A new medium-sized rupicolous Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) of the C. bangara clade from granite boulder habitats in Krishnagiri, Tamil Nadu India Author Agarwal, Ishan 0000-0001-9734-5379 Thackeray Wildlife Foundation, Mumbai, 400051, India. & National Centre for Biological Sciences, Tata Institute of Fundamental Research, Bengaluru, 560065, India. & ishan. agarwal @ gmail. com; http: // orcid. org / 0000 - 0001 - 9734 - 5379 ishan.agarwal@gmail.com Author Thackeray, Tejas 0000-0002-9981-8763 Thackeray Wildlife Foundation, Mumbai, 400051, India. & tejasthackeray @ gmail. com; https: // orcid. org / 0000 - 0002 - 9981 - 8763 tejasthackeray@gmail.com Author Khandekar, Akshay 0000-0002-7956-089X Thackeray Wildlife Foundation, Mumbai, 400051, India. & National Centre for Biological Sciences, Tata Institute of Fundamental Research, Bengaluru, 560065, India. & akshaykhandekar 555 @ gmail. com; https: // orcid. org / 0000 - 0002 - 7956 - 089 X akshaykhandekar555@gmail.com text Zootaxa 2021 2021-05-11 4969 2 351 366 journal article 6507 10.11646/zootaxa.4969.2.7 e78e91d5-6e7b-4be5-b5f9-81c827e0d139 1175-5326 4749151 408CE900-0929-451C-AF92-1C6E4E62B700 Cnemaspis krishnagiriensis sp. nov. ( Figures 2–6 ; Tables 3 & 4 ) Holotype . NRC-AA-1120 (AK 896), SVL 40 mm , adult male, from behind Kugai Periya Mariamman Temple, near Krishnagiri fort, ( 12.544 , 78.224 ca. 550 m asl .), Krishnagiri District , Tamil Nadu , India , collected by Akshay Khandekar , Ishan Agarwal and Swapnil Pawar on 16 October 2019 . FIGURE 1. Elevation map of south-eastern peninsular India showing the distribution of members of the bangara clade, and inset, maximum likelihood phylogeny (ND2) with numbers at nodes indicating % bootstrap support/ posterior probability. Prepared by Ishan Agarwal. Paratypes . NRC-AA-1121 (AK 897) SVL 39.5 mm , adult female, BNHS 2798 (AK 898) SVL 27.2 mm , subadult female, same collection data as holotype . Etymology. The specific epithet is a toponym for Krishnagiri in Krishnagiri District of Tamil Nadu , the type and only known locality for this species. Suggested common name. Krishnagiri dwarf gecko. Diagnosis. A medium-sized Cnemaspis , maximum SVL 40 mm (n=3). Dorsal pholidosis heterogeneous, weakly keeled, granular scales intermixed with large, strongly keeled, conical tubercles, seven or eight longitudinal rows of dorsal tubercles, 13 or 14 tubercles in paravertebral rows, spine-like tubercles absent on flank; ventral scales on belly smooth, imbricate, 23–25 scales across the belly, 120–123 longitudinal scales between mental to anterior border of cloaca; subdigital scansors smooth, mostly entire, unnotched, lamellae under digit IV of pes 22–25; male with four femoral pores on each thigh, separated on either side by nine or 10 poreless scales from a continuous series of three precloacal pores; dorsal pholidosis of tail heterogeneous, weakly keeled, granular scales intermixed with enlarged, strongly keeled, conical tubercles forming four whorls on only anterior portion, six tubercles in first whorl, four tubercles in second to fourth whorls, enlarged tubercles restricted to only paravertebral region on rest of the tail, lacking lateral and ventrolateral rows, median row of sub-caudals smooth and distinctly enlarged. A single large central black ocellus anterior to forelimb insertions flanked by smaller spots on each side; dark dorsal markings with fine yellow speckles; tail tip orange in adults, distal half of tail orange in juveniles; gular region with a broken dark streak on each side. FIGURE 2. Cnemaspis krishnagiriensis sp. nov. (holotype, NRC-AA-1120), dorsal view (A), and ventral view (B). Scale bars 10 mm; photos by Akshay Khandekar. Comparison with members of the bangara clade. C. krishnagiriensis sp. nov. can be distinguished from its closest relatives C. bangara , C. graniticola and C. yelagiriensis (opposing character states in parentheses) in having seven or eight longitudinal rows of dorsal tubercles ( versus dorsal tubercles irregularly arranged in C. bangara ); 13 or 14 tubercles in paravertebral rows ( versus 7–9 in C. bangara and 14–16 in C. yelagiriensis ); 23–25 scales across the belly ( versus 20 in C. bangara ); 120–123 longitudinal scales between mental to anterior border of cloaca ( versus 116–120 in C. bangara , 115–121 in C. graniticola and 125–137 in C. yelagiriensis ); four femoral pores on each thigh, separated on either side by nine or 10 poreless scales from a continuous series of three precloacal pores ( versus three or four femoral pores separated on either side by 9–11 poreless scales from a continuous series of two precloacal pores which may be separated by a single poreless scale in C. yelagiriensis ; two femoral pores separated on either side by 11 poreless scales from a continuous series of two precloacal pores in C. bangara ; three femoral pores separated on either side by eight or nine poreless scales from a continuous series of four precloacal pores in C. graniticola ). Additionally, C. krishnagiriensis sp. nov . is 9.2 % divergent from C. yelagiriensis , 16.9 % divergent from C. bangara , and 17.6 % divergent from C. graniticola in ND2 sequence data ( Table 2 ). FIGURE 3. Cnemaspis krishnagiriensis sp. nov. (holotype, NRC-AA-1120), dorsal view of head (A), ventral view of head (B), lateral view of head (C), view of cloacal region showing precloacal and femoral pores (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 krishnagiriensis sp. nov. (holotype, NRC-AA-1120): (A) dorsal view and (B) ventral view. Scale bars 10 mm; photos by Akshay Khandekar. Description of the holotype . Adult male in good state of preservation except for a 4.4 mm long skin injury on dorsal surface of the tail base, a 2.7 mm long incision in the sternal region for tissue collection and half of the tail missing ( Figure 2 ). SVL 40 mm , head short (HL/SVL 0.25), wide (HW/HL 0.61), not strongly depressed (HD/HL 0.38), distinct from neck. Loreal region marginally inflated, canthus rostralis not prominent. Snout just half than head length (ES/HL 0.50), 2.5 X eye diameter (ED/ES 0.45); scales on snout and canthus rostralis large, roughly circular, smooth, juxtaposed; more than twice the size than those on forehead and interorbital region; occipital and temporal region with much smaller granules ( Figure 3A ). Eye small (ED/HL 0.22); with round pupil; orbit with extra-brillar fringe scales that are largest anteriorly; supraciliaries not elongate. Ear-opening deep, vertical, small (EL/HL 0.06); eye to ear distance greater than diameter of eye (EE/ED 1.34) ( Figure 3C ). Rostral much wider ( 1.7 mm ) than long ( 0.7 mm ), incompletely divided dorsally by a strongly developed rostral groove and by inner internasal scale for more than half of its length; two enlarged supranasal on each side, slightly larger than postnasals, separated from each other by two slightly smaller internasals and two much smaller scales on the snout; rostral in contact with supralabial I, nasal, lower postnasal, inner supranasal and inner internasal on the left and supralabial I, nasal, inner supranasal and inner internasal on the right side; nostrils oval, each surrounded by upper and lower postnasal, inner supranasal and rostral; two rows of scales separate the orbit from the supralabials ( Figure 3A ). Mental enlarged, subtriangular, slightly wider ( 1.9 mm ) than long ( 1.6 mm ); two pairs of postmentals, inner pair large, roughly rectangular, bordered by mental, infralabial I, outer postmentals and two enlarged chin shields on each side; inner postmentals separated from each by a single, smaller, gular scale below mental; outer postmentals slightly smaller than inner postmentals, roughly rectangular, bordered by infralabials I and II, inner postmentals and four enlarged chin scales 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, slightly smaller than outermost postmentals, rest small, flat, smooth. Infralabials bordered below by a row of slightly enlarged scales, decreasing in size posteriorly ( Figure 3B ). Eight supralabials up to angle of jaw and seven at midorbital position on each side; supralabial I largest, decreasing in size posteriorly; seven infralabials to angle of jaw and six up to midorbital position on each side; infralabial I largest, decreasing in size posteriorly ( Figure 3B ). Seven extra-brillar fringe scales on each side, eight interorbital scale rows across narrowest point of frontal; 30 or 31 scale rows between left and right supraciliaries at midorbit ( Figure 3A ). Body relatively slender (BW/SVL 0.16), trunk less than half of SVL (AGL/SVL 0.40) without ventrolateral folds or spine-like tubercles on flank. Dorsal scales on trunk heterogeneous, weakly keeled, granular scales intermixed with much larger, strongly keeled, conical tubercles; approximately eight tubercles in longitudinal rows at midbody; 14 tubercles in paravertebral row from above forelimb insertion to hindlimb insertion ( Figure 4A ). Scales on nape slightly smaller than those on paravertebral rows, smaller still on occiput ( Figure 3A ). Scales on flank slightly larger than those on dorsum, roughly circular, smooth. Ventral scales more than thrice the size of granular scales on dorsum, those on belly smooth, imbricate, subequal from chest to vent; midbody scale rows across belly 24; 123 scales from mental to anterior border of cloaca ( Figure 4B ). Scales on throat and pectoral region slightly smaller than those on belly, flat and imbricate; gular region with much smaller, flattened scales, those on chin bordering postmentals, enlarged, juxtaposed and flattened ( Figure 3B ). Scales on femoral region marginally larger than those on belly; four femoral pores on each thigh, separated by 10 poreless scales on left and nine on right side from three continuous precloacal pores ( 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 with scales much larger than dorsal granules, weakly keeled, subimbricate; those near forelimb insertion much smaller; dorsal aspect of forearm and elbow with scales smaller than those on upper arm, weakly keeled, flat, roughly rounded; dorsal aspect of hand predominantly bearing large, flattened, smooth, imbricate scales. Ventral aspect of upper arm with smooth, roughly rounded granules; scales on forearm and wrist larger, smooth, weakly imbricate. Scales on dorsal aspect of thigh much larger than those on dorsal granules, weakly keeled, imbricate except those near hindlimb insertion which are much smaller, conical granules; scales on dorsal aspect of knee and shank slightly smaller than those on dorsum of thigh, subimbricate, weakly keeled; dorsal aspect of foot predominantly bearing small, flattened, smooth, imbricate scales. Scales on ventral aspect of thigh marginally larger than those on mid-body ventrals; scales on shank slightly smaller than those on thigh, smooth, subimbricate ( Figure 3D ). Fore and hind limbs moderately long, slender (LAL/SVL 0.17); (CL/SVL 0.22); 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: 2-5-5-7-5 (right manus; Figure 3E ), 2-7-8-9-8 (right pes; Figure 3F ), 2-6-6-8-6 (left manus), 2-6- 8-9-8 (left pes); distal lamellae series: 11-12-16-15-13 (right manus; Figure 3E ), 10-13-16-16-15 (right pes; Figure 3F ), 11-12-15-15-12 (left manus), 11-13-15-15-16 (left pes). Relative length of digits (measurements in mm in parentheses): IV (4.1)> III (4.0)> V (3.6)> II (3.5)> I (2.6) (right manus); IV (5.3)> III (4.8)> V (4.5)> II (4.0)> I (2.2) (right pes). Tail original, not entire, approximately half of its length missing, sub-cylindrical, relatively slender, flattened beneath, base distinctly swollen when viewed ventrally and laterally, tail length 28.4 mm ( Figure 2 ). Dorsal scales at tail base granular, weakly keeled, similar in size and shape to those on mid-body dorsum, gradually becoming larger, flatter, subimbricate posteriorly, intermixed with enlarged, strongly keeled, conical tubercles forming four whorls on only anterior portion, six tubercles in first whorl, four tubercles in second to fourth whorls, enlarged tubercles restricted to only paravertebral region on rest of the tail, lacking lateral and ventrolateral rows. Scales on ventral aspect of tail much larger than those on dorsal, imbricate, smooth, with a series of three enlarged subcaudal scales of which the median series is almost twice the size of adjunct two rows, roughly hexagonal; those on tail base much smaller, imbricate and smooth. A single enlarged, smooth postcloacal spur on each side ( Figure 2B ). FIGURE 5. Cnemaspis krishnagiriensis sp. nov. in life: (A) male, holotype NRC-AA-1120, (B) female, paratype NRC-AA- 1121, and (C) juvenile female, paratype BNHS 2798; photos by Ishan Agarwal. FIGURE 6. Type series of Cnemaspis krishnagiriensis sp. nov. : BNHS 2798 & NRC-AA-1121. Scale bar 10 mm; photos by Akshay Khandekar. Colouration in life. ( Figure 5A ) Dorsal ground colour of head, body, limbs and tail pale brown; head and dorsum with darker speckling throughout, brille dull yellow. Dark brown preorbital streak runs from nostril to orbit; upper labials dull white with dark blotches. Two dark brown postorbital streaks, upper one merging with its counterpart from the other orbit to form an indistinct)-shaped band on occiput; lower one continuing until earopening. A single large central black ocellus anterior to forelimb insertions, flanked by slightly smaller spots on each side; all dark dorsal markings with fine yellow speckles. Four indistinct dark bands from axilla to tail base alternating with lighter, elongate mid-dorsal blotches; dorsum of forelimbs and hindlimbs with dark blotches and bands, dark blotches speckled with yellow scales, digits with alternating dark and light bands; dorsum of half broken tail with four alternating brown and light grey bands. Ventral surfaces dull white, gular region with a broken dark streak on each side, both running along lower labials up to lower postmentals, underside of tail marbled with grey. Pupil black, iris golden. Variation and additional information from type series. Mensural and meristic data for the type specimens is given in Table 3 & 4 respectively. There are adult and sub-adult females ranging in size from 27.2 mm to 39.5 mm . All specimens resemble the holotype except as follows: two internasal scales and a much smaller scale separate supranasals on snout in BNHS 2798 . Outer postmentals bordered by four gular scales on left and five on right side in NRC-AA-1121; outer postmentals bordered by three gular scales on left and four on right side in BNHS 2798 ; outer postmentals bordered by infralabials I & II on left and infralabials II & on right in NRC-AA-1121. A few subdigital lamellae divided; in NRC-AA-1121 fourth proximal on digit IV of right manus, fourth proximal on digit III of right pes and sixth, seventh and ninth on digit IV of right pes and sixth and eighth on digit IV of left pes; in BNHS 2798 eighth proximal on digit IV of right pes, sixth proximal on left; 11 th and 12 th distal on digit II of pes right. Both paratypes —NRC-AA-1121 and BNHS 2798 , with original and complete tails slightly longer than body (TL/ SVL 1.26 and 1.17 respectively). Both the paratypes match the holotype in overall colouration except tail. The adult female paratype NRC-AA-1121 with only the extreme tail tip dull orange; the juvenile paratype BNHS 2798 with posterior half of the tail bright orange ( Figure 6 ) . TABLE 3. Mensural (mm) data for the type series of Cnemaspis krishnagiriensis sp. nov. Abbreviations are listed in Materials and Methods. * = tail incomplete

Type	Holotype	Paratype
Specimen no.	NRC-AA-1120	NRC-AA-1121	BNHS 2798
Sex	Male	Female	Female
SVL	40.0	39.5	27.2
TL	28.4*	50.1	32.0
TW	3.9	3.3	2.2
LAL	7.0	7.2	4.4
CL	9.1	9.3	5.6
AGL	16.0	15.1	10.3
BH	3.4	4.0	1.9
BW	6.5	7.2	4.6
HL	10.1	9.8	7.1
HW	6.2	6.0	4.3
HD	3.9	3.7	2.8
ED	2.3	2.2	1.5
EE	3.1	3.0	2.2
ES	5.1	5.0	3.6
EN	4.1	4.1	2.8
IN	1.0	1.0	0.8
IO	2.5	2.5	1.7
EL	0.7	0.6	0.4

TABLE 4. Meristic data for type series of Cnemaspis krishnagiriensis sp. nov. Abbreviations are listed in Materials and Methods. L & R = Left & Right; abs. = Absent.

Type	Holotype	Paratypes
Specimen number	NRC-AA-1120	NRC-AA-1121	BNHS 2798
SL L & R	8 & 8	8 & 8	9 & 8
IL L & R	7 & 7	7 & 7	7 & 7
SL M L & R	7 & 7	7 & 7	7 & 6
IL M L & R	6 & 6	5 & 5	5 & 6
PVT	14	14	13
DTR	8	8	7
MVSR	24	23	25
VS	123	123	120
LamF1 L & R	13 & 13	13 & 13	12 & 12
LamF4 L & R	23 & 22	22 & 22	20 & 20
LamT1 L & R	12 & 13	13 & 13	12 & 12
LamT4 L & R	24 & 25	25 & 25	24 & 22
LamT5 L & R	24 & 23	21 & 23	22 & 22
PP	3	abs.	abs.
FP L & R	4 & 4	abs.	abs.
SB PP&FP L & R	10 & 9	abs.	abs.
Number of postcloacal tubercles L & R	1 & 1	1 & 1	1 & 1
Anterior extra-brillar fringe scales enlarged (1) or not enlarged (0)	1	1	1
Ventral scales keeled (1) or smooth (0)	0	0	0
Gular scales keeled (1) or smooth (0)	0	0	0
Pectoral scales keeled (1) or smooth (0)	0	0	0
Precloacal pores continuous (1) or separated (0)	1	abs.	abs.
Precloacal pores elongate (1) or round (0)	1	abs.	abs.
femoral pores elongate (1) or round (0)	1	abs.	abs.
Dorsal pholidosis homogeneous (1) or heterogeneous (0)	0	0	0
Dorsal tubercles keeled (1) or not keeled (0)	1	1	1
Tubercles linearly arranged (1) or more random (0)	1	1	1
Spine-like tubercles on flank present (1) or absent (0)	0	0	0
Lateral caudal furrows present (1) or absent (0)	0	0	0
Sub-caudals keeled (1) or smooth (0)	0	0	0
Single median row of sub-caudals keeled (1) or smooth (0)	0	0	0
Caudal tubercles encircle tail (1) or not (0)	0	0	0
Enlarged median sub-caudal scale row (1) or not (0)	1	1	1
Enlarged femoral scales present (1) or absent (0)	0	0	0
Subtibial scales keeled (1) or smooth (0)	0	0	0

Distribution and natural history. Cnemaspis krishnagiriensis sp. nov. is so far known only from a hillock behind Kugai Periya Mariamman Temple, near Krishnagiri fort, Krishnagiri District, Tamil Nadu , at an elevation of ca. 550 m asl. ( Figure 1 ). The species was encountered on a granite hillock with large boulders, predominantly covered by sparse scrub vegetation ( Figure 7 ). Like other members of its clade, the new species seems to be diurnal as we observed individuals coming out of their hiding spots in the morning (0730–0845 hrs.) All three individuals were observed on the underside of granite boulders that formed a cave at a height of ~ 30–120 cm above the ground. Sympatric geckos at the type locality include Cnemaspis cf. otai , Calodactylodes aureus (Beddome) , Hemidactylus graniticolus Agarwal, Giri & Bauer , and H. giganteus Stoliczka. Cnemaspis cf. otai is largely crepuscular and found either at the base of boulders (< 30 cm above the ground) or under rocks on the ground, and is also considerably smaller in size (SVL < 32 mm ). The other three sympatric geckos are all much larger (SVL 40–120 mm ) and are strictly nocturnal. FIGURE 7. Habitat photo of Cnemaspis krishnagiriensis sp. nov. from near Kugai Periya Mariamman Temple, Krishnagiri fort, Krishnagiri District, Tamil Nadu, India; photos by Akshay Khandekar. Nomenclatural note on the recently described Cnemaspis stellapulvis Khandekar, Thackeray & Agarwal. Cnemaspis stellapulvis is a recently described species, with the original etymology stating that the name was an adjective formed from the Latin words stella and pulvis (Khandekar et al. 2020). This was an error made by the original authors, and the name should be treated as a compound noun in apposition.