Cnemaspis heteropholis Bauer, 2002

Khandekar, Akshay, Thackeray, Tejas, Pal, Saunak & Agarwal, Ishan, 2020, A new large-bodied, rupicolous Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) allied to Cnemaspis heteropholis Bauer, 2002 from the Central Western Ghats of Karnataka, India, Zootaxa 4801 (1), pp. 57-84 : 62-67

publication ID	https://doi.org/ 10.11646/zootaxa.4801.1.2
publication LSID	lsid:zoobank.org:pub:0E8B717D-FFBA-406D-ABB9-8C285536B7AC
persistent identifier	https://treatment.plazi.org/id/04071322-090C-FFBE-1E9A-E46DFC91FCFE
treatment provided by	Plazi
scientific name	Cnemaspis heteropholis Bauer, 2002
status

Treatment

( Figures 2–4 View FIGURE 2 View FIGURE 3 View FIGURE 4 ; 5A, B View FIGURE 5 ; 6A View FIGURE 6 ; Tables 3 View TABLE 3 , 4)

Holotype. R06158 ( ZMH), SVL 40.2 mm, adult female, from Gund (15.167° N, 74.667° E), Uttar Kannada District , Karnataka, India collected by G. A. Von Maydell on 20 January 1956, during German Indian Expedition of 1955–1958. View Materials GoogleMaps

Additional material. CES L692 , SVL 33.4 mm, adult male from Gund (15.089°N, 74.524°E; ca. 450 m asl.) GoogleMaps and CES L802 , SVL 31.3 mm, sub-adult male, from near Castlerock (15.424°N, 74.325°E ca. 530 m asl.), Uttar Kannada District, Karnataka , India collected by Saunak Pal and Varun Torsekar on 27 th August and 4 th September 2012 respectively GoogleMaps ; CES L693 , SVL 38.5 mm, adult female, from Ram Dongar (15.014°N, 74.234°E ca. 520 m asl.) GoogleMaps and CES L700 , SVL 41.9 mm, adult female, from near Verlem (15.051°N 74.252°E; ca. 580 m asl.), South Goa District, Goa, collected by Saunak Pal and Varun Torsekar on 29 th and 31 st August 2012 respectively GoogleMaps ; CES L864 , SVL 44.4 mm, adult female, from near Ulavi , (15.000°N, 74.524°E; ca. 490 m asl.) Uttar Kannada District, Karnataka, India, collected by Saunak Pal on 24 th November 2013 GoogleMaps .

Diagnosis. A medium-sized Cnemaspis , snout-vent length upto 45 mm; dorsal scales on trunk heterogeneous, small, weakly keeled, oval, granular scales intermixed with enlarged, irregularly arranged, strongly keeled, conical tubercles; tubercles on dorsolateral aspect of flank more pronounced and conical than those on paravertebral region; spine-like scales absent on flanks. Ventral scales on belly smooth, subimbricate, 20–25 scales across mid-body, 119–123 longitudinal scales between mental to anterior border of cloaca. Subdigital scansors smooth, entire except for three or four proximal and 1–3 distal scansors on digit I–V of both manus and pes which are divided, unnotched; 20–22 lamellae under digit IV of manus, and 24 or 25 lamellae under digit IV of pes. Males with five or six femoral pores on each thigh, separated by 16–18 poreless scales; precloacal pores absent; dorsal pholidosis of tail homogeneous, composed of small, smooth, flattened, roughly circular, regularly arranged, subimbricate scales; without whorls of enlarge tubercles; median row of sub-caudals smooth, enlarged; enlarged median row is irregularly arranged with a large scale alternating with two slightly smaller scales on the posterior portion of the tail; the larger scales on the anterior portion divided; postcloacal spur small, indistinct, not conical.

Comparison with Indian congeners. Cnemaspis heteropholis can be distinguished from all other Indian congeners on the basis of the following differing or non-overlapping characters: a medium-sized Cnemaspis SVL upto 45 mm (versus small-sized Cnemaspis SVL <37 mm in C. andersonii (Annandale) , C. aaronbaueri Sayyed, Grismer, Campbell & Dileepkumar , C. adii Srinivasulu, Kumar & Srinivasulu , C. agarwali Khandekar , C. ajijae Sayyed, Pyron & Dileepkumar , C. amba, Khandekar, Thackeray & Agarwal , C. amboliensis Sayyed, Pyron & Dileepkumar , C. australis Manamendra-Arachchi, Batuwita & Pethiyagoda , C. assamensis Das & Sengupta , C. flaviventralis Sayyed, Pyron & Dahanukar , C. gracilis (Beddome) , C. girii Mirza, Pal, Bhosale & Sanap , C. goaensis Sharma , C. indica (Gray) , C. koynaensis Khandekar, Thackeray & Agarwal , C. limayei Sayyed, Pyron & Dileepkumar , C. littoralis (Jerdon) , C. mahabali Sayyed, Pyron & Dileepkumar , C. monticola Manamendra- Arachchi, Batuwita & Pethiyagoda, C. mysoriensis (Jerdon) , C. otai Das & Bauer , C. avasabinae Agarwal, Bauer & Khandekar , C. shevaroyensis Khandekar, Gaitonde & Agarwal , C. yercaudensis Das & Bauer and C. wicksii (Stoliczka) ; large-sized Cnemaspis SVL> 50 mm in C. anamudiensis Cyriac, Johny, Umesh, & Palot , C. beddomei (Theobald) , C. maculicollis Cyriac, Johny, Umesh, & Palot and C. sisparensis (Theobald)) ; spine-like scales absent on flanks (versus spine-like scales present on flanks in C. andersonii , C. amboliensis , C. anandani Murthy, Nitesh, Sengupta & Deepak , C. assamensis , C. flaviventralis , C. goaensis , C. jerdonii (Theobald) , C. koynaensis , C. littoralis , C. monticola , C. mysoriensis , C. nilagirica Manamendra-Arachchi, Batuwita & Pethiyagoda , C. otai , C. wicksii , and C. yercaudensis ); scales on dorsal aspect of trunk heterogeneous (versus scales on dorsal aspect of trunk homogeneous in C. adii , C. assamensis , C. boiei (Gray) , C. indica , C. jerdonii , C. kolhapurensis Giri, Bauer & Gaikwad , C. littoralis , C. nilagirica , C. sisparensis , and C. wynadensis (Beddome)) ; dorsal pholidosis of tail homogeneous, lacking enlarged tubercles forming whorls (versus dorsal pholidosis of tail heterogeneous, having enlarged, conical tubercles forming whorls in C. aaronbaueri , C. adii , C. agarwali , C. ajijae , C. amba , C. amboliensis , C. anandani , C. australis , C. flaviventralis , C. girii , C. goaensis , C. gracilis , C. jerdonii , C. koynaensis , C. limayei , C. littoralis , C. mahabali , C. monticola , C. mysoriensis , C. nilagirica , C. ornata (Beddome) , C. otai , C. avasabinae , C. shevaroyensis , C. thackerayi Khandekar, Gaitonde & Agarwal , C. yercaudensis ); tail with median row of sub-caudal scales smooth, irregularly arranged with a large scale alternating with two slightly smaller scales on the posterior portion of the tail, the larger scales on the anterior portion divided (versus median row of sub-caudal scales smooth and not enlarged in C. amba , C. ajijae , C. flaviventralis , C. girii , C. limayei , and C. koynaensis ; median row of subcaudal scales slightly enlarged in C. amboliensis , C. goaensis , C. mahabali , C. mysoriensis , C. otai , C. avasabinae , and C. yercaudensis ); median row of sub-caudal scales smooth (versus median row of sub-caudal scales keeled in C. amboliensis , C. australis , C. goaensis and C. monticola ); males lacking precloacal pores and having five or six femoral pores on each side, separated by 16–18 poreless scales (versus males with only precloacal pores present in C. aaronbaueri , C. anamudiensis , C. beddomei , C. maculicollis , C. nairi Inger, Marx & Koshy , C. ornata , C. avasabinae ; males with both femoral and precloacal pores present in C. andersonii , C. adii , C. agarwali , C. amboliensis , C. australis , C. bangara Agarwal, Thackeray, Pal & Khandekar , C. goaensis , C. gracilis , C. graniticola Agarwal, Thackeray, Pal & Khandekar C. mysoriensis , C. otai , C. shevaroyensis , C. thackerayi , C. yelagiriensis Agarwal, Thackeray, Pal & Khandekar , C. yercaudensis and C. wicksii ; males with a continuous series of 26–28 precloacalfemoral pores in C. kolhapurensis ; males without femoral and precloacal pores in C. assamensis , C. boiei ; males with only femoral pores present, three or four in C. ajijae , two or three in C. amba , three in C. flaviventralis , three or four in C. girii , eight in C. jerdonii , three or four in C. koynaensis , 14–18 in C. littoralis , three or four in C. mahabali and seven or eight in C. sisparensis ).

C. heteropholis closely resembles C. kottiyoorensis Cyriac & Umesh , and C. wynadensis . However, it can be distinguished from both by having 20–22 lamellae under digit IV of manus, and 24 or 25 lamellae under digit IV of pes (versus 19 or 20 lamellae under digit IV of manus, and 20 or 21 lamellae under digit IV of pes in C. kottiyoorensis ; 15 or 16 lamellae under digit IV of manus, and 18 or 19 lamellae under digit IV of pes in C. wynadensis ); males with five or six femoral pores separated by 16–18 poreless scales (versus males with four or five femoral pores separated by 13–15 poreless scales in C. wynadensis ); 119–122 longitudinal scales from mental to cloaca (versus 113-118 longitudinal scales from mental to cloaca in C. wynadensis ; 138 longitudinal scales from mental to cloaca in C. kottiyoorensis ).

Topotypic C. heteropholis can be distinguished from the Cnemaspis heteropholis from Agumbe ( Ganesh et al. 2011) by having 20–22 lamellae under digit IV of manus, and 24 or 25 lamellae under digit IV of pes (versus 25–26 lamellae under digit IV of manus, and 31–32 lamellae under digit IV of pes); 119–122 longitudinal scales from mental to cloaca (versus 138 longitudinal scales from mental to cloaca) and 20–25 scales across mid-body (versus 30 scales across mid-body).

Description of male (based on CESL 692): SVL 33.4 mm, head short (HL/SVL 0.30), wide (HW/HL 0.68), not strongly depressed (HD/HL 0.40), distinct from neck. Loreal region marginally inflated, canthus rostralis not prominent. Snout just less than half head length (ES/HL 0.46), more than twice eye diameter (ED/ES 0.42); scales on snout and canthus rostralis smooth, round, weakly conical and slightly larger than those on the forehead and interorbital region; occipital and temporal region with small, round, weakly conical granules intermixed with few slightly larger, round and conical tubercles ( Figure 3A View FIGURE 3 ). Eyes small (ED/HL 0.19), with round pupil; extra-brillar fringe scales largest anteriorly, decreasing in size posteriorly; supraciliaries not elongate. Ear-opening deep, vertical, small (EL/HL 0.08); eye to ear distance greater than diameter of eye (EE/ED 1.78) ( Figure 3C View FIGURE 3 ). Rostral half as long as wide, partially divided by a median grove; single enlarged, protuberant, supranasal on each side, twice the size of the postnasals; nostrils oval, each surrounded by postnasal, supranasal, rostral and supralabial I; two rows of scales separate the orbit from the supralabials ( Figure 3C View FIGURE 3 ). Mental enlarged, subtriangular, almost as long (1.78 mm) as wide (1.84 mm); two pairs of postmentals, inner pair much smaller than mental, roughly squarish, bordered by mental, infralabial I, outer postmentals and three small chin scales on each side; inner pair of postmental separated from each other by diastema of three chin scales; outer postmentals slightly smaller than inner postmentals, roughly oval, bordered by infralabials I and II, inner postmentals and four enlarged chin scales on each side; outer postmentals separated from each other by diastema of nine or ten gular scales; chin shields bordering postmentals round, smooth, slightly protuberant, smaller than outermost postmentals, rest small, round, smooth, conical. Infralabials bordered below by a row of slightly enlarged scales, decreasing in size posteriorly ( Figure 3B View FIGURE 3 ). Six supralabials upto angle of jaw and five at midorbital position on each side; supralabial I largest, decreasing in size posteriorly; six infralabials to angle of jaw on left and seven on right; five infralabials upto midorbital position on each side; infralabial I largest, decreasing in size posteriorly ( Figure 3C View FIGURE 3 ). Extra-brillar fringe scales 15 or 16 on each side, 13 interorbital scale rows across narrowest point of frontal; 29 scale rows between left and right supraciliaries at midorbit ( Figure 3A, C View FIGURE 3 ).

Body relatively robust, (BW/SVL 0.18), trunk marginally less than half of SVL (AGL/SVL 0.46) without ventrolateral folds or spine-like scales on flank. Dorsal scales on trunk heterogeneous, small, weakly keeled, oval, granular scales intermixed with enlarged, irregularly arranged, strongly keeled, conical tubercles; tubercles on dorsolateral aspect of flank more pronounced and conical than those on vertebral and paravertebral region ( Figure 5A View FIGURE 5 ). Scales on nape round, slightly smaller than those on paravertebral region, smaller still on occiput ( Figure 2A View FIGURE 2 ). Ventral scales twice the size of granular scales on dorsum, those on belly smooth, subimbricate, subequal from chest to vent; 24 mid-body scale rows; 121 scales from mental to anterior border of cloaca ( Figure 5B View FIGURE 5 ). Scales on throat and pectoral region slightly smaller than those on belly, flat and subimbricate; gular region with much smaller, round, conical, granular scales, those on chin bordering postmentals, slightly enlarged, juxtaposed and flattened ( Figure 2B View FIGURE 2 ). Scales on femoral region slightly larger than those on belly; five femoral pores on each thigh, separated on either side by 16 poreless scales ( Figure 3D View FIGURE 3 ).

Scales on palm and sole smooth, flat, roughly circular, subimbricate; scales on dorsal aspect of manus and pes heterogenous in size and shape, upper arm and forearm with slightly smaller scales than dorsal granules, weakly keeled, round at the base, slightly conical; those near forelimb insertion much smaller, smooth, subimbricate; dorsal aspect of hand predominantly bearing large, flattened, smooth, imbricate scales. Ventral aspect of upper arm with smooth, roughly rounded, conical granules; scales on forearm and wrist larger, flattened, smooth, subimbricate. Scales on dorsal aspect of thigh, knee and shank much larger than those on dorsal granules, weakly keeled, conical except those near hindlimb insertion which are much smaller, weakly conical granules; dorsal aspect of foot predominantly bearing small, weakly keeled, subimbricate scales; scales on ventral aspect of thigh and shank slightly larger than those on mid-body ventrals, smooth, subimbricate ( Figure 2B View FIGURE 2 ). Fore and hind limbs moderately long, slender (LAL/SVL 0.16); (CL/SVL 0.19); digits long, with a strong, recurved claw, distinctly inflected, distal portions laterally compressed conspicuously. Series of paired and unpaired lamellae on basal portion of digits, separat- ed from narrower distal lamellae by a single large scale at the inflection; proximal lamellae series: 2–4–4–5–4 (right manus; Figure 3E View FIGURE 3 ), 2–4–5–8–4 (right pes; Figure 3F View FIGURE 3 ), 2–3–4–5–3 (left manus), 2–5–5–7–3 (left pes); distal lamellae series: 10–14–16–17–14 (right manus; Figure 3E View FIGURE 3 ), 8–14–17–17–18 (right pes; Figure 3F View FIGURE 3 ), 10–15–16–16–15 (left manus), 10–13–15–14–15 (left pes). Relative length of digits (measurements in mm in parentheses): IV (3.6)> III (2.9)> V (2.8)> II (2.4)> I (1.9) (right manus); V (5.1)> IV (4.9)> III (4.5)> II (4.3)> I (2.6) (right pes).

Tail incomplete, cylindrical, relatively slender, flattened beneath, base distinctly swollen when viewed ventrally and laterally, tail length 17.9 mm ( Figure 2C, D View FIGURE 2 ). Dorsal scales on tail base heterogenous, similar to those on mid-body dorsum; scales on rest of the tail homogenous, composed of small, smooth, regularly arranged, flattened, roughly circular, subimbricate scales, without any enlarged tubercles ( Figure 2C View FIGURE 2 ). Scales on ventral aspect of the tail much larger than those on dorsal, smooth, imbricate with a series of three enlarged sub-caudal scales of which the median series is twice the size of adjunct two rows roughly sub pentagonal, enlarged median series is irregularly arranged with a large scale alternating with two slightly smaller scales on the posterior portion of the tail, the larger scales on the anterior portion divided; those on tail base slightly smaller, smooth and subimbricate; a single, small, indistinct postcloacal spur on each side ( Figure 2D View FIGURE 2 ).

Colouration in life. ( Figure 4A View FIGURE 4 ): Dorsal ground colour of head, body, limbs and tail olive brown, speckled with lighter blotches along head and trunk; six or seven black spots longitudinally along mid vertebral from neck to tail base alternated with brown patches, continuing on the tail as light and dark blotches forming obscure broken bands towards the tail end. Head brown with a lighter broken stripe running from the nostril to lower margin of eye, lighter streak continuing backwards from posterior dorsal corner of eye to above ear; supralabials glossy amber with darker edges; a broken pale buff band from last labial to the ear; infralabials amber bordered by greyish band on the top which continues till below the ear. Pupil black outlined by thin, brick red iris. Limbs mottled with lighter blotches and scattered olive yellow scales; digits alternating with light and dark bands. Ventrals pale grey, throat darker, speckled with yellow spots; infralabials, mental, postmentals and two or three rows bordering infralabials dirty yellow. Ventral surface of the tail dirty grey, slightly darker than ventrals.

Variation and additional information from referred specimens. Mensural and meristic data for the studied specimens is given in Table 3 View TABLE 3 & 4 respectively. There are two male and three female specimens ranging in size from 31.3 mm to 44.4 mm. All specimens resemble the male topotype (CESL 692) except as follows: a single gular scale below mental separates the inner pair of postmental in CESL 802. Outer postmentals bordered by five gular scales on left and four on right in CESL 802; outer postmentals bordered by infralabials I on left and right in CESL 802. CESL 693 and CESL 864, gravid females with two eggs inside stomach. All specimens closely agree with the CESL 692 in colouration except CESL 693 overall with brighter yellow markings along the head and tubercles and prominent darker spots. CESL 802 is much darker with dark grey ventrals.

Distribution and Natural history. Cnemaspis heteropholis is so far known from the type locality, Gund, and nearby localities near Ulavi and Castle Rock in Uttar Kannada, Karnataka and two adjacent localities in Goa ( Figure 1 View FIGURE 1 ). All the locations lie in a fairly contiguous stretch of moist deciduous to semi-evergreen forest along the border of Goa and Karnataka. At Gund, an adult male (CESL 692) was collected from a mud cliff along forest path and another sub-adult male (CESL 802) from a tree hole, approximately 1 m above the ground. Both the individuals were seen active and collected in the evening hours, between 18:00 hrs and 20:30 hrs. An adult female (CESL 700) was found moving on a horizontal rock near forest stream in the evening (ca. 19:00 hrs) at Verlem, Netravali Wildlife Sanctuary, Goa. A gravid female (CESL 693) was found hiding under a decaying log on the forest floor in the afternoon (15:30 hrs) at Ram Dongar, Cotigao Wildlife Sanctuary, Goa on 29 th August 2012. Another gravid female (CESL 864) was collected from the forest floor near a cluster of rocks next to a stream in the evening (ca. 19:30 hrs) near Ulavi, Uttar Kannada, Karnataka on 24 th November 2013. All the individuals were observed in well-shaded forests with dense vegetation and leaf litter on forest floor.