Cnemaspis anaimalaiensis, Khandekar & Thackeray & Agarwal, 2024

Khandekar, Akshay, Thackeray, Tejas & Agarwal, Ishan, 2024, Five new species of the Cnemaspis beddomei clade (Squamata: Gekkonidae) from high elevation, evergreen forests of the Southern Western Ghats, India, Zootaxa 5469 (1), pp. 1-70 : 33-43

Cnemaspis anaimalaiensis sp. nov.

urn:lsid:zoobank.org:act:FB148E29-1071-4113-80B6-0AA8F6429C87

( Figs 15–19 ; Tables 9 –11)

Holotype. NRC-AA-8392 ( AK-R 1948 ), adult male, near Kamatchi Amman Temple (10.3901°N, 76.9970°E; ca. 1410 m asl.), on Pollachi to Valparai road, Anaimalai Tiger Reserve , Coimbatore District, Tamil Nadu State, India; collected by Akshay Khandekar, Ishan Agarwal, Swapnil Pawar and team on 18 th May 2022.

Paratypes. NRC-AA-8393 ( AK-R 1949 ), NRC-AA-8394 ( AK-R 1950 ), adult females, same data as holotype ; NRC-AA-8395 ( AK-R 1951 ), NRC-AA-8397 ( AK-R 1986 ), adult males, NRC-AA-8396 ( AK-R 1952 ), adult female, elsewhere near Kamatchi Amman Temple (10.3823°N, 76.9983°E; ca. 1500 m asl.) ; NRC-AA-8398 (AK- R 1987), NRC-AA-8400 ( AK-R 1989 ), adult males, NRC-AA-8399 ( AK-R 1988 ), adult female, from Pollachi to Valparai road (10.3971°N, 76.9918°E; ca. 1300 m asl.); same collection data as holotype .

Etymology. The specific epithet is a toponym for the Anaimalai Hills, the mountainous landscape that contains the type locality of the new species.

Suggested common name. Anaimalai dwarf gecko.

Diagnosis. A medium-sized Cnemaspis , snout to vent length up to 50 mm (n = 9). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with irregularly arranged rows of enlarged, weakly keeled, conical tubercles; 10–13 rows of dorsal tubercles at midbody, tubercles in paravertebral rows irregular; ventral scales subequal from chest to vent, smooth, subcircular and subimbricate with weakly pointed end; 28–33 scales across belly at midbody, 155–168 longitudinal scales from mental to cloaca; subdigital scansors smooth, unnotched, some divided and others entire; 12–15 lamellae under digit I of manus and 12–17 under digit I of pes, 18–21 lamellae under digit IV of manus and 20–25 lamellae under digit IV of pes; males with continuous series of two precloacal pores (n = 3/5) but sometime precloacal pores separated by single poreless scale (n = 2/5); scales on non-regenerated tail dorsum homogeneous; composed of enlarged, smooth, slightly elongate, flattened, imbricate scales that are becoming slightly more pronounced on lateral sides; median row of subcaudals smooth and slightly enlarged. Dorsum khaki with a discontinuous straw-yellow to orange mid-vertebral streak and about seven dark markings, each made up of a pair of black longitudinal streaks; tail with alternating black and light grey spots; no apparent sexual dimorphism.

Comparisons with members of beddomei clade. Cnemaspis anaimalaiensis sp. nov. can be easily distinguished from all described members of the beddomei clade as well as from C. boiei by a combination of the following differing or non-overlapping characters: males with continuous series of two precloacal pores, but sometime precloacal pores separated by single poreless scale (versus males with continuous series of seven or eight precloacal pores in C. aaronbaueri , C. nairi , C. sathuragiriensis and in C. smaug , 6–8 precloacal pores in C. agamalaiensis sp. nov., C. azhagu , C. beddomei , C. regalis and C. rubraoculus , eight or nine in C. anuradhae sp. nov. and C. sundara , seven in C. galaxia , 10 in C. maculicollis , six or seven in C. nigriventris , C. ornata and C. vangoghi , 4–6 in C. nimbus , 7–11 in C. rashidi and eight in C. wallaceii ); a medium-sized Cnemaspis , snout to vent length up to 50 mm (versus small-sized Cnemaspis , snout to vent length <40 mm in C. aaronbaueri , C. azhagu , C. boiei , C. galaxia , C. nigriventris , C. regalis , C. sathuragiriensis , C. sundara and C. vangoghi ; large-sized Cnemaspis , snout to vent length> 50 mm, 58 mm in C. anamudiensis , 53 mm C. beddomei , 53 mm C. maculicollis and 56 mm C. smaug ; maximum SVL 46 mm in C. ornata and 43 mm in C. nairi ); scales on non-regenerated tail dorsum homogeneous, lacking enlarged tubercles (versus scales on non-regenerated tail dorsum heterogeneous, with the presence of enlarged tubercles in C. aaronbaueri , C. azhagu , C. galaxia , C. nairi , C. nigriventris , C. ornata , C. rashidi , C. regalis , C. rubraoculus , C. sathuragiriensis , C. smaug , C. sundara , C. vangoghi and C. wallaceii ); median row of subcaudals smooth and slightly enlarged (versus median row of subcaudals smooth and slightly enlarged with condition of two slightly enlarged scales alternating with a divided scale in C. aaronbaueri , C. azhagu , C. beddomei , C. galaxia , C. nairi , C. nigriventris , C. ornata , C. regalis , C. sathuragiriensis , C. smaug , C. vangoghi , median row of subcaudals smooth and enlarged with irregularly arranged scales in C. maculicollis , and median row with individual large scale alternating with slightly smaller scale in C. nimbus ); enlarged tubercles on midbody dorsum irregularly arranged (versus enlarged tubercles on midbody dorsum absent in C. boiei , enlarged tubercles on midbody dorsum regularly arranged in C. aaronbaueri , C. azhagu , C. galaxia , C. maculicollis , C. nairi , C. nigriventris , C. ornata , C. rashidi , C. regalis , C. smaug , C. vangoghi , C. wallaceii ); 155–168 longitudinal scales from mental to cloaca (versus 135– 141 longitudinal scales from mental to cloaca in C. aaronbaueri , 137–149 in C. agamalaiensis sp. nov., 140–156 in C. anuradhae sp. nov., 117 in C. boiei , 143–147 in C. nairi , 134–141 in C. nimbus , 170–172 in C. rashidi , 148–154 in C. regalis , 122–146 in C. rubraoculus , 130–137 in C. sathuragiriensis , 141–151 in C. smaug and 125–140 in C. vangoghi ); tubercles in paravertebral rows irregular (versus 16–24 tubercles in paravertebral rows in C. agamalaiensis sp. nov., 20–25 in C. anuradhae sp. nov., 14 in C. galaxia , 16 or 17 in C. nimbus , 21–23 in C. ornata , 12 in C. regalis , 12–17 in C. rubraoculus , 21–27 in C. smaug , 7–14 in C. vangoghi and 18–20 in C. wallaceii ); 10–13 rows of dorsal tubercles at midbody (versus only a few enlarged scattered tubercles at midbody dorsum in C. anamudiensis , two or three rows of dorsal tubercles at midbody in C. azhagu , eight in C. galaxia , 16–18 in C. nairi , 7–9 in C. regalis , 6–8 in C. sathuragiriensis , 18–20 in C. smaug , six in C. sundara and 14 or 15 in C. wallaceii ); 28–33 scales across belly at midbody (versus 22–26 in C. agamalaiensis sp. nov.,> 38 in photographs of the holotype of C. anamudiensis , 34–44 in C. azhagu , 38–40 in C. nigriventris , 26 or 27 in C. nimbus , 40–44 in C. regalis and 35 or 36 in C. sundara ); ventral scales smooth (versus ventral scales weakly keeled in C. beddomei and C. rubraoculus ); a distinct white ocellus on ventrolateral sides of neck absent in both sexes (versus a distinct white ocellus on ventrolateral sides of neck present in males in C. galaxia , C. nairi , C. nigriventris , C. ornata , C. rashidi , C. sathuragiriensis , C. sundara and C. vangoghi ); a distinct of white ocelli or streak on shoulder and behind the forearm insertions absent (versus a distinct white ocelli or streak on shoulder and behind the forearm insertion present in C. smaug , and a distinct white ocelli or streak on occiput, neck and shoulder present in C. maculicollis ). Cnemaspis anaimalaiensis sp. nov. is diagnosed against the other new species as part of their descriptions below.

Description of the holotype. Adult male in good state of preservation except a continuous fold of skin on vertebral region from behind the neck to hind limb insertion, tail tip bent towards left, a 6.3 mm and 2.5 mm longitudinal skin injury on flank and anterolateral thigh respectively on the left side, and a 3.8 mm long incision in sternal region for tissue collection ( Fig. 15A–C ). SVL 43.2 mm, head short (HL/SVL 0.26), wide (HW/HL 0.72), not strongly depressed (HD/HL 0.43), distinct from neck. Loreal region marginally inflated, canthus rostralis indistinct. Snout half of head length (ES/HL 0.46), more than two times eye diameter (ES/ED 2.5); scales on snout and canthus rostralis elongate, subequal, weakly keeled, strongly conical, much larger than those on forehead and interorbital region; scales on forehead similar to those on snout and canthus rostralis except slightly smaller; scales on interorbital region even smaller, granular; scales on occipital, and temporal region heterogeneous, composed of weakly keeled granular scales intermixed with enlarged, weakly keeled, conical tubercles ( Fig. 16A ). Eye small (ED/HL 0.18), with round pupil; supraciliaries short, largest anteriorly; 13 interorbital scale rows across narrowest point of frontal; 40 scale rows between left and right supraciliaries at mid-orbit ( Fig. 16A, C ). Ear-opening deep, oval, small (EL/HL 0.06); eye to ear distance almost two times greater than diameter of eye (EE/ED 1.95) ( Fig. 16C ). Rostral much wider (2.0 mm) than high (1.3 mm), incompletely divided dorsally by a strongly developed rostral groove for more than half of its height; a single enlarged, roughly rectangular supranasal on each side, more than two times larger than upper postnasal, and in contact with each other behind rostral but separated posteriorly but two or three much smaller scales on the snout; internasal scales absent; rostral in contact with supralabial I, lower postmental, nostril, and supranasal on either side; nostrils oval, surrounded by two postnasals, supranasal, and rostral on either side; two enlarged postnasals on either side, upper one largest, roughly circular; lower postnasal much elongate, slightly smaller than upper postnasal; two single row of scales separate orbit from supralabials ( Fig. 16C ). Mental enlarged, subtriangular, slightly wider (2.8 mm) than high (2.2 mm); two pairs of postmentals, inner pair roughly rectangular, much shorter (0.9 mm) than mental, separated from each other below mental by a single enlarged median chin shield; inner pair bordered by mental, infralabial I, outer postmental, median chin shield and two enlarged chin shields on either side; outer postmentals roughly rectangular, square smaller (0.6 mm) than inner pair, bordered by inner postmentals, infralabial I & II, and four enlarged chin shields on left and three on right side; five enlarged gular scales (including median chin shield) between left and right outer postmentals; all chin shields bordering postmentals protrudent, subcircular, smooth, and smaller than outermost postmentals; scales on rest of throat granular, much smaller, conical, weakly keeled and subcircular, ( Fig. 16B ). Infralabials bordered below by a row or two of slightly enlarged, much elongated scales, decreasing in size posteriorly. Ten supralabials on left and 11 on right side up to angle of jaw and six at midorbital position on each side; supralabial I largest, gradually decreasing in size posteriorly; nine infralabials on left and eight on right side up to angle of jaw, and five at midorbital position on either side; infralabial I largest, gradually decreasing in size posteriorly ( Fig. 16C ).

Body relatively slender (BW/AGL 0.44), trunk slightly less than half of SVL (AGL/SVL 0.40) without spinelike tubercles on flank ( Fig. 17A–C ). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with irregularly arranged rows of enlarged, weakly keeled, conical tubercles; granular scales gradually increasing in size towards each flank, largest on mid-flank; granular scales on occiput and temporal region much smaller than paravertebral granules; enlarged tubercles in approximately 13 longitudinal rows at midbody; tubercles in paravertebral rows irregular ( Fig. 17A ). Ventral scales much larger than granular scales on dorsum, subequal from chest to vent, smooth, subcircular and subimbricate with weakly pointed end; scales on precloacal region and four or five rows on lower thigh distinctly enlarged; midbody scale rows across belly 28; 168 scales from mental to anterior border of cloaca ( Fig. 17B ). Two precloacal pores, femoral pores absent ( Fig. 16D ).

Scales on palm and soles, smooth, subcircular, flattened and subimbricate; scales on dorsal aspects of limbs homogenous; composed of small, weakly keeled, weakly conical, subimbricate scales; scales on ventral aspect of upper arm smooth, granular, much smaller than granular scales on body dorsum, scales on ventral aspect of lower arm with much larger scales than those on upper arm, weakly keeled, subcircular, weakly pointed, and subimbricate; ventral aspect of thigh and shank with enlarged, smooth, flattened, subimbricate scales, much larger than body ventrals ( Fig. 15A, B ). Forelimbs and hindlimbs 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. Digits with both paired and unpaired lamellae, separated into a basal and narrower distal series by single enlarged lamella at inflection; one or two most basal paired on basal series and 1–4 paired lamellae above the inflection; basal lamellae series: (4-5-4-5-5 right manus, Fig. 16E ; 2-7-6-8-4 right pes, Fig. 16F ), (4-4-5-5-5 left manus, 3-6-7-9-3 left pes); distal lamellae series: (10-12-15-15-13 right manus, Fig. 16E; 10-12-16-15-15 right pes, Fig. 16F ), (10-12-15-14-12 left manus, 11-13-16-16-15 left pes). Relative length of digits (measurements in mm in parentheses): IV (4.9)> III (4.6)> V (4.5)> II (4.2)> I (3.2) (right manus); IV (6.2)> V (5.3) = III (5.3)> II (4.2)> I (2.8) (right pes).

Tail original, subcylindrical, slender, entire, almost equal to body length (TL/SVL 1.03) ( Fig. 15A–C ). Dorsal pholidosis on tail homogeneous; composed of enlarged, smooth, slightly elongate, flattened, imbricate scales that are becoming slightly more pronounced on lateral sides ( Fig. 15A ). Scales on tail venter much larger than those on dorsal aspect, smooth, roughly subcircular, flattened, subimbricate; median series smooth, slightly enlarged than rest ( Fig. 15B ). Scales on tail base much smaller, smooth, subimbricate; a single, enlarged smooth postcloacal tubercle on either side ( Fig. 16D ).

Colouration in life ( Fig. 18A ). Dorsal ground colouration of head, body, limbs and tail khaki. Indistinct dark preorbital streak and dark postorbital streak runs till ear opening; head with few dark markings except a small black longitudinal streak on occiput flanked by two small straw yellow spots; nape with a pair of similar black streaks with a single straw-yellow spot in the middle. Dorsum with about seven similar sets of dark markings till hindlimb insertions and 13 dark markings on tail, discontinuous straw-yellow vertebral streak alternates with dark markings and continues onto tail as fine spots. Limbs with peach spots, more prominent on femur and digits of pes. Ventral colouration dirty white; throat with a few rows of dark scales bordering infralabials; subcaudals with some dark grey or black scale edges; precloacal and femoral regions cream. Pupil black, iris red.

Variation and additional information from the paratype series ( Figs 18A , 19A, B ). Mensural, meristic and additional character state data for the type series is given in Tables 9 –11 respectively. There are four adult males and four adult females ranging in size from 41.0– 49.6 mm ( Fig. 19A, B ). All paratypes resemble the holotype except as follows: supranasals separated from each other behind rostral by much smaller one or two internasals in all the paratypes except for NRC-AA-8393. Inner postmentals bordered by mental, outer postmental, infralabial I and an enlarged median chin shield in all paratypes (except for NRC-AA-8395 and NRC-AA- 8398 in which enlarged median chin shield is not bordering inner postmental), additionally, bordered by a single chin shield on either side in NRC-AA-8394 and NRC-AA-8400; inner postmental separated from each other below mental by an enlarged median chin shield in all paratypes, additionally separated by two smaller chin shields in NRC-AA-8395 and NRC- AA-8398. Outer postmentals bordered by inner pair and infralabial I & II in all paratypes, additionally, bordered by four small chin shields on either side in NRC-AA-8394, NRC-AA-8399 and NRC-AA-8400, three on either side in NRC-AA-8397 and NRC-AA-8398; outer postmentals separated from each other by an enlarged median chin shield in all paratypes, additionally, by two smaller chin shields in NRC-AA-8394 and NRC-AA-8400, five in NRC- AA-8395. Four paratypes —NRC-AA-8393, NRC-AA-8394, NRC-AA-8396 and NRC-AA-8399 with original and complete tail, slightly smaller or marginally longer than body (TL/SVL 0.81, 1.02, 0.90 and 1.15 respectively), NRC-AA-8397 with complete tail but tip regenerated, slightly shorter than body (TL/SVL 0.77); NRC-AA-8395 and NRC-AA-8398 with complete but fully regenerated tail, slightly shorter than body (0.72 and 0.73 respectively); tail completely missing with minor regenerated growth in NRC-AA-8400; NRC-AA-8398 with partially everted hemipenis on either side ( Fig. 19A, B ). The mid-vertebral streak varies in colour from straw-yellow to orange or red and is continuous on the forebody of NRC-AA-8395, NRC-AA-8396 and NRC-AA-8397, and is least prominent in NRC-AA-8400. No apparent sexual dimorphism ( Fig. 18B ).

Distribution and natural history. Cnemaspis anaimalaiensis sp. nov. is known only from around its type locality (Pollachi to Valparai road, Anaimalai Tiger Reserve, Coimbatore District, Tamil Nadu) ( Fig. 1 ). The new species was recorded in evergreen forest patches at elevations of 1300–1500 m asl. ( Fig. 20A ) active during the daytime (0930–1100 hrs) on rocks <3 m high from the base in dense forest patches ( Fig. 20B ). During a heavy downpour at night (2030 hr), we also observed this species inactive on the same rocks. During summer, a large number of sightings (n => 15) were obtained at all the three sampled locations indicating high abundance. Individuals of the new species were observed to be relatively sluggish when approached. The only other lizard sympatric with the new species we found was Cnemaspis sp. (another new species described below) at the type locality.

TABLE 11. Additional morphological characters of Cnemaspis anaimalaiensis sp. nov. M = male, F = female, A =

absent, / = data unavailable.

Figures

FIGURE 1. Elevation map of the southern Western Ghats, India showing sampled localities for the anamudiensis (triangles) and beddomei (circles) subclades of the beddomei clade.

FIGURE 2. Maximum Likelihood phylogeny of the beddomei clade of South Asian Cnemaspis based on concatenated ND2 and 16S data with subclades marked by horizontal lines. Numbers at nodes indicate bootstrap support/ posterior probability (values <85/ 0.99 not shown), outgroups not shown.

FIGURE 3. Cnemaspis agamalaiensis sp. nov. (holotype, NRC-AA-8374): (A) dorsal view of body, (B) ventral view of body, (C) dorsal view of tail, (D) ventral view of tail, (E) lateral view of tail. Scale bars 10 mm; photos by Akshay Khandekar.

FIGURE 4. Cnemaspis agamalaiensis sp. nov. (holotype, NRC-AA-8374): (A) dorsal view of head, (B) ventral view of head, (C) lateral view of head on right, (D) view of precloacal region showing precloacal pores, (E) ventral view of right manus, (F) ventral view of right pes. Scale bars 5 mm; photos by Akshay Khandekar.

FIGURE 5. Cnemaspis agamalaiensis sp. nov. (holotype, NRC-AA-8374): (A) dorsal view of midbody, (B) ventral view of midbody, (C) lateral view of midbody. Scale bars 10 mm; photos by Akshay Khandekar.

FIGURE 6. Cnemaspis agamalaiensis sp. nov. in life: (A) adult male (holotype, NRC-AA-8374), (B) adult male (paratype, NRC-AA-8381), and (C) subadult female (paratype, NRC-AA-8376). Photos by Akshay Khandekar.

FIGURE 7. Paratype series of Cnemaspis agamalaiensis sp. nov., from left to right, NRC-AA-8375–8384: (A) dorsal view and (B) ventral view. Scale bars 10 mm; photos by Akshay Khandekar.

FIGURE 8. Habitat photos of Cnemaspis agamalaiensis sp. nov.: (A) general view of evergreen forest and (B) microhabitat photo showing the boulder located inside the evergreen forest patch from where the new species was collected; both from the type locality, Agamalai in Theni District, Tamil Nadu. Photos by Akshay Khandekar.

FIGURE 15. Cnemaspis anaimalaiensis sp. nov. (holotype, NRC-AA-8392): (A) dorsal view of body, (B) ventral view of body, (C) lateral view of tail. Scale bars 10 mm; photos by Akshay Khandekar.

FIGURE 16. Cnemaspis anaimalaiensis sp. nov. (holotype, NRC-AA-8392): (A) dorsal view of head, (B) ventral view of head, (C) lateral view of head on right, (D) view of precloacal region showing precloacal pores, (E) ventral view of right manus, (F) ventral view of right pes. Scale bars 5 mm; photos by Akshay Khandekar.

FIGURE 17. Cnemaspis anaimalaiensis sp. nov. (holotype, NRC-AA-8392): (A) dorsal view of midbody, (B) ventral view of midbody, (C) lateral view of midbody. Scale bars 10 mm; photos by Akshay Khandekar.

FIGURE 18. Cnemaspis anaimalaiensis sp. nov. in life: (A) adult male (holotype, NRC-AA-8392) and (B) adult female (paratype, NRC-AA-8393). Photos by Akshay Khandekar.

FIGURE 19. Paratype series of Cnemaspis anaimalaiensis sp. nov., from left to right, NRC-AA-8393–8400: (A) dorsal view and (B) ventral view. Scale bars 10 mm; photos by Akshay Khandekar.

FIGURE 20. Habitat photos of Cnemaspis anaimalaiensis sp. nov. and C. valparaiensis sp. nov.: (A) general view of evergreen forest and (B) microhabitat photo showing the boulders located inside the evergreen forest patch from where the new species were collected; both from the type locality, near Valparai in Anaimalai Tiger Reserve, Coimbatore District, Tamil Nadu. Photos by Akshay Khandekar.