Lepidoblepharis nukak, Calderón-Espinosa, Martha Lucia & Medina-Rangel, Guido Fabian, 2016

Calderón-Espinosa, Martha Lucia & Medina-Rangel, Guido Fabian, 2016, A new Lepidoblepharis lizard (Squamata: Sphaerodactylidae) from the Colombian Guyana shield, Zootaxa 4067 (2), pp. 215-232 : 217-229

Lepidoblepharis nukak sp. nov.

( Fig. 2 ).

Holotype. ICN-R 12305. An adult male, 27.87 mm SVL, 29.76 mm TL, collected by the authors on 27th October 2012 in Puerta de Orión, vereda La Pizarra on the way from Bogotá to San José del Guaviare (2°30'38.42"N, 72°42'24.44"W; datum Bogotá WGS 84), Guaviare department, Colombia, at approximately 217 m elevation in tropical wet forest.

Paratypes. Twenty adult and five juvenile specimens collected from several localities near the type locality in Guaviare department, Colombia, at approximately 216 m elevation in tropical wet forest. These individuals correspond to five males (ICN-R 12303-04, 12310, 12359, 12466), one female (ICN-R 12516), and one juvenile individual (ICN-R 12515), 27.87 mm SVL, collected on 20-27th October 2012 (12303-4), April 16th and October 3rd 2013, and March 19th 2014 in Puentes Naturales, vereda El Retiro on the way to San José del Guaviare (2°32'24.55"N, 72°42'48.95"W); four males (ICN-R 12313, 12351, 12373, 12467), eight females (ICN-R 12306, 12309, 12329, 12463-65, 12374, 12514), and one juvenile (ICN-R 12360) collected in Puerta de Orion, Vereda El Retiro, (2°30'48.19"N, 72°44'33.95"W) on October 9th, 10 th and 27th 2012, April 14th 2013, and March 20th 2014; two juveniles (ICN-R 12311-12) from Ciudad de Piedra, Serranía La Lindosa (2°27'55.64"N, 72°42'10.96"W), collected on April 18th and October 9th 2013; one male (ICN-R 12366) and one juvenile (12307) from Playa Güío (2°34'24.70"N, 72°42'48.99"W) collected on October 9th 2012 and October 27th 2013; and one female (ICN-R 12468) from Cerro Azul (2°34'51.96"N, 72°49'51.85"W), collected on March 22–23rd 2014.

Diagnosis. A small Lepidoblepharis (adult size 27.8–29.3 mm SVL) with the following characteristics: 10–13 lamellae beneath the fourth toe; dorsal and lateral body scales granular or subconical at midpoint between levels of axilla and groin; escutcheon on belly and thighs present in adult males; snout short and rounded; mental large, convex or slightly convex (backward projected) at its posterior edge, which is also indented and followed by two to five (usually four) large anterior postmentals, 20–31 scales across snout at lst/2nd supralabial suture ( Fig. 3 ).

Comparison with other species. Lepidoblepharis nukak sp. nov. is distinct from other Lepidoblepharis , except from L. xanthostigma (Noble) , in having a posterior edge of mental convex or slightly convex; besides, it belongs to species with body size of less than 30 mm SVL (including L. buchwaldi Werner , L. microlepis (Noble) , L. miyatai Lamar , L. montecanoensis Markezich & Taphorn , L. peracae Boulenger and L. sanctaemartae (Ruthven)) ( Table 1).

From species recorded or potentially distributed in the Colombian Amazonia (Cis- Andean region) ( L. festae (Peracca) , L. heyerorum Vanzolini and L. hoogmoedi Avila-Pires ), L. nukak differs in the following characteristics (data for L. nukak sp. nov. is given first): by its smaller size (29.3 mm maximum SVL versus 42, 35 and 38 mm, respectively in the others species); presence of escutcheon in both abdomen and thighs versus only on the abdomen in the other three species; from L. festae in the size of postmental scales (larger than vs. same size as those further back), and number of subdigital lamellae (10–13 versus 14–15), and in the number of escutcheon scales on the abdomen (34–52 versus 60); from L. heyerorum by its number of ventral scales (first count) (36–45 versus 48–49 in L. heyerorum ), and finally, from L. hoogmoedi , in the number of scales around midbody (80–91 versus 92–93), in the number of escutcheon scales on the abdomen (34–52 versus 56), and it also differs by the absence of a light arc on posterior part of head.

Lepidoblepharis nukak sp. nov. differs from Andean and Trans Andean species of Lepidoblepharis ( L. buchwaldi , L. colombianus Mechler, L. conolepis Avila-Pires , L. duolepis Ayala & Castro , L. grandis Miyata , L. intermedius Boulenger , L. microlepis , L. miyatai , L. montecanoensis , L. oxycephalus (Werner) , L. peraccae , L. ruthveni Parker, L. sanctaemartae , L. williamsi Ayala & Serna, and L. xanthostigma ), in the following combination of characters: from L. sanctaemartae and L. miyatai , by its larger body size (29.3 vs. 25 and 23 mm SVL, respectively), having dorsal scales granular (approximately conical), versus flat, smooth and imbricate, and by its higher counts of scales around midbody ( Table 1); it further differs from L. miyatai by having fewer ventral scales (both counts), more dorsal scales between first supralabial suture, and larger postmental scales; and from L. sanctaemartae in having more lamellae and by the presence of escutcheon scales on the abdomen and thighs versus only on the abdomen; from L. ruthveni and L. duolepis by its homogeneous dorsal scales, versus heterogeneous in the other two, fewer ventral scales (first count, Table 1), and maximum body size (29.3 vs 32 mm in L. ruthveni and 46 mm in L. duolepis ); it further differs from L. ruthveni by having fewer lamellae, fewer ventral scales (second count), and fewer midventral scales; and from L. duolepis by having the escutcheon on abdomen and thighs versus only in the abdomen; from L. intermedius by fewer ventral scales (both counts, Table 1), presence of escutcheon on thighs and abdomen versus only on the abdomen, fewer escutcheon scales, and a shorter versus a very long and pointed snout in L. intermedius ; from L. conolepis and L. grandis by the shape and size of the dorsal scales (smaller and subconical versus larger and conical in the other two), its smaller body size (29.3 vs. 44 and 56 in the two species, respectively), fewer subdigital lamellae, fewer ventral scales (longitudinal row scales along a midventral line between anterior level of fore- and hind limbs), fewer midventral scales (Table 2), and more dorsal scales between first supralabial suture, and presence of escutcheon on thighs and abdomen versus only on the abdomen; from L. grandis in having fewer lamellae, postmental scales and loreal scales, and more dorsal scales between first supralabial suture; from L. conolepis in having fewer ventral scales (first count), fewer scales around midbody, and fewer escutcheon scales; from L. montecanoensis by its larger body size (29.3 vs 21 mm), the presence of escutcheon on thighs and abdomen versus only in the abdomen and fewer escutcheon scales; from L. oxycephalus by the size of the postmental scales (larger versus same size as posterior scales of chin); from L. xanthostigma , by its smaller body size (29.3 vs. 38 mm), number of ventrals (count 2; Table 1); from L. peraccae by having fewer ventral scales (first count), more subdigital lamellae which are not expanded, and by the presence of escutcheon on thighs and abdomen versus only on abdomen; from L. microlepis by the lower number of loreal scales; from L. emberawoundule Batista, Ponce, Vesely, Mebert, Hertz, Köhler, Carrizo & Lotzkat , L. rufigularis Batista, Ponce, Vesely, Mebert, Hertz, Köhler, Carrizo & Lotzkat , and L. victormartinezi Batista, Ponce, Vesely, Mebert, Hertz, Köhler, Carrizo & Lotzkat by its larger body size; from L. emberawoundule and L. victormartinezi by having more subdigital lamellae; from L. rufigularis and L. victormartinezi by fewer abdominal escutcheon scales, from L. victormartinezi by having escutcheon scales on abdomen and thighs versus only on abdomen; and from L. rufigularis by having more dorsal scales between first supralabial suture; and finally from L. microlepis and L. buchwaldi by its larger body size (29.3 vs. 23 and 27 mm, respectively).

Description of holotype. Rostral large and high, with a median cleft, central depression sharply indented; transverse posterior margin in contact with two large supranasals, separated at midline by one flat, pentagonal like scale; granules on top of snout polygonal, prominent, becoming gradually smaller and tubercular and granular (approximately conical) toward the posterior region of the head; 14 granular or subconical scales (viewed dorsally) in a transverse line between anterior notches of superciliary flaps; nine granules bordering posterior edge of postnasals from labial to labial ( Fig. 3 B); nostril indents rostral; two large postnasals, one pentagonal and one rectangular; four supralabials decreasing in size posteriorly, suture between third and fourth beneath center of eye ( Fig. 3 A); teeth short and conical; superciliary flap in a double line with three enlarged scales on the top side (second scale of the top side largest) ( Figs. 3 A, B); posterior half of the superciliary region covered by 10–12 small conical laterally projecting scales; mental large, nearly trapezoid, convex (posteriorly projected), transverse posterior margin bordered by three flat, enlarged scales; four infralabials, decreasing in size posteriorly; first infralabial large, nearly equal to first supralabial ( Fig. 3 C). A row of flat, slightly enlarged scales bordering infralabials; other gulars small and granular ( Fig. 3 C). Dorsal scales (from shoulder region to hind limb insertion) larger than those on head and neck, granular or subconical and juxtaposed; fairly homogeneous in shape, arrangement, and size; scales at base of tail large, smooth and flat ( Fig. 3 I). Transition between gulars and ventrals abrupt ( Fig. 2 D), at the midpoint between ear and shoulder. Ventrals flat, smooth, and imbricate, arranged in diagonal rows ( Fig. 3 H). Ventrals with rounded border, increasing in size posteriorly; escutcheon discernible on abdomen and thighs (40 scales on the abdomen, 6–9 on thighs) ( Figs. 3 and 4 ), scales of this region have angular borders, appear thicker, and the dark pigment concentrates on the border of each scale; 15 ventrals in diagonal row at midbody. Flat, smooth, imbricate scales cover dorsal surface of upper arm, forearm, and carpus, granules elsewhere; palm with flat scales becoming subconical towards the base of fingers ( Fig. 3 D); digits 3-4-2-5- 1 in size order; subdigital lamellae (from first to fifth finger): 5-7-9-9-7. Hind limb with flat, smooth, imbricate scales on anterior half of thigh, ventral aspect of thigh, tibia and dorsal surface of tarsus, cone-like or granular elsewhere; plantar surface covered with granules ( Fig. 3 F); toes 4-5-3-2- 1 in size order; subdigital lamellae (from first to fifth toe): 4-8-10-12-10. Approximately half of the tail is regenerated; dorsal scales on proximal portion of tail smooth, flat, imbricate and round to slightly diamond-shaped; ventrally lateral scales are smaller than central scales.

L. nukak sp. nov. L. heyerorum L. hoogmoedi L. festae ^

Geοgraphic Distributiοn Cοlοmbia, Guaviare department, 9‾11 Brazil and French Guiana. Brazil and Peru. Amazοnian, Ecuadοr, Peru and Brazil. Amazοnian Km. frοm San Jοs. del Guaviare. Amazοnian, between 50‾ 200 m. belοw 50 m. and Andean slοpe, belοw 2400 m. Guyana Shield, between 190‾ 230 m.

Dοrsal scales Hοmοgeneοus, small, larger than scales Hοmοgeneοus, small, slightly Hοmοgeneοus, small, slightly Hοmοgeneοus οr subequals, small, οn tοp οf head granular, ϳuxtapοsed. larger than scales οn tοp οf larger than scales οn tοp οf slightly smaller than scales οn tοp οf head, granular, ϳuxtapοsed. head, granular, ϳuxtapοsed. head, granular, smοοth οr cοnical, ϳuxtapοsed.

Fοurth tοe lamellae number 10‾13 11‾14 12‾14 14 ̶ 15

…… continued on the next page …… continued on the next page L. emberawoundule ^ L. grandis ^ L. intermedius L. microlepis ^

Geοgraphic Distributiοn Eastern Panama, 227‾ 773 m. Ecuadοr, Pichincha. Between 600‾ Cοlοmbia and Ecuadοr. Pacific Cοlοmbia. Pacific regiοn, belοw 200

1400 m. lοwlands, belοw 100 m. m.

Dοrsal scales Hοmοgeneοus, small, granular, and Hοmοgeneοus, granular, Hοmοgeneοus, larger than scales οn Hοmοgeneοus, tiny, slightly large ϳuxtapοsed, large than scales οn tοp (apprοximately cοnical, sligthly tοp οf the head, granular, ϳuxtapοsed. than scales οn tοp οf head, granular, οf head small and granular wider than high), large than scales strοngly keeled οr strοngly οn tοp οf head granular, sligthly tubercular, ϳuxtapοsed. larger tοward flanks, ϳuxtapοsed.

Fοurth tοe lamellae number 6̶9 16̶20 11‾18 (usually 12‾14) (expanded) 10 οr less.*

ventral scales (lοngitudinal rοw Nο data 43‾48 51̶59 Nο data

scales in a straight line between

thrοat tο pοsteriοr bοrder οf clοaca)

ventral scales (lοngitudinal rοw Nο data 36̶40 35̶43 Nο data

scales alοng a midventral line

between anteriοr level οf fοre‾and

‾limbs) ***

Midventral scales (diagοnal rοws at 16‾20 20̶22 18‾26 16 ‾ 20

midbοdy, between small lateral

granular scales)

Scales arοund midbοdy 69‾92 87‾92 66‾95 Nο data

Bοrder mental shape in its pοsteriοr Concave, edge a vaguely M̶ Concave, edge M̶shaped, with or Concave, edge M̶shaped, with Concave, edge M̶shaped, without

margin shaped, with two large without short clefts at one or both two large clefts. or with two clefts. paramedian clefts at both anterior anterior tips of the "M".

tips of the "M".

Pοstmental scales (size) Sligthly larger than thοse medial and Hardly larger than pοsteriοr scales First pοstmental scales larger than Enlarged scales οf which the median

pοsteriοr gular scales. οn chin, althοugh median οnes may thοse farther back. is the largest. be slightly larger.

Pοstmental scales(number) 3‾7 6̶9 3‾4 3 ‾ 5

Lοreal scales (number) 5‾7 11̶12 8‾10 (rarely 7) 10

Dοrsal scales between first 18‾21 14̶15 19‾24 Nο data

supralabial suture

Escutcheοn pοsitiοn Abdοmen and thigts Abdomen Abdomen Abdοmen and thigts

Escutcheοn scales (number in adult 38‾61⁄4‾5 30‾60 167̶275 (juvenile 47) 28‾110⁄3‾4 (abdοmen⁄thighs)

males)

max. (mm) 30 56 32 23

…… continued on the next page …… continued on the next page L. rufigularis ^ L. ruthveni L. sanctaemartae …… continued on the next page L. victormartinezi ^ L. williamsi L. xanthostigma Color in life. From color photographs and field notes; apparently there are two color patterns among adult males as well as among adult females and juveniles, one lighter than the other ( Fig. 2 ). Adult males dorsally Grayish Olive (274) or Smoke Grey (267) with small and dark irregular Jet Black (300) markings and very small Cyan White (155) stipples over head, back and tail, more conspicuous in some males; background dorsal color of females Raw Umber (22) or Sepia (279), with irregular Burnt Umber (48) or Jet Black (300) spots; Smoke White (261) stipples that also cover fore and hind limbs, stipples being more conspicuous in females than in males; dorsal surfaces of head and body of juveniles mottled with Smoke Gray (266) and Sepia (286), or Dusky Brown (285); head and neck region Dark Salmon Color (59) or Light Pratt´s Rufous (71) in males, Grayish Horn Color (268) or Burnt Umber (48) in females; mottled similar to head and dorsum in young individuals; females with a stippled pattern on the head, males with a more homogeneous background; a dorsolateral series of Medium Chrome Orange (75) splotches or Drab (19) continuous line Black Jet (300) bordered in young individuals (similar to that found in some adult females, see below); a Drab (19) line from posterior border of eye to base of tail in some adult males and females; this line appears well defined and bordered by Black Jet (300) in most females; this dorsolateral line, when present, ends with a transversal line at the base of the tail (“H” shape) ( Fig. 2 ). These dorsolateral lines become wider and are Robin Rufous (29) in females, and Raw Umber (23) in males from the base of the tail towards its tip; tail with Medium Chrome Orange (75) and Dusky Brown (285) in juveniles. Lateral side of the head and body and back with small Cyan White (155) or white stipples; stipples being more abundant in females; mottled or as their back in young individuals; posterior lateral side of thighs with irregular and large ocelli, the one closest to the base of the tail largest, Light Yellow Ocher (13) colored. Lips strongly barred (mainly those labials below the eye) with Dusky Brown (285) and Smoky White (261) in some females and juveniles; barred is faint in other males and females. Throat in males Medium Chrome Orange (75). Gular region of some females and juveniles with or without Dusky Brown (285) bars, originating from the infralabial scales and extending onto the neck; gular region of some females diffusely mottled with Dusky Brown (285) and Smoky White (261). Venter Smoke Gray (266), lighter toward chest. Palmar surfaces Hair Brown (277), darker than venter; iris Raw Umber (23) with Orange Yellow (8) line bordering the pupil.

Color in preservative. The general pattern similar to specimens in life; background dorsal color is dark brown; orange color from lateral side of neck, dorsal, lateral and ventral surface of the head is lost; gular region turns cream greyish in those specimens. Whitish tiny dots that cover most of the body surface are hardly noted. Palmar surfaces are dark grey.

Variation. Variation of the type series in meristic characteristics and body size is presented in Table 1. Color variation was observed between males and females, between young and adult specimens, and among adult males as well as among adult females, and is described in color description section.

Distribution and natural history. Lepidoblepharis nukak sp. nov. is known only from the type locality and neighboring localities in the Guaviare department, in remnants of tropical wet forest. The area is part of a transition zone between Amazonian forests and Orinoquia grassland vegetation ( Prance 1996; Huber 2006). Annual average monthly temperature is around 34 C° and average monthly rainfall around 201 mm (Rangel-Ch. & Aguilar-P. 1987). All specimens were collected during the day in leaf litter on the forest floor, below flat stones, or within dead bracts of palm trees, and on vertical surfaces of outcropping rocks covered with tree roots, which were used as refuges by some individuals; those microhabitats exhibited particularly high humidity conditions ( Fig. 5 ). One gravid female collected in April 2013 (ICN-R 12329), and three during March 2014 (ICN-R 12465, 12468, 12514), each had one large egg, while juvenile individuals were collected in April 2013 and October 2014, suggesting probably an extended reproductive activity of females.

Etymology. The name of this new taxon is given in honor of the nomadic Amazonian people Nukak . This word means: the true people or people that speak the Nukak language, and is how these people are selfdenominated.

Geographic distribution. Lepidoblepharis nukak sp. nov. is known only from localities in the vicinity of San José del Guaviare, in the Guaviare department, Colombia ( Fig. 1 ).

Figures

FIGURE 1. Map of distribution of Lepidoblepharis nukak sp. nov. in Colombia; red star corresponds to the holotype ICN-R 12305 locality, solid black square corresponds to the collection localities of others specimens. INEGI. © 2015 Google – Map (QGIS Development Team 2015).

FIGURE 2. Adult male (A—ICN-R 12305 and B) and female (C—ICN-R 12306 and D—ICN-R 12464), juvenile (E—ICN-R 12515 and F—ICN-R 12312 specimens of Lepidoblepharis nukak sp. nov., and gular region of males (G—ICN-R 12305) and females (H—ICN-R 12306).

FIGURE 3. Details of head, body and limbs of the holotype of Lepidoblepharis nukak sp. nov. ICN-R 12305. Lateral (A), dorsal (B) and ventral (C) view of head; ventral surface of forelimb (D) and hind limb (F); detail of dorsal view of fourth finger (E) and fourth toe (G); ventral view of abdomen and thighs (H) and tail (I).

FIGURE 4. Scales of belly and thighs showing escutcheon area of male Lepidoblepharis nukak sp. nov. ICN-R 12313.

FIGURE 5. Habitats of Lepidoblepharis nukak sp. nov. near to San José del Guaviare in Guaviare department. Tropical rainforest in the foothills of Cerro Azul, Serranía la Lindosa (A); savannah vegetation between the rocky formations of Ciudad de Piedra (B); vegetation between rocky outcrops in Puerta de Orión (C); vegetation at the base of the rocks where the species is usually found (D); and detail of microsites occupied by lizards found on the outcrops (E).