small spinulate calyces. Due to its incomplete eversion, it is not possible to determinate whether the hemipenis is as bilobate as those of other species of the genus described and/or pictured herein ( Fig. 6 ). Nevertheless, it is apparent that the hemipenis of Lepidoblepharis rufigularis does not bear a conspicuous basal third lobule, in contrast to the other two species described herein.

Coloration in preservative (alcohol 70%). Dorsal ground color Hair Brown (277); posterior margin of orbit with a pale Beige (254) line; dorsum of head suffused with Beige (254); infra- and supralabials with alternating Sepia (279) and white bars; chin and throat with Sepia (286) reticulations on a dirty white background; venter pigmented with Vandyke Brown (281); escutcheon scales unpigmented in the center, with Olive Brown (278) borders.

Coloration in life ( Fig. 11 ). Dorsal ground color Grayish Horn Color (268), with patches of Lavender Blue (195) and Vandyke Brown (281) scales, same pattern on neck and head; a straight Light Sky Blue (191) postorbital line, running up to level of ear; infra- and supralabials with alternating Sepia (279) and Burnt Orange (10) bars; chin and throat with Sepia (279) reticulations on a Burnt Orange (10) background; ventral parts suffused with Dusky Brown (285) and Light Sky Blue (191); escutcheon scales Smoke Gray (267), with Dusky Brown (285) scale tips; tail with a dorsolateral Salmon Color (59) line; toes and fingers suffused with Brick Red (36).

Distribution and habitat. Lepidoblepharis rufigularis is known only from the type locality in southeastern Darién province. Given the close proximity of the type locality to Colombia, it likely occurs there as well. The habitat at the type locality of L. rufigularis is part of the eastern Panamanian montane forests ( Fund 2011), in the Darién mountain range, around 1000 m elev. Most probably, L. rufigularis lives in the leaf-litter and feeds on small invertebrates like other Lepidoblepharis spp. ( Vitt et al. 2005). The specimen probably resided between the leaf-litter on a large fallen log, and was uncovered and collected after one of the local supporters in the group stepped over the log.

Etymology. The name rufigularis is a compound word that comes from the Latin rufus (red) and gula (throat) referring to the bright orange throat color in this species in life.

Figures

FIGURE 3. Morphological distinction among Lepidoblepharis from Lower Central America; images from left to right: dorsals, ventral views of foot, tail, and mental region; scale bars equal 1 mm. (A) L. emberawoundule; (B) L. rufigularis; (C) L. sanctaemartae; (D) L victormartinezi; (E) L. xanthostigma.

FIGURE 4. Feet and hands of Panamanian Lepidoblepharis. Lepidoblepharis emberawoundule: right foot of SMF 81954 and right hand of MHCH 2957; L. rufigularis (SMF 50659): right foot and right hand; L. sanctaemartae: left foot of SMF 97419 and right hand of MHCH 2948; L. victormartinezi: left foot and left hand of SMF 89963; L. xanthostigma: left foot of SMF 90190 and left hand of SMF 89576. Scale bars equal 1 mm, tip of fourth finger and toe marked with an asterisk.

FIGURE 5. Escutcheon scales in males of Panamanian Lepidoblepharis. (A) L. emberawoundule: ventral escutcheon with 46 scales, 7 scales long x 11 wide, and subfemoral escutcheon with 4 scales per side, of SMF 81951 (SVL = 25 mm); (B) L. rufigularis: ventral escutcheon with 62 scales, 7 scales long x 13 wide, and subfemoral escutcheon with 3 and 4 scales per side, of SMF 50659 (SVL = 25 mm); (C) L. sanctaemartae: ventral escutcheon of SMF 97419 (SVL = 22 mm) with 92 scales, 10 scales long x 12 wide; (D) L. victormartinezi: ventral escutcheon of SMF 89963 (SVL = 27 mm) comprising 57 intact scales (an original total of 61 is assumed considering the symmetrical scale arrangement of the non-damaged portions), 7 scales long x 14 wide; (E – F) L. xanthostigma: (E) ventral escutcheon with 75 scales, 12 scales long x 11 wide, and subfemoral escutcheon with 2 scales per side (no additional scales hidden under strings), of SMF 90189 (SVL = 35 mm), and (F) ventral escutcheon with 36 scales, 6 scales long x 8 wide, and no subfemoral escutcheon of AB 1250 (SVL = 24 mm). All scale bars equal 1 mm.

FIGURE 6. Hemipenis morphology of four Lepidoblepharis species from Panama. Sulcate-asulcate view, (A – B) L. emberawoundule SMF 50968; (C – D) L. sanctaemartae SMF 97419 (E – F) L. victormartinezi SMF 50950; (G – H) L. xanthostigma SMF 91558. All scale bars equal 1 mm.

FIGURE 11. Holotype of Lepidoblepharis rufigularis (male SMF 50659) in life. (A – B) lateral view; (C) gular and chest region.