Cnemaspis uttaraghati, Khandekar & Thackeray & Agarwal, 2021

Khandekar, Akshay, Thackeray, Tejas & Agarwal, Ishan, 2021, A novel small-bodied rupicolous Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from the northern Western Ghats, Maharashtra, India, with comments on the status of C. indraneildasii, Bauer 2000, Zootaxa 4969 (2), pp. 331-350 : 336-345

Cnemaspis uttaraghati sp. nov.

( Figures 2–6 ; Tables 3 & 4)

Holotype. NRC-AA-1122 (AK 1233), adult male, from Kedareshwar caves on Harishchandragad mountain in Kalsubai-Harishchandragad Wildlife Sanctuary, (19.392° N, 73.779° E; ca. 1200 m asl.), Ahmednagar district, Maharashtra state, India, collected by Santosh Saste, Vaibhav Patil and Yogesh Satam on 27 October 2019.

Paratypes. BNHS 2799 (AK 1234), adult male, NRC-AA-1123 (AK 1235) , BNHS 2800 (AK 1236), adult females, same collection and data as holotype .

Etymology. The specific epithet is used as a noun in apposition and is derived from the Sanskrit ‘uttara’ for north and ‘ghat’ for mountains, as this mountain locality represents the currently known northern limit of Cnemaspis distribution in the Western Ghats.

Suggested Common Names. Northern dwarf gecko or Harishchandragad dwarf gecko.

Diagnosis: A small-sized Cnemaspis , snout to vent length less than 39 mm (n =4). Dorsal pholidosis heterogeneous; weakly keeled, granular scales intermixed with large, weakly keeled, regularly arranged tubercles, enlarged tubercles gradually increasing in size towards flank, lowest row largest, spine-like; eight or nine rows of dorsal tubercles at mid-body, 7–10 paravertebral tubercles between forelimb and hindlimb insertions; ventral scales smooth, sub-imbricate, sub-equal from chest to vent, 27–32 scales across belly at mid-body, 124–132 longitudinal scales from mental to cloaca; subdigital scansors smooth, mostly entire, unnotched; 10–12 lamellae under digit I of manus and 10 or 11 lamellae under digit I of pes, 17 or 18 lamellae under digit IV of manus and 19–22 lamellae under digit IV of pes; males with four femoral pores on each thigh separated medially by 22–24 poreless scales (n =2); dorsal pholidosis of tail with strongly keeled, granular scales, similar in size and shape to granular scales on mid-body dorsum, gradually becoming larger, flattened, imbricate posteriorly, intermixed with enlarged, strongly keeled, distinctly pointed, conical tubercles forming whorls, six tubercles in first eight whorls; median row of subcaudals smooth, slightly enlarged, regularly arranged with condition of two slightly larger scales alternating with a large divided scale. Dorsal colouration greenish-brown with six dark vertebral blotches running from the occiput to the sacrum, speckled with fine saffron granules, limbs with light yellow bands.

Comparison with peninsular Indian congeners. Cnemaspis uttaraghati sp. nov. can be distinguished from all other Indian congeners on the basis of the following differing or non-overlapping characters: small-sized Cnemaspis SVL 36–39 mm (versus medium-sized Cnemaspis SVL 40–50 mm in C. anandani Murthy, Anandan, Sengupta & Deepak , C. bangara Agarwal, Thackeray, Pal & Khandekar , C. boiei (Gray) , C. chengodumalaensis Cyriac, Palot, Deuti & Umesh , C. jerdonii (Theobald) , C. kolhapurensis Giri, Bauer & Gaikwad , C. heteropholis Bauer , C. kottiyoorensis Cyriac & Umesh , C. nairi Inger, Marx & Koshy , C. nilagirica Manamendra-Arachchi, Batuwita & Pethiyagoda , C. graniticola Agarwal, Thackeray, Pal & Khandekar , C. yelagiriensis Agarwal, Thackeray, Pal & Khandekar , C. wynadensis (Beddome) , C. ornata (Beddome) , and C. thackerayi Khandekar, Gaitonde & Agarwal ); large-sized Cnemaspis SVL > 50 mm in C. anamudiensis Cyriac, Johny, Umesh, & Palot , C. beddomei (Theobald) , C. maculicollis Cyriac, Johny, Umesh, & Palot , C. magnifica Khandekar, Thackeray, Pal & Agarwal , C. sisparensis (Theobald) , C. zacharyi Cyriac, Palot, Deuti & Umesh ); spine-like tubercles present on flanks (versus spine-like tubercles absent on flanks in C. aaronbaueri Sayyed, Grismer, Campbell & Dileepkumar , C. adii Srinivasulu, Kumar & Srinivasulu , C. agarwali Khandekar , C. ajijae , C. amba , C. anamudiensis , C. australis Manamendra-Arachchi, Batuwita & Pethiyagoda , C. bangara , C. beddomei , C. boiei , C. chengodumalaensis , C. girii , C. graniticola , C. heteropholis , C. indica (Gray) , C. kolhapurensis , C. kottiyoorensis , C. limayei , C. maculicollis , C. magnifica , C. mahabali , C. nairi , C. ornata , C. palakkadensis Sayyed, Cyriac & Dileepkumar , C. ranganaensis Sayyed & Sulakhe , C. shevaroyensis Khandekar, Gaitonde & Agarwal , C. sisparensis , C. thackerayi , C. wynadensis , and C. yelagiriensis , C. zacharyi ); scales on dorsal aspect of trunk heterogeneous (versus scales on dorsal aspect of trunk homogeneous in C. adii , C. assamensis Das & Sengupta , C. boiei , C. indica , C. jerdonii , C. kolhapurensis , C. littoralis , C. nilagirica , C. palakkadensis , C. sisparensis , C. wynadensis , and C. zacharyi ); caudal tubercles enlarged, strongly keeled, distinctly pointed, forming whorls (versus scales on dorsal aspect of tail smooth, without whorls of enlarged tubercles in C. anamudiensis , C. beddomei , C. boiei , C. chengodumalaensis , C. heteropholis , C. indica , C. kolhapurensis , C. kottiyoorensis , C. maculicollis , C. magnifica , C. sisparensis , C. wynadensis , C. zacharyi ); sub-caudal scales smooth, median row slightly enlarged, scales in median row regularly arranged with condition of two slightly larger scale alternating with a large divided scale (versus sub-caudal scales smooth, median row not enlarged in, C. amba , C. ajijae , C. australis , C. flaviventralis , C. girii , C. limayei and C. koynaensis ; subcaudal scales smooth, median row distinctly enlarged in C. aaronbaueri , C. agarwali , C. anandani , C. bangara , C. boiei , C. chengodumalaensis , C. gracilis (Beddome) , C. graniticola , C. heteropholis , C. indica , C. jerdonii , C. kolhapurensis , C. magnifica , C. nairi , C. nilagirica , C. ornata , C. shevaroyensis , C. sisparensis , C. thackerayi , C. wynadensis , C. yelagiriensis , C. zacharyi ; median row of sub-caudal scales slightly enlarged, without slightly larger scale alternating with a large divided scale in C. amboliensis , C. avasabinae Agarwal, Bauer & Khandekar , C. goaensis , C. mahabali , C. monticola , C. mysoriensis (Jerdon) , C. otai Das & Bauer , C. rishivalleyensis Agarwal, Thackeray & Khandekar , C. stellapulvis Khandekar, Thackeray &Agarwal and C. yercaudensis Das & Bauer ); males with four femoral pores on each thigh, separated medially by 22–24 poreless scales (versus femoral pores absent, only precloacal pores present in C. aaronbaueri , C. anamudiensis , C. beddomei , C. maculicollis , C. nairi , and C. ornata ; males with both femoral and precloacal pores present in C. adii , C. agarwali , C. amboliensis , C. australis , C. avasabinae , C. bangara , C. goaensis , C. gracilis , C. graniticola , C. mysoriensis , C. otai , C. ranganaensis , C. rishivalleyensis , C. shevaroyensis , C. stellapulvis , C. thackerayi , C. yelagiriensis , and C. yercaudensis ; both femoral and precloacal pores absent in C. assamensis and C. boiei ; a continuous series of 26–28 precloacal-femoral pores in C. kolhapurensis ).

Cnemaspis uttaraghati sp. nov. closely resembles members of girii clade ( C. ajijae , C. amba , C. flaviventralis , C. girii , C. koynaensis , C. limayei , C. mahabali ). However, it can be further distinguished from all by having 7–10 paravertebral tubercles between forelimb and hindlimb insertions (versus 17–22 paravertebral tubercles between forelimb and hindlimb insertions in C. ajijae and C. amba ; 18 in C. flaviventralis , 20–22 in C. girii , 18–25 in C. koynaensis and 13–15 in C. limayei ); eight or nine rows of dorsal tubercles at mid-body (versus 11–14 rows of dorsal tubercles at mid-body in C. amba , 12 or 13 in C. flaviventralis , 12–14 in C. girii , and 10–14 in C. koynaensis ); 27–32 ventral scales across belly at mid-body (versus 22–24 ventral scales across belly at mid-body in C. amba , 20–26 in C. koynaensis , and 26 in C. mahabali ); 124–132 longitudinal ventral scales from mental to cloaca (versus 141–149 longitudinal ventral scales from mental to cloaca in C. amba , and 133–139 in C. girii ); 17 or 18 lamellae under digit IV of manus (versus 15 or 16 lamellae under digit IV of manus in C. ajijae , C. flaviventralis and C. mahabali , 15 in C. limayei ); SVL up to 36–39 mm (versus maximum SVL <34 mm in C. amba , C. girii , C. koynaensis , C. limayei , C. mahabali ).

Description of the holotype. Adult male in good state of preservation except tail tip slightly bent towards right and a 2.5 mm long incision in the sternal region for tissue collection ( Figure 2 ). SVL 37.0 mm, head short (HL/SVL 0.25), wide (HW/HL 0.66), not strongly depressed (HD/HL 0.41), distinct from neck. Loreal region slightly inflated, canthus rostralis not prominent. Snout half the head length (ES/HL 0.45), 2.5 times eye diameter (ED/ES 0.39); scales on snout and canthus rostralis large, roughly circular, smooth, somewhat conical; much larger than those on forehead and interorbital region; occipital and temporal region with much smaller granules intermixed with slightly larger conical tubercles ( Figure 3A ). Eye small (ED/HL 0.17); with round pupil; orbit with 9–12 extra-brillar fringe scales, largest scales on anterior side; supraciliaries not elongate; interorbital scale rows across narrowest point of frontal seven or eight; 26 scale rows between left and right supraciliaries at mid-orbit ( Figure 3A, C ). Earopening deep, vertical, small (EL/HL 0.05); eye to ear distance greater than diameter of eye (EE/ED 1.82) ( Figure 3C ). Rostral much wider (1.7 mm) than high (0.7 mm), incompletely divided dorsally by a strongly developed rostral groove about more than half of its height; a single enlarged supranasal on each side, right supranasal twice the size of left, separated from each other by a single enlarged internasal and two much smaller scales on the snout; two enlarged postnasals on left and single on right side, slightly smaller than the supranasals; rostral in contact with supralabial I, nasal, supranasal and internasal; nostrils oval, surrounded by upper postnasal, lower postnasal, supranasal, rostral and supralabial I on left and lower postnasal, supranasal, rostral and supralabial I on right side; two rows of scales separate the orbit from the supralabials ( Figure 3A, C ). Mental enlarged, subtriangular, marginally wider (2.3 mm) than long (1.6 mm); two pairs of postmentals, inner pair slightly larger than outer pair, roughly rectangular, separated from each other by a single enlarged chin shield below mental; inner pair bordered by mental, infralabial I, outer postmentals and three enlarged chin shields; outer postmentals roughly rectangular, bordered by inner postmentals, infralabial I and II, four enlarged chin shields on left and three on right side; three enlarged gular scales prevent contact of left and right outer postmentals; chin shields bordering postmentals flat, smooth, smaller than outermost postmentals, rest small, smooth ( Figure 3B ). Infralabials bordered below by a row of slightly enlarged scales, decreasing in size posteriorly. Nine supralabials on left and eight on right side up to angle of jaw; seven supralabials on left and six on right at mid-orbital position; supralabial I largest, decreasing in size posteriorly; seven infralabials up to angle of jaw and six at midorbital position on both sides; infralabial I largest, infralabials decreasing in size posteriorly ( Figure 3C ).

Body relatively slender, trunk less than half of SVL (AGL/SVL 0.41) without ventrolateral folds. Dorsal pholidosis heterogeneous; weakly keeled, granular scales intermixed with about four or five regularly arranged rows of large, weakly keeled tubercles on each flank, enlarged tubercles gradually increasing in size towards flank, lowest row largest, weakly keeled, spine-like, four or five spine-like tubercles in last row; nine rows of dorsal tubercles at mid-body, seven paravertebral tubercles between forelimb and hindlimb insertions. ( Figure 4A, B ). Scales on occiput slightly smaller than those on nape and paravertebral rows; scales on flank slightly larger than those on dorsum, weakly keeled, conical. Ventral scales much larger than granular scales on dorsum, those on belly smooth, sub-imbricate, sub-equal from chest to vent; scales above cloaca and thigh slightly larger than those on midbody ventral; mid-body scale rows across belly 27; 125 scales from mental to anterior border of cloaca ( Figure 4B ). Scales on throat slightly smaller than those on belly; gular region with much smaller, flattened scales with those on chin bordering postmentals, enlarged, juxtaposed and flattened ( Figure 3B ). Femoral pores elongate, four on each thigh, separated medially by 22 poreless scales ( Figure 3D ).

Scales on palm and sole smooth, flat and roughly circular; scales on dorsal aspect of manus and pes heterogenous in size, upper arm and thigh with scales much larger than dorsal granules, strongly keeled, imbricate; those near limb insertions much smaller, granular, smooth; dorsal aspect of lower arm and shank with scales much smaller than those on upper arm and thigh, weakly keeled, imbricate; dorsal aspect of hand and foot predominantly bearing large, flattened, weakly keeled, imbricate scales ( Figure 2A ). Ventral aspect of upper arm with smooth, roughly rounded, sub-imbricate scales; scales on lower arm and wrist slightly larger than those on upper arm, smooth, sub-imbricate; scales on ventral aspect of thigh and shank larger than those on mid-body ventrals, smooth, subimbricate ( Figure 2B ). Fore-limbs and hind-limbs slightly long, slender (LAL/SVL 0.14); (CL/SVL 0.18); digits long, with a strong, recurved claw, distinctly inflected, distal portions laterally compressed conspicuously. Series of unpaired lamellae on basal portion of digits, separated from narrower distal lamellae by a single large scale at the inflection; proximal lamellae series: 2-4-6-6-5 (right manus; Figure 3E ), 2-5-7-9-7 (right pes; Figure 3F ), 2-5-6-6- 6 (left manus), 2-5-6-8-7 (left pes); distal lamellae series: 8-10-12-11-10 (right manus; Figure 3E ), 8-10-11-13-13 (right pes; Figure 3F ), 9-10-12-11-10 (left manus), 8-10-12-13-12 (left pes). Relative length of digits (measurements in mm in parentheses): IV (4.2)> III (4.0)> V (3.7)> II (3.6)> I (2.5) (right manus); IV (5.4)> III (4.7)> V (4.5)> II (4.1)> I (2.5) (right pes).

Tail original, entire, sub-cylindrical, relatively slender, flattened beneath, slightly longer than snout-vent length (TL/SVL 1.15) ( Figure 2C, D ). Dorsal scales on tail base strongly keeled, granular, similar in size and shape to granular scales on mid-body dorsum, gradually becoming larger, flattened, imbricate posteriorly, intermixed with enlarged, strongly keeled, distinctly pointed, conical tubercles forming whorls; six tubercles each on first eight whorls ( Figure 2C ). Scales on ventral aspect of original tail much larger than those on dorsal aspect, sub-imbricate, smooth; median series slightly larger than rest, roughly circular, regularly arranged with two slightly larger scales alternating with a large divided scale; scales on tail base slightly smaller, sub-imbricate, smooth, a single and smooth enlarged postcloacal spur on each side ( Figure 2D ).

Colouration in life. ( Figure 5A ) Dorsal ground colour of head, body, limbs, and tail light brown; head with numerous dark brown blotches, brille dirty yellow. Dark brown preorbital streak from orbit forming an indistinct preorbital streak on snout; yellow and dark grey alternating bands on upper labials; two dark brown postorbital streaks, upper one small, half the length of lower, which continues until ear-opening. Dorsum with six dark vertebral blotches, each followed by an indistinct light grey blotch speckled with scattered saffron granules; vertebral blotches flanked by same number of blotches forming indistinct wavy cross-bars, minutely speckled with dirty saffron between forelimb and hind insertions; spine-like tubercles dirty yellow. Dorsum of limbs with light yellow bands; digits with alternating dark and dirty yellow markings; dorsum of original portion of tail with alternating dark brown and ashy bands, dark brown bands speckled with saffron, tail tip orangish ( Figure 5A, C ). Gular, pectoral, abdominal region, and underside of limbs and tail off white with no dark markings; postcloacal spur and conical tubercles in lateral and ventrolateral rows on tail close to the cloacal opening light yellow. Pupil black, iris golden.

Variation and additional information from type series. Mensural and meristic data for the type series is given in Table 3 & 4 respectively. There are two male and two female specimens ranging in size from 36.5 mm to 38.9 mm. All specimens resemble the holotype except as follows: two internasal scales separate supranasals on snout in NRC-AA-1123 and BNHS 2800 ; a two postnasals on each side in BNHS 2799 , NRC-AA-1123 and BNHS 2800 ; outer postmentals bordered by four gular scales on each side in BNHS 2799 outer postmental bordered by four gular scales on right and three on left in NRC-AA-1123 and BNHS 2800 . Two paratypes —NRC-AA-1123 and BNHS 2800 with original and complete tails, slightly longer than body (TL/ SVL 1.13 and 1.12 respectively); BNHS 2799 with tail incomplete and regenerated tail tip, detached from the body, slightly shorter than body (TL/ SVL 0.76) ( Figure 6A ). All paratypes closely agree with the holotype in overall colouration except NRC-AA-1123 with dorsal ground colour of mid-body dorsum olive greenish, vertebral blotches and either side of flank heavily speckled with fine saffron granules ( Figure 5B ); BNHS 2800 is slightly paler than holotype ( Figure 5C ); colouration of the regenerated tail of BNHS 2799 is brown, lacking bands ( Figure 6A ) .

Distribution and Natural history. Cnemaspis uttaraghati sp. nov. is so far known only from the type locality, Harishchandragad mountain, in Kalsubai-Harishchandragad Wildlife Sanctuary,Ahmednagar district of Maharashtra at an elevation of ca. 1200 m asl. ( Figure 1 ). The individuals of the new species were encountered during a single day of fieldwork in the afternoon (1330–1445 hrs.). All four individuals were seen active, only on vertical basaltic rocks <2m height, inside man-made caves ( Figure 7B ). A communal egg-laying site containing 11 hatched eggs and a single dark coloured unhatched egg (probably unviable) were seen on a horizontal rock face at approximately 3m height ( Figure 7C ). These caves are located on a high elevation (~ 1200 m asl.) basaltic plateau, predominantly covered by sparse scrubs and grasses on the top and semi-evergreen forest patches on the slopes ( Figure 7A ). Hemidactylus maculatus Duméril & Bibron , and H. murrayi Gleadow are the only two other gecko species that were observed in sympatry with the new species.

Figures

FIGURE 1. Elevation map showing type localities of Cnemaspis spp. distributed in the northern Western Ghats: 1, Cnemaspis uttaraghati sp. nov.; 2, C. mahabali; 3, C. ajijae; 4, C. girii; 5, C. koynaensis; 6, C. amba; 7, C. kolhapurensis; 8, C. limayei; 9, C. ranganaensis; 10, C. amboliensis and C. flaviventralis; 11, C. goaensis; 12, C. indraneildasii. Prepared by Ishan Agarwal.

FIGURE 2. Cnemaspis uttaraghati sp. nov. (holotype, NRC-AA-1122): (A) dorsal view of body, (B) ventral view of body, (C) dorsal view of tail, and (D) ventral view of tail. Scale bars 10 mm; photos by Akshay Khandekar.

FIGURE 3. Cnemaspis uttaraghati sp. nov. (holotype, NRC-AA-1122), dorsal view of head (A), ventral view of head (B), lateral view of head (C), view of cloacal region showing femoral pores and post cloacal spurs (D), ventral view of right manus (E), and ventral view of right pes (F). Scale bars 5 mm; photos by Akshay Khandekar.

FIGURE 4. Details at mid-body of Cnemaspis uttaraghati sp. nov. (holotype, NRC-AA-1122): (A) dorsal view and (B) ventral view. Scale bars 5 mm; photos by Akshay Khandekar.

FIGURE 5. Cnemaspis uttaraghati sp. nov. in life: (A) male, holotype NRC-AA-1122, (B) female, paratype NRC-AA-1123, and (C) female, paratype BNHS 2800; photos by Tejas Thackeray.

FIGURE 6. Type series of Cnemaspis uttaraghati sp. nov. from left to right: BNHS 2799, NRC-AA-1123 and BNHS 2800. Scale bar 10 mm; photos by Akshay Khandekar.

FIGURE 7. A) Habitat of Cnemaspis uttaraghati sp. nov. near Kedareshwar Caves on Harishchandragad, Ahmednagar district, Maharashtra state, India; B) entrance of one of the caves at type locality; C) hatched communal egg clutch with a single dark unhatched egg inside the cave; photos by Y. Satam.