NRC-AA-9672 (AK-R 603), adult female, from near Wood House (8.5220°N, 77.5040°E; ca. 230 m asl.), Kalakad Range, Kalakad-Mundanthurai Tiger Reserve (KMTR), Tirunelveli District, Tamil Nadu State, India, collected by Akshay Khandekar, Swapnil Pawar and team, on 29^th April 2021.

Figure 3. 

Cyrtodactylus (Geckoella) kalakadensis sp. nov. (holotype, NRC-AA-9672): A dorsal view of body, B ventral view of body, C dorsal view of head, D ventral view of head, and E right side lateral view of head. Scale bars 10 mm (A, B) and 5 mm (C–E); photos by Akshay Khandekar.

Figure 4. 

Cyrtodactylus (Geckoella) kalakadensis sp. nov. (holotype, NRC-AA-9672): A dorsal view of midbody, B ventral view of midbody, C ventral view of left manus, D ventral view of left pes. Scale bars 5 mm; photos by Akshay Khandekar.

Figure 5. 

Colouration in life and habitat of Cyrtodactylus (Geckoella) kalakadensis sp. nov.: A holotype, female (NRC-AA-9672), B paratype, male (NRC-AA-9672), C paratype, male (NRC-AA-9674), D paratype, male (NRC-AA-9675), E paratype, female (NRC-AA-9676), F paratype, male (NRC-AA-9678), and G general habitat at the type locality, near Sengaltheri, KMTR, Tamil Nadu. Photos by Akshay Khandekar and Saunak Pal (F).

Figure 6. 

Paratypes of Cyrtodactylus (Geckoella) kalakadensis sp. nov. from left to right, NRC-AA-9673–9678: A dorsal view, B ventral view. Scale bars 10 mm; photos by Akshay Khandekar.

NRC-AA-9673 (AK-R 602), NRC-AA-9674 (AK-R 604), adult males, same collection data as holotype; NRC-AA-9675 (AK-R 665), adult male, NRC-AA-9676 (AK-R 979), NRC-AA-9677 (AK-R 980), adult females, from Kalakad Range (8.5274°N, 77.4868°E; ca. 320 m asl.) collected by Akshay Khandekar, Ishan Agarwal, Swapnil Pawar and team on 30th March 2022; NRC-AA-9678 (CESL 510), adult male, from Manimuthar, Mundanthurai Range (8.6244°N, 77.4109 °E; ca. 130 m asl.) collected by Saunak Pal on 18^th December 2011; all from KMTR, Tirunelveli District.

A small-sized Cyrtodactylus (Geckoella), snout to vent length up to 55 mm (n = 7); tail shorter than SVL. Dorsal pholidosis on trunk homogeneous; covered with smooth, subcircular, weakly conical granular scales; ventral scales slightly larger than granular scales on dorsum, smooth, oval, subimbricate with rounded end, subequal from chest to vent; 15–19 ventral scales at midbody contained within one eye diameter, 30–44 ventral scales across belly at midbody, 67–88 longitudinal scales between axilla to groin, 185–202 longitudinal scales from mental to cloaca; subdigital scansors smooth, unnotched, and mostly entire; 10–13 lamellae under digit I of manus and pes, 12–15 lamellae under digit IV of manus and 17–21 lamellae under digit IV of pes; absence of femoral and precloacal pores; absence of enlarged precloacal or femoral scales, no precloacal groove. Ground colouration khaki, three brown black-edged cross-bars between occiput and hindlimb insertions that may be notched or broken into spots; post-occipital collar contiguous, notched or separated mid-vertebrally into pair of spots, no regular spots on flanks, darker markings cover > 50 % of dorsum between limb insertions.

The small body size and original tail shorter than SVL, besides the lack of femoral or precloacal pores in males, easily differentiate the new species from all other Cyrtodactylus, and its homogeneous dorsal pholidosis without rows of enlarged, rounded tubercles from all Cyrtodactylus (Geckoella) apart from the C. collegalensis complex. The new species can be distinguished by the absence of a few, enlarged dorsal tubercles on the dorsum (versus the presence of these tubercles in C. chengodumalaensis) and by the absence of a patch of enlarged, roughly hexagonal scales on the canthus rostralis and beneath the angle of the lower jaw (versus the presence of these enlarged scales in C. varadgirii). The dorsal pattern consisting of three dark brown cross-bars (sometimes notched or forming paired spots) from behind nape to hindlimb insertions of C. kalakadensis sp. nov. easily distinguishes it from the species with a banded dorsal pattern — C. aravindi (single dark broad band and one or two single dark spots), C. rishivalleyensis Agarwal, 2016 and C. speciosus (two broad dark bands), and C. yakhuna (usually two broad dark bands or paired spots on dorsum); as well as the spotted species with four or more rows of spots from behind nape to hindlimb insertions: C. chengodumalaensis and C. varadgirii (4–6 pairs of dark spots), C. irulaorum Agarwal, Thackeray & Khandekar, 2023 and C. relictus Agarwal, Thackeray & Khandekar, 2023 (four pairs of dark spots). Both C. collegalensis and C. srilekhae Agarwal, 2016 also have three pairs of dark dorsal markings from nape to hindlimb insertions, however C. kalakadensis sp. nov. may be distinguished by the condition of the dark markings on dorsum between nape and hindlimb insertions — which consist of bands that continue laterally onto the flanks where they flare out and may meet, first band entire/ mid-vertebrally notched and second band entire/ notched/ rarely just separated into two spots (versus first and second dark marking consisting of paired spots that do not extend onto flank and may be fused centrally in C. collegalensis; and first dark marking a pair of spots that may be fused and second row of dark markings usually consisting of a small central spot and two or more larger indistinct blotches laterally in C. srilekhae). Cyrtodactylus kalakadensis sp. nov. is diagnosed against C. shenbagathoppuensis sp. nov. after the description of the latter.

Adult female in good state of preservation except tail slightly bent towards left, a 2.9 mm longitudinal skin injury on precloacal region, and a 7.9 mm vertical incision in the sternal region for liver tissue collection (Fig. 3A, B). SVL 55.0 mm, head short (HL/SVL 0.26), wide (HW/ HL 0.70), not strongly depressed (HD/HL 0.44), slightly narrower than body (HW/BW 0.91), and distinct from neck (Fig. 3C–E). Loreal region slightly inflated, canthus rostralis not prominent. Snout less than half head length (ES/HL 0.42) and less than two times eye diameter (ES/ED 1.82); scales on snout, canthus rostralis, and loreal region large, subcircular, smooth and, flattened; much larger than granular scales on forehead and interorbital region; occipital and temporal region with much smaller, smooth granules (Fig. 3C). Eye small (ED/HL 0.23); pupil vertical, dilated with indistinct crenulate margins; supraciliaries short, larger anteriorly, not elongate; 14 interorbital scale rows across narrowest point of frontal; 40 scale rows between left and right supraciliaries at mid-orbit (Fig. 3C, E). Ear-opening oval, small (EL/HL 0.05); eye to ear distance more than eye diameter (EE/ED 1.39). Rostral roughly 1.7 times wider (2.5 mm) than deep (1.5 mm), incompletely divided dorsally by weakly developed rostral groove for slightly more than half of its height; a single enlarged supranasal on each side, more than two times larger than the postnasals, separated from each other by a single enlarged internasal on the snout; three subequal postnasals, much smaller than supranasals; rostral in contact with nostril, supralabial I, supranasals, and internasals on either side; nostrils oval, directed somewhat outwards, covering most of the nasal scale; surrounded on either side by supralabial I, rostral, supranasals, and postnasals; two single row of smaller scales separate the orbit from the supralabials (Fig. 3E). Mental enlarged, triangular, wider (2.3 mm) than long (1.6 mm); two pairs of postmentals; inner pair in strong contact with each other, roughly pentangular, and slightly shorter than mental (1.9 mm length); bordered by mental, infralabial I, outer postmental, three chin scales on left and single on right side; outer postmentals separated from each other by inner postmentals, roughly rectangular and almost half the size (1.1 mm length) of inner pair; bordered by inner postmentals, infralabial I and II, and three enlarged chin shields on either side; chin shields bordering postmentals and infralabials flat, smooth, smaller than outermost postmentals; rest flattened, even smaller, smooth; three or four rows of enlarged elongated scales separating gular scales from infralabials (Fig. 3D). Ten supralabials up to angle of jaw, and eight at midorbital position on each side (supralabial four divided into two scales on right); six infralabials up to angle of jaw, and five infralabials at midorbital position on both sides (Fig. 3E).

Body relatively slender (BW/AGL 0.44), trunk less than half of SVL (AGL/SVL 0.46) without ventrolateral folds. Dorsal pholidosis on trunk homogeneous; covered with smooth, subcircular, weakly conical granular scales; 14 dorsal midbody scale rows across trunk contained within one eye diameter (Fig. 4A). Granular scales on occiput and nape slightly smaller than those on body dorsum; granular scales on flank equally larger than those on dorsum. Ventral scales slightly larger than granular scales on dorsum, subequal from chest to vent, smooth, oval, and subimbricate with rounded end; 17 ventral scale rows at midbody contained within one eye diameter; 42 midbody scale rows across belly (Fig. 4B). Scales on throat slightly smaller than those on belly; gular region with much smaller granular scales, those on chin bordering postmentals, enlarged, juxtaposed and flattened (Fig. 3D). No enlarged precloacal or femoral scales, no precloacal or femoral pores; no precloacal groove (Fig. 3B).

Scales on palm and soles granular, smooth, rounded to oval; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small, granules similar to dorsum and many smooth flattened and subimbricate scales which are much larger than granules on the body dorsum, largest on anterior aspect of the hands and feet; lateral and ventral aspects of fore-limbs with small granular scales; ventral aspect of thigh with scales similar to midbody ventrals except slightly smaller and gradually becoming even smaller and granular on posterior aspect; ventral aspect of shank with large, imbricate scales (Fig. 3A, B). Fore-limbs and hind-limbs slightly long, slender (LAL/SVL 0.14; CL/SVL 0.16); digits short, slender, with a strong, recurved claw, moderately inflected, distal portions laterally compressed. Series of unpaired lamellae on basal portion of digits except one or two which on some digits are paired, separated from narrower distal lamellae by a single large lamella at the inflection, unpaired except one or two which are divided; basal lamellae series: (4-5-5-5-4 right manus, Fig. 4C; 4-7-6-8-7 right pes, Fig. 4D), (4-6-5-5-4 left manus; 3-6-7-8-6 left pes); distal lamellae series: (7-8-8-8-7 right manus, Fig. 4C; 8-8-10-10-10 right pes, Fig. 4D), (7-7-9-8-7 left manus; 8-8-11-10-11 left pes). Relative length of digits (measurements in mm in parentheses): III (3.1) > IV (2.6) > II (2.8) > V (2.4) > I (1.9) (right manus); III (4.0) > IV (3.8) > V (3.7) = II (3.5) > I (2.1) (right pes).

Tail original, circular in cross section with indistinct median dorsal furrow, relatively thick, tapering gradually to tip, unsegmented, shorter than snout-vent length (TL/SVL 0.64). Scales on dorsal aspect of tail base heterogenous, a few scattered enlarged, smooth to feebly keeled tubercles intermixed with much smaller and smooth granular scales, enlarged tubercles occasionally extending onto groin; scales on dorsal aspect of tail large flat, subcircular, smooth, and imbricate, becoming slightly larger towards lateral aspect, largest on ventral side, but not forming median row of transversely enlarged subcaudal scales. Small, smooth, subequal, conical postcloacal spurs, three on either side of tail base; no hemipenial swelling. Tail slightly constricted at the base (Fig. 3A, B).

Dorsal ground colour of body, head and limbs khaki, three pairs of dark brown, black-edged cross-bars from nape to hindlimb insertions and one above tail base, second bar notched mid-vertebrally and third divided into paired spots; bars widen on flanks; scattered small black spots on light interspaces of dorsum. Dark brown dorsal markings edged by black on the outer 2–5 rows of granules; ventrolateral portion of flanks heavily pigmented from the dorsal bands with a few scattered and irregular fine black spots. Tail dorsum with about four paired spots following pair of spots on tail base; dorsum of limbs with fine spots, digits with alternating pale and brown bands. Post-occipital collar broken mid-vertebrally, composed of two elongate spots contiguous with postorbital streak; interparietal streak and other head markings broken into spots (Fig. 5A).

There are four adult male, and two adult female paratypes, SVL ranging from 37.9–53.9 mm (Tables 4, 5). All specimens resemble the holotype except for the following variation: inner postmental bordered by mental, infralabial I, outer postmental in all paratypes; additionally, bordered by one chin scale on left and three on right in NRC-AA-9673, three chin scales on left and two on right in NRC-AA-9674 and NRC-AA-9678, two chin scales on either side in NRC-AA-9675 and NRC-AA-9677, two chin scales on left and one on right in NRC-AA-9676. Outer postmental bordered by inner postmentals, infralabial I and II; additionally, bordered by three chin scales on left and four on right in NRC-AA-9674 and NRC-AA-9675. Two paratypes (NRC-AA-9675 and NRC-AA-9676) have an original and complete tail, much shorter than body (TL/SVL 0.69, 0.70); and remaining paratypes (NRC-AA-9673, NRC-AA-9674, NRC-AA-9677, NRC-AA-9678) are with short and entirely regenerated tails (0.52, 0.55, 0.16 and 0.48 respectively) (Fig. 6A, B). There is considerable variation in dorsal colour pattern of body and head in the type series (Fig. 5B–F); NRC-AA-9676 and NRC-AA-9677 are the most pigmented with the dark dorsal bands covering most of dorsum; A NRC-AA-9673 and NRC-AA-9677 have the first dark marking on dorsum notched; NRC-AA-9673, NRC-AA-9675, NRC-AA-9676, and NRC-AA-9678 have the second dark marking on dorsum broken into spots or deeply notched while the other paratypes have a solid band; NRC-AA-9673, NRC-AA-9678 have the third dark marking on dorsum broken into spots like the holotype while the other paratypes have a solid band.

Cyrtodactylus (Geckoella) kalakadensis sp. nov. is known from only three closely-spaced localities about 15 km apart at elevations of ~130–320 m (Fig. 1). These low elevation localities consist of moist deciduous forest with good canopy cover and leaf litter (Fig. 5G). The species was seen active after dark (~1830–2100 hrs) on forest floor especially in areas with high leaf litter. A total of 13 individuals were observed in three different trips. The new species is sympatric with the KMTR endemics Cnemaspis sp. (Khandekar et al. unpub. data), Hemidactylus acanthopholis Mirza & Sanap, 2014, and Dravidoseps kalakadensis Agarwal, Thackeray & Khandekar, 2024; beside common lizard species like H. leschenaultii Duméril & Bibron, 1836, H. frenatus Duméril & Bibron, 1836, Eutropis carinata (Schneider, 1801), E. cf. macularia, Riopa cf. albopunctata, Calotes calotes (Linnaeus, 1758), C. versicolor (Daudin, 1802), and Psammophilus cf. blanfordanus.

The specific epithet is a toponym for the type and other known localities of the new species, in Kalakad Forest Range, KMTR (Kalakad-Mundanthurai Tiger Reserve), Tirunelveli District, Tamil Nadu. We suggest the common name Kalakad or KMTR Geckoella.

NRC-AA-9669 (AK-R 1461), adult male, from Shenbagathoppu (9.5709°N, 77.5625°E; ca. 320 m asl.), Srivilliputhur-Megamalai Tiger Reserve (SMTR), Virudhunagar District, Tamil Nadu state, India; collected by Akshay Khandekar, Ishan Agarwal, Swapnil Pawar and team on 21^th April 2022.

Figure 7. 

Cyrtodactylus (Geckoella) shenbagathoppuensis sp. nov. (holotype, NRC-AA-9669): A dorsal view of body, B ventral view of body, C dorsal view of head, D ventral view of head, and E right side lateral view of head. Scale bars 10 mm (A, B) and 5 mm (C–E); photos by Akshay Khandekar.

Figure 8. 

Cyrtodactylus (Geckoella) shenbagathoppuensis sp. nov. (holotype, NRC-AA-9669): A dorsal view of midbody, B ventral view of midbody, C ventral view of left manus, D ventral view of left pes. Scale bars 5 mm; photos by Akshay Khandekar.

Figure 9. 

Colouration in life and habitat of Cyrtodactylus (Geckoella) shenbagathoppuensis sp. nov.: A holotype, male (NRC-AA-9669), B paratype, female (NRC-AA-9670), C paratype, subadult male (NRC-AA-9671), D uncollected specimen, and E general habitat at the type locality, Shenbagathoppu, SMTR, Tamil Nadu. Photos by Ishan Agarwal, Gopal Murali (D), Akshay Khandekar (E).

Figure 10. 

Paratypes of Cyrtodactylus (Geckoella) shenbagathoppuensis sp. nov. from left to right, NRC-AA-9670 and NRC-AA-9671: A dorsal view, B ventral view. Scale bars 10 mm; photos by Akshay Khandekar.

NRC-AA-9671 (AK-R 1312), subadult male, same data as holotype and NRC-AA-9670 (AK-R 1462), adult female, same data as holotype except collected on 15^th April 2022 from ~ 1.5 km SW of type locality (9.5580°N, 77.5560°E; ca. 230 m asl.).

A small-sized Cyrtodactylus (Geckoella), snout to vent length up to 53.4 mm (n = 3); tail shorter than SVL. Dorsal pholidosis on trunk homogeneous; covered with smooth, subcircular, weakly conical granular scales; ventral scales much larger than granular scales on dorsum, smooth, subcircular, subimbricate with rounded end, subequal from chest to vent; 15 ventral scales at midbody contained within one eye diameter, 31–39 ventral scales across belly at midbody, 61–71 longitudinal scales between axilla to groin, 164–174 longitudinal scales from mental to cloaca; subdigital scansors smooth, unnotched, and mostly entire; 10–12 lamellae under digit I of manus and 11–14 lamellae under digit I of pes, 12–15 lamellae under digit IV of manus and 17 or 18 lamellae under digit IV of pes; absence of femoral and precloacal pores; absence of enlarged precloacal or femoral scales, no precloacal groove. Ground colouration tan, with five or six dark brown black-edged spots between nape and hindlimb insertions, spots well-separated along mid-vertebral line; post-occipital collar separated mid-vertebrally into pair of spots, no regular spots on flanks, darker markings cover ~ 50 % of dorsum between limb insertions.

The small body size and original tail shorter than SVL, besides the lack of femoral or precloacal pores in males, easily differentiate the new species from all other Cyrtodactylus, and its homogeneous dorsal pholidosis without rows of enlarged, rounded tubercles from all Cyrtodactylus (Geckoella) apart from the C. collegalensis complex. The new species can be distinguished by the absence of a few, enlarged dorsal tubercles on the dorsum (versus the presence of these tubercles in C. chengodumalaensis) and by the absence of a patch of enlarged, roughly hexagonal scales on the canthus rostralis and beneath the angle of the lower jaw (versus the presence of these enlarged scales in C. varadgirii).The dorsal pattern consisting of five or six pairs of dark brown spots from behind nape to hindlimb insertions of C. shenbagathoppuensis sp. nov. easily distinguishes it from the species with a banded dorsal pattern — C. aravindi (single dark broad band and one or two single dark spots), C. rishivalleyensis and C. speciosus (two broad dark bands), C. kalakadensis sp. nov. (three dark brown dorsal cross-bars that are sometimes notched or form paired spots) and C. yakhuna (usually two broad dark bands or paired spots on dorsum); as well as the species with four or less rows of spots from behind nape to hindlimb insertions: C. collegalensis (three pairs of dark spots that may be fused forming horizontal figure 8-shaped markings), C. irulaorum and C. relictus (four pairs of dark spots), and C. srilekhae (three rows of irregular dark blotches). The new species is most similar in colour pattern to C. chengodumalaensis and C. varadgirii (4–6 pairs of dark spots from behind nape to hindlimb insertions) but can be distinguished by the absence of a few, enlarged dorsal tubercles on the dorsum (versus the presence of these tubercles in C. chengodumalaensis) and by the absence of a patch of enlarged, roughly hexagonal scales on the canthus rostralis and beneath the angle of the lower jaw (versus the presence of these enlarged scales in C. varadgirii); as well as by its body size up to at least 53.7 mm SVL (versus up to 48 mm in C. aravindi and C. chengodumalaensis). Cyrtodactylus shenbagathoppuensis sp. nov. occupies a distinct position in multivariate morphospace (Fig. 2) and is distinguished from its sister species, C. aravindi on a number of morphometric variables (Table 3), colour pattern and body size (up to at least 53.7 mm SVL versus up to 48 mm in C. aravindi); and from the other Southern Western Ghats species described above in number of ventral scales between mental and cloaca (164–174 versus 185–202 in C. kalakadensis sp. nov.).

Adult male in good state of preservation except head bent towards left, tail sigmoid, and a 4.9 mm vertical incision in the abdominal region for liver tissue collection (Fig. 7A, B). SVL 48.4 mm, head short (HL/SVL 0.27), wide (HW/ HL 0.70), not strongly depressed (HD/HL 0.43), slightly broader than body (HW/BW 1.05), and distinct from neck (Fig. 7C–E). Loreal region slightly inflated, canthus rostralis not prominent. Snout marginally less than half head length (ES/HL 0.43), slightly less than twice eye diameter (ES/ED 1.72); scales on snout, canthus rostralis, and loreal region large, subcircular, smooth and, flattened; much larger than granular scales on forehead and interorbital region; occipital and temporal region with much smaller, smooth granules (Fig. 7C, E). Eye small (ED/HL 0.25); pupil vertical, dilated with indistinct crenulate margins; supraciliaries short, larger anteriorly, not elongate; 13 interorbital scale rows across narrowest point of frontal; 39 scale rows between left and right supraciliaries at mid-orbit (Fig. 7E). Ear-opening oval, small (EL/HL 0.07); eye to ear distance slightly more than eye diameter (EE/ED 1.12). Rostral roughly 1.7 times wider (2.4 mm) than deep (1.4 mm), incompletely divided dorsally by weakly developed rostral groove for slightly more than half of its height; a single enlarged supranasal on each side, more than three times larger than the postnasals, separated from each other by a single enlarged internasal on the snout; three subequal postnasals, much smaller than supranasals; rostral in contact with nostril, supralabial I, supranasals, and internasals on either side; nostrils oval, directed somewhat outwards, covering most of the nasal scale; surrounded on either side by supralabial I, rostral, supranasals, and postnasals; a single row of smaller scales separate the orbit from the supralabials (Fig. 7E). Mental enlarged, triangular, wider (1.9 mm) than long (1.4 mm); two pairs of postmentals; inner pair in strong contact with each other, and longer than mental (2.2 mm); bordered by mental, infralabial I, outer postmental and two enlarged chin shields on either side; outer postmentals separated from each other by inner postmentals, roughly hexagonal and almost half the length (1.15 mm) of inner pair; bordered by inner postmentals, infralabial I and II, and three enlarged chin shields on either side; chin shields bordering postmentals and infralabials flat, smooth, smaller than outermost postmentals; rest flattened, even smaller, smooth; two or three rows of enlarged elongated scales separating gular scales from infralabials (Fig. 7D). Ten supralabials up to angle of jaw on left and eleven on right, and seven at midorbital position on each side (supralabial four divided into two scales on right); six infralabials up to angle of jaw on left and seven on right, and five infralabials at midorbital position on both sides (Fig. 7E).

Body relatively slender (BW/AGL 0.41), trunk less than half of SVL (AGL/SVL 0.45) without ventrolateral folds. Dorsal pholidosis on trunk homogeneous; covered with smooth, subcircular, weakly conical granular scales; 20 dorsal midbody scale rows across trunk contained within one eye diameter (Fig. 8A). Granular scales on occiput and nape slightly smaller than those on body dorsum; granular scales on flank slightly larger than those on dorsum. Ventral scales much larger than granular scales on dorsum, subequal from chest to vent, smooth, subcircular, and subimbricate with rounded end; 15 ventral scale rows at midbody contained within one eye diameter; 31 ventral midbody scale rows across belly (Fig. 8B). Scales on throat slightly smaller than those on belly; gular region with much smaller granular scales, those on chin bordering postmentals, enlarged, juxtaposed and flattened (Fig. 7D). No enlarged precloacal or femoral scales, no precloacal or femoral pores; no precloacal groove (Fig. 7B).

Scales on palm and soles granular, smooth, rounded to oval; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small, granules similar to dorsum and many smooth flattened and subimbricate scales which are much larger than granules on the body dorsum, largest on anterior aspect of the hands and feet (Fig. 7A, B). Scales on ventral aspect of fore-limbs granular, slightly smaller than those on dorsal aspect; ventral aspect of thigh with scales similar to midbody ventrals except slightly smaller and gradually becoming even smaller and granular on posterior aspect; ventral aspect of shank with large, imbricate scales. Fore-limbs and hind-limbs slightly long, slender (LAL/SVL 0.15; CL/SVL 0.17); digits short, slender, with a strong, recurved claw, moderately inflected, distal portions laterally compressed. Series of unpaired lamellae on basal portion of digits except one or two which on some digits are paired, separated from narrower distal lamellae by a single large lamella at the inflection, unpaired except one or two which are divided; basal lamellae series: (3-6-5-6-4 right manus, Fig. 8C; 4-6-7-8-5 right pes, Fig. 8D), (3-7-6-6-4 left manus; 3-6-7-8-5 left pes); distal lamellae series: (8-8-10-9-9 right manus, Fig. 8C; 10-9-10-10-11 right pes, Fig. 8D), (5*-8-10-9-8 left manus; 11-10-11-10-11 left pes). Relative length of digits (measurements in mm in parentheses): III (3.0) > IV (2.7) > II (2.4) > V (2.0) > I (1.7) (right manus); III (4.2) > IV (3.8) > V (3.7) = II (3.5) > I (2.0) (right pes).

Tail original, circular in cross section with indistinct median dorsal furrow, relatively thick, tapering gradually to tip, unsegmented, slightly shorter than snout-vent length (TL/SVL 0.76). Scales on dorsal aspect of tail heterogenous, a few scattered enlarged, smooth to feebly keeled tubercles intermixed with much smaller and smooth granular scales, enlarged tubercles sometimes extend onto groin; scales on dorsal aspect of tail large flat, subcircular, smooth, and imbricate, becoming slightly larger towards lateral aspect, largest on ventral side, but not forming median row of transversely enlarged subcaudal scales. Small, smooth, subequal, conical postcloacal spurs, two on left and three on right side of tail base; prominent hemipenial swelling with barely everted hemipenis on either side, flap of skin covering cloacal aperture. Tail slightly constricted at the base (Fig. 7A, B).

Dorsal ground colour of body, head and limbs tan, five pairs of dark brown spots from behind nape to hindlimb insertions and a smaller pair above tail base; a row of four small spots between pairs three and four; all spots well separated mid-vertebrally except those on tail base just meeting; third pair of spots slightly larger than rest and fifth smallest and subequal to pair on tail base. Spots edged by black on the outer 2–5 rows of granules; ventrolateral portion of flanks with a few scattered and irregular black spots or blotches. Tail dorsum with eight dark brown black-edged crossbars following pair of spots on tail base; dorsum of limbs with scattered dark cross-bars and spots, digits with alternating dark brown bands. Post-occipital collar broken mid-vertebrally, composed of two elongate spots contiguous with parietal spots and separated from postorbital streak and broken interparietal streak. Crown sightly darker than trunk with dark brown markings that have a darker border flanked by a finer light border; a pre-frontal spot subequal in size to two interorbital spots which are broader and shorter than the longer part of the (broken) interparietal streak. Brille similar in colour to light scales on crown. Postocular streak runs from posterior edge of eye to beyond tympanum, separated from dorsolateral markings of collar and first pair of dorsal spots on neck; preocular streak extends till nostril. Labials with a few thick dark streaks, unmarked scales finely spotted with black. Ventral aspects dirty white with mottling under limbs and belly and fine spots along margin of belly, thick streaks and small spots on gular region, ventral aspect of tail dark with indistinct banding (Fig. 9A).

The subadult male SVL 34.0 mm and adult female SVL 53.4 mm paratypes resemble the holotype except for the following: inner postmental bordered by mental, infralabial I, outer postmental, and two enlarged chin scales on either side in NRC-AA-9670; in both paratypes – inner postmental bordered by mental, outer postmental, two enlarged chin scales on either side, and infralabial I on right and infralabial I and II on left side in NRC-AA-9671. Outer postmental bordered by inner postmentals on either side in both the paratypes; additionally, bordered by infralabial II and four chin scales on left and infralabial I and II and five on right side in NRC-AA-9671; and bordered by infralabial I and II, and four enlarged chin shields on either side in NRC-AA-9670. NRC-AA-9670 with original and complete tail, slightly shorter than body (TL/SVL 0.82); and NRC-AA-9671 with short and entirely regenerated tail (Fig. 10A, B). The two paratypes are generally similar in colouration and pattern to the holotype except that the post-occipital collar (spots) is separated from the parietal spots while the prefrontal and interorbital spots are fused in NRC-AA-9670 (Fig. 9B, C).

Cyrtodactylus shenbagathoppuensis sp. nov. is only known from its type locality which has moist deciduous forest at elevations of 230–320 m asl. (Figs 1, 9E). All specimens were spotted on the ground in leaf-litter after dark, the type was found about two hours after sunset at the base of a large boulder.

The species has been previously observed in the months of March, April, and September (2020 and 2021), with most sightings soon after dark (~18:00–20:00 h) and as late as midnight (Satheeshkumar pers. obs.). These localities span an elevational range of ~150–600 m asl., and individuals were spotted on rocks, leaf litter and the long staircase leading up to the Shenbagathoppu Kaatalagar Temple. A dead individual was recorded from farmlands based on photographic evidence provided by a third-party observer.

The specific epithet is a toponym for the type locality of the new species, Shenbagathoppu, in Srivilliputhur-Megamalai Tiger Reserve, Virudhnagar District, Tamil Nadu. The Tamil name means Shenbaga = champaca [= Magnolia champaca (Linnaeus)] and Thoppu = grove. We suggest the common name Shenbagathoppu Geckoella.

NRC-AA-9679 (IAG 081), adult male, from near Dongargaon (20.6323°N, 80.3651°E; ca. 280 m asl.), Gadchiroli District, Maharashtra, India, collected by Tarun Khichi, NS Achyuthan & Ishan Agarwal on 29^th April 2015.

Figure 11. 

Cyrtodactylus (Geckoella) satpuraensis sp. nov. (holotype, NRC-AA-9679): A dorsal view of body, B ventral view of body, C dorsal view of head, D ventral view of head, and E right side lateral view of head. Scale bars 10 mm (A, B) and 5 mm (C–E); photos by Akshay Khandekar.

Figure 12. 

Cyrtodactylus (Geckoella) satpuraensis sp. nov. (holotype, NRC-AA-9679): A dorsal view of midbody, B ventral view of midbody, C ventral view of left manus, D ventral view of left pes. Scale bars 5 mm; photos by Akshay Khandekar.

Figure 13. 

Colouration in life and habitat of Cyrtodactylus (Geckoella) satpuraensis sp. nov.: A holotype, male (NRC-AA-9679), B paratype, male (NRC-AA-9694), C paratype, female (NRC-AA-9693), D paratype, male (NRC-AA-9691), E paratype, female (NRC-AA-9688), F paratype, subadult female (NRC-AA-9685), and G general habitat at Toranmal, Dhule District, Maharashtra — the westernmost extent of the Satpuras. Photos by Ishan Agarwal.

Figure 14. 

Paratypes of Cyrtodactylus (Geckoella) satpuraensis sp. nov. from left to right, NRC-AA-9680–9694: A dorsal view, B ventral view. Scale bars 10 mm; photos by Akshay Khandekar.

NRC-AA-9681 (IAG 079), subadult female, NRC-AA-9682 (IAG 080), adult male, same locality and collection data as holotype; NRC-AA-9680 (IAG 064), adult male, from near Chikhaldara (21.4076°N, 77.2973°E; ca. 1080 m asl.), Amravati District, Ma­ha­rashtra, collected by Tarun Khichi, NS Achyuthan and Ishan Agarwal on 21^th April 2015; NRC-AA-9688, adult female, NRC-AA-9683 (IAG 124), NRC-AA-9684 (IAG 125), NRC-AA-9689, NRC-AA-9690, NRC-AA-­9691, adult males, from near Kumhedin (23.4354°N, 81.8369°E; ca. 580 m asl.), Shahdol District, Madhya Pradesh, collected by Tarun Khichi, Ishan Agarwal and Akshay Khandekar on 06^th May 2015; NRC-AA-9685, NRC-AA-9687, subadult females, NRC-AA-9686, adult male, from near Bhimlat (22.1276°N, 80.6892°E; ca. 570 m asl.), Balaghat District, Madhya Pradesh, collected by Ishan Agarwal, on 08^th April 2015; NRC-AA-9692, adult male, CES09/1377 (tissue only) from near Chhawada (22.24073°N, 78.52270°E; ca. 980 m asl.), Chhindwara District, Madhya Pradesh, collected by Ishan Agarwal, on 12th April 2015; NRC-AA-9693, adult female, NRC-AA-9694, adult male, from Toranmal (21.8570°N, 74.4632°E; ca. 1030 m asl.), Nandurbar District, Maharashtra, collected by Ishan Agarwal and Sayana Pawara, on 04^th April 2015.

A small-sized Cyrtodactylus (Geckoella), snout to vent length up to 53.5 mm (n = 16). Dorsal pholidosis heterogeneous; smooth granular scales intermixed with more or less regularly arranged rows of enlarged, feebly keeled, blunt to weakly conical tubercles; 12 or 13 rows of dorsal tubercles at midbody, 25–32 tubercles in paravertebral rows; ventral scales smooth, subcircular, and subimbricate with rounded end, subequal from chest to vent; 8–14 ventral scales at midbody contained within one eye diameter, 33–40 ventral scales across belly at midbody, 44–51 longitudinal scales between axilla to groin, 124–145 longitudinal scales from mental to cloaca; subdigital scansors smooth, unnotched, and mostly entire; 9–13 lamellae under digit I of manus and pes, 11–15 lamellae under digit IV of manus and 15–18 lamellae under digit IV of pes. 5–7 pairs of dark rounded blotches from nape to hindlimb insertions which may meet on the midline forming horizontal ‘8’ markings or uniform bands; post-occipital collar prominent except rarely separated on neck, postocular streak confluent with first pair of dorsal markings; no regular spots on flanks, darker markings cover > 50 % of dorsum between limb insertions. Ground colouration varied shades of brown; dorsal markings dark brown in center, edged with black on 1–3 rows of scales and light yellow on the posterior one or two rows.

The small body size and original tail shorter than SVL, besides the lack of femoral or precloacal pores in males, easily differentiate the new species from all other Cyrtodactylus, and its dorsal pholidosis of smooth granular scales intermixed with more or less regularly arranged rows of enlarged, feebly keeled, blunt to weakly conical tubercles from all Cyrtodactylus (Geckoella) apart from the C. nebulosus complex. Cyrtodactylus (Geckoella) satpuraensis sp. nov. can be differentiated by having 12 or 13 rows of dorsal tubercles at midbody (versus 16–18 rows of dorsal tubercles at midbody in C. teraiensis); 124–145 longitudinal scales from mental to cloaca (versus 150–157 in C. teraiensis); 33–40 (mean 37.6) ventral scales across belly at midbody (versus 30–32 in C. teraiensis); 5–7 pairs of dark blotches between neck and hindlimb insertions that may form broken or continuous bands, dark markings cover > 50 % of dorsum (versus 4–7 dark dorsal markings between neck and hindlimb insertions that vary from irregular paired blotches to cross-bars alternating with 1–3 small paired spots/ cross-bars that cover < 50 % of dorsum in C. nebulosus and four or five paired spots between neck and hindlimb insertions alternating with two or three much smaller paired spots that cover < 50 % of body dorsum in C. teraiensis); post-occipital collar extends to posterior margin of eye (versus post-occipital collar does not extend up to eye in C. teraiensis); post-orbital streak extends onto back and is confluent with first or first and second dorsal markings (versus post-orbital streak terminates before tympanum in C. teraiensis); pre-orbital streaks do not meet on internasals (versus pre-orbital streaks meet on internasals in C. teraiensis).

All three members of the C. nebulosus complex are well separated in PCA and C. satpuraensis sp. nov. is separated from C. nebulosus in a number of morphometric and some meristic characters and from C. teraiensis in LAL and a number of meristic characters (Fig. 2, Tables 3, 8).

Adult male in good state of preservation except tail slightly bent towards left, and a 10.3 mm vertical incision in the sternal region for liver tissue collection (Fig. 11A, B). SVL 43.7 mm, head short (HL/SVL 0.27), wide (HW/ HL 0.70), not strongly depressed (HD/HL 0.43), slightly narrower than body (HW/BW 0.90), and distinct from neck (Fig. 11C–E). Loreal region slightly inflated, canthus rostralis not prominent. Snout less than half head length (ES/HL 0.43) and more than two times eye diameter (ES/ED 2.04); scales on snout, canthus rostralis, and loreal region large, hexagonal to subcircular, smooth and, flattened; much larger than granular scales on forehead and interorbital region; occipital and temporal region with heterogenous scalation consisting of much smaller, smooth granular scales intermixed with slightly enlarged, smooth, round tubercles (Fig. 11C, E). Eye small (ED/HL 0.21); pupil, dilated, vertical with crenulate margins; supraciliaries short, larger anteriorly, not elongate; 13 interorbital scale rows across narrowest point of frontal; 38 scale rows between left and right supraciliaries at mid-orbit (Fig. 11E). Ear-opening oval, small (EL/HL 0.09); eye to ear distance more than eye diameter (EE/ED 1.40). Rostral roughly 1.8 times wider (2.2 mm) than deep (1.2 mm), incompletely divided dorsally by weakly developed rostral groove for slightly more than half of its height; a single enlarged supranasal on each side, more than two times larger than the postnasals, separated from each other by a single enlarged internasal on the snout; three subequal postnasals, much smaller than supranasals; rostral in contact with nostril, supralabial I, supranasals, and internasals on either side; nostrils oval, directed somewhat outwards, covering most of the nasal scale; surrounded on either side by supralabial I, rostral, supranasals, and postnasals; a single row of smaller scales separate the orbit from the supralabials (Fig. 11C). Mental enlarged, rectangular, wider (1.7 mm) than long (1.3 mm); two pairs of postmentals; inner pair in strong contact with each other, roughly pentangular, and as long as mental is wide (1.7 mm length); bordered by mental, infralabial I, outer postmental, three chin scales on either side; outer postmentals separated from each other by inner postmentals, roughly rectangular and almost half the size (1.0 mm length) of inner pair; bordered by inner postmentals, infralabial I and II, and three enlarged chin shields on left and four on right; chin shields bordering postmentals and infralabials flat, smooth, smaller than outermost postmentals; rest flattened, even smaller, smooth; four rows of enlarged elongated scales separating gular scales from infralabials (Fig. 11D). Thirteen supralabials up to angle of jaw, and seven at midorbital position on each side; nine infralabials up to angle of jaw on left and 10 on right, and six infralabials at midorbital position on both sides (Fig. 11E).

Body relatively slender (BW/AGL 0.50), trunk less than half of SVL (AGL/SVL 0.42) without ventrolateral folds. Dorsal pholidosis on trunk heterogeneous; smooth granular scales intermixed with more or less regularly arranged rows of enlarged, feebly keeled, blunt to weakly conical tubercles; granular scales gradually increasing in size towards each flank, largest on mid-flank; granular scales on occiput slightly larger than paravertebral granular scales; enlarged tubercles in approximately 13 longitudinal rows at midbody; 31 (left) and 32 (right) tubercles in paravertebral rows (Fig. 12A). Ventral scales much larger than granular scales on dorsum, subequal from chest to vent, smooth, subcircular, and subimbricate with rounded end; 38 ventral midbody scale rows across belly (Fig. 12B). Scales on throat slightly smaller than those on belly; gular region with much smaller granular scales, those on chin bordering postmentals, enlarged, juxtaposed and flattened (Fig. 11D). No enlarged precloacal or femoral scales, no precloacal or femoral pores and no precloacal groove (Fig. 11B).

Scales on palm and soles granular, smooth, rounded to oval; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small granules, similar to those on body dorsum and many smooth flattened and subimbricate scales on anterior aspects, largest on anterior aspect of the hands and feet; lateral and ventral aspects of fore-limbs with small granular scales; ventral aspect of thigh with scales similar to midbody ventrals except slightly smaller and gradually becoming even smaller and granular on posterior aspect; ventral aspect of shank with large, imbricate scales (Fig. 11A, B). Fore-limbs and hind-limbs slightly long, slender (LAL/SVL 0.15; CL/SVL 0.18); digits short, slender, with a strong, recurved claw, moderately inflected, distal portions laterally compressed. Series of unpaired lamellae on basal portion of digits except one or two which on some digits are paired, separated from narrower distal lamellae by a single large lamella at the inflection, unpaired except one or two which are divided; basal lamellae series: (4-6-7-5-5 right manus, Fig. 12C; 4-5-7-7-6 right pes, Fig. 12D), (4-6-6-6-6 left manus; 3-5-7-7-5 left pes); distal lamellae series: (8-8-10-9-7 right manus, Fig. 12C; 9-9-11-10-11 right pes, Fig. 12D), (8-8-10-9-8 left manus; 9-9-11-11-11 left pes). Relative length of digits (measurements in mm in parentheses): III (3.2) > IV (2.7) > II (2.5) > V (2.4) > I (1.8) (right manus); III (4.0) > IV (3.7) = V (3.7) > II (2.9) > I (2.3) (right pes).

Tail original, circular in cross section with indistinct median dorsal furrow, relatively thick, tapering gradually to tip, unsegmented, shorter than snout-vent length (TL/SVL 0.82). Scales on dorsal aspect of tail base similar to body dorsum; scales on dorsal aspect of tail large flat, subcircular, smooth, and imbricate, becoming slightly larger towards lateral aspect, largest on ventral side, but not forming median row of transversely enlarged subcaudal scales. Small, smooth, subequal (one on right distinctly larger), conical postcloacal spurs, two on either side of tail base; prominent hemipenial swelling with partially everted hemipenis on either side, flap of skin covering cloacal aperture. Tail slightly constricted at the base (Fig. 11A, B).

Dorsal ground colour khaki, four pairs of dark brown spots from behind occiput to hindlimb insertions and one pair above tail base, separated mid-vertebrally and extending onto lateral edge of body as streaks; roughly subequal, first spot and that on tail base smallest; spots with a fine black and finer outer cream border; lighter interspaces between spots less than thickness of spots. Tail dorsum sightly lighter than body dorsum with faint yellow tint; roughly seven dark crossbars on tail, last two light bands suffused with orange. Dorsum of limbs and digits with dark or grey reticulations. Post-occipital collar narrower than spots on dorsum, posteriorly darker with a fine light border, not in contact with post-orbital streak. Head dorsum slightly duller than trunk with irregular brown markings and a few scattered spots, a roughly V-shaped marking on snout-tip formed by pre-ocular streaks which do not meet in internasal region, prominent postocular streak merges with first pair of spots. Brille similar in colour to crown. Labials with few dark streaks, and a few unmarked scales finely spotted with black. Ventral aspects dirty white with numerous short streaks and spots on infralabials and gular region; ventral aspect of tail with thick dark reticulations and light markings that are less prominent on anterior half (Fig. 13A).

There are 10 adult male, two adult and three subadult female paratypes, SVL ranging from 36.9–53.5 mm (Tables 6, 7). All paratypes resemble the holotype except for the following variation: inner postmental bordered by mental, infralabial I, outer postmental in all paratypes; additionally, bordered by two chin scale on left and three on right in NRC-AA-9680 and NRC-AA-9691, two chin scale on left and four on right in NRC-AA-9692, five chin scale on left and two on right in NRC-AA-9683; three chin scale on left and four on right in NRC-AA-9684; four chin scales on either side in NRC-AA-9687 and NRC-AA-9690; two chin scales on either side in NRC-AA-9689 and NRC-AA-9692. Outer postmental bordered by inner postmentals, infralabial I and II in all paratypes; additionally, boarded by four chin scales on left and three on right in NRC-AA-9680, NRC-AA-9682, and NRC-AA-9694; three chin scales on left and a single scale on right in NRC-AA-9681; three chin scales on either side in NRC-AA-9683, NRC-AA-9686, NRC-AA-9688, NRC-AA-9691, NRC-AA-9692, and NRC-AA-9693; four chin scales on left and two on right in NRC-AA-9684; two chin scales on left and three on right in NRC-AA-9685; three chin scales on left and two on right in NRC-AA-9681; three chin scales on left and four on right in NRC-AA-9689; and two chin scales on either side in NRC-AA-9690. Ten paratypes (NRC-AA-9682, NRC-AA-9684, NRC-AA-9685, NRC-AA-9687, NRC-AA-9688, NRC-AA-9689, NRC-AA-9692, NRC-AA-9693, and NRC-AA-9694) have an original and complete tail, slightly shorter than body except for NRC-AA-9691, which has marginally shorter tail than body (TL/SVL 0.84, 0.85, 0.74, 0.85, 0.85, 0.82, 0.90, 0.84, 0.80, and 0.76 respectively); NRC-AA-9681 with partially regenerated tail, much shorter than body (TL/SVL 0.62), NRC-AA-9683 and NRC-AA-9686 with fully regenerated tails, much shorter than body (TL/SVL 0.54 and 0.68 respectively), NRC-AA-9680 and NRC-AA-9690 without tail (Fig. 14A, B). The type series have 5–7 pairs of dark blotches between neck and hindlimb insertions that may form broken or continuous bands with variation in size and arrangement of dorsal spots and overall colouration (Figs 13B–F, 14A, 14B).

Cyrtodactylus satpuraensis sp. nov. is known from multiple localities across the Satpura-Maikal landscape and lower elevations in eastern Maharashtra that span a maximum distance of > 770 km between elevations of ~270–1200 m in Madhya Pradesh and Maharashtra (Fig. 1). The species was observed in deciduous forest, with abundant leaf litter during summer, and most are at mid to high elevations (Fig. 13G). Cyrtodactylus satpuraensis sp. nov. was spotted emerging from under rocks or logs and crevices as darkness fell, on forest paths and in leaf litter after dark, and occasionally under rocks in the day. There are photographic records from Achanakmar-Amarkantak Biosphere Reserve, Anuppur District (Ingle 2020); Pachmarhi, Satpura Tiger Reserve, Hoshangabad District (IA pers. obs.), and Kanha Tiger Reserve, Mandla District (https://www.inaturalist.org/observations/238382363; https://www.inaturalist.org/observations/251979530); all from Madhya Pradesh; and it likely occurs in Pench Tiger Reserve (Agarwal 2007) and further east into Chhattisgarh.

The specific epithet is a toponym for the Satpura mountain range across which the new species is distributed. We suggest the common name Satpura Geckoella.