Cnemaspis kalakadensis, Khandekar & Thackeray & Agarwal, 2022

Khandekar, Akshay, Thackeray, Tejas & Agarwal, Ishan, 2022, Three more novel species of South Asian Cnemaspis Strauch, 1887 (Squamata, Gekkonidae) from Kalakad Mundanthurai Tiger Reserve, Tamil Nadu, India, Vertebrate Zoology 72, pp. 385-422 : 385

Cnemaspis kalakadensis sp. nov.

(Figs 15 , 16 , 17 , 18 , 19 , 20

Holotype.

NRC-AA-1180 (AK-R 648), adult male, from near Sengaltheri forest guesthouse, Kalakad forest range (8.5340°N, 77.4502°E; ca. 960 m asl.), Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu state, India; collected by Akshay Khandekar, and team on 30 April 2021.

Paratypes.

NRC-AA-1182 (AK-R 644), BNHS 2826 (AK-R 646), adult males; NRC-AA-1181 (AK-R 643), NRC-AA-1183 (AK-R 645), BNHS 2827 (AK-R 647), BNHS 2828 (AK-R 654), adult females. AK-R 644 and AK-R 646 collected from same locality and other details as holotype; AK-R 643 and AK-R 645 collected from 8.5379°N, 77.4536°E, ca. 910 m asl.; AK-R 647 and AK-R 654 collected from 8.5290°N, 77.4467°E, ca. 1060 m asl; all from Kalakadu reserve forest, Kalakad forest range, Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu state, India, collectors and date same as holotype.

Etymology.

The specific epithet is a toponym for the Kalakad forest range of Kalakad-Mundanthurai Tiger Reserve in Tirunelveli district of Tamil Nadu, the type and currently only known locality for this species.

Suggested Common Name.

Kalakad dwarf gecko.

Diagnosis.

A small-sized Cnemaspis , snout to vent length less than 33 mm (n =7). Dorsal pholidosis heterogeneous; smooth, subcircular, weakly conical granular scales intermixed with irregularly arranged rows of enlarged, smooth, laterally compressed, spine-like scales on flanks; four or five rows of dorsal tubercles at mid-body, enlarged scales or tubercles absent in paravertebral region; ventral scales subcircular, smooth, subimbricate, subequal from chest to vent; 28-34 scales across belly at mid-body, 122-134 longitudinal scales from mental to cloaca; subdigital scansors smooth, unpaired, unnotched; basal scansors distinctly enlarged, plate like; 11 or 12 lamellae under digit I of manus and pes, 16-19 lamellae under digit IV of manus and 19-22 lamellae under digit IV of pes; males (n =3/7) with a series of 12-14 femoral pores on each thigh separated medially by 10 or 11 poreless scales; tail dorsum with enlarged, smooth, flattened scales only in the paravertebral rows, not forming whorls; 16-18 scales in paravertebral rows on either side, only 2-4 enlarged scales in the lateral row on either side; median row of subcaudals smooth, roughly subcircular, and distinctly enlarged. Dorsum grey-brown with indistinct dark bands; single longitudinally placed hourglass shaped black marking anterior to forelimb insertions; tail with ~12 alternating dark and pale bands; underside of body, limbs and tail in males pale yellow with precloacal and femoral region off-white, gular region bright yellow; ventral surfaces in females off-white to cream.

Comparison with members of C. littoralis clade.

Cnemaspis kalakadensis sp. nov. is a member of the Cnemaspis littoralis clade and can be easily distinguished from all four members of the clade by combination of following differing or non-overlapping characters: four or five rows of dorsal tubercles at mid-body restricted to flanks, laterally compressed, spine-like (versus few scattered tubercles on the flank, reduced, subconical in C. flavigularis ; a few scattered tubercles on flanks, spine-like tubercles only in the lowermost row on flanks in C. littoralis ; conical or spine-like tubercles absent on flanks in C. palakkadensis ; a few scattered spinose tubercles on the flanks in C. palanica Pal, Mirza, Dsouza and Shanker, 2021); 28-34 scales across belly at mid-body (versus 21-23 scales across belly at mid-body in C. flavigularis , and 16-18 in C. palanica ); 122-134 longitudinal scales from mental to cloaca (versus 104-108 longitudinal scales from mental to cloaca in C. flavigularis , 143-157 in C. littoralis , and 103-106 in C. palanica ); 19-22 lamellae under digit IV of pes (versus 16-18 lamellae under digit IV of pes in C. flavigularis , 16-18 in C. littoralis , 14-17 in C. palakkadensis , and 17 or 18 in C. palanica ); males with a series 12-14 femoral pores on each thigh separated medially by 10 or 11 poreless scales (versus males with a series of 15-18 femoral pores separated by 14-16 poreless scales in C. littoralis , and 15 or 16 femoral pores separated by at least 15 poreless scales in C. palakkadensis ).

Description of the holotype.

Adult male in good state of preservation except head marginally bent towards right and fully everted hemipenis on left side (Fig. 15A, B ) and digit V of left manus incomplete (indicated by *). SVL 29.6 mm, head short (HL/SVL 0.25), slightly wider (HW/HL 0.61), not strongly depressed (HD/HL 0.38), distinct from neck. Loreal region marginally inflated, canthus rostralis indistinct. Snout marginally shorter than half the head length (ES/HL 0.46), 2.5 times larger than eye diameter (ES/ED 2.50); scales on snout and canthus rostralis smooth, subcircular, subequal, hemispherical and protrudent, slightly larger than those on forehead and almost twice the size than those on interorbital region and occiput; scales on forehead similar to those on snout and canthus rostralis except slightly smaller and weakly protrudent; scales on interorbital region, occipital and temporal region smooth, still smaller, granular (Fig. 16A ). Eye small (ED/HL 0.18); with round pupil; supraciliaries short, larger anteriorly; seven interorbital scale rows across narrowest point of frontal bone; 28 or 29 scale rows between left and right supraciliaries at mid-orbit (Fig. 15A ). Ear-opening deep, oval, small (EL/HL 0.05); eye to ear distance greater than diameter of eye (EE/ED 1.78) (Fig. 16C ). Rostral marginally more than twice as wide (1.3 mm) as high (0.6 mm), incompletely divided dorsally by a strongly developed rostral groove for more than half of its height; a single enlarged supranasal on each side, almost twice the size of upper postnasal, separated from each other by a smaller, elongated internasal on the snout; rostral in contact with supralabial I, nostril, supranasal, internasal and a lower postnasal on either side; nostrils subcircular, surrounded by two postnasals, supranasal, and rostral on either side; two postnasals on either side, similar in size to each other, both almost half the size than supranasal; one or two rows of scales separate orbit from supralabials (Fig. 16C ). Mental enlarged, subtriangular, slightly wider (1.7 mm) than high (1.2 mm); two pairs of postmentals, inner pair roughly rectangular, much shorter (0.5 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 and three enlarged chin shields on left side and bordered by mental, infralabial I & II, outer postmental and three enlarged chin shields on right side; outer postmentals roughly square, much smaller (0.3 mm) than inner pair, bordered by inner postmentals, infralabial I, and two enlarged chin shields on left side and bordered by inner postmentals, infralabial I & II, and two enlarged chin shields on right side; five enlarged, gular scales between left and right outer postmentals; all chin scales bordering postmentals hemispherical and protrudent, subcircular, smooth, smaller or equal in size to outermost postmentals; scales on rest of the gular with smooth, subcircular, hemispherical, and protrudent scales decreasing in size posteriorly (Fig. 16B ). Infralabials bordered below by a row or two of slightly enlarged, somewhat circular scales, decreasing in size posteriorly and laterally. Eleven supralabials up to angle of jaw on left and 10 on right side and eight at midorbital position on each side; supralabial I largest, gradually decreasing in size posteriorly; 10 infralabials up to angle of jaw on left side and nine or right, and seven at midorbital position on left and six on right side; infralabial I largest, gradually decreasing in size posteriorly (Fig. 16C ).

Body relatively slender (BW/AGL 0.40), trunk less than half of SVL (AGL/SVL 0.42) without ventrolateral folds (Fig. 17B, C ). Dorsal pholidosis heterogeneous; smooth, subcircular, weakly conical granular scales intermixed with irregularly arranged rows of enlarged, smooth, laterally compressed, spine-like scales on flank; approximately five longitudinal rows spine-like scales at mid-body; enlarged scales absent on paravertebral region; granular scales on nape similar in size to those on paravertebral region, slightly smaller on occiput (Fig. 17A ). Ventral scales much larger than granular scales on dorsum, subcircular, smooth, subimbricate, subequal from chest to vent; mid-body scale rows across belly 30; 122 scales from mental to anterior border of cloaca (Fig. 15B ). Scales on throat marginally smaller than those on belly, gular region with still smaller, smooth and protrudent scales, those on chin bordering postmentals, enlarged, smooth, subcircular, and protrudent (Fig. 16B ). A series of fourteen femoral pores each thigh, separated from each other 11 poreless scales (Fig. 16D ). Scales on palm and soles small, smooth, subimbricate; scales on dorsal aspects of limbs heterogeneous is size, those on upper arm and thigh larger, smooth, subcircular and subimbricate except those near limb insertions are much smaller, smooth and granular; posterolateral aspect of thigh with small, and smooth granular scales; dorsal aspect of lower arm and shank with scales smaller than those on upper arm and thigh, smooth and subimbricate; ventral aspect of upper arm with small, granular scales, lower arm with slightly larger, smooth, subimbricate scales; ventral aspect of thigh with scales similar to mid-body ventrals, scales on shank marginally larger than those on thigh, smooth, flattened and subimbricate (Fig. 15A, B ).

Forelimbs and hindlimbs slightly long, slender (LAL/SVL 0.13); (CL/SVL 0.16); digits long, with a strong, recurved claw, distinctly inflected, distal portions laterally compressed.

Digits with unpaired lamellae (except 1-3 paired lamellae at the base), separated into a basal and narrower distal series by single enlarged lamella at inflection, basal lamellae much enlarged, plate like; basal lamellae series: (2-3-3-4-4 right manus, 2-4-4-6-5 right pes), (2-3-3-4-4 left manus, Fig. 16E ; 2 - 5-5 - 6 - 5 left pes, Fig. 16F ); distal lamellae series: (9-9-12-12-10 right manus, 9-11-14-13-13 right pes), (9-10-12-12-10* left manus, Fig. 16E ; 9 - 10 - 13-13-13 left pes, Fig. 16F ). Relative length of digits (measurements in mm in parentheses): IV (2.5)> III (2.4)> V (2.0) = II (2.0)> I (1.5) (left manus); IV (3.5)> V (3.1) = III (3.1)> II (2.7)> I (1.5) (left pes).

Tail original, entire, subcylindrical, slender, marginally longer than snout-vent length (TL/SVL 1.12; Fig. 15C-E ). Dorsal pholidosis on tail heterogeneous; small, smooth, flattened, subcircular and subimbricate scales (becoming slightly larger and elongated posteriorly) intermixed with enlarged, smooth, flattened scales only in the paravertebral rows, not forming whorls; 16 scales in paravertebral rows on either side, only two enlarged scales in the lateral row on either side (Fig. 15C, E ). Scales on tail venter much larger than those on tail dorsum, smooth, flattened, and sub-imbricate; with a series of three enlarged subcaudal scales of which the median series almost twice the size of adjunct two rows, roughly subcircular (Fig. 15D ). Scales on tail base small, smooth and subimbricate; a single enlarged, smooth, weakly conical postcloacal spur on each side (Fig. 15D, E ).

Colouration in life (Fig. 18A).

Dorsal ground colour of head, body, limbs and tail pale grey-brown; finely speckled with darker markings. Anterior edge of brille and snout tip faintly suffused with yellow; labials dull white to yellow, finely speckled with dark markings, dark markings prominent on outer margins. Indistinct dark brown preorbital streak runs from nostril to orbit, postorbital streaks indistinct. A single longitudinally placed, hourglass-shaped black marking anterior to forelimb insertions, flanked by a poorly defined lighter blotch anteriorly and posteriorly. About four poorly defined dark bands between axilla and tail base of which the first is most prominent, narrowest in vertebral region and flaring out on flanks, alternating with lighter, elongate mid-dorsal blotches; dorsum of forelimbs, hindlimbs and digits mottled with small dark blotches and bands; dorsum of original tail with about 12 alternating dark and pale bands. Underside of body, limbs and tail pale yellow with a few scattered grey scales, only precloacal and femoral region off-white; gular region almost entirely covered by a bright yellow patch mottled with a few black scales and very few larger grey blotches. Pupil black, iris greyish.

Variation and additional information from type series.

Mensural, meristic and additional character state data for the type series is given in Tables 9 , 10 & 11 respectively. There are two adult males and four adult females ranging in size from 27.4-32.8 mm (Fig. 19 ). All paratypes resemble the holotype except as follows: supranasals marginally in contact with each other anteriorly on the snout in NRC-AA-1181 and NRC-AA-1183. Upper postmentals separated medially by two small chin scales below mental in BNHS 2826 and BNHS 2827; upper postmentals bordered by mental, infralabial I, outer postmental, and three chin scales on left side and two on right side in NRC-AA-1181, NRC-AA-1182; upper postmentals bordered by infralabial I & II on right and infralabial I on left side, and by mental, outer postmental, and three chin scales on either side in NRC-AA-1183; upper postmentals bordered by mental, infralabial I, outer postmental, and two chin scales on either side in BNHS 2826 and BNHS 2827; upper postmental bordered by infralabial I & II and two chin scales on left and infralabial I and three chin scales on right side in BNHS 2828. Outer postmental bordered by inner postmental, infralabials I & II, and four chin scales on left and three on right side in NRC-AA-1181; outer postmental bordered by inner postmental, infralabials II on left and infralabial I & II on right side, and three chin scales on either side in NRC-AA-1182 and NRC-AA-1183; outer postmental bordered by inner postmental, infralabials I & II, and three chin scales on either side in BNHS 2826 and BNHS 2827; outer postmental bordered by inner postmental, infralabials II and two chin scales on right and infralabial I & II and three chin scales on left side in BNHS 2828. Outer postmental separated from each other below mental by four enlarged chin scales in BNHS 2827 and BNHS 2828. A single paratype (BNHS 2827) has original and complete tail, marginally longer than body (TL/SVL 1.08); two paratypes (NRC-AA-1183 and BNHS 2828) have complete but partially regenerated tail, marginally shorter and longer than body respectively (TL/SVL 0.97 and 1.08 respectively); tail partially detached from the body and lost in NRC-AA-1181, NRC-AA-1182, and BNHS 2826. All paratypes agree with the holotype in colouration except for the four female paratypes which differ in ventral colouration, lacking the yellow seen in males: with ventral surfaces off-white with faint, scattered grey stippling; belly and underside of tail light cream. Dark markings on dorsum poorly defined in NRC-AA-1182, BNHS 2826 and prominent in NRC-AA-1181, BNHS 2828 (Figs 18B & 19 ).

Distribution and Natural history.

Cnemaspis kalakadensis sp. nov. is currently known only from in and around its type locality (Sengaltheri in Kalakadu reserve forest, ca. 900-1060 m asl.) in Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu (Fig. 1 ). Individuals of the new species were seen active during the daytime (1030-1600 hrs) on tree trunks (2-5 m above the ground) inside wet evergreen forest patches (Fig. 20A-C ). Cnemaspis kalakadensis sp. nov. appears to be a tree specialist as they were only observed on tree trunks, some individuals were seen effortlessly climbing above approximately 4 m on trees when disturbed. They were observed to be fairly common (n =>15) at all three collection sites indicating good abundance. The new species were recorded in good numbers only at places with high canopy cover and were seen only during the daytime. Sympatric lizards encountered on tree trunks at the locality include Cnemaspis australis Manamendra-Arachchi, Batuwita and Pethiyagoda, 2007; Dravidogecko douglasadamsi Chaitanya, Giri, Deepak, Datta-Roy, Murthy and Karanth, 2019; Hemidactylus frenatus Duméril and Bibron, 1836; Dasia johnsinghi Harikrishnan, Vasudevan, De Silva, Deepak, Kar, Naniwadekar, Lalremruata, Prasoona and Aggarwal, 2012; Draco dussumieri Duméril and Bibron, 1837; and Monilesaurus ellioti ( Günther, 1864).

Figures

Figure 1. Elevation map of southern Kerala and Tamil Nadu showing the distribution of all known Cnemaspis species from the region: circles, Cnemaspis beddomei clade; squares, Cnemaspis gracilis clade; triangle, Cnemaspis littoralis clade; pentagon, Cnemaspis australis clade.

Figure 2. Maximum likelihood (ML) phylogeny of South Asian Cnemaspis based on the mitochondrial ND 2 gene. Bootstrap support> 70 shown at nodes; Cnemaspis beddomei, Cnemaspis gracilis and Cnemaspis littoralis clades marked by text. Roman numerals indicate other broad clades: I, Cnemaspis australis; II, Cnemaspis mysoriensis; III, Cnemaspis goaensis; IV, girii; V, Cnemaspis bangara; VI, Cnemaspis indica; VII, Cnemaspis podihuna; VIII, Cnemaspis assamensis; IX, Cnemaspis wynadensis. Top left inset is a 16 S ML phylogeny of the Cnemaspis littoralis clade (outgroups not shown).

Figure 5. Cnemaspis azhagu sp. nov. (holotype, NRC-AA- 1170): A dorsal aspect of mid-body, B ventral aspect of mid-body, and C right side lateral aspect of mid-body. Scale bars 5 mm; photos by Akshay Khandekar and Satpal Gangalmale.

Figure 6. Colouration in life of Cnemaspis azhagu sp. nov.: A adult male (holotype, NRC-AA- 1170), and B adult female (paratype, BNHS 2821). Photos by Akshay Khandekar and Satpal Gangalmale.

Figure 9. Cnemaspis mundanthuraiensis sp. nov. (holotype, NRC-AA- 1175): A dorsal aspect of body, B ventral aspect of body, C dorsal aspect of tail, D ventral aspect of tail, and E lateral aspect of tail. Scale bars 10 mm; photos by Akshay Khandekar and Satpal Gangalmale.

Figure 10. Cnemaspis mundanthuraiensis sp. nov. (holotype, NRC-AA- 1175): A dorsal aspect of head, B ventral aspect of head, C lateral aspect of right side head, D aspect of cloacal region showing precloacal and femoral pores, E ventral aspect of left manus, and F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar and Satpal Gangalmale.

Figure 13. Paratype series of Cnemaspis mundanthuraiensis sp. nov. from left to right, NRC-AA- 1176, NRC-AA- 1177, NRC-AA- 1178, NRC-AA- 1179, BNHS 2823, BNHS 2824, BNHS 2825, BNHS 2822. Scale bar 10 mm; photo by Akshay Khandekar and Satpal Gangalmale.

Figure 15. Cnemaspis kalakadensis sp. nov. (holotype, NRC-AA- 1180): A dorsal aspect of body, B ventral aspect of body, C dorsal aspect of tail, D ventral aspect of tail, and E lateral aspect of tail. Scale bars 10 mm; photos by Akshay Khandekar and Satpal Gangalmale.

Figure 16. Cnemaspis kalakadensis sp. nov. (holotype, NRC-AA- 1180): A dorsal aspect of head, B ventral aspect of head, C lateral aspect of right side head, D aspect of cloacal region showing femoral pores, E ventral aspect of left manus, and F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar and Satpal Gangalmale.

Figure 17. Cnemaspis kalakadensis sp. nov. (holotype, NRC-AA- 1180): A dorsal aspect of mid-body, B ventral aspect of mid-body, and C right side lateral aspect of mid-body. Scale bars 5 mm; photos by Akshay Khandekar and Satpal Gangalmale.

Figure 18. Colouration in life of Cnemaspis kalakadensis sp. nov.: A adult male (holotype, NRC-AA- 1180), and B adult female (paratype, BNHS 2828). Photos by Akshay Khandekar and Satpal Gangalmale.

Figure 19. Paratype series of Cnemaspis kalakadensis sp. nov. from left to right, NRC-AA- 1181, NRC-AA- 1182, NRC-AA- 1183, BNHS 2826, BNHS 2827, BNHS 2828. Scale bar 10 mm; photo by Akshay Khandekar and Satpal Gangalmale.

Figure 20. Habitat at the type locality of Cnemaspis kalakadensis sp. nov.: A general habitat showing the wet evergreen forest, and B & C microhabitat showing the trees and a stream inside forested area from where the type series were collected. Photos by Akshay Khandekar.