Search

Hemidactylus xericolus sp. nov. Figs. 5–8, Table 3 Holotype. National Centre for Biological Sciences (NCBS) NRC-AA-1110, adult male; collected from near Marrigudda, District Nalgonda, Telangana, India (16.96270° N, 78.85869° E, 430 m asl) on 23August 2015. Collected by Aparna Lajmi, Taneraw Singh and Maitreya Sil. Paratypes. CES 16170 adult male, NRC-AA-1111 adult female. Collection details are same as the holotype. Etymology. The specific epithet is a masculine adjective derived from the Greek xeros (= dry) and the Latin cola (= inhabitant of) and is descriptive of the arid, scrub habitats that the new species and many other endemic Indian Hemidactylus inhabit. The name also seeks to bring attention to the neglected and biodiverse Indian dry zone. Suggested common name. Nalgonda yellow-tailed brookiish gecko Diagnosis. A small sized Hemidactylus, snout-vent length up to at least 44.7 mm (n=3). Dorsal pholidosis heterogeneous composed of subcircular granular scales intermixed with moderately enlarged, circular, flattened, feebly keeled tubercles extending from occiput to tail and in 6–8 irregularly arranged longitudinal rows at midbody. Ventrolateral folds indistinct, about 21–26 scale rows across venter. Digits with slightly enlarged, divided scansors; lamellae in oblique series, seven (manus) and six or seven (pes) beneath fourth digit and four (manus and pes) beneath first digit; 15 or 16 precloacofemoral pores on each side separated by a single poreless scale in males (n=2). Original tail slightly depressed, verticillate, oval in transverse section with indistinct median dorsal furrow; dorsal tail pholidosis heterogenous with rounded, smooth, subimbricate scales intermixed with two to three enlarged, conical keeled tubercles at the base of every whorl, on either side of median dorsal furrow; subcaudals much enlarged; a pair of slightly enlarged postcloacal spurs on either side. Dorsal colouration of transversely arranged, pale grey to ashy markings on a pale, mustard-brown background; nuchal collar indicated by spots, not in contact with postorbital streak. The tail is distinctly yellow in adults of both sexes and juveniles. * indicates tail incomplete, L & R = left and right; numbers in parentheses indicate either number of poreless scales separating femoral pore series, number of notched or undivided lamellae, supralabial at midorbital position. ...Continued on the next page Comparison with other members of the H. brookii group. Hemidactylus xericolus sp. nov. can be distinguished from many members of the H. brookii group by its small adult size (maximum SVL 44.7 mm versus 55.8 in H. brookii, 51.5 mm in H. chikhaldaraensis, 74.2 mm in H. chipkali, 56.2 mm in H. cf. gleadowi, 51.4 mm in H. kushmorensis, 65.0 mm in H. murrayi (Lajmi et al. 2016), 53.8 mm in H. malcolmsmithi, 52.3 mm in H. parvimaculatus (Lajmi et al. 2016), 62.5 mm in H. rishivalleyensis, 50.8 mm in H. sankariensis, 61.7 mm in H. subtreidroides, 70.2 mm in H. treutleri, and 61.9 mm in H. varadgirii). Hemidactylus xericolus sp. nov. can be distinguished from the species that are less than 55 mm SVL by the presence of dorsal tubercles in 6–8 irregularly arranged rows versus 15–17 fairly regularly arranged rows in H. chikhaldaraensis; 19 or 20 fairly regularly arranged longitudinal rows in H. kushmorensis; 15–20 fairly regularly arranged longitudinal rows in H. malcolmsmithi; 15–18 fairly regularly arranged longitudinal rows in H. parvimaculatus; 15 fairly regularly arranged longitudinal rows in H. sankariensis. The new species can be easily distinguished from members of the ground dwelling clade by the presence of 15 or 16 precloacofemoral pores on each side in males (versus H. albofasciatus Grandison & Soman, H. gracilis Blanford, H. imbricatus, H. reticulatus Beddome, H. sataraensis Giri & Bauer and H. vijayraghavani Mirza). Hemidactylus xericolus sp. nov. can be distinguished from H. gleadowi by the lower number of scales across the belly (21–26 versus 32–34) and fewer dorsal tubercle rows (6–8 versus 17 or 18). The yellow colouration of the tail in life, the dorsal colour pattern and the very low number of dorsal tubercle rows are unique among Indian Hemidactylus. Hemidactylus xericolus sp. nov. is most closely related to its sister species, Hemidactylus flavicaudus sp. nov. but is genetically deeply divergent (Table 2) and can be diagnosed from based on fewer dorsal tubercle rows at midbody (6–8 versus 11–14) as well as in dorsal colour pattern (upper preorbital streaks on each side almost meeting at rostral versus meeting at rostral; nuchal collar indicated by spots and usually not meeting postorbital streak laterally versus complete nuchal collar meeting postorbital streak laterally. Holotype description. The holotype is generally in good condition with some minor exceptions; all artifacts of preservation: the body shape is somewhat dorsoventrally flattened, tail partly regenerated, is curved in a sigmoid manner towards the right. Head short (HL/ SVL 0.27), slightly elongate (HW /HL 0.71), not strongly depressed (HH/HL 0.36), relatively broad (HW / BW 0.83), distinct from neck. Loreal region slightly inflated, canthus rostralis not prominent. Snout short (SE /HL 0.41); slightly longer than eye diameter (OD / SE 0.56); scales on snout, canthus rostralis and forehead region homogeneous in shape and slightly heterogeneous in size, rounded, not flat and juxtaposed; distinctively larger than those on interorbital and occipital region; scales on interorbital and occipital region homogeneous in shape, slightly heterogeneous in size, granular, those on occipital region smallest. Eye small (OD / HL 0.23); pupil vertical with crenate margins; supraciliaries small, mucronate, increasing in size from midorbital position with those on the anterior end of orbit largest. Ear opening oval (greatest diameter 1.20 mm); eye to ear distance greater than diameter of eye (EE / OD 1.34). Rostral wider (1.9 mm) than deep (1.1 mm), completely divided dorsally by weakly developed rostral groove; two enlarged internasals in contact medially, one supranasal on each side which is half the size of internasal; two postnasals on either side, slightly smaller than supranasal; both postnasals on right side are of similar size; rostral in contact with nasal, supralabial I and internasal; nostrils large, slightly oval, directed upwards, covering most of the nasal scale; nasal on either side surrounded by supranasal, internasal, rostral, supralabial I and postnasals. Mental almost equal in length (1.9 mm) and width (1.8 mm) triangular, two well-developed postmentals, the inner pair slightly shorter (1.5 mm) than mental and not in contact with each other behind mental, outer twice in length (1.5 mm) than the inner pair (0.7 mm), separated from each other by inner pair. Inner postmentals bordered by mental, infralabial I, slightly infralabial II, outer postmental and two gular scales; outer postmental bordered by inner postmental, infralabial II, and four gular scales. Infralabials bordered by a single row of enlarged and elongated scales starting from centre of infralabial III to infralabial VI on right and from below infralabial III to infralabial VII on left. Eight supralabials (to midorbital position) on either side; 10 supralabials (to angle of jaw) on either side; seven infralabials (to angle of jaw) on the right and eight on the left side. Body slender, trunk not elongate (TRL / SVL 0.48), with indistinct ventrolateral folds without denticulate scales. Dorsal pholidosis heterogeneous, composed of conical, granular scales intermixed with irregularly arranged, longitudinal rows of 6–8 slightly larger, rounded, feebly keeled, indistinct tubercles at midbody, extending from shoulder to tail; those on parasagittal rows slightly smaller, gradually increasing in size laterally, being slightly larger on flanks; roughly two to three times longer than adjacent granules, surrounded by a rosette of 6–8 small granules, 2–6 granules between two adjacent enlarged tubercles. Ventral scales much larger than granular scales on dorsum, roughly hexagonal, smooth, imbricate, more or less similar in size; midbody scale rows across belly 24 or 25; gular region with still smaller, smooth, rounded, subimbricate scales, increasing in size anteriolaterally, those below mental, postmentals and infralabials are largest. Femoral and precloacal pores 16 on each side, separated by a single non-pored scale. Scales on the palm and sole smooth, granular, rounded; scales on forelimb homogeneous, dorsal aspect with rounded, smooth, subimbricate scales which are slightly smaller than largest enlarged tubercles on dorsum, those on ventral portion are smaller and granular; scales on hindlimbs heterogenous in size and shape, dorsal part of thigh and shank are similar like dorsum, with granular scales, intermixed with scattered, enlarged, rounded, slightly conical, feebly keeled tubercles, which are denser on shank than thigh, anterior portion of thighs and ventral aspect of hindlimbs with much enlarged, smooth, imbricate scales. Fore- and hind limbs relatively slender; forearm short (FL / SVL 0.13); tibia short (CL/ SVL 0.15); digits moderately long, strongly clawed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, half or more than half as long as associated toepad; scansors beneath each toe in oblique series divided, except a distal and one or two basal scansors in all digits single; scansors from proximalmost at least twice the diameter of palmar scales to distalmost single scansor: 4-6-7-7-6 (left manus) 4-6-7-7-6 (right manus), 4-6-7-7-6 (left pes) 4-7-7-6-6 (right pes). The first digit on all limbs shorter, slightly less than half the length of second digit. Relative length of digits (measurements in mm in parentheses): III(3.4)> IV(3.2)> V (3.1)> II(2.8)> I(1.9) (left manus); IV(4.1)> V (3.8)> III(3.2)> II(3.0)> I(2.1) (left pes). Tail slightly depressed, verticillate, oval in transverse section with indistinct median dorsal furrow; dorsal tail pholidosis heterogenous with smaller, rounded, smooth, subimbricate scales, gradually increasing in size laterally, intermixed with four to six enlarged, conical, weakly keeled tubercles on every whorl, two to three on either side of median dorsal furrow; ventral scales enlarged, imbricate, median row (subcaudal plates) covering almost entire base of the tail, bordered laterally by two or three rows of larger, smooth, imbricate scales; those close to vent small, smooth, flat and imbricate (Fig. 6 B). A pair of slightly enlarged postcloacal spurs on either side. Colouration in life (Fig. 5 B). Dorsal ground colour of head, body, limbs and tail beige with scattered indistinct lighter blotches. Two distinct dark preorbital streaks enclosing a yellow streak, upper narrower and converging from each side at rostral but not meeting, lower terminating at supralabials three and four; distinct dark postorbital streak flanked by narrow yellow markings which is broader than the lower preorbital streak and extends from behind eye until the nuchal collar with a break just after the tympanum; anterior portion of brille yellow. Head dorsum mottled and suffused with yellow, interorbital region slightly bluish, three large spots forming a transverse series on occiput in line with the end of the postorbital streak; nuchal collar indicated by three dark spots, the central largest; a large dark spot above forelimb insertions; labials with fine black spots and some yellow on anterior supralabials, rostral yellowish. Dorsum with four sets of dark markings between limb insertions; the first three consist of a central spot flanked on four corners by four smaller or subequal spots (forming the arms of an X) and the fourth has a central spot flanked by one spot on either side, the outer spots flanked by another row of similar sized spots on the flanks. Tibia, tarsus, femur and digits with a few dark cross bars; tail suffused with saffron-yellow, five dark cross bars on original portion, regenerated portion similar to rest of tail ground colour except with less yellow. Venter offwhite, immaculate. Variation and additional information from type series (Fig. 8, Table 3). There is one male and one female specimen that are very similar in SVL to the holotype (44.7. 44.5 mm). The paratypes resemble the holotype except as follows: supralabials nine on the right side of NRC-AA-1111; inner pair of postmental is in contact with each other in NRC-AA-1111; lamellae six on right side of fourth digit of pes; femoral pores 15 of left side of NRC-AA- 1110. The overall colour pattern is consistent across the type series, though the size and shape of dorsal markings is variable, there are 13 dark bands on the complete original tail of CES 16169, and the nuchal collar in CES 16169 is indicated by only two spots. Distribution. Hemidactylus xericolus sp. nov. is known only from its type locality. Habitat and Natural History. The type locality is surrounded by hills with rocky boulders and scrub vegetation (Fig. 9). The vegetation is denser than at localities where Hemidactylus flavicaudus sp. nov. is known from and included stunted trees. The specimens were collected from horizontal surfaces of large boulders or on vertical rock surfaces at low heights of under four feet. Sympatric congeners in the rocky habitat included H. cf. treutleri, H. cf. giganteus, and an undescribed large tuberculated species (denoted by “ Hemidactylus species 2” from Lajmi & Karanth 2020). Hemidactylus cf. triedrus, H. cf. reticulatus, and H. cf. gleadowi were also seen in the adjoining scrub at the base of the hillock.

Hemidactylus flavicaudus sp. nov. Figs. 2–5, Table 3 Holotype. National Centre for Biological Sciences (NCBS) NRC-AA-1105, adult female; collected from near Guddeguda village, Mahabubnagar District, Telangana, India (16.68069° N, 77.82587° E, 416 m asl) on 22August 2015 by Aparna Lajmi, Aniruddha Datta Roy, Deepak Veerappan and Praveen Karanth. Paratypes. CES 17028 adult female, CES 17029, CES 17031 adult males (same collection data as holotype); NRC-AA-1106, NRC-AA-1108, adult females (same collection data as holotype except collected on 20 April 2015 by Aparna Lajmi, Taneraw Singh and Maitreya Sil; NRC-AA-1107, CES 16129 adult females, NRC-AA-1109 adult male collected from Manyamkonda, Mahabubnagar District, Telangana, India (16.6504° N, 77.888° E, 560 m asl) on 20 April 2015 by Aparna Lajmi, Taneraw Singh and Maitreya Sil. Etymology. The specific epithet is an adjective derived from the Latin flavus (= yellow) and cauda (= tail), for the yellow tail of the new species. Suggested common name. Mahabubnagar yellow-tailed brookiish gecko Diagnosis. A small sized Hemidactylus, snout-vent length up to at least 41.8 mm (n=9). Dorsal pholidosis heterogeneous composed of subcircular granular scales intermixed with moderately enlarged, circular, flattened, feebly keeled tubercles extending from occiput to tail and in 11–14 irregularly arranged longitudinal rows at midbody. Ventrolateral folds indistinct, about 22–26 scale rows across venter. Digits with slightly enlarged, divided scansors; lamellae in oblique series, six or seven (manus) and seven or eight (pes) beneath fourth digit and four (manus) and four or five (pes) beneath first digit; 15–17 precloacofemoral pores on each side separated by one or two poreless scales in males (n=3). Original tail slightly depressed, verticillate, oval in transverse section with indistinct median dorsal furrow; dorsal tail pholidosis heterogenous with rounded, smooth, subimbricate scales intermixed with two to three enlarged, conical keeled tubercles at the base of every whorl, on either side of median dorsal furrow; subcaudals much enlarged; a pair of slightly enlarged postcloacal spurs on either side. Dorsal colouration of transversely arranged, pale grey to ashy markings on a pale, mustard-brown background; thick nuchal collar which fuses with postorbital streaks laterally. Tail distinctly yellow in adults of both sexes and juveniles. Comparison with other members of the H. brookii group. Hemidactylus flavicaudus sp. nov. can be distinguished from all other members of the Indian radiation, except H. xericolus sp. nov. based on its conspicuous yellow tail. It can be further distinguished from many members of the H. brookii group by its small adult size (maximum SVL 41.8 mm versus 55.8 in H. brookii, 51.5 mm in H. chikhaldaraensis Agarwal, Bauer, Giri & Khandekar, 74.2 mm in H. chipkali Mirza & Raju, 56.2 mm in H. cf. gleadowi, 51.4 mm in H. kushmorensis, 65.0 mm in H. murrayi (Lajmi et al. 2016), 53.8 mm in H. malcolmsmithi (Constable), 52.3 mm in H. parvimaculatus Deraniyagala (Lajmi et al. 2016), 62.5 mm in H. rishivalleyensis Agarwal, Thackeray & Khandekar, 50.8 mm in H. sankariensis Agarwal, Bauer, Giri & Khandekar, 61.7 mm in H. subtreidroides, 70.2 mm in H. treutleri Mahony, and 61.9 mm in H. varadgirii Chaitanya, Agarwal, Lajmi & Khandekar). Hemidactylus flavicaudus sp. nov. can be distinguished from the species that are less than 55 mm SVL by the presence of moderately enlarged, flattened, feebly keeled dorsal tubercles in 11–14 irregularly arranged rows versus strongly enlarged and keeled, pointed tubercles in 15–17 fairly regularly arranged longitudinal rows in H. chikhaldaraensis; enlarged and keeled conical dorsal tubercles in 19 or 20 fairly regularly arranged longitudinal rows in H. kushmorensis; slightly enlarged, keeled, conical tubercles in 15–20 fairly regularly arranged longitudinal rows in H. malcolmsmithi; slightly enlarged, keeled, conical tubercles in 15–18 fairly regularly arranged longitudinal rows in H. parvimaculatus; strongly enlarged and keeled, pointed tubercles in 15 fairly regularly arranged longitudinal rows in H. sankariensis. The new species can be easily distinguished from members of the ground dwelling clade by the presence of 15–17 precloacofemoral pores on each side in males (versus less than eight precloacal pores in males of H. albofasciatus Grandison & Soman, H. gracilis Blanford, H. imbricatus, H. reticulatus Beddome, H. sataraensis Giri & Bauer and H. vijayraghvani Mirza). Hemidactylus flavicaudus sp. nov. can be distinguished from H. gleadowi by the lower number of scales across the belly (22–26 versus 32–34) and fewer dorsal tubercle rows (11–14 versus 17 or 18). The yellow colouration of the tail in life as well as the dorsal colour pattern are unique among Indian Hemidactylus except for Hemidactylus xericolus sp. nov., the diagnosis against which is in the species description. Holotype description. The holotype is generally in good condition with some minor exceptions; all artefacts of preservation: the body shape is somewhat dorsoventrally flattened, the partly regenerated tail is curved in a sigmoid manner towards the right. Head short (HL/ SVL 0.28), slightly elongate (HW /HL 0.70), not strongly depressed (HH/ HL 0.39), relatively broad (HW / BW 0.82), distinctly broader than neck. Loreal region slightly inflated, canthus rostralis not prominent. Snout short (SE /HL 0.44); slightly longer than eye diameter (OD / SE 0.54); scales on the snout, canthus rostralis and forehead region homogeneous in shape and slightly heterogeneous in size, rounded, flat and juxtaposed; distinctively larger than those on interorbital and occipital region; scales on interorbital and occipital region homogeneous in shape, slightly heterogeneous in size, granular, those on occipital region smallest. Eye small (OD /HL 0.24); pupil vertical with crenate margins; supraciliaries small, mucronate, increasing in size from mid-orbital position with those on the anterior end of orbit largest. Ear opening oval (greatest diameter 1.00 mm); eye to ear distance greater than eye diameter (EE / OD 1.30). Rostral wider (1.6 mm) than deep (0.9 mm), completely divided dorsally by weakly developed rostral groove; two enlarged internasals in contact medially, one supranasal on each side which is half the size of internasal; two postnasals on either side, slightly smaller than supranasal; both postnasals on right side are of similar size; on left, lower postnasal is smaller than the upper; rostral in contact with nasal, supralabial I and internasal; nostrils large, slightly oval, directed upwards, covering most of the nasal scale; nasal on either side surrounded by supranasal, internasal, rostral, supralabial I and postnasals. Mental almost equal in length (1.8 mm) and width (1.7 mm) triangular, two well-developed postmentals, the inner pair slightly shorter (1.4 mm) than mental and narrowly in contact with each other (0.5 mm) behind mental, outer pair slightly smaller (1.0 mm) than the inner pair, separated from each other by inner pair. Inner postmentals bordered by mental, infralabial I, barely infralabial II, outer postmental and four gular scales; outer postmental bordered by inner postmental, infralabial II, and four gular scales. Infralabials bordered by a single row of enlarged and elongated scales starting from centre of infralabial III to infralabial VI on right and from below infralabial III to infralabial VII on left. Eight supralabials (to midorbital position) on either side; 11 supralabials (to angle of jaw) on the right and 10 on the left; eight infralabials (to angle of jaw) on the either side. Body slender, trunk not elongate (TRL / SVL 0.46), with indistinct ventrolateral folds without denticulate scales. Dorsal pholidosis heterogeneous, composed of conical, granular scales intermixed with irregularly arranged, longitudinal rows of 12 or 13 slightly larger, rounded, keeled tubercles at midbody, extending from nape to tail; those on nape and two parasagittal rows more or less of similar size, smallest, gradually increasing in size laterally, being largest on flanks; roughly two to three times longer than adjacent granules, surrounded by a rosette of eight or nine small granules, 2–5 granules between two adjacent enlarged tubercles. Ventral scales much larger than granular scales on dorsum, roughly hexagonal, smooth, imbricate, more or less similar in size; midbody scale rows across belly 24 or 25; gular region with still smaller, smooth, rounded, subimbricate scales, increasing in size anterolaterally, those below mental, postmentals and infralabials are largest. Femoral and precloacal pores absent. Scales on the palm and sole smooth, granular, rounded; scales on forelimb homogeneous, dorsal aspect with rounded, smooth, subimbricate scales which are slightly smaller than largest enlarged tubercles on dorsum, those on ventral portion are smaller and granular; scales on hindlimbs heterogeneous in size and shape, dorsal part of thigh and shank are similar to the dorsum, with granular scales, intermixed with scattered, enlarged, rounded, slightly conical, feebly keeled tubercles, which are denser on shank than thigh, anterior portion of thighs and ventral aspect of hindlimbs with much enlarged, smooth, imbricate scales. Fore and hind limbs relatively slender; forearm short (FL / SVL 0.14); tibia short (CL/ SVL 0.15); digits moderately long, strongly clawed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, half or more than half as long as associated toepad; scansors beneath each toe in oblique series divided, except a distal and one or two basal scansors in all digits single; scansors from most proximal at least twice the diameter of palmar scales to most distal single scansor: 4-6-6-7-6 (left manus), 4-6-6-7-6 (right manus; Fig. 3 E), 4-7-7-7-6 (left pes) 4-7-8-8-6 (right pes; Fig. 3 F). The first digit on all limbs shorter, slightly less than half the length of second digit. Relative length of digits (measurements in mm in parentheses): III (2.9)> IV (2.8)> V (2.6)> II (2.4)> I (1.7) (left manus); IV(3.5)> V (3.2)> III(3.1)> II(3.0)> I(1.8) (left pes). Tail slightly depressed, verticillate, oval in transverse section with indistinct median dorsal furrow; dorsal tail pholidosis heterogenous with smaller, rounded, smooth, subimbricate scales, gradually increasing in size laterally, intermixed with four to six enlarged, conical, weakly keeled tubercles on every whorl, two to three on either side of median dorsal furrow; scales on regenerated tail dorsum subimbricate; ventral scales enlarged, imbricate, median row (subcaudal plates) covering almost entire base of the tail except regenerated portion, bordered laterally by two or three rows of larger, smooth, imbricate scales; those close to vent small, smooth, flat and imbricate (Fig. 2 B). Postcloacal spurs indistinct. Colouration in preservative (Fig. 2). Dorsal ground colour of head, body, limbs and tail beige with scattered indistinct light brown blotches. Two distinct dark preorbital streaks, upper narrower and converging from each side and meeting at rostral, roughly ‘V’ shaped when viewed from above; lower bordering supralabials 3 to 6; distinct postorbital streak broader than the lower preorbital streak and extends from behind eye until above forelimb insertions. Head dorsum mottled, three large spots forming a transverse series on occiput; distinct dark nuchal collar extending across base of neck and fusing with postorbital streak on either side anterior to forelimb insertions; labials with fine black spots. Dorsum with four dark approximately X-shaped markings between limb insertions. The first lies just posterior to forelimb insertions and consists of a central longitudinally elongate rectangular spot, flanked on its four corners by four smaller or subequal spots. The other three sets of markings are less distinct but always include the central elongate spot, and may be similar to the first set of markings (third set) or consist of the central marking flanked by a single spot on either side (second and fourth set). Forelimbs and femur mottled with light and dark splotches, tibia and digits with dark crossbars; tail with seven dark cross bars on original portion, regenerated portion darker than rest of tail ground colour. Venter off-white, immaculate. Colouration in life (Fig. 5A; based on paratype CES 16128). Dorsal ground colour of head, body, limbs and tail beige with light tinge of yellow and scattered indistinct light brown blotches. Two distinct dark preorbital streaks enclosing a yellow streak, upper narrower, starting as a dark spot, converging from each side and meeting at rostral, roughly ‘V’ shaped when viewed from top, lower terminating at supralabials third and fourth; distinct postorbital streak flanked by diffuse yellow markings which is broader than the lower preorbital streak and extends from behind eye until above forelimb insertions; anterior portion of brille yellow. Head dorsum mottled, snout suffused with yellow, rostral yellow; interorbital region slightly bluish; a large and small dark spot and pale blotch on occiput; distinct dark nuchal collar extending across base of neck and fusing with postorbital streak on either side anterior to forelimb insertions; labials with fine black spots, anterior more strongly spotted and suffused with yellow. Dorsum with scattered light blotches and an indistinct light mid-ventral line; four dark approximately X-shaped markings between limb insertions. The first lies just posterior to forelimb insertions and consists of a central longitudinally elongate rectangular spot, flanked on its four corners by four smaller or subequal spots. The other three sets of markings are less distinct but always include the central elongate spot, and may be similar to the first set of markings (third set) or consist of the central marking flanked by a single spot on either side (second and fourth set). Forelimbs and femur mottled with light and dark splotches, tibia and digits with dark crossbars; tail yellow with 11 dark cross bars those are distinct towards the base becoming pale and indistinct towards the tail tip. Venter off-white, immaculate. Variation and additional information from type series (Fig. 4, Table 3). There are three male and six female specimens ranging in size from 31.0 – 41.8 mm (average 39.2 mm). The paratypes resemble the holotype except as follows: the number of lamellae on digit IV of manus is 7 except CES 17029 which has 6. The number of supralabials is 9–11 except CES 17031 has 8 on left side. The inner pair of postmental is in contact with each other except NRC-AA-1107 and CES 17031 not in contact. Femoral pores on either side are separated by two unpored scales but CES 17029 has single. The overall colour pattern is consistent across the type series, though the size and shape of dorsal markings are variable, the first mark on the back behind forelimb insertion is distinctly ‘X’ shaped and other two bands on back are distinct in NRC-AA-1107, CES 16129 and NRC-AA-1109. There are 13 dark bands on the complete original tail of 16128 and CES 16129 and 11 on NRC-AA-1109, but the regenerated tail lacks these markings. The nuchal collar in 16129 is indicated by only two spots and CES 17029 and CES 17031 by three spots. Distribution. Hemidactylus flavicaudus sp. nov. is currently known only from Guddeguda, the type locality and Oblaipalle, ~ 7–8 km away, both in Mahabubnagar District, Telangana, India. Habitat and Natural History. The type locality of Hemidactylus flavicaudus sp. nov. is a small rocky outcrop close to Guddeguda village characterised by large rocky boulders interspersed with scrub vegetation (Fig. 9), as is the other known locality Oblaipalle. All individuals of the new species were found at night on vertical surfaces of boulders and usually at a relatively low height of 90–120 cm from the ground. The most common species of Hemidactylus on the boulders were H. cf. treutleri followed by H. flavicaudus sp. nov. Other sympatric rupicolous congeners included H. cf. giganteus Stoliczka and a large tuberculated undescribed species (denoted by “ Hemidactylus species 3” from Lajmi & Karanth 2020). The terrestrial Hemidactylus cf. gleadowi was also noted in the vicinity of the rocky outcrop, on the soil surface. CES16128 and CES16129 collected on 20 April were noted to be carrying eggs, indicating this to be the breeding period (Fig. 5 A).

Dravidogecko smithi sp. nov. (Figs 10 A���D, 13C, 15A; Table 7) Holotype. BNHS 2350, an adult male, Ponmudi Hills (8.7570 ��N, 77.1145 ��E; ca. 920 m asl.), Tiruvananthapuram District, Kerala, collected by Jafer Palot and RC on 25 th November, 2017. Paratypes. Details of collection same as the holotype. ZSIK 2981, adult female. Type locality. Ponmudi Hills, Tiruvananthapuram District, Kerala. Summarized description and diagnosis. Snout-vent length up to 49.1 mm (n=2); one scale between internasals; two pairs of well-developed postmentals, inner pair longer than the outer but shorter than mental, bordered posteriorly by 2 or 3 gular scales; ventral scales counted at midbody, 29���32; precloacofemoral pores, 48 (n=1); subdigital lamellae under digit IV of manus, 8 or 9 and under digit IV of pes, 10 or 11; supralabials, 9 or 10 and infralabials, 7 or 8 on each side. Dravidogecko smithi sp. nov. can be distinguished from other congeners based on the following characters: number of precloacofemoral pores (PcFP 48 versus 45 or 46 in D. anamallensis, 52���56 in D. septentrionalis sp. nov., 36���38 in D. meghamalaiensis sp. nov. & 42 or 43 in D. douglasadamsi sp. nov.); postmentals shorter in length than mental (ML /1PML 1.07���1.12 versus longer, 0.74���0.81 in D. anamallensis); one scale separating internasals (versus two in D. septentrionalis sp. nov.). Genetic divergence (p-distance). Dravidogecko smithi sp. nov. exhibits 0.2% intraspecific variation for the mitochondrial ND2 gene, while it is 10.8% ���17.0% divergent from all other congeners. Despite the proximity in range with D. douglasadamsi sp. nov. (straight line distance of ca. 50 kms), D. smithi sp. nov. exhibits 11.3% divergence from the former (Table 9). Description of holotype. The holotype is generally in good condition (Fig 10A). Hemipenes everted, and visible on both sides when viewed dorsally. Posterior half of tail regenerated, tip of which is curved upwards, fourth and fifth fingers on right forelimb curved upwards���both artefacts of preservation (Fig 10A). Adult male, SVL 49.1 mm. Head short (HL/SVL 0.27), slightly elongate (HW/HL 0.61), slightly depressed (HH/HW 0.55), distinct from neck. Loreal region slightly inflated, canthus rostralis indistinct (Fig 10C). Snout short (SE/HL 0.36), longer than orbital diameter (OD/SE 0.64); scales on snout, canthus rostralis, inter-orbital region, forehead, occipital and nuchal regions granular and rounded with those on the snout and canthus rostralis being larger (Fig 10B). Eye small (OD/HL 0.23); pupil vertical with crenulated margins; supraciliaries small, rounded, directed outwards, increasing in size anteriorly. Ear opening roughly elliptical (longer diameter 0.6 mm); eye to ear distance longer than diameter of eye (EE/OD 1.15). Rostral wider than deep (RL/RW 0.33), rostral groove distinct but extending only marginally downwards from the suturing with the internasals, medially; two large, roughly circular internasals, separated by a smaller scale, all in broad contact with rostral; two postnasals on either side, slightly smaller than the internasals, the lower in contact with supralabial I; rostral in contact with nasal, supralabial I, internasals and the smaller scale separating the internasals; nostrils about the size of the lower postnasal, roughly circular with nasal pad visible posteriorly; nasal surrounded by internasal, rostral, two postnasals and supralabial I on either side; 2 or 3 rows of scales separate orbit from supralabials around mid-orbital position. Supralabials roughly rectangular, increasing in length anteriorly. Supralabials (to midorbital position) 7 (right), 7 (left); supralabials (to angle of jaw) 9 (right), 9 (left); infralabials (to angle of jaw) 7 (right), 8 (left). Mental triangular; two pairs of smaller postmentals, the inner pair slightly shorter (1.0 mm) than the mental (1.2 mm), and in strong contact with each other (0.7 mm) behind mental; outer pair shorter still (0.8 mm), separated from each other by two gular scales that are smaller than postmentals (Fig 10D). Inner postmentals bordered by mental, infralabial I, outer postmentals and the two smaller gular scales that separate the outer postmentals; outer postmentals bordered by infralabials I and II, inner postmentals, and four smaller gular scales each of dissimilar sizes. Body dorsoventrally flattened, relatively slender, elongate (TRL/SVL 0.46). Dorsal pholidosis composed of small, rounded granules that are juxtaposed in arrangement throughout, becoming slightly larger at the lateral aspects; Ventral scales larger than dorsals, largely homogeneous in shape and size, smooth, flat, weakly pointed and sub-imbricate; gular region with smaller, granular, juxtaposed scales, anterior-most gular scales visibly larger, flatter; scales on sacral and femoral regions larger than those on chest; precloacal scales larger still; midbody scale rows across belly 31 or 32; Non-lamellar scales in the palmar and plantar regions heterogeneous in size, flat, rounded, sub-imbricate; scales on dorsal aspect of upper arm larger than granules on dorsum, flat, weakly pointed, sub-imbricate and smooth; dorsal aspect of forearm with smaller, sub-imbricate scales intermixed with a few rounded granules around the elbow; scales on dorsal aspect of hand and digits larger than those on forearm, flat, weakly pointed and imbricate; scales on anterior aspect of thigh large, flat, sub-imbricate and weakly pointed; rest of the dorsal scales on hindlimb smaller, granular and rounded. Scales on dorsal aspect of feet and toes larger than those on shank, flat, weakly pointed and imbricate. Forearm (FL/SVL 0.11) and tibia short (CL/SVL 0.11); digits moderately short with relatively long terminal phalanges, strongly clawed; all digits of manus and digits I���IV of pes indistinctly webbed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, more than half as long as associated toepad; scansors beneath each toe undivided throughout, in a straight transverse series: 6-6-7-8-7 (left manus), 5-6- 7-8-8 (right manus), 6-8-9-10 -8 (left pes), 5-9-10-10 -8 (right pes). Relative length of digits (measurements in mm in parentheses): IV (3.9)> III (3.8)> II (3.3)> V (3.1)> I (2.7) (left manus); IV (4.7)> III (4.3)> II (4.0)> V (3.8)> I (3.2) (left pes). Tail rounded at the base with distal half regenerated, flat beneath, tapering posteriorly, covered above uniformly with round, smooth, flat, sub-imbricate scales that become slightly larger laterally; basal portion of tail with six or seven rows of flat, weakly pointed, sub-imbricate scales; subsequent subcaudal scales larger, with an undivided median series of enlarged scales extending to tail tip. An uninterrupted series of 48 precloacofemoral pores, that are only faintly visible towards the knee (Fig 13C). Variation in paratype. Rostral groove absent in ZSIK 2981. Inner postmentals bordered posteriorly by three gular scales; outer postmentals bordered by 3 gulars on left in ZSIK 2981. Other morphological variations are listed in Table 7. Colour in preservative. Dorsum uniformly greyish-brown, mottled with darker, discontinuous horizontal streaks in the trunk (Fig 10A). Similar mottling faintly visible on dorsal aspect of limbs. Occipital region with a dark, longitudinal streak, flanked anteriorly by two dark spots. Two discontinuous lines emanate from the eye, breaking posteriolaterally at the head, following the contour of the cranium laterally and extending beyond the forearm insertion. Inter-orbital region with a scattering of dark spots, with a distinct dark blotch bordering the supraciliary region on either side. Labials paler than the rest of the head with a faint, pattern-less scattering of darker spots on each labial. A dark, roughly rectangular streak emanates from eye upto the region above the third supralabial on the right side and the nostril on the left. Limbs no different from rest of the dorsum. Tail of similar ground colour to dorsum with alternating pale-dark longitudinal bands, the first of which is roughly saddle-shaped, up to the regenerated portion. Regenerated portion of tail uniformly greyish throughout with a scattering of darker longitudinal streaks. Ventral region cream coloured with a scattering of three to five dark spots on each ventral scale. Ventral surface of tail pale, with scattered mid-brown speckling in the hemipenial region followed by alternating pale-dark bands up to the regenerated portion. Colouration (in life). Dorsum mid-brown in life (Fig 15A). Distinct yellow blotches visible across dorsal aspect of head, trunk and original portion of tail. Snout predominantly yellow. Lateral aspect with a series of pale yellow spots. Iris dark green with darker venations. Pupil black, with indistinctly crenulated margins. Other patterns and markings in accordance with the description of colour in preservative. Etymology. The specific epithet is an eponym honouring British herpetologist Malcolm Arthur Smith for establishing the genus Dravidogecko in the year 1933. His seminal work on Indian herpetology, resulting in the text ���The fauna of British India, including Ceylon and Burma ��� in three volumes, is still considered the bedrock of reptilian taxonomy in India. Suggested Common name. Smith���s Dravidogecko. Distribution. Dravidogecko smithi sp. nov. is currently restricted in distribution to the Ponmudi Hills in Thiruvananthapuram District, Kerala. The habitat chiefly constitutes tropical evergreen rainforests (Champion & Seth 1968). The Agastyamalai Hill Range just south of Ponmudi has similar habitats in which Dravidogecko might be found. Habitat and natural history. The type-series of Dravidogecko smithi sp. nov. was collected in the Ponmudi Hills at an altitude of ca. 900 m asl. These geckos are found occupying human structures that are scattered along the road to the Ponmudi Hills. Other lizards found in sympatry with Dravidogecko in the region were Hemidactylus cf. frenatus, Cnemaspis sp. and Eutropis cf. carinata, which was also abundant in the adjoining shola grasslands., Published as part of Chaitanya, R., Giri, Varad B., Deepak, V., Datta-Roy, Aniruddha & Karanth, Praveen, 2019, Diversification in the mountains: a generic reappraisal of the Western Ghats endemic gecko genus Dravidogecko Smith, 1933 (Squamata: Gekkonidae) with descriptions of six new species, pp. 1-56 in Zootaxa 4688 (1) on pages 27-30, DOI: 10.11646/zootaxa.4688.1.1, http://zenodo.org/record/3514770, {"references":["Champion, H. G. & Seth, S. K. (1968) A revised survey of the forest types of India. Government of India, Press, New Delhi, 404 pp., XXVII pls."]}

Dravidogecko tholpalli sp. nov. (Figs 11 A���D, 13G, 15B; Table 7) Hoplodactylus anamallensis: Boulenger, 1885 Hoplodactylus anamallensis [non Gecko anamallensis G��nther, 1875] ��� Boettger, 1893. Hemidactylus anamallensis: Bauer & Russell, 1995 Hemidactylus anamallensis [non Gecko anamallensis G��nther, 1875] ��� Ganesh, 2010; Holotype. BNHS 2351, an adult male, Kodaikanal town (10.2334 ��N, 77.4910 ��E; ca. 2110 m asl.), Dindigul District, Tamil Nadu, collected by R. Venkitesan and RC on 17 th December, 2016. Paratypes. Details of collection same as the holotype. BNHS 2352, BNHS 2353, ZSIK 2982, ZSIK 2984, ZSIK 2985, ZSIK 2986 ���adult males; BNHS 2354, BNHS 2355 and ZSIK 2983 ���adult females. Type locality. Kodaikanal, Dindigul District, Tamil Nadu. Summarized description and diagnosis. Snout-vent length up to 52.2 mm (n=10); internasals separated by one smaller scale; two pairs of well-developed postmentals, inner pair longer than the outer; ventral scales counted at midbody, 25���31; precloacofemoral pores, 38���40 (n=7); subdigital lamellae under digit IV of manus, 7 or 8 and under digit IV of pes, 9���11; supralabials 8���11 and infralabials, 8���10 on each side. Dravidogecko tholpalli sp. nov. can be distinguished from other congeners based on the following characters: number of precloacofemoral pores (PcFP 38���40 versus 45 or 46 in D. anamallensis, 52���56 in D. septentrionalis sp. nov., 36���38 in D. meghamalaiensis sp. nov., 42 or 43 in D. douglasadamsi sp. nov. & 48 in D. smithi sp. nov.); one smaller scale separating the internasals (versus two in D. septentrionalis sp. nov.); first pair of postmentals much longer than the second (2PML/1PML 0.41���0.67 versus only slightly longer, 0.82���0.96 in D. meghamalaiensis sp. nov.). Genetic divergence (p-distance). Dravidogecko tholpalli sp. nov. exhibits 0.3% intraspecific variation, while it is 16.8% ���21.4% divergent from all other congeners (Table 9). Description of holotype. The holotype is in good condition except, head is slightly tilted towards the right���an artefact of preservation (Fig 11A). Body is dorsoventrally flattened with the posterior 3/4 th of tail regenerated.Adult male, SVL 50.9 mm. Head short (HL/SVL 0.26), slightly elongate (HW/HL 0.68), not depressed (HH/HW 0.61), distinct from neck. Loreal region slightly inflated, canthus rostralis indistinct (Fig 11C). Snout short (SE/HL 0.40), longer than orbital diameter (OD/SE 0.53); scales on snout, canthus rostralis, inter-orbital region, forehead, occipital and nuchal regions granular and rounded with those on the snout and canthus rostralis being larger (Fig 11B). Eye small (OD/HL 0.21); pupil vertical with crenulated margins; supraciliaries small, rounded, directed outwards and uniform in size. Ear opening elliptical (longer diameter 0.7 mm); eye to ear distance longer than diameter of eye (EE/OD 1.46). Rostral wider than deep (RL/RW 0.37), with a distinct rostral groove extending halfway through the scale medially; two large internasals, separated by a smaller, subequal scale, all in broad contact with rostral; two postnasals on either side, slightly smaller than the internasals, the lower in contact with supralabial I; rostral in contact with nasal, supralabial I, internasals and the smaller scale separating the internasals; nostrils about the size of the lower postnasal, roughly circular with nasal pad visible posteriorly; nasal surrounded by internasal, rostral, two postnasals and a small scale separating it from supralabial I on either side; 2���4 rows of scales separate orbit from supralabials at mid-orbital position. Supralabials roughly rectangular, increasing in length anteriorly. Supralabials (to midorbital position) 8 (right), 8 (left); supralabials (to angle of jaw) 10 (right), 10 (left); infralabials (to angle of jaw) 8 (right), 8 (left). Mental triangular; two pairs of well-developed postmentals, the inner pair slightly shorter (1.0 mm) than the mental (1.1 mm), and in strong contact with each other (0.5 mm) behind mental; outer pair distinctly shorter (0.6 mm) than the inner pair, separated from each other by five gular scales that are smaller than postmentals (Fig 11D). Inner postmentals bordered by mental, infralabial I & II (barely touching on both sides), outer postmentals and five smaller gular scales; outer postmentals bordered by infralabials I (barely touching on the right) and II, inner postmentals, and smaller gular scales each of dissimilar sizes, four on the right and two on the left sides. Body relatively slender, elongate (TRL/SVL 0.47). Dorsal pholidosis composed of small, rounded granules that are juxtaposed in arrangement, becoming slightly larger, flatter, weakly pointed and sub-imbricate laterally; Ventral scales larger than dorsals, largely homogeneous in shape and size, smooth, flat, sub-imbricate; gular region with smaller, granular, juxtaposed scales; anterior gular scales visibly larger, flatter; scales on femoral region larger than those on chest; precloacal scales larger than scales on femoral region; midbody scale rows across belly 26���28. Non-lamellar scales in the palmar and plantar regions heterogeneous in size, rounded, juxtaposed on palm and sole; scales on dorsal aspect of upper arm larger than granules on dorsum, flat, pointed, sub-imbricate and smooth; dorsal aspect of forearm with smaller, sub-imbricate scales intermixed with a few rounded granules around the elbow; scales on dorsal aspect of hand and digits larger than those on forearm, flat, weakly pointed and imbricate; scales on anterior aspect of thigh large, flat, imbricate and weakly pointed; rest of the dorsal scales on hindlimb smaller, granular and rounded. Scales on dorsal aspect of foot larger than those on shank, flat, weakly pointed and imbricate. Forearm (FL/SVL 0.10) and tibia short (CL/SVL 0.13); digits moderately short with relatively long terminal phalanges, strongly clawed; all digits of manus and digits I���IV of pes indistinctly webbed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, more than half as long as associated toepad; scansors beneath each toe undivided throughout, in a straight transverse series: 6-7-8-7-7 (left manus), 6- 7-7-8-7 (right manus), 6-8-9-9-7 (left pes), 6-8-9-10 -7 (right pes). Relative length of digits (measurements in mm in parentheses): IV (4.0)> III (3.8)> II (3.5)> V (3.0)> I (2.6, claw broken) (left manus); IV (5.2)> III (4.7)> V (4.6) = II (4.6)> I (3.3) (left pes). Tail partially regenerated, rounded at the base, flat beneath, tapering posteriorly, covered above uniformly with round, smooth, flat, sub-imbricate scales that become slightly larger laterally; subcaudal scales larger, with an undivided median series of enlarged scales. An uninterrupted series of 38 precloacofemoral pores that are only faintly visible towards the knee (Fig 13G). Variation in paratypes. Inner postmentals in contact with only infralabial I on both sides in all other paratypes. Inner postmentals bordered posteriorly by three gular scales in BNHS 2352, BNHS 2353, ZSIK 2982, ZSIK 2983 and ZSIK 2985, and by five gulars in BNHS 2354. Right inner postmental bordered by a small gular scale laterally in BNHS 2355 and ZSIK 2984. Outer postmentals bordered by 3 gulars in BNHS 2352 (R), 5 in BNHS 2354 (R) and ZSIK 2984 (R) and 6 in BNHS 2355 (R) and ZSIK 2983 (R). Outer postmentals not in contact with infralabials BNHS 2355 (R), ZSIK 2983 (R) and ZSIK 2984 (R). Outer postmentals in contact with both infralabial I and II on both sides in BNHS 2352, BNHS 2353, BNHS 2354, ZSIK 2982, ZSIK 2985 and ZSIK 2986. Other morphological variations are listed in Table 7. Colour in preservative. Dorsum predominantly light brown mottled with darker, discontinuous streaks from the snout to the base of tail (Fig 11A). Similar mottling faintly visible on dorsal aspect of limbs. Neck with a roughly circular, dark blotch flanked by 2 longitudinal streaks on either side. Posterior part of head demarcated by a disctinct saddle-shaped horizontal streak (Fig 11B). Inter-orbital region slightly darker than rest of the body with scattered dark-brown granules. Labials paler than rest of the head with faint, darker spots bordering each labial. Supralabials bordered by a dark, roughly triangular streak from nostril to eye. Limbs no different from rest of the dorsum. Tail predominantly grey with darker, longitudinal markings in the regenerated portion. Ventral region uniformly cream coloured. Ventral surface of tail pale, with scattered mid-brown speckling throughout. Colouration (in life) (based on photographs of an uncollected topotype). Dorsal markings more evident in life (Fig 15B). Dorsum pale-brown with darker streaks throughout. Head dorsum pale-brown, snout darker, with a dark streak emanating from snout to eye. Yellow dots on each labial with a scattering of these in the loreal region. Forehead ground colour, interspersed by darker spots. A dark, discontinuous streak emanates from eye up to the forelimb insertion. A dark saddle shaped collar in the occipital region. Six dark streaks along the vertebral region after the collar, followed posteriorly by two saddle shaped markings in the sacral region. Limbs of ground colour with dark spots sprinkled all over. Tail distinctly banded with alternating light and dark portions. Bands more conspicuous after the first three segments. Iris marbled, golden, suffused with prominent dark-brown venation; pupil black with crenulated margins. Etymology. The specific epithet is a compound noun formed by the combination of two Tamil words from the Sangam era (3 rd century BC���3 rd century AD) that alludes to the ancient divergence and colonization of these geckos in peninsular India. The stem word, ��� thol ��� (pronounced /����l/) is an archaic Tamil word for ���ancient��� and ��� palli ��� (pronounced /p��ll��/) an ancient word still in common parlance, is the Tamil for ���gecko���. Suggested Common name. Kodaikanal Dravidogecko. Distribution. Dravidogecko tholpalli sp. nov. is presently restricted in distribution to Kodaikanal town and its outskirts in the Palani Hills of the southern Western Ghats. They are found in large numbers around the Kodaikanal Lake in the centre of the town, which is surrounded by disturbed evergreen forests. The habitat in the Palani Hills chiefly constitutes moist deciduous and southern tropical wet evergreen forests (B. Balaguru et al. 2016). These habitats are at an altitude of 1600���2000 m asl and receive an average annual rainfall of 1500 mm (Bhupathy et al. 2009). Other areas in the Palani Hills such as Perumalmalai and Vattakanal are likely to harbour populations of D. tholpalli sp. nov. Habitat and natural history. The type-series of Dravidogecko tholpalli sp. nov. was collected in Kodaikanal town from abandoned buildings and stone walls near forested areas. Kodaikanal falls under a special case of the Madurai-Pollachi rainfall regime with 0���4 dry months and slightly more (~92) rainy days annually (Pascal 1982). Other lizards found in sympatry were Cnemaspis sp., Kaestlea cf. palnica and Salea anamallayana., Published as part of Chaitanya, R., Giri, Varad B., Deepak, V., Datta-Roy, Aniruddha & Karanth, Praveen, 2019, Diversification in the mountains: a generic reappraisal of the Western Ghats endemic gecko genus Dravidogecko Smith, 1933 (Squamata: Gekkonidae) with descriptions of six new species, pp. 1-56 in Zootaxa 4688 (1) on pages 30-34, DOI: 10.11646/zootaxa.4688.1.1, http://zenodo.org/record/3514770, {"references":["Boulenger, G. A. (1885) Catalogue of Lizards in the British Museum. Vol. 1. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae, Second Edition. Trustees of the British Museum (Natural History), London, 436 pp., XXXII pls.","Gunther, A. (1875) Second report on collections of Indian reptiles obtained by the British Museum. Proceedings of the Zoological Society of London, 43 (1), 224 - 234.","Boettger, O. (1893) Katalog der Reptilien-Sammlung im Museum der Senckenbergishen naturforschenden Gesellschaft in Frankfurt am Main. I Teil, (Rhynchocephalen, Schildkroten, Krokodile, Eidechsen, Chamaleons). Gebruder Knauer, Frankfurt, 140 pp.","Bauer, A. M. & Russell, A. P. (1995) The systematic relationships of Dravidogecko anamallensis (Gunther 1875). Asiatic Herpetological Research, 6, 30 - 35. https: // doi. org / 10.5962 / bhl. part. 7983","Ganesh, S. R. (2010) Catalogue of Indian herpetological specimens in the collection of the Government Museum Chennai, India. Hamadryad, 35, 46 - 63.","Balaguru, B., Soosairaj, S., Nagamurugan, N., Ravindran, R. & Khaleel, A. A. (2016) Native vegetation pattern and the spread of three invasive species in Palani Hill National Park, Western Ghats of India. Acta Ecologica Sinica, 36 (5), 367 - 376. https: // doi. org / 10.1016 / j. chnaes. 2016.05.005","Bhupathy, S., Srinivas, G. & Sathishkumar, N. (2009) A study on herpetofaunal communities of the Upper Vaigai Plateau, Western Ghats, India. Final Technical Report submitted to Ministry of Environment and Forests, Government of India. Salim Ali Centre for Ornithology and Natural History, Coimbatore, 75 pp.","Pascal, J. P. (1982) Explanatory notes on the Bioclimate maps of the Western Ghats. Institut francais de Pondichery, Pondicherry, 4 pp."]}

Chaitanya, R., Giri, Varad B., Deepak, V., Datta-Roy, Aniruddha, Karanth, Praveen (2019): Diversification in the mountains: a generic reappraisal of the Western Ghats endemic gecko genus Dravidogecko Smith, 1933 (Squamata: Gekkonidae) with descriptions of six new species. Zootaxa 4688 (1): 1-56, DOI: https://doi.org/10.11646/zootaxa.4688.1.1

Dravidogecko douglasadamsi sp. nov. (Figs 9 A–D, 13B, 14C; Table 6) Hoplodactylus anamallensis: Boulenger, 1885 Hoplodactylus anamallensis [non Gecko anamallensis Günther, 1875]— Boulenger, 1885 [partim]; Boulenger, 1890 [partim]. Dravidogecko anamallensis: Smith, 1933 Dravidogecko anamallensis [non Gecko anamallensis Günther, 1875]— Smith, 1935 [partim]; Murthy, 1993; Sharma, 2002 [partim]. Hemidactylus anamallensis: Bauer & Russell, 1995 Hemidactylus anamallensis [non Gecko anamallensis Günther, 1875]— Johnsingh, 2001; Srinivasulu, Srinivasulu & Molur, 2014 [partim]; etc. Holotype. BNHS 2349, an adult male, Manjolai (8.5514 °N, 77.3597 °E; ca. 1300 m asl.), Tirunelveli District, Tamil Nadu, collected by R. Venkitesan on 10 th June, 2017. Referred specimens (Topotypes). BMNH 82.5.22.79, Adult female, BMNH 82.5.22.81, juvenile male, BMNH 82.5.22.80 & BMNH 82.5.22.82, Adult male—collected by Colonel Beddome from “Tinnevely” (now Tirunelveli) and deposited in the NHMUK. Type locality. Manjolai, Tirunelveli District, Tamil Nadu. Summarized description and diagnosis. Snout-vent length up to 48.5 mm (n=5); two pairs of well-developed postmentals, inner pair of comparable length to the outer postmentals and mental, bordered posteriorly by 2 or 3 gular scales; ventral scales counted at midbody, 31 or 32; precloacofemoral pores, 42 or 43; subdigital lamellae under digit IV of manus, 9 or 10 and under digit IV of pes, 10–12; supralabials 10–12 and infralabials, 8–10 on each side. Dravidogecko douglasadamsi sp. nov. can be distinguished from other congeners based on the following characters: number of precloacofemoral pores (PcFP 40–43 versus 45 or 46 in D. anamallensis, 52–56 in D. septentrionalis sp. nov. & 36–38 in D. meghamalaiensis sp. nov.); postmentals of comparable length with mental (ML /1PML 0.98–1.05 versus much longer, 0.74–0.81 in D. anamallensis). Genetic divergence (p-distance). Dravidogecko douglasadamsi sp. nov. is 11.0% –16.5% divergent from other previously described congeners. Description of holotype. The holotype is in good condition (Fig 9A), except for an incision of about 2.1 mm at mid-trunk region, made to extract liver tissue. Posterior portion of tail curved in a sinusoidal manner, fifth finger on left forelimb curved upwards—both artefacts of preservation. Adult male, SVL 48.5 mm. Head short (HL/ SVL 0.27), slightly elongate (HW /HL 0.64), slightly depressed (HH/ HW 0.57), distinct from neck. Loreal region slightly inflated, canthus rostralis indistinct (Fig 9C). Snout short (SE /HL 0.36), longer than orbital diameter (OD / SE 0.66); scales on snout, canthus rostralis, inter-orbital region, forehead, occipital and nuchal regions granular and rounded with those on the snout and canthus rostralis being larger (Fig 9B). Eye small (OD /HL 0.24); pupil vertical with crenulated margins; supraciliaries small, rounded, directed outwards, increasing in size anteriorly. Ear opening elliptical (longer diameter 0.8 mm); eye to ear distance longer than diameter of eye (EE / OD 1.19). Rostral wider than deep (RL / RW 0.32), rostral groove absent; two large, roughly circular internasals, separated by two smaller, subequal scales, all in broad contact with rostral; two postnasals on either side, slightly smaller than the internasals, the lower in contact with supralabial I; rostral in contact with nasal, supralabial I, internasals and the two smaller scales separating the internasals; nostrils about the size of the lower postnasal, roughly circular with nasal pad visible posteriorly; nasal surrounded by internasal, rostral, two postnasals and supralabial I on either side; 2–4 rows of scales separate orbit from supralabials around mid-orbital position. Supralabials roughly rectangular, increasing in length anteriorly. Supralabials (to midorbital position) 9 (right), 9 (left); supralabials (to angle of jaw) 12 (right), 12 (left); infralabials (to angle of jaw) 10 (right), 10 (left). Mental triangular; two pairs of postmentals, smaller but roughly the same length as the mental; the inner pair slightly shorter (1.1 mm) than the mental (1.2 mm), and in strong contact with each other (0.7 mm) behind mental; outer pair shorter still (1.0 mm), separated from each other by two gular scales that are smaller than postmentals (Fig 9D). Inner postmentals bordered by mental, infralabial I, infralabials II (barely touching on right), outer postmentals and the two smaller gular scales that separate the outer postmentals; outer postmentals bordered by infralabials I (barely touching only on the left) and II, inner postmentals, and smaller gular scales each of dissimilar sizes, three on the right and four on the left sides. Body dorsoventrally flattened, relatively slender, elongate (TRL / SVL 0.46). Dorsal pholidosis homogenous, composed of small, rounded granules throughout, becoming slightly larger at the lateral aspects; Ventral scales larger than dorsals, largely homogeneous in shape and size, smooth, flat, weakly pointed and sub-imbricate; gular region with smaller, flat, rounded, juxtaposed scales; anteriormost gular scales visibly larger, flatter; scales on sacral and femoral regions larger than those on chest; precloacal scales largest; midbody scale rows across belly 31 or 32; Non-lamellar scales in the palmar and plantar regions heterogeneous in size, flat, rounded, juxtaposed on palm and sub-imbricate on sole; scales on dorsal aspect of upper arm much larger than granules on dorsum, flat, weakly pointed, sub-imbricate and smooth; dorsal aspect of forearm with smaller, sub-imbricate scales intermixed with a few rounded granules around the elbow; scales on dorsal aspect of hand and digits larger than those on forearm, flat, weakly pointed and imbricate; scales on anterior aspect of thigh large, flat, sub-imbricate and weakly pointed; rest of the dorsal scales on hindlimb smaller, granular and rounded; scales on dorsal aspect of feet and toes larger than those on shank, flat, weakly pointed and imbricate. Forearm (FL / SVL 0.11) and tibia short (CL/ SVL 0.12); digits moderately short with relatively long terminal phalanges, strongly clawed; all digits of manus and digits I–IV of pes indistinctly webbed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, more than half as long as associated toepad; scansors beneath each toe undivided throughout, in a straight transverse series: 6-8-9-9-7 (left manus), 6-8- 9-9-7 (right manus), 7-9-10-12 -9 (left pes), 7-9-10-12 -9 (right pes). Relative length of digits (measurements in mm in parentheses): IV (3.8)> III (3.6)> II (3.2)> V (2.6)> I (2.5) (left manus); IV (4.6)> III (4.5)> II (4.3)> V (3.8)> I (3.3) (left pes). Tail entire, rounded at the base, flat beneath, tapering posteriorly, covered above uniformly with round, smooth, flat, sub-imbricate scales that become slightly larger laterally; pygal region containing the hemipenal bulge with six or seven rows of flat, weakly pointed, sub-imbricate scales; subsequent subcaudal scales larger, with an undivided median series of enlarged scales extending to tail tip. An uninterrupted series of 43 precloacofemoral pores that are indistinct towards the knee (Fig 13B). Variation in referred specimens. Internasals separated by one smaller scale in BMNH 82.5.22.83. Inner postmentals bordered posteriorly by three gular scales in BMNH 82.5.22.79 and BMNH 82.5.22.80. Outer postmentals bordered by 4 gulars on right and 5 on left in BMNH 82.5.22.79, BMNH 82.5.22.80, BMNH 82.5.22.83. Outer postmentals not in contact with infralabials in BMNH 82.5.22.80 (L), and in contact with both infralabials I and II in BMNH 82.5.22.79 and BMNH 82.5.22.83. Other morphological variations are listed in Table 6. Colouration in preservative. Dorsum uniformly brown, mottled with darker, discontinuous streaks from the snout to the base of tail (Fig 9A). Similar mottling faintly visible on dorsal aspect of limbs. Neck with a dark, discontinuous longitudinal streak, flanked at the break by two dark lines at a forty-five degree angle. Two discontinuous lines emanate from the eye, following the contour of the cranium posteriorly and extending beyond the forearm insertion. Inter-orbital region with a scattering of dark spots, witha distinct dark blotch bordering the supraciliary region on either side. Labials paler than the rest of the head with a faint, pattern-less scattering of darker spots bordering each labial. A dark, roughly rectangular streak emanates from eye up to the region above the third supralabial on the right side and the nostril on the left. Limbs no different from rest of the dorsum. Tail of similar ground colour to dorsum with alternating pale-dark longitudinal bands, the first of which is roughly saddle-shaped. Ventral region cream coloured with a scattering of two or three dark spots on each ventral scale. Ventral surface of tail pale, with scattered mid-brown speckling in the hemipenal region followed by alternating pale-dark bands in the distal half. Colouration (in life) (based on photographs of an uncollected topotype). Dorsum mid-brown with faint, darker streaks throughout (Fig 14C). Head dorsum ground colour, snout slightly darker with a mottling of yellow scales throughout. A dark streak emanates from above the first supraocular and extends to the eye. Forehead with a scattering of spots that are either paler or darker. A longitudinal streak from the occiput extending into forehead, is flanked by a roughly inverted ‘V’ shaped marking posteriorly. Two dark spots follow, at and just beyond the forelimb insertion. Six irregular, roughly transverse markings follow, until the sacral region. Trunk with four or five rows of transversely arranged pale spots. Limbs of ground colour with irregular dark spots. Digits interspersed with yellow spots. Tail distinctly banded with alternating light and dark portions, more pronounced posteriorly. Etymology. The specific epithet is a patronym honouring the English author and satirist, Douglas Noel Adams. Adams was also a renowned environmental activist. His radio documentary on critically endangered animals for the British Broadcasting Corporation (BBC) titled “Last Chance to See” and its accompanying book influenced the thinking of a whole generation of wildlife biologists. The etymology also alludes to the number ‘ 42 ’—the number of precloacofemoral pores that most specimens of this species possess. The number 42 incidentally is also the answer to the “ ultimate question of Life, The Universe and Everything ” according to Adams’ seminal book “The Hitchhikers Guide to the Galaxy”. Suggested Common name. Adams’ Dravidogecko. Distribution. Dravidogecko douglasadamsi sp. nov. is presently restricted in distribution to Manjolai and its environs in Tirunelveli district, south of the Shencottah gap in the southern Western Ghats. Similar habitats are seen in various parts of Kalakkad Mundanthurai Tiger Reserve, around which populations of this species might be found. Habitat and natural history. The habitat in Manjolai and the adjoining Kalakkad- Mundanthurai forests where Dravidogecko is found, is chiefly comprised of southern- tropical semi-evergreen (700 m asl) and southern tropical wet evergreen forests (800–1500 m asl). These habitats receive an average annual rainfall of ca. 1600 mm (Ayyanar & Ignacimuthu 2008). This species was seen occupying walls of a tea estate building during the night. There were no other geckos in sympatry, though a species of Eutropis was seen in the habitat during the daytime.

Dravidogecko meghamalaiensis sp. nov. (Figs 8 A–D, 13D, 14B; Table 6) Hemidactylus anamallensis: Bauer & Russell, 1995 Hemidactylus anamallensis [non Gecko anamallensis Günther, 1875]— Chandramouli & Ganesh, 2010; Holotype. BNHS 2345, an adult male, Meghamalai (9.6925 °N, 77.3992 °E; ca. 1480 m asl.), Theni District, Tamil Nadu, collected by RC on 30 th May, 2016. Paratypes. Details of collection same as the holotype. BNHS 2346, BNHS 2347, BNHS 2348, BNHS 2349, ZSIK 2977, ZSIK 2979 – adult females; ZSIK 2978 and ZSIK 2980 adult males. Type locality. Approximately 8 km southwest of Meghamalai village, en route to the Highwavy Mountains in Theni District, Tamil Nadu. Summarized description and diagnosis. Snout-vent length up to 48.7 mm (n=9); two pairs of well-developed postmentals, inner pair only slightly longer than the outer (2PML/1PML 0.82–0.96), and of comparable length to the mental; ventral scales counted at midbody 28–34; precloacofemoral pores, 36–38 (n=3); subdigital lamellae under digit IV of manus 7–9 and under digit IV of pes, 9 or 10; supralabials, 9–11 and infralabials 8–10 on each side. Dravidogecko meghamalaiensis sp. nov. can be distinguished from other congeners based on the following characters: number of precloacofemoral pores (PcFP 36–38 versus 45 or 46 in D. anamallensis & 52–56 in D. septentrionalis sp. nov.); inner postmentals comparable in length to mental (ML /1PML 0.95–1.23 versus much longer, 0.74–0.81 in D. annamallensis); fewer subdigital lamellae under digit IV of pes (9 or 10 versus 11 or 12 in D. annamallensis). Genetic divergence (p-distance). Dravidogecko meghamalaiensis sp. nov. exhibits 0.4% intraspecific variation while it is 13.1% –13.8% divergent from D. anamallensis and 13.0%–13.7% divergent from D. septentrionalis sp. nov. (Table 9). Description of holotype. The holotype is in good condition (Fig 8A). The head is slightly tilted towards the right, tail curved towards left and two distinct folds of skin just beneath the forearm insertion—all artefacts of preservation. Body is dorsoventrally flattened with the distal half of tail regenerated. Adult male, SVL 45.1 mm. Head short (HL/SVL 0.28), slightly elongate (HW/HL 0.67), not depressed (HH/HW 0.56), distinct from neck. Loreal region slightly inflated, canthus rostralis indistinct (Fig 8C). Snout short (SE/HL 0.39), longer than orbital diameter (OD/SE 0.57); scales on snout, canthus rostralis, inter-orbital region, forehead, occipital and nuchal regions granular and rounded with those on the snout and canthus rostralis being larger (Fig 8B). Eye small (OD/HL 0.22); pupil vertical with crenulated margins; supraciliaries small, roughly triangular, pointed upwards and gradually increasing in size anteriorly. Ear opening elliptical (longer diameter 0.8 mm); eye to ear distance longer than diameter of eye (EE/OD 1.37). Rostral wider than deep (RL/RW 0.30), rostral groove distinct but extending only marginally downwards from the suturing with internasals, medially; two large internasals, separated by two smaller, subequal scales, all in broad contact with rostral; two postnasals on either side, slightly smaller than the internasals, the lower in contact with supralabial I; rostral in contact with nasal, supralabial I, internasals and the two smaller scales separating the internasals; nostrils about the size of the lower postnasal, roughly circular with nasal pad visible posteriorly; nasal surrounded by internasal, rostral, two postnasals and a small scale separating it from supralabial I on either side; 2–4 rows of scales separate orbit from supralabials at mid-orbital position. Supralabials roughly rectangular, increasing in length anteriorly. Supralabials (to midorbital position) 6 (right), 7 (left); supralabials (to angle of jaw) 9 (right), 9 (left); infralabials (to angle of jaw) 8 (right), 8 (left). Mental triangular; two pairs of welldeveloped postmentals, the inner pair slightly shorter (0.9 mm) than the mental (1.1 mm), and in strong contact with each other (0.5 mm) behind mental; outer pair similar in size to inner pair, separated from each other by two gular scales that are only smaller than postmentals (Fig 8D). Inner postmentals bordered by mental, infralabial I, outer postmentals and two smaller gular scales; outer postmentals bordered by infralabials I (only on the left) and II, inner postmentals, and five smaller gular scales each of dissimilar sizes on either side. Body relatively slender, elongate (TRL/SVL0.49). Dorsal pholidosis homogenous, composed of small, rounded granules, becoming slightly larger, flatter, weakly pointed and sub-imbricate laterally; Ventral scales larger than dorsals, largely homogeneous in shape and size, smooth, flat, sub-imbricate; gular region with smaller, granular, juxtaposed scales; anterior gular scales visibly larger, flatter; scales on femoral region larger than those on sacrum and chest with some precloacal scales being largest; midbody scale rows across belly 28–29. Non-lamellar scales in the palmar and plantar regions heterogeneous in size, rounded and juxtaposed on palm and sole; scales on dorsal aspect of upper arm larger than granules on dorsum, flat, pointed, sub-imbricate and smooth; dorsal aspect of forearm with smaller, sub-imbricate scales intermixed with a few rounded granules around the elbow; scales on dorsal aspect of hand and digits larger than those on forearm, flat, weakly pointed and imbricate; scales on anterior aspect of thigh large, flat, imbricate and weakly pointed; rest of the dorsal scales on hindlimb smaller, granular and rounded. Scales on dorsal aspect of foot larger than those on shank, flat, weakly pointed and imbricate. Forearm (FL/SVL 0.11) and tibia short (CL/SVL 0.14); digits moderately short with relatively long terminal phalanges, strongly clawed; all digits of manus and digits I–IV of pes indistinctly webbed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, more than half as long as associated toepad; scansors beneath each toe undivided throughout, in a straight transverse series: 6-6-7-7-7 (left manus), 6- 7-7-7-6 (right manus), 6-7-8-9-7 (left pes), 6-7-8-9-7 (right pes). Relative length of digits (measurements in mm in parentheses): IV (4.1)> III (3.9)> II (3.6)> V (3.5)> I (2.9, claw broken) (left manus); IV (5.0)> III (4.5)> V (4.3)> II (4.1)> I (3.3) (left pes). Tail long (TL/SVL 1.06), rounded at the base, flat beneath, tapering posteriorly, covered above uniformly with round, smooth, flat, sub-imbricate scales that become slightly larger laterally; subcaudal scales larger, with an undivided median series of enlarged scales that continue until the regenerated portion. An uninterrupted series of 36 precloacofemoral pores that are only faintly visible towards the knee (Fig 13D). Variation in paratypes. Rostral groove extends halfway through the scale in BNHS 2346, BNHS 2347, BNHS 2348, ZSIK 2978, ZSIK 2979. Internasals separated by one smaller scale in 2346, BNHS 2347, ZSIK 2976 and ZSIK 2979. Inner postmentals in contact with infralabials I and II in BNHS 2347 (L), BNHS 2348 (L), ZSIK 2978 (R) and ZSIK 2980 (R). Inner postmentals bordered posteriorly by three gular scales in BNHS 2346, BNHS 2348, ZSIK 2978 and ZSIK 2979 and four in ZSIK 2976 and ZSIK 2977. Outer postmentals bordered by 6 gulars in BNHS 2346 (L) and ZSIK 2979 (R) and 4 in ZSIK 2976 (L), ZSIK 2977 (L) and ZSIK 2978 (L,R). Outer postmentals in contact only with infralabial II in ZSIK 2978 (R) and ZSIK 2980 (R) and only with infralabial I in BNHS 2347 (L) and BNHS 2348. Other morphological variations are listed in Table 6. Colour in preservative. Dorsum predominantly light brown, mottled with darker, discontinuous streaks from the snout to the base of tail (Fig 8A). Similar mottling visible on dorsal aspect of limbs. Neck with a roughly circular, dark blotch flanked by 2 longitudinal streaks on either side. Posterior part of head demarcated by a disctinct horizontal streak. Inter-orbital region slightly darker than rest of the body with scattered vague dark-brown blotches. Labials appear paler than rest of the head with faint spots that are darker. Supralabials bordered by a dark, roughly triangular streak from nostril to eye. Limbs no different from rest of the dorsum. Tail predominantly grey with darker, faint, saddle shaped markings. Venter predominantly cream coloured. Ventral surface of tail pale, with scattered midbrown speckling throughout until the regenerated portion, which is predominantly mid-grey. Colouration (in life). Dorsum pale with dark-brown streaks throughout that are bordered by one or two rows of yellowish scales (Fig 14B). Head dorsum ground colour, posterior part of snout predominantly with scattered yellow scales. Irregularly arranged dark spots in the inter-orbital region and forehead. A dark streak emanates from loreal region up to the eye and continues posteriorly into the lateral aspect of the neck. A discontinuous, roughly W shaped collar followed by a dark spot in the occipital region. Six dark, transverse streaks across the vertebral region until the sacrum. Limbs of ground colour with irregular dark streaks. Tail lighter than dorsum, with seven irregular, dark streaks. Tip of tail regenerated. Etymology. The specific epithet is an adjectival toponym referring to the Meghamalai Hills, where the type series was collected. Suggested Common name. Meghamalai Dravidogecko. Distribution. Dravidogecko meghamalaiensis sp. nov. is presently restricted in distribution to the Meghamalai Hills in the southern Western Ghats. Similar habitats are seen in the Vellimalai Range within the Meghamalai Wildlife Sanctuary and in many parts of the Srivilliputtur Grizzled Squirrel Wildlife Sanctuary, where this species could also possibly occur. Habitat and natural history. The type-series of Dravidogecko meghamalaiensis sp. nov. was collected en route to the Highwavy Mountains within the Meghamalai Wildlife Sanctuary, where the habitat chiefly constitutes moist mixed deciduous forests (Bhupathy & Babu 2013). Individuals were found on trees and abundantly in unoccupied buildings. Sub-adults (SVL et al. 2009).

Smithophis atemporalis gen. et. sp. nov. (Figs. 1–7; Table 2) Rhabdops bicolor (Blyth, 1854): Das (2015: photograph on p. 65), Laltanpuia et al. (2008: 119, in part; fig. 16). Diagnosis. A Smithophis lacking temporal shields. Holotype. BNHS 3523 (Figs. 2, 3), male, collected from Mizoram University Campus, Aizawl, India (23.76338°N, 93.09916°E, 833 m elevation) by H.T. Lalremsanga on 10 July 2014. See map in Fig. 4. Paratypes (n = 6). BNHS 3524 — BNHS 3527, BNHS 3529, males and BNHS 3528, female, collected from Mizoram University Campus, Aizawl on 14 August 2014 by H.T. Lalremsanga. Referred specimens. BNHS 3530 male, collected from Model Veng, Aizawl, India (23.71083°N, 92.93194°E, 1014 m elevation) on 18 September 2015 and BNHS 3531 male, collected from Luangmual, Aizawl, India (24.45750°N, 92.69953°E, 1025 m elevation) on 20 October 2014 by S. Lalrounga. These are referred rather than paratypic material because we have not generated detailed morphometric and meristic data. Description of holotype. See Table 2 for morphometric and meristic data. Good condition, slightly dehydrated; ca. 20 mm longitudinal ventral incision into ventral surface of base of tail (through which hemipenis observed). Body somewhat laterally compressed (1.5 times as high as wide at midbody) with flattened venter and arched dorsum, widest at midbody, tapering posteriorly and more substantially anteriorly. Head elongate, broader than tall, slightly wider than anterior of body. In dorsal view head slightly ovate, sides very slightly convex, gently converging anteriorly. Front of snout not pointed, truncated. In lateral view head tapers very gently from back to prefrontal, more strongly tapered in front of prefrontal. Paired shields on top of head (= parietals only) abutting along midline rather than imbricate/overlapping. In dorsal view rostral approximately three times broader than long, substantially shorter than distance between it and frontal; projects beyond tip of lower jaw; ventrally with transverse concavity, notched (C-shaped) at margin of mouth. Frontal subpentagonal, lateral edges anteriorly diverging; shorter and smaller than each parietal, shorter than distance between it and snout tip. Internasal single, smaller than single prefrontal. Nasals subquadrangular, squarish, notably larger than squarish loreal. Single supraocular, preocular and postocular each side; supraocular slightly larger than preocular, itself slightly larger than postocular. Pre- and postocular each kidney shaped; supraocular elongate, longer than wide. External naris on dorsolateral-facing surface, seen as C-shaped slit in front of flap, in broad, shallow depression slightly posterodorsal of centre of nasal. External naris visible anteriorly as well as dorsally and (mostly) laterally. Five supralabials (SLs) on each side; SL 5 largest, longest; SLs 1 and 4 subequal, smallest; SLs 2 and 3 subequal. SL 1 contacts rostral and nasal, SL 2 contacts loreal, SL3 contacts preocular and eye, SL 4 contacts postocular and parietal, SL 5 contacts parietal). Eye contacts third supralabial only. Eye lateral; pupil subcircular, very slightly elliptical (long axis vertical). No temporal shields. Parietals longer than wide, larger than other head scales. Midline interparietal suture a little more than two thirds length of each parietal; parietals barely projecting posterior to suture; each parietal more than twice as long as frontal; each parietal contacts frontal, supraocular, postocular, and fourth and fifth supralabial, plus five other scales. Posterior margin of each parietal rounded, somewhat scalloped. Mental small, subtriangular, wider than long. Infralabials 7,7; first pair in midline contact; second pair smallest; fourth largest. On right, fourth infralabial longest, on left seventh as long. Two pairs of genials; first pair largest, longer than broad, in long midline contact; second pair not in contact, separated by one small, midline scale. Anterior genials contact infralabials (ILs) 1–4; posterior genials contact IL 4 and 5 (left) or 4 only (right). Anteriormost ventral separated from each posterior genial by four scales, separated from each posteriormost infralabial by six scales. Teeth largely obscured by gingivae, but estimated as 10 or 11 marginals on each side of upper jaw. Macroscopically and under low magnification (using a light dissecting microscope) body scales smooth. No keels or apical pits. Vertebral scale row not different from adjacent dorsal scale row. Exposed parts of dorsal body scales generally evenly sized on dorsum and along body except for those involved in dorsal scale row reductions and for lowest dorsal scales level with first few ventrals which are lower, more elongate. Dorsal scales closer to vent slightly smaller than at midbody; those at anterior of body smallest. Dorsal scale rows 17 at level of first ventral, maintained to vent except for loss of third dorsal scale row on left, six ventrals anterior to vent. First ventral divided/paired. Posteriormost ventral with slight posteromedial extension. Anals paired (right overlapping left), each slightly larger than posteriormost ventral. Each anal overlaps 6, 7 small scales in addition to first subcaudal. Tail subtriangular in cross section, flattened ventrally. Dorsal tail scales more heterogenous in size than on body, without clear patterns. Subcaudals paired/divided throughout, terminal scale (scute) conical, approximately as wide as long, pointed. Macroscopically bicoloured, darker black-grey above, whitish below. Body scales iridescent. Dorsum dark. Alternating (at least asymmetric) longer dark and shorter pale markings along body and tail. On body 36, 35 dark ‘bands’, on tail 20, 16. On last three quarters of body, ventrally projecting dark trapezoids extend onto lateral edges of ventrals (leaving broad, entirely pale band along most of venter), these dark trapezoids anteriorly steadily less ventrally extensive. Anteriormost dark trapezoid (two head lengths behind head) extends onto second dorsal scale row. Whitish inverted-Vs extend dorsally between each dark trapezoid up to the fourth dorsal scale row, though partly continues as mottled pale grey scales up to vertebral scale or even one scale row onto opposite side and often (mostly on posterior half of body) connecting with grey dorsal part of inverted pale Vs from opposite side. Along the first quarter of body the border between pale-dark markings wavy, rapidly becoming increasingly sharply zigzag. Ventrally projecting dark marks extend as far as midventer on tail from behind vent, forming alternating (dark-pale) transverse bands; where pale bands approach midventer (only on anterior half of tail, where they are longer) they remain separated by narrow midventral dark line. Whitish ventrals with fine pale grey speckles on posterior half of body. Anals each with pale grey blotches as well as speckles. Head dark dorsally, brown-grey, slightly paler than body dorsum, generally uniform, rostral slightly paler. Dark of dorsum extends onto upper margins of SLs, SLs otherwise pale (whitish, as body venter) with very few (more on SL1) pale grey specks. Dark scales on dorsum (including head) are mottled under low power dissecting microscope. First pair of ILs and anterior genials with pale grey speckles (fewer on mental). Underside of head otherwise pale, whitish. Inside of mouth pale. Variation among paratypes. The heads of the paratypes are illustrated in Fig. 5. See Table 2 for variation in meristic and morphometric features. Paratypes generally in moderate to good condition, with following exceptions. BNHS 3524 dehydrated, contorted, yellowed ventrally with dorsolateral longitudinal incision on posterior half; BNHS 3525 soft posterior half, with 10 mm tail incision and missing terminal scute of tail; BNHS 3526 with couple of breaks along body, distorted mouth, marginal and palatal tooth-bearing bones removed from upper jaw, long tail incision with both hemipenes removed; BNHS 3527 a little dehydrated with two ventral longitudinal incisions on body, ventral tail incision with one hemipenis removed; BNHS 3528 dehydrated in small parts, soft in others, ventral incision in tail; BNHS 3529 a little dehydrated with ventral incision on anterior of tail. Paratypes typically match holotype description except where noted here. Notch on ventral surface of rostral shield very shallow in BNHS 3524, more U- than C-shaped in BNHS 3526. Posterior margins of parietals not scalloped in BNHS 3524, 3525, 3526 and 3527, and more pointed than rounded in BNHS 3526, 3527, 3528 and 3529. Supraocular and postocular ‘fused’ on left of BNHS 3524; left pro- and postocular subequal in size in BNHS 3529; six infralabials on left of BNHS 3524; two postoculars on left of BNHS 3524. Each parietal contacts four rather than five scales in addition to head shields in all paratypes except for six on left of BNHS 3526, five on left of BNHS 3527 and 3529. Only first (not first and second) SLs contact nasal shield on each side in BNHS 3524. Fourth IL invariably longest and largest. Only IL 4 contacts posterior genial in BNHS 3527, 3528 and 3529, otherwise IL 5 also makes contact on one side only. Anterior and posterior genials subequal in BNHS 3524, 3526 and 3528; posterior genials slightly larger in BNHS 3529. First ventral entire rather than divided in all paratypes except BNHS 3529. Three rather than four scales between posterior genials and first ventral in BNHS 3526 (and on right of BNHS 3528 and 3529). Upper jaw marginal tooth counts approximately 10 or 11 on each side in all paratypes. Eighteen (rather than 17) dorsal scale rows at level of first ventral in BNHS 3526, though reducing to 17 by fourth ventral; 17 rows maintained to vent in BNHS 3524, 3526, 3527, 3528 and 3529; only BNHS 3525 with reduction to 16 rows (like holotype) shortly anterior to vent (level with third ventral anterior to vent). Each anal overlaps 4–7 small scales in addition to anteriormost subcaudals. Tail with subcircular rather than subtriangular cross section posteriorly in all paratypes except BNHS 3524 and 3528. Terminal scute slightly upturned in BNHS 3527 and 3528. Dark bands on dorsum of body 27–37, on tail 13–18 (Table 2). Whitish part of inverted Vs on body extend dorsally to third (rather than fourth) dorsal scale row in all paratypes except BNHS 3525 where some reach the fourth. Few of the grey speckled apices of inverted Vs extend beyond midvertebral row or connect with opposite inverted Vs in most paratypes; where they occur, connections typically confined to midbody (BNHS 3528) or posteriorly (BNHS 3524, 3525, 3529). Grey speckles on ventrals sparsely on anterior as well as posterior of BNHS 3525; only at midbody and some of last few ventrals on BNHS 3528; concentrated into very narrow, broken midventral line (as far forwards as second ventral) in BNHS 3529; absent except for single posterior speck in BNHS 3527. Anals variably without grey blotches (BNHS 3527) or speckles (left of BNHS 3526) or both (BNHS 3524). Midventral darker line on posterior of tail absent in BNHS 3526, thick but mostly broken in BNHS 3528, well developed in BNHS 3525 and 3529. Dark bands do not meet midventrally on anterior two thirds of tail in BNHS 3527. Head blackish dorsally, not paler than anterior of body in BNHS 3525. In addition to rostral, internasal (and to a lesser degree prefrontal, frontal) and anterior half of nasal and internasal paler than rest of head shields dorsally in BNHS 3524 and 3527, respectively. First SL no more speckled than other SLs in BNHS 3525, 3526, 3527 and 3528. Speckles extending beyond first ILs to second (BNHS 3524), fourth (BNHS 3526, 3527) or sixth (BNHS 3525); no speckles on left first IL of BNHS 3529. Mouth with tiny specks of pigment under tongue and posterolaterally in BNHS 3526. Colour in life. Based on uncollected specimens (Fig. 6). Head, body and tail predominantly glossy black above, creamish-white below. More precisely: upper scales of head (frontal, prefrontal, internasal, nasals, loreals, preoculars, oculars, supraoculars, postoculars, parietals and all scales bordering posterior margin of parietals) uniform glossy black. Rostral predominantly black, creamish-white on lower portion. Supralabials black above, creamish-white below; the proportions between the two shades vary from about half and half, to almost entirely creamish-white; the border between the two is indistinct and either heavily mottled, or speckled with black. Eye with round pupil; pupil and iris black. Venter creamish-white. Under the head and neck and to approximately three head-lengths behind head, the creamish-white is tinted with yellow (less markedly posteriorly) and this pale region forms an undulating line where it meets the glossy black dorsum on the lower lateral edges of the snake. After approximately three head-lengths, the pale venter colour forms upward-pointing triangles. From approximately midbody the triangles become narrower and taller, sometimes even reaching the uppermost dorsal scale row and forming, or giving the appearance of faint bands. Etymology. The specific epithet is in reference to the lack of temporal shields, a diagnostic feature of the new species. For nomenclatural purposes, the species epithet is considered a noun in apposition. Suggested common name. Mizo rain snake or Narrow-headed smithophis (English). Mizo Ruahrul (Mizo). Distribution, natural history and conservation. Smithophis atemporalis gen. et. sp. nov. is presently known with certainty only from in and around the type locality of the campus of Mizoram University and some nearby localities. The precise limits of this species’ distribution are unclear, partly because the species has sometimes previously been confused with S. bicolor, such that anecdotal reports need reassessment. Reassessment of distribution is one of the issues that need to be dealt with before the conservation assessment of this species can be adequately assessed. In Aizawl, S. atemporalis gen. et. sp. nov. is commonly seen during the monsoons, especially in, or close to streams and other wet places, close to human habitation and after rain showers. Locally this species is called ‘ ruahlawmrul ’, a rain-loving snake. Smithophis atemporalis gen. et. sp. nov. appears to have a tolerance for human dominated landscapes, though traffic is likely a threat given that these snakes have been seen on paved roads (Fig. 7). Few sightings have been made in forest habitats, though systematic survey data are not available. Smithophis atemporalis gen. et. sp. nov. have been seen in the wild feeding on earthworms and juvenile skinks, and one individual was observed in captivity eating the eggs of the frog Fejervarya asmati (HTL, SL pers. obs.). One other S. atemporalis gen. et. sp. nov. individual laid three eggs in captivity (HTL, SL pers. obs.).

Smithophis gen. nov. urn:lsid:zoobank.org:act: 444965CF-9756-4EF1-83B5-41051AE0F256 Type species. Smithophis atemporalis gen. et. sp. nov., by original designation. Diagnosis. Natricine snakes with the following combination of characters: (1) a single (���fused��� or unpaired) internasal shield and a single prefrontal shield, (2) internasal very broad and not or only slightly tapered anteriorly, (3) smooth, unkeeled dorsal scales, (4) fewer than six pairs of supralabial shields, (5) 17 rows of dorsal scales at midbody, (6) C-shaped, valvular external nares, (7) eyes with rounded or elliptical pupils, and (8) eye diameter substantially smaller than distance between eye and naris. Remarks. The new genus is clearly monophyletic and most closely related to a monophyletic Opisthotropis in our molecular phylogenetic trees, and the two genera are similar phenotypically in having a single prefrontal shield (paired in a few species of Opisthotropis). Molecular phylogenetic support for Opisthotropis monophyly is not yet compelling though clearly stronger than for its non-monophyly, but we have elected to erect a new genus given also that (1) Smithophis gen. nov. is distinct from Opisthotropis in also having a single internasal shield (which, in concert with the single prefrontal, is a unique condition among natricines), in having fewer than 6 supralabial shields on each side, and in having internasal(s) that are not or only slightly anteriorly tapered, and (2) because the phylogenetic relationships of several key taxa remain unclear given incomplete taxon sampling thus far in phylogenetic analyses���in this context we are thinking particularly of the type species of Opisthotropis (O. ater G��nther, 1872), and any representatives of any of the at least superficially similar (David et al. 2015) monotypic Isanophis David et al. 2015, and Paratapinophis Angel, 1929 (though it might be noted that the latter three taxa also differ from Smithophis gen. nov. in having paired internasals that are not notably broad and which taper anteriorly, and in having more than 6 supralabials). All specimens of Smithophis we have observed have a single internasal shield and, to the best of our knowledge, all specimens of Opisthotropis have paired internasals. Species of all synonyms of Opisthotropis have paired rather than single internasals. Single internasals are derived, providing additional evidence to support the monophyly of the new genus. Wall (1908: 322���323) considered the unusual head shields of Rhabdops bicolor distinctive enough that it was ���extremely probable that it will have to be separated and relegated to a genus by itself���. Content. Two nominal species; the type species (described below) and S. bicolor (Blyth, 1854). Etymology. Named in honour of Malcolm A. Smith (1875���1958) in recognition of his many contributions to herpetology and especially to the knowledge of Asian snakes. Distribution. Northeast India, from Meghalaya state, northeast India east to Myanmar and possibly western Yunnan, China (Anderson 1879; Pope 1935; Smith 1943; Dowling & Jenner 1988; Wallach et al. 2014)., Published as part of Giri, Varad B., Gower, David J., Das, Abhijit, Lalremsanga, H. T., Lalronunga, Samuel, Captain, Ashok & Deepak, V., 2019, A new genus and species of natricine snake from northeast India, pp. 241-264 in Zootaxa 4603 (2) on page 245, DOI: 10.11646/zootaxa.4603.2.2, http://zenodo.org/record/2682351, {"references":["Wall, F. (1908) Notes on a collection of snakes from the Khasi Hills, Assam. Journal of Bombay Natural History Society, 18 (2), 312 - 337.","Anderson, J. (1879) Reptilia and amphibia. In: Anderson, J. (Ed.), Anatomical and zoological researches, comprising an account of the zoological results of the two expeditions to western Yunnan in 1868 and 1875. Fol. 1. B. Quaritch, London, pp. 809 - 810.","Pope, C. H. (1935) The Reptiles of China, Turtles, Crocodilians, Snakes, Lizards. Natural History of Central Asia, 10, i-xvii + 1 - 604, 25 pls., 78 figs.","Smith, M. A. (1943) The fauna of British India, Ceylon and Burma including the whole of the Indo-Chinese sub-region. Reptilia and Amphibia. Fol. III. Serpentes. Taylor and Francis, London, xii + 583 pp.","Dowling, H. G. & Jenner, J. V. (1988) Snakes of Burma: Checklist of reported species and bibliography. Smithsonian Herpetological Information Service, 76, 1 - 19.","Wallach, V., Williams, K. L. & Boundy, J. (2014) Snakes of the World: A catalogue of living and extinct Species. CRC Press, Taylor & Francis Group, London, 1227 pp. https: // doi. org / 10.1201 / b 16901"]}

Phylogenetic datasets

Cyrtodactylus bhupathyi sp. nov. Fig. 4, 5, and 6A Holotype. BNHS 2255, adult female, near Bagdogra (26.88004��N 88.47187��E; ca. 200 m asl.), Darjeeling District, West Bengal state, India, collected by A. Datta-Roy and I. Agarwal, 28 October 2010. Paratype. BNHS 2256, adult female, same data as holotype. Etymology. The specific epithet is a patronym honouring the late Subramaniam Bhupathy for his contributions to Indian herpetology. Dr. Bhupathy also led herpetological research projects on elevational patterns in reptile diversity in Sikkim (Chettri et al. 2010). Dr. Bhupathy passed away in 2014 in an accident while conducting fieldwork in the southern Western Ghats, India. Suggested common name. Bhupathy���s bent-toed gecko. Diagnosis and comparison with regional congeners. A small sized Cyrtodactylus, snout-vent length to 61.5 mm; body and limbs relatively slender, digits relatively short; two pairs of well-developed postmentals, inner pair longer than outer and either in broad contact behind mental or separated by a large scale; 24���25 rows of circular tubercles, bluntly conical and feebly keeled throughout; 51���55 paravertebral tubercles; 37 or 38 mid-body ventral scale rows; no precloacal groove; 4���7 weakly pitted femoral scales on each side separated by 17 or 18 smaller nonpored scales from a single series of 10 or 11 pitted precloacal scales on females; male condition unknown. Five or six basal and 13 apical subdigital lamellae beneath Digit IV of pes, with three intervening rows of non-lamellar granular scales between basal and distal lamellae series. Subcaudal scalation of original tail without clearly defined median series of enlarged scales. Dorsal pattern of 8���10 paired transversely arranged dark blotches/cross-bars, which may be more or less irregular, occasionally alternating with lighter blotches. Original tail with 11���13 alternating dark and light bands, distal-most light bands almost white. The condition of females with precloacal and femoral pores separated by a diastema in the new species is unique within the Indo-Burma Cyrtodactylus clade for which females are known. Cyrtodactylus bhupathyi sp. nov. differs from the following species in having 24 or 25 dorsal tubercle rows: C. annandalei (16���18 DTR), C. cayuensis (18 DTR), C. fasciolatum (15���17 DTR), C. gubernatoris (20 or 21 DTR), Cyrtodactylus himalayicus comb. nov. (19���21 DTR), C. mandalayensis (18 DTR), C. markuscombaii (14 or 15 DTR), C. russelli (22 DTR), C. tamaiensis (21 DTR), the new species from Tripura described below (19���21 DTR), and C. brevidactylus (27 DTR). The small body size of C. bhupathyi sp. nov. (SVL to 61.5 mm) and the number of mid-body ventral scale rows (37 or 38) distinguishes it from the large bodied species C. slowinskii (SVL to 108 mm, MVSR 27���32). Cyrtodactylus bhupathyi sp. nov. differs from the following species by possessing femoral pores separated from precloacal pores in females: C. ayeyarwadyensis (no pores on females, continuous series of 10���28 Pcp to PcFP pores on males), C. gansi (14 small PcP on females, 16���29 PcP on males), C. khasiensis (no pores on females, 10���12 PcP on males), C. martinstolli (no pores on females, 0���8 PcP on males), C. wakeorum (12 PcP on female holotype). Cyrtodactylus bhupathyi sp. nov. is most similar morphologically to C. gubernatoris from which it differs in the number of paravertebral tubercles (51���55 vs. 46���48), dorsal tubercle rows (24 or 25 vs. 20 or 21), and by the presence of femoral pores/partially pitted femoral scales in females (vs. absence on females). Description of holotype. Holotype in generally good condition except for minor folds of skin on thighs, head slightly kinked toward left, constriction anterior to hindlimb insertions (from tying a previously attached specimen tag), all artefacts of preservation; tail tip removed as tissue sample for molecular analyses (Fig. 4A). Adult female, SVL 61.5 mm. Head moderately long (HL/SVL ratio 0.27), and wide (HW/HL ratio 0.64), dorsoventrally depressed (HH/HW ratio 0.57), distinct from neck; loreal region slightly inflated, interorbital area flat, canthus rostralis not prominent; snout moderately short (SE/HL ratio 0.40), almost twice as long as OD (OD/SE ratio 0.55); scales on forehead, canthus rostralis and snout homogenous, small, rounded and granular; scales on snout and canthus rostralis slightly larger than those on forehead; scales of interorbital and occipital region homogenous, granular, those in occipital region mixed with slightly larger, rounded, conical tubercles (Fig. 5A). Eye small (OD/ HL ratio 0.22); pupil vertical with crenulated margins; supraciliaries mucronate, decreasing in size towards posterior edge of orbit; ear opening oval, obliquely oriented, large; eye to ear distance slightly more than eye diameter. Rostral wider (2.7 mm) than deep (1.4 mm), partially divided dorsally by weakly developed rostral groove; single large supranasal on either side, separated by two small scales (internasals), which are approximately twice the size of enlarged granular scales on snout; rostral in contact with SL I, nasals, supranasals and an internasal; nostrils semicircular, laterally oriented, posterior half covered by nasal pad, each in broad contact with rostral and also surrounded by supranasal, SL I, and three or four postnasals; three or four scale rows separate orbits from supralabials; mental slightly wider (2.0 mm) than long (1.7 mm), triangular, two pairs of welldeveloped postmentals, inner pair longer (maximum length 1.5 mm) than and separating outer pair (maximum length 0.8 mm), separated from each other behind mental by a large gular scale which is subequal in size to outer postmental pair; inner postmentals bordered by mental, IL I, outer postmental and six gular scales; outer postmental bordered by inner postmental, IL I and IL II, and two gular scales on either side; supralabials 10 (8) on each side, bordered by a row of large, flat, slightly elongated scales (Fig. 5C); infralabials 9/9, IL II to IL VII bordered by one row of chinshields, largest anteriorly; interorbital scale rows across narrowest point of frontal bone approximately 30. Body moderately slender, relatively short (TrL/SVL ratio 0.47) with weak ventrolateral folds; dorsal scales heterogeneous, mostly rounded granular, intermixed with irregularly arranged small (2���4 times granule size) circular tubercles, bluntly conical and feebly keeled throughout (Fig 4), becoming more conical and smaller towards flanks, tubercles extend from frontal region to proximal one third of tail length; tubercles on nape smaller than those of dorsum, largest on flanks; enlarged tubercles on tail completely flat and weakly pointed and keeled; tubercles in approximately 24 irregular longitudinal rows at mid-body; 51���53 paravertebral tubercles; ventral scales much larger than dorsal scales, smooth, cycloid, imbricate to subimbricate, 37 or 38 mid-body ventral scale rows; gular scales smaller than ventrals and granular except a few rows of larger, flat and juxtaposed scales, including a single row of chinshields bordering mental, postmentals and infralabials (Fig 5C). Eleven pitted precloacal scales in a continuous series with 17 or 18 slightly smaller poreless scales followed by four weakly pitted femoral scales on either side; no precloacal groove. Three or four rows of enlarged scales between pitted precloacal scales and vent, as large as the largest ventrals and much larger than pitted precloacal scales. Tail original, dorsoventrally depressed, without distinct median furrow, tapering; tail tip removed for molecular analyses. Dorsal scales at base of tail granular, gradually becoming flatter, subimbricate posteriorly, increasing in size on lateral aspect, intermixed with 6���8 slightly enlarged tubercles near base of tail and reducing to two by fourth transverse row of tubercles (Fig 4); ventral scales larger that dorsal scales, imbricate, median row comprises irregularly enlarged subcaudals in one or two rows; two enlarged postcloacal tubercles at base of tail. Fore and hindlimbs relatively slender; forearm (FL/SVL ratio 0.15) and crus (CL/SVL ratio 0.18) relatively short; digits relatively short, strongly inflected at each joint, all bearing robust, recurved claws; subdigital lamellae widened beneath basal phalanx; basal lamellae series on Digits I-V: 4-4-4-5-3 (right manus) and 5-6-6-6-6 (right pes); apical lamellae series (intervening rows of non-lamellar granular scales between basal and distal lamellae series in parentheses) on Digits I-V: 6(3)-8(2)-10(1)-8(3)-7(3) (right manus) and 8(1)-8(3)-9(3)-9(3)-9(3) (right pes); interdigital webbing absent on manus, rudimentary between Digits I���V of pes; relative length of digits (measurements in mm in parentheses): I (3.7) Colouration in preservative. Ground colouration light brown, dorsum with darker irregular broken up transverse markings with an almost unmarked mid-vertebral line, approximately 8���10 irregular cross-bars on dorsum anterior to hindlimb insertions, fused together between forelimb insertions and neck. Occipital markings separated from post-orbital streak, which continues indistinctly anterior to eye. Dorsum of head unicolour except for indistinct paired dark markings anterior to orbits. Limbs mottled with dark markings, flanks with irregular light and dark markings. Venter off-white with dark abdomen (artefact of preservation). Tail colouration similar to ground colour with 11 darker bands on original tail, lighter bands fading toward tail-tip (Fig. 4). Colouration in life. Ground colouration pinkish-brown with darker brown-black transverse markings, head darker than ground colour of body and lighter than dark dorsal markings. Anterior portion of brille yellow, pupil outlined by red, iris dull greenish-gold (Fig. 6A). Variation. This species is known from only two female specimens, the holotype and paratype. The paratype (BNHS 2256) closely agrees with holotype (Table 3) except that it has seven femoral pores on each side; the inner pair of postmentals are in broad contact posterior to the mental; 11 (9) supralabials on each side; 11 infralabials on the left side and on the right; two to three rows of enlarged scales between precloacal pore-bearing scales and vent, as large as the largest ventrals and much larger than pore-bearing scales; dorsal cross-bars more irregular. Molecular Data. Cyrtodactylus bhupathyi sp. nov. is more than 16% divergent from Cyrtodactylus ayeyarwadyensis, C. khasiensis, and the new species from Tripura (for both ND2 and cyt b), and 13.2% (cyt b), and 16.7% (ND2) divergent from its closest sampled sister species, C. gubernatoris (Table 4). Agarwal et al. (2014) incorrectly identified this species as Cyrtodactylus gubernatoris (CES10/1235= BNHS 2255). Distribution and Natural History. This species was encountered during approximately one-half an hour of fieldwork an hour after sunset along a national highway. The geckos were spotted by eyeshine TABLE ��. Measurements (mm) anđ meristic đata for the type series of Cyrtodactylus bhupathyi sp. nov. anđ Cyrtodactylus tripuraensis sp. nov., as well as the holotypes anđ referređ material of Cyrtodactylus gubernatoris, Cyrtodactylus himalayicus comb. nov. anđ C. khasiensis. Abbreviations as in materials anđ methođs. * inđicates incomplete/regenerateđ tail, x inđicates measurement coulđ not be taken on that specimen. Numbers in parentheses inđicate supralabial at miđorbital position. ������continued on the next page TABLE ��. (Continueđ), Published as part of Agarwal, Ishan, Mahony, Stephen, Giri, Varad B., Chaitanya, R. & Bauer, Aaron M., 2018, Two new species of bent toed geckos, Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Northeast India with comments on name-bearing types from the region, pp. 334-356 in Zootaxa 4420 (3) on pages 342-347, DOI: 10.11646/zootaxa.4420.3.2, http://zenodo.org/record/1250911, {"references":["Agarwal, I., Bauer, A. M., Jackman, T. R. & Karanth, K. P. (2014) Insights into Himalayan biogeography from geckos: A molecular phylogeny of Cyrtodactylus (Squamata: Gekkonidae). Molecular Phylogenetics and Evolution, 80, 145 - 155. https: // doi. org / 10.1016 / j. ympev. 2014.07.018"]}

Cyrtodactylus himalayicus (Annandale, 1906) comb. nov. Fig. 2E and F Gymnodactylus himalayicus Annandale, 1906: 287; 1907: pl. 5, fig. 1. Holotype [by monotypy]. ZSIK 15716, male, from “ Kurseong, Darjeeling dist. (5,000 ft)” [= Kurseong subdivision (ca. 1524 m asl.), Darjeeling district, West Bengal state, India], collected by N. Annandale, ca. 1906 (Annandale 1906). Additional material. ZSIK 19546, adult female, from “Gopaldhara, Darjeling dist.” (Smith 1935; Ahmed & Dasgupta 1992) [=Gopaldhara Tea Estate, near Mirik town, Darjeeling district, West Bengal state, India], collected by H. Stevens, 24 April 1946. Diagnosis and comparisons with regional congeners. A small sized Cyrtodactylus, snout-vent length to 64.5 mm; body relatively slender; limbs and digits moderately long; two or three pairs of well-developed postmentals, inner pair longer than outer two and in broad contact behind mental; 19–21 rows of smooth weakly raised to bluntly conical, rounded dorsal tubercles; 33 or 34 [35 count in Annandale 1906] ventral scales between outermost ventral scale rows enlarged relative to granular flank scales; ventrolateral folds absent; no precloacal groove; 10 precloacal pores in a single series on the male and an equivalent 10 fine pores/ pitted scales on the female, femoral pores absent on both sexes; scales bordering precloacal pore row are approximately 3x enlarged relative to porebearing scales. Nine basal subdigital lamellae and 11 distal lamellae on Digit IV of pes [n=1: toe tips broken off on ZSIK 15716]. Subcaudal scalation of original tail without single median series of enlarged plates, though subcaudals are moderately enlarged relative to dorsal caudal scales. Dorsal pattern of approximately eight paired transversely arranged small dark spots on trunk and sacrum of male, or female with four almost continuous longitudinal brown stripes on trunk. Original tail with approximately 13 dark bands. Cyrtodactylus himalayicus comb. nov. differs from the following species in the presence of only precloacal pores on males (vs. PcP + femoral pores): C. annandalei Bauer, C. fasciolatum, C. gubernatoris, C. russelli and C. slowinskii. Cyrtodactylus himalayicus comb. nov. differs from the following species by possessing 10 precloacal pores: C. brevidactylus Bauer (8 PcP), C. cayuensis (6–9 PcP), C. mandalayensis (8 PcP + single enlarged porebearing scale posterior to the series), C. markuscombaii (7 PcP), C. martinstolli (0–8 PcP); and from the following species by possessing precloacal pores only (vs. precloacofemoral pores): C. gansi (16–29 PcFP), and C. tamaiensis (40 PcFP). Cyrtodactylus himalayicus comb. nov. differs from the following species by having 19–21 dorsal tubercle rows: C. brevidactylus (27 DTR), C. mandalayensis (18 DTR), C. markuscombaii (14–15 DTR), and C. wakeorum Bauer (24 DTR). Cyrtodactylus himalayicus comb. nov. lacks a ventrolateral fold, which distinguishes it from C. ayeyarwadyensis, C. cayuensis, C. mandalayensis and C. martinstolli. Cyrtodactylus himalayicus comb. nov. differs from C. khasiensis by having enlarged plate-like scales posteriorly bordering the pore-bearing scale series (equivalent scales not enlarged on C. khasiensis), and snout to vent length to 64.5 mm (to 83.3 mm for C. khasiensis). Refer to the diagnosis section of new taxa for comparisons with Cyrtodactylus himalayicus comb. nov. Notes. Smith (1935) examined available specimens of C. “ khasiensis ” in the collections of ZSIK and BMNH, to which he referred a specimen from “Mogok, Upper Burma ” (BMNH 1900.9.20.1), which has eight precloacal pores. Having examined the holotype and referred specimen (ZSIK 19546) of Gymnodactylus himalayicus, Smith (1935) noted that this taxon had 10 preanal pores, which fell within the morphological range of his redefined C. khasiensis. As a result, Smith (1935) referred G. himalayicus to the synonymy of C. khasiensis. Cyrtodactylus himalayicus comb. nov. is still known from only two poorly preserved specimens and all pertinent taxonomic characters could not be collected from these. Our recent examination of these specimens demonstrate that this taxon is sufficiently distinct from C. khasiensis, as well as all other species in the region, and is herein raised to the status of a valid species.

Cyrtodactylus tripuraensis sp. nov. Figs. 6B, 7���9 Holotype. BNHS 2244, adult male, from Chobimura (23.55039��N 91.62054��E; ca. 36 m asl.), Gomati district, Tripura state, India, collected by A. Datta-Roy, T. Khichi, NPI Das and I. Agarwal, 10 November 2010. Paratypes. All from Tripura state, India, same collectors as holotype: BNHS 2230, adult female, from Rowa Wildlife Sanctuary (24.29062��N 92.16496��E; ca. 56 m asl.), North Tripura district, 0 2 October 2010; BNHS 2236��� 2238, BNHS 2242, and BNHS 2243, five adult males, BNHS 2239���2241, three adult females, Sepahijala Wildlife Sanctuary (23.65906��N 91.33167��E; ca. 29 m asl.), Sepahijala district, 06���08 November 2010; BNHS 2245, adult male, Gumti (23.42661��N 91.81937��E; ca. 40 m asl.), Gomati district, 11 November 2010. Etymology. The species epithet is a toponym for Tripura, the Indian state in which all currently known populations are found. Suggested common name. Tripura bent-toed gecko. Diagnosis and comparison against regional congeners. A moderate sized Cyrtodactylus, snout-vent length to 70.7 mm; body and limbs relatively slender, digits short; one pair of enlarged postmental scales in broad contact behind mental; 19���21 rows of circular tubercles at mid-dorsum, bluntly conical and weakly keeled throughout; 44��� 54 paravertebral tubercles; 35���43 mid-body ventral scale rows; no precloacal groove; males with 29���37 precloacofemoral pores in a single series, females with 19���29 pitted/pored scales. Four to nine basal and 11���16 apical subdigital lamellae beneath Digit IV of pes with one or two intervening rows of non-lamellar granular scales between basal and distal lamellae series. Subcaudals larger than dorsal tail scales, not in an enlarged median series (on original tail). Dorsal pattern of 7���10 paired transversely arranged dark spots that may be irregular and broken up, occasionally alternating with lighter spots. Original tail with 11���13 alternating dark and light bands, last few light bands almost white. Cyrtodactylus tripuraensis sp. nov. differs from the following species by the presence of a continuous series of precloacofemoral pores (vs. precloacal pores separated from femoral pores by a diastema of multiple poreless scales): C. annandalei, Cyrtodactylus bhupathyi sp. nov., C. fasciolatum, C. gubernatoris, C. russelli, and C. slowinskii. The large number of precloacofemoral pores on males of the new species (29���37) separates it from C. brevidactylus (8 PcP), C. cayuensis (6���9 PcP), Cyrtodactylus himalayicus comb. nov. (10 PcP), C. khasiensis (10��� 12 PcP), C. mandalayensis (8 PcP + single enlarged pore-bearing scale at the posterior apex of PcP), C. markuscombaii (7 PcP), C. martinstolli (0���8 PcP), and C. wakeorum (12 PcP). Cyrtodactylus tripuraensis sp. nov. differs from C. tamaiensis by having slightly fewer precloacofemoral pores (29���37 vs. 40), male body size (SVL 55.4���65.0 mm vs. 90.0 mm), longer trunk (TrL/SVL mean 0.49, range 0.46���0.52 vs. 0.41) and dorsal pattern (8���10 irregular paired dark spots vs. irregular dark and light reticulations), and from C. gansi in the absence of a precloacal groove (vs. presence of precloacal groove on males). Cyrtodactylus tripuraensis sp. nov. is most similar to C. ayeyarwadyensis from which it differs by a slightly higher number of precloacofemoral pores on males (29��� 37 vs. 10���28), the presence of pores in females (vs. absence of pores), and fewer dorsal tubercle rows (19���21 vs. 22���24). Cyrtodactylus tripuraensis sp. nov. differs from C. ayeyarwadyensis sensu lato (Mahony et al. 2009a) by the presence of precloacofemoral pores on females (vs. absence of pores). Description of holotype. Holotype in generally good condition, except minor folds on skin of abdomen, slightly protruding tongue, constriction anterior to hindlimb insertions (from tying previously attached specimen tag), all artefacts of preservation; tail tip removed for molecular tissue sampling (Fig. 7A). Adult male, snout to vent length 63.7 mm. Head moderately long (HL/SVL ratio 0.27), slightly wide (HW/HL ratio 0.62), somewhat depressed (HH/HW ratio 0.58), distinct from neck; loreal region slightly inflated, interorbital area flat, canthus rostralis not prominent; snout moderately short (SE/HL ratio 0.40), almost twice as long as OD (OD/SE ratio 0.56); scales on forehead homogenous, small, rounded, granular; those on canthus rostralis and snout are homogenous, slightly larger than those on forehead, roughly hexagonal, juxtaposed; distinctively larger than those on interorbital region and occipital region; scales on interorbital and occipital region homogenous, granular, those on occipital region smallest (Figure 8A). Eye small (OD/HL ratio 0.23); pupil vertical with crenulated margins; supraciliaries small, slightly mucronate, decreasing in size towards posterior edge of orbit, last three or four more strongly mucronate; ear opening oval, obliquely oriented, large (EL/HL ratio 0.09); eye to ear distance slightly more than eye diameter (OD/EE ratio 0.72). Rostral wider (2.7 mm) than deep (1.8 mm), divided dorsally by weakly developed rostral groove; single large supranasal on either side, separated by eight smaller scales arranged in three rows completely filling space formed by dorsal bifurcation of rostral; rostral in contact with SL I, nasals, supranasals, and three scales separating supranasals; nostrils semicircular, laterally oriented, posterior half covered by nasal pad, each in broad contact with rostral and also surrounded by supranasal, SL I, and three to four postnasals; three to four scale rows separate orbits from supralabials; mental wider (2.5 mm) than long (1.8 mm), triangular, two well developed postmentals on either side, inner pair much larger (maximum length 2.0 mm) than outer pair and broadly in contact with each other (forming 1.0 mm long suture) behind mental, outer postmental pair 0.8 mm long, separated from each other by inner postmental pair; inner postmentals bordered by mental, IL I, outer postmental and six gular scales; outer postmental bordered by inner postmental, IL I and II, and three gular scales on either side (Figure 8C); supralabials 10 (7) (right)���12 (9) (left), bordered by two rows of large, flat, slightly elongated scales; nine infralabials on each side, bordered by one row of chinshields, largest anteriorly; interorbital scale rows across narrowest point of frontal bone ~25. Body moderately slender, relatively short (TrL/ SVL ratio 0.52) with weak ventrolateral folds; dorsal scales heterogeneous, mostly rounded granular, intermixed with irregularly arranged small (2���4 times granule size) circular tubercles, bluntly conical and weakly keeled throughout, becoming more conical and strongly keeled posteriorly towards tail, tubercles extend from frontal region to proximal portion of tail; tubercles on nape considerably smaller than those of dorsum and those of tail completely flat and elongate (Figures 7A, 7C); tubercles in approximately 21 non-linear longitudinal rows at midbody; 46 tubercles in paravertebral row from occiput to mid-sacrum; ventral scales much larger than dorsal scales, smooth, cycloid, imbricate to subimbricate, 37 mid-body ventral scale rows; posterior most row under thighs, and those bordering pored scales posteriorly and medially, enlarged; gular region with smaller granular scales, grading into large, flat and juxtaposed scales anterolaterally, those posterior to chinshields largest. Twenty-nine precloacofemoral pores in a continuous series, most prominent near precloacal region; no precloacal groove; a patch of enlarged scales border pored scales posteriorly and medially; prominent hemipenal bulge. Tail base original, remaining portion regenerated, rounded, without distinct median furrow, tail tip removed. Dorsal scales on original portion of tail granular, gradually becoming flatter and subimbricate posteriorly, intermixed with six to eight slightly enlarged, flat tubercles; ventral scales larger than dorsal scales, subimbricate; no median row of enlarged subcaudals; regenerated portion covered with uniform, small, flat subimbricate scales throughout; three enlarged postcloacal tubercles at base of tail on each side. Fore and hindlimbs relatively slender; forearm (FL/SVL ratio 0.14) and crus (CL/SVL ratio 0.16) relatively short; digits relatively short, strongly inflected at each joint, all bearing robust, recurved claws; subdigital lamellae widened beneath basal phalanx; basal lamellae series on Digits I-V: 4-4-4-5-3 (right manus) and 4-4-5-7-5 (right pes); apical lamellae series (intervening rows of non-lamellar granular scales between basal and distal lamellae series in parentheses) on Digits I-V: 7(1)-9(0)-10(0)-9(1)-8(1) (right manus) and 7(2)-8(2)-10(1)-11(1)-11(1) (right pes); interdigital webbing absent on manus, rudimentary between Digits I���V of pes; relative length of digits (measurements in mm in parentheses): I (3.5) Colouration in preservative. Ground colour mottled brown, dorsum with six darker irregular paired blotches from hindlimb insertion to nape, one each on occipital region and tail base. Head and limbs slightly rufous, heavily mottled with darker markings. Venter off-white with darker abdomen (artefact of preservation). Colour of regenerated tail similar to dorsum, with dark speckling (Fig. 7). Colouration in life. Ground colouration in life mottled brown with darker brown-black transverse markings, head with indistinct markings. Pupil outlined by bronze, remaining iris dull greenish-gold (Fig. 6B). Variation. Relative size and shape of dark blotches on the back variable, sometimes very broken up (Fig. 9). There are six males and four females in the type series, with 29���37 PcFPs on males and 19���29 pitted scales on females. The largest specimen is 70.7 mm SVL (BNHS 2230). The number of mid-body ventral scale rows ranges from 35���43, the number of dorsal tubercle rows from 19���21, supralabials from 9���12, infralabials 8���10, and the number of postcloacal spurs from 2���4 (Table 3). Molecular Data. Cyrtodactylus tripuraensis sp. nov. is more than 12% divergent from C. gubernatoris, C. khasiensis and Cyrtodactylus bhupathyi sp. nov., with a minimum divergence of 7.3/ 9.8% (ND2/ cyt b) from its closest sampled sister taxon, C. ayeyarwadyensis (Table 4). Animals from Sepahijala (BNHS 2236, 2238) and Gumti (BNHS 2245) are less than 1% divergent from each other, and are 4.6���5.0% divergent from BNHS 2230 from Rowa, North Tripura (ND2 data). Agarwal et al. (2014) referred to this species as Cyrtodactylus sp. Tripura. Distribution and Natural History. This is one of the few widely distributed Cyrtodactylus species in the Indo-Burma region (Agarwal et al. 2014), found across the lowlands of Tripura. These geckos were always collected C. tripuraensis sp. nov. should be found in neighbouring regions of eastern and southeastern Bangladesh, and southern Assam (e.g., Karimganj district) in India. This species, though widely distributed, was not observed by us in the same abundance as other Cyrtodactylus species found in more rocky habitats in other parts of Northeast India (including a high elevation habitat in Tripura). Cyrtodactylus tripuraensis and C. ayeyarwadyensis are the only Cyrtodactylus in India and Myanmar with wide distributions that have been confirmed by genetic data (Agarwal et al. 2014), and these two species both live in lowland habitats at elevations below 300 m asl. Remarks. Southeastern Bangladesh populations referred to as Cyrtodactylus sp. (Mahony & Reza 2008; Mahony et al. 2009b) were later considered to represent Cyrtodactylus ayeyarwadyensis based on overall morphological similarity (Mahony et al. 2009a). The northern most Bangladesh population, from Kaptaimukh Beat, is located ca. 100 km south of Gumti Wildlife Sanctuary in the same continuous north���south running valley. These Bangladesh populations therefore require further molecular confirmation of taxonomic affinity in light of the morphologically similar new species, Cyrtodactylus tripuraensis sp. nov. described here., Published as part of Agarwal, Ishan, Mahony, Stephen, Giri, Varad B., Chaitanya, R. & Bauer, Aaron M., 2018, Two new species of bent toed geckos, Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Northeast India with comments on name-bearing types from the region, pp. 334-356 in Zootaxa 4420 (3) on pages 348-353, DOI: 10.11646/zootaxa.4420.3.2, http://zenodo.org/record/1250911, {"references":["Mahony, S., Ahmed, M., Hossain, M. K., Kabir, M. M. & Hasan, M. K. (2009 a) Cyrtodactylus ayeyarwadyensis BAUER, 2003 (Reptilia: Gekkonidae) in Bangladesh with habitat details of new collection localities and a discussion of morphological variation. Salamandra, 45 (4), 245 - 250.","Agarwal, I., Bauer, A. M., Jackman, T. R. & Karanth, K. P. (2014) Insights into Himalayan biogeography from geckos: A molecular phylogeny of Cyrtodactylus (Squamata: Gekkonidae). Molecular Phylogenetics and Evolution, 80, 145 - 155. https: // doi. org / 10.1016 / j. ympev. 2014.07.018","Mahony, S. & Ali Reza, A. H. M. (2008) A herpetofaunal collection from the Chittagong Hill Tracts, Bangladesh, with two new species records for the country. Hamadryad, 32 (1), 45 - 56.","Mahony, S., Hasan, M. K., Kabir, M. M., Ahmed, M. & Hossain, M. K. (2009 b) A catalogue of amphibians and reptiles in the collection of Jahangirnagar University, Dhaka, Bangladesh. Hamadryad, 34 (1), 80 - 94."]}

Cyrtodactylus kazirangaensis sp. nov. Figs. 6���9. Holotype. Adult male (BNHS 2148, field number CES09/1137) collected from near Hatikhuli Tea Estate, Golaghat district, Assam state, India (26.57810��N, 93.40701��E, 100 m asl.) by Ishan Agarwal on 11 November 2009. Paratypes. Adult males (BNHS 2147, BNHS 2149) bear the same collection data as the holotype. Etymology. The specific epithet is a toponym for Kaziranga National Park, which is adjacent to the type locality. Kaziranga, a World Heritage Site, is best known for having most of the world���s surviving Indian onehorned rhinoceros, though it has high biodiversity across taxonomic groups. Diagnosis. Cyrtodactylus kazirangaensis sp. nov. can be distinguished from all congeners by its moderate body size (SVL to at least 80.0 mm); 11 or 12 supralabials; 9���11 infralabials; 22 or 23 longitudinal rows of rounded, feebly keeled dorsal tubercles; tubercles not extending beyond first segment of tail; 36���38 paravertebral tubercles; 37���43 ventral scales between ventrolateral folds; no precloacal groove; 10 or 11 precloacal pores and no femoral pores in males; 14���19 total subdigital lamellae beneath toe IV of pes; subcaudal scalation of original tail without enlarged plates; dorsal pattern composed of six or seven irregular light and dark crossbars; tail with alternating dark and light bands. Description of holotype. Holotype in good preservation condition: tail curved toward left and tail tip removed as tissue voucher, fourth digit of right pes broken at inflection, constriction just anterior to hind limb insertion due to overzealous tag tying. Adult male, SVL 79.0 mm. Head long (HL/SVL 0.27; HW/HL 0.69), somewhat depressed (HD/HW 0.55), distinct from neck; loreal region somewhat inflated, interorbital region flat, canthus rostralis not prominent; snout short (SE/HL 0.45), twice as long as orbital diameter (ED/SE 0.49); scales on snout and canthus rostralis granular, juxtaposed, homogenous in shape, with scales on snout being slightly larger; scales on interorbital region, dorsal surface of eyelids, forehead and occipital region slightly smaller, granular, juxtaposed, with those on forehead and interorbital region interspersed with slightly larger tubercles that are rounded, smooth and two to three times the size of adjacent granules; enlarged tubercles sparse on forehead and dense on occipital region. Eye small (ED/HL 0.22); pupil vertical with crenulated margins; supraciliaries large, decreasing in size towards posterior and anterior ends of orbit, largest about one-third the way from anterior edge of orbit; dorsal surface of eyelids covered with granular scales, row of tubercles bordering supraciliaries absent. Ear opening oval, obliquely oriented, large; eye to ear distance slightly more than eye diameter (ED/EE 0.73). Rostral much wider (3.2 mm) than deep (1.4 mm), partially divided dorsally by weakly developed rostral groove; single much enlarged supranasal on either side, separated by one slightly smaller, circular internasal, which is twice the size of enlarged scales on snout; rostral in contact with supralabial I, nostrils, supranasals and internasals; nostrils oval, laterally oriented, posterior half covered by a nasal pad, each in broad contact with rostral and surrounded by supranasal, supralabial I, and four smaller, granular postnasal scales; seven rows of granular scales between midorbital position and supralabials; mental wider (2.8 mm) than deep (2.0 mm), triangular, two well-developed postmentals on either side, inner pair in broad contact (1.5 mm) behind mental, larger (2.2 mm) than and separating outer pair (1.3 mm); each inner postmental bordered by mental, infralabial I, outer postmental and two enlarged gular scales; outer postmentals bordered by inner postmentals, infralabials I and II, a large, roughly hexagonal scale posteriorly and four gular scales; eight supralabials to midorbital position and 12 to angle of jaw on each side, bordered by a row of flat, somewhat elongated scales, slightly larger than their adjacent granules; 10 infralabials on each side. Body slender (BW/TRL 0.42), short (TRL/SVL 0.44); dorsal scales heterogeneous, mostly rounded granules, intermixed with irregularly arranged enlarged tubercles that are rounded and feebly keeled, flatter and more strongly keeled towards vertebral region; tubercles extend from forehead posteriorly onto base of tail; tubercles on nape smaller than those of dorsum; 22 dorsal tubercles across mid dorsum; 36���38 paravertebral tubercles; ventrolateral folds covered by scattered enlarged, smooth scales; ventral scales much larger than dorsals, smooth, rounded, juxtaposed; smaller under thighs; two rows of enlarged subimbricate scales along posterior border of precloacal pores; 39 or 40 ventral scales between ventrolateral folds; gular region with much smaller granular scales throughout, except two rows of larger, flat, juxtaposed scales originating from the posterior margin of the outer postmentals and bordering infralabials. Eleven precloacal pores in a continuous series; femoral pores absent; no precloacal groove; hemipenal bulge distinct. Fore and hind limbs slender; forearm (FL/SVL 0.15) and tibia (CL/SVL 0.17) short; digits short, strongly inflected at each joint, all bearing robust recurved claws; five subdigital lamellae in basal series and 10 in distal series with four non-lamellar granules at inflection (digit IV, right manus); eight basal and 11 distal lamellae with four non-lamellar granules at inflection (digit IV, right pes); inter-digital webbing absent on manus, rudimentary between toes I to IV; relative length of digits: I Colouration in life (for paratype BNHS 2149, Fig. 6). Dorsal ground colour of body, limbs and tail brown; top of head pale brown with scattered dark brown streaks; labials darker than dorsum of head; a preorbital streak and two light yellow narrow postorbital streaks that do not extend as far as ear opening; occiput with light yellowbuff blotches; neck with light tan blotches forming indistinct wavy cross-bar flanked by a pair of dark blotches, light mid-dorsal marking extends from occiput to tail base, seven light and seven dark bands from fore limb insertion to tail base; ventral surfaces immaculate; limbs with indistinct crossbars and a few lighter blotches; regenerated tail light brown with dark spots dorsally, light grey and tan ventrally; iris green-brown with darker reticulations, pupil outlined in orange-red Colouration in preservative (Figs. 7 & 8). Dorsal ground colour of body, limbs and tail brown; top of head mottled brown; labials about as dark as dorsum of head; two indistinct light narrow postorbital streaks that do not extend as far as ear opening, preorbital streak indistinct; occiput with light blotches; neck with light tan blotches forming indistinct cross-shaped marking bar flanked by a pair of dark blotches, light mid-dorsal marking discontinuous from occiput to tail base, seven light and seven dark bands from fore limb insertion to tail base; ventral surfaces immaculate; limbs with indistinct crossbars and a few lighter blotches; regenerated tail light brown with dark spots dorsally, light grey and tan ventrally. Variation (Fig. 9). Dorsal colour pattern of six or seven irregular light and dark crossbars on types; BNHS 2147 darker in ground colouration, without clear mid-dorsal markings; crown-shaped light marking on nape; regenerated portion of tail dirty brown; ventral surfaces dirty white, subcaudals dark with some lighter, and yellow spots. Both paratypes have a continuous series of 10 precloacal pores (Table 4). Distribution and Natural History. Cyrtodactylus kazirangaensis sp. nov. is known only from its type locality, which lies at the northern edge of the Mikir Hills. The entrance to Kaziranga National Park is less than 5 km away, and it is likely that this species also occurs in the park itself. The type series was collected together emerging from drainage holes in a concrete wall on the edge of a patch of degraded deciduous forest/plantations, facing a small stream, about an hour after sunset. Comparisons. Cyrtodactylus kazirangaensis sp. nov. is a member of the lowland clade and differs from other species by 14.7���19.7 % uncorrected genetic distance. Cyrtodactylus kazirangaensis sp. nov. can be diagnosed by the presence of 10 or 11 precloacal pores and absence of femoral pores on males from C. ayeyarwadyensis (10���28 PcP���PcFP), Cyrtodactylus guwahatiensis sp. nov. (26 PcFPs in a discontinuous series), and C. tripuraensis (29���37 PcFP). Cyrtodactylus kazirangaensis sp. nov. can be further diagnosed by having more ventral scales (37���43 MVSR) than C. ayeyarwadyensis (32���37 MVSR). Cyrtodactylus kazirangaensis sp. nov. is very similar morphologically to C. khasiensis from which it differs in the ear to eye distance being proportionately less than snout to eye distance (EE/SE 0.67���0.69 vs. 0.70���0.77), and ear to eye distance proportionately less than nostril to eye distance (EE/NE 0.81���0.88 vs. 0.90���1.01). Cyrtodactylus kazirangaensis sp. nov. is diagnosed against C. septentrionalis sp. nov. after its description. Major diagnostic characters for the new species and other regional congeners are summarized in Table 3. Remarks. Wood et al. (2012) included a specimen of C. ��� khasiensis ��� (MFA 50083) from ���Kaziranga, Assam ��� and the ND2 sequence (JX440543) was found to be 99% similar to the sequence (KM255170) of the paratype of Cyrtodactylus kazirangaensis sp. nov. in a BLAST search (results not shown), and thus is referable to this species. This sequence was also included in Brennan et al ���s (2017) phylogenetic study as C. ��� khasiensis ���., Published as part of Agarwal, Ishan, Mahony, Stephen, Giri, Varad B., Chaitanya, R. & Bauer, Aaron M., 2018, Six new Cyrtodactylus (Squamata: Gekkonidae) from northeast India, pp. 501-535 in Zootaxa 4524 (5) on pages 510-514, DOI: 10.11646/zootaxa.4524.5.1, http://zenodo.org/record/2610704, {"references":["Wood, P. L. Jr., Heinicke, M. P., Jackman, T. R. & Bauer, A. M. (2012) Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of diversification. Molecular Phylogenetics and Evolution, 65, 992 - 1003. https: // doi. org / 10.1016 / j. ympev. 2012.08.025","Brennan, I. G., Bauer, A. M., Tri, N., Wang, Y. Y., Wang, W. Z., Zhang, Y. P. & Murphy, R. W. (2017) Barcoding utility in a megadiverse, cross-continental genus: keeping pace with Cyrtodactylus geckos. Scientific Reports, 7 (5592), 1 - 11, Suppliment Info. https: // doi. org / 10.1038 / s 41598 - 017 - 05261 - 9"]}

Cyrtodactylus guwahatiensis sp. nov. Figs. 3���5. Holotype. Adult male (BNHS 2146, field number CES09/1127), collected from Hengrabari Road, Guwahati City, Kamrup Metropolitan district, Assam state, India (26.16193��N, 91.78298��E, elevation Diagnosis. Cyrtodactylus guwahatiensis sp. nov. can be distinguished from all congeners by: its moderate body size (SVL up to at least 67.8 mm); 10 supralabials; nine infralabials; 23 or 24 longitudinal rows of bluntly conical, feebly keeled dorsal tubercles; tubercles extending till third segment of tail; 35 paravertebral tubercles; ~34 ventral scales between ventrolateral folds; no precloacal groove; 26 precloacofemoral pores in a discontinuous series interspersed by 11 unpored scales; a row of enlarged scales above and below precloacal pore-bearing scales, slightly larger than pore-bearing scales; 16 total subdigital lamellae beneath toe IV of pes; subcaudal scalation of original tail without enlarged plates; dorsal pattern of 8���10 transverse rows of two or three pale buff blotches outlined by thick dark reticulations. Tail with alternating dark and lighter bands. Etymology. The specific epithet is a toponym for the type locality of the species, Guwahati, the largest city in Assam and northeast India. Description of holotype. Holotype in generally good preservation condition except folds on abdomen, head bent towards right, tail tip removed as tissue voucher, constriction just anterior to hind limb insertion due to overzealous tag tying. Adult male, SVL 67.8 mm. Head long (HL/SVL 0.26; HW/HL 0.67), dorsoventrally depressed (HD/HW 0.58), distinct from neck; loreal region somewhat inflated, interorbital area flat, canthus rostralis not prominent; snout short (SE/HL 0.43), twice as long as orbit diameter (ED/SE 0.51); scales on forehead, canthus rostralis and snout homogeneous, those on forehead smaller, roughly hexagonal, juxtaposed, weakly pointed; scales on interorbital and occipital region heterogeneous, granular, intermixed with larger, rounded, conical tubercles which are two to three times larger than adjacent granules, enlarged tubercles sparse on interorbital region and dense on occiput, single row of enlarged pointed tubercles on the eyelids, one or two scale rows from supraciliaries on each side. Eye small (ED/HL 0.22); pupil vertical with crenulated margins; supraciliaries small, mucronate, decreasing in size towards posterior and anterior end of orbit, largest about onethird the way from anterior edge of orbit; ear opening oval, obliquely oriented; eye to ear distance approximately half eye diameter (ED/SE 0.51). Rostral wider (2.7 mm) than deep (1.7 mm), partially divided dorsally by weakly developed rostral groove; single much enlarged supranasal on either side, separated by single small internasal, approximately same size as enlarged scales on snout; rostral in contact with supralabials I, nostrils, supranasals and an internasal; nostrils semicircular, laterally oriented, posterior half covered by nasal pad, each in broad contact with rostral and surrounded by supranasal, supralabial I, and 3���4 postnasals; three or four rows of scales separate orbit from supralabials; mental wider (2.3 mm) than long (2.0 mm), triangular, two well-developed postmentals on either side, inner pair in broad contact (1.2 mm) behind mental, twice the size (2.1 mm) of and separating outer pair (1.0 mm); inner postmentals bordered by mental, infralabial I, outer postmental and eight gular scales; outer postmental bordered by inner postmental, infralabials I and II, and five gular scales of dissimilar sizes on either side; eight supralabials to midorbital position and 10 supralabials to angle of jaw on each side, bordered by a row of large, flat, keeled, elongate scales; nine infralabials on each side, infralabials II to VI bordered by one or two rows of enlarged scales, increasing in size anteriorly. Body slender, short (TRL/SVL 0.47) with ventrolateral folds; dorsal scales heterogeneous, mostly rounded granules, intermixed with irregularly arranged, enlarged tubercles, bluntly conical and keeled throughout, becoming more conical and slightly smaller towards flanks, ventrolateral folds with single row of scattered enlarged, smooth tubercles; tubercles extend from occipital region posteriorly onto tail base; tubercles on nape smaller than those of dorsum; enlarged tubercles on tail flat, weakly pointed and feebly keeled; 23 or 24 dorsal tubercles across mid-dorsum; 35 paravertebral tubercles; ventral scales much larger than dorsals, smooth, cycloid, imbricate to subimbricate; slightly smaller in size under thighs, two rows of enlarged scales between precloacal pores and vent; ~34 ventral scales between ventrolateral tubercle rows; gular region with small granular scales throughout except a few anterior scales bordering the postmentals that are larger, juxtaposed. Eleven distinct porebearing precloacal scales in a continuous series, separated on each side by one unpored scale from a discontinuous series of smaller pore-bearing femoral scales; six femoral pores on the right and nine on the left with five and four unpored scales in each series, respectively; no precloacal groove; hemipenal bulge distinct. Fore and hind limbs slender, short (FL/SVL 0.14, CL/SVL 0.16); digits short, strongly inflected at each joint, all bearing robust recurved claws; five subdigital lamellae in basal series and nine in distal series with two nonlamellar granules at inflection (digit IV, right manus); seven basal and nine distal lamellae with three non-lamellar granules at inflection (digit IV, right pes), inter-digital webbing absent on manus, rudimentary between toes I to IV; relative length of digits: I Colouration in life (Fig. 3) Dorsal ground colour of body, limbs and tail dark-brown, head brown; top of head with indistinct light yellow blotches forming larger indistinct paired markings on occiput; labials lighter than dorsum of head; two light yellow narrow postorbital streaks not extending as far as ear opening; neck with pair of light tan spots, 8���10 transverse rows of two or three indistinct pale buff coloured blotches outlined by thick dark reticulations extending from above fore limb insertions to tail base, mid-dorsal line confluent with central light spots; limbs with indistinct crossbars and a few lighter blotches; eight dark and seven light caudal bands, dark bands approximately twice as wide as light bands; iris grey-silver with dark reticulations, pupil bordered by brown. Colouration in preservative (Figs. 4 & 5). Dorsal ground colour of body, limbs and tail light-brown, head brown; top of head with indistinct light blotches forming larger indistinct paired markings on occiput; labials similar in shade to dorsum of head; two light narrow postorbital streaks not extending as far as ear opening; neck with pair of light tan spots, 8���10 transverse rows of two or three indistinct pale buff coloured blotches outlined by thick dark reticulations extending from above fore limb insertions to tail base, mid-dorsal line confluent with central light spots; limbs with indistinct crossbars and a few lighter blotches; eight dark and seven light caudal bands, dark bands approximately twice as wide as light bands. Distribution and Natural History. This species is only known from a degraded moist deciduous forest on a small hill, completely surrounded by the urban sprawl of Guwahati, Assam. The holotype and only known specimen was collected near a seepage on a muddy wall along a hill road about 1.5 hours after sunset. Comparisons. Cyrtodactylus guwahatiensis sp. nov. is a member of the lowland clade and is sister to C. septentrionalis sp. nov., the clade also including C. ayeyarwadyensis, C. tripuraensis, C. khasiensis and C. kazirangaensis sp. nov. The p- distance analysis demonstrated that the ND2 gene of the new species is 11.3% divergent from that of C. septentrionalis sp. nov. and 13.0���19.7% divergent from others in the clade (Table 2). Cyrtodactylus guwahatiensis sp. nov. can be diagnosed by the total number of precloacofemoral pores (26 PcFP) on males from C. khasiensis (10���12 PcP) and C. tripuraensis (29���37 PcFP); further differs from C. tripuraensis by having slightly more DTR (23 or 24) and fewer MVSR (34) vs. (19��21 DTR, 35���43 MVSR). The new species differs in colouration from C. ayeyarwadyensis (no white punctuations or narrow band following dark dorsal markings) Cyrtodactylus guwahatiensis sp. nov. is diagnosed from C. kazirangaensis sp. nov. and C. septentrionalis sp. nov. after the descriptions of the latter two species. Major diagnostic characters for the new species and other regional congeners are summarized in Table 3., Published as part of Agarwal, Ishan, Mahony, Stephen, Giri, Varad B., Chaitanya, R. & Bauer, Aaron M., 2018, Six new Cyrtodactylus (Squamata: Gekkonidae) from northeast India, pp. 501-535 in Zootaxa 4524 (5) on pages 503-509, DOI: 10.11646/zootaxa.4524.5.1, http://zenodo.org/record/2610704

Cyrtodactylus montanus sp. nov. Figs. 9 and 15–17 Holotype. Adult female (BNHS 2235, field number CES09/1215), collected from near Phuldungsei village in the southern Jampui Hills, Kanchanpur subdivision, North Tripura district, Tripura state, India (23.82257°N, 92.26034°E, 850 m asl.) by Tarun Khichi, N.P.I. Das, Aniruddha Datta-Roy and Ishan Agarwal on 3 November 2010. Paratypes. Adult males (BNHS 2232–2234) and an adult female (BNHS 2231), bear the same collection data as the holotype. Etymology. The specific epithet is a nominative, masculine, singular, Latin adjective meaning “pertaining to a mountain” as this species is a member of the mountain clade, and is restricted to a mountainous region in northwestern Tripura. Diagnosis. Cyrtodactylus montanus sp. nov. can be distinguished from all congeners by its moderate body size (SVL to at least 78.2 mm); eight or 10 supralabials; 8–10 infralabials; 21–23 longitudinal rows of rounded, conical, feebly keeled dorsal tubercles; tubercles extending posteriorly to at least third tail segment; 37–43 paravertebral tubercles; 21–23 ventral scales between ventrolateral folds; no precloacal groove; 8–10 precloacal pores in males, no femoral pores; a row of enlarged scales posterior to precloacal pore-bearing scales, slightly larger than porebearing scales; 13–18 total subdigital lamellae beneath toe IV of pes; subcaudal scalation lacking enlarged median plates; dorsal colour pattern of thick dark reticulations enclosing lighter blotches; tail with alternating dark and lighter bands. Description of holotype. Holotype in good preservation condition: tail tip removed for tissue voucher, toe V of pes missing, constriction just above hind limb insertion due to overzealous tag tying, small abrasion on gular region, and folds of skin on dorsal surface of head, all artefacts of preservation and storage. Adult female, SVL 58.8 mm. Head long (HL/SVL 0.28), slightly wide (HW/HL 0.66), dorsoventrally depressed (HD/HW 0.58), distinct from neck; loreal region inflated, interorbital region flat, canthus rostralis not prominent; snout short (SE/ HL 0.39), twice as long as orbital diameter (ED/SE 0.57); scales on snout and canthus rostralis, granular, juxtaposed, homogenous in shape with scales on snout being slightly larger; scales on interorbital region, forehead and occipital region slightly smaller, granular, juxtaposed; forehead and occipital region interspersed with larger tubercles that are rounded, smooth and two to four times size of adjacent granules; dense enlarged tubercles slightly smaller on forehead and larger on occipital region. Eye small (ED/HL 0.22); pupil broad, vertical with crenulated margins; supraciliaries only slightly larger than adjacent scales, weakly pointed; upper eyelids covered with small granular scales, row of enlarged tubercles bordering supraciliaries absent. Ear opening oval, obliquely oriented, large; eye to ear distance less than twice diameter of eye (ED/EE 0.69). Rostral wider (2.3 mm) than deep (1.1 mm), partially divided dorsally by a weakly developed rostral groove; single enlarged rounded supranasal on either side, separated from each other by two rows of two smaller granular internasal scales; rostral in contact with supralabials I, nostrils, supranasals and two small internasals separating supranasals; nostrils oval, laterally oriented, posterior half covered by a distinct nasal pad, each in broad contact with rostral and surrounded by supranasal, supralabial I, and five small granular postnasal scales; five rows of granular scales between mid-orbit and supralabials; mental wider (1.8 mm) than deep (1.2 mm), triangular, two well-developed postmentals on either side, inner pair in broad contact (1.4 mm) behind mental, twice as large (1.8 mm) and separating outer pair from each other (0.9); each inner postmental bordered by mental, infralabial I, outer postmental and a row of two or three gular scales; outer postmentals bordered by inner postmental, infralabial I and II, a large, roughly hexagonal scale posteriorly and three gular scales that are twice size of other gular scales; seven supralabials at midorbital positionand 10 to angle of jaw, bordered by a row of flat, somewhat elongated scales slightly larger than their adjacent granules; eight infralabials on left side, ten on right side, infralabials I–V bordered by one or two rows of enlarged scales, anteriormost largest. Body slender (BW/TRL 0.39), short (TRL/SVL 0.49); dorsal scales heterogeneous, mostly rounded granules intermixed with enlarged tubercles that are irregularly arranged, rounded, feebly keeled to slightly conical, and more or less homogenous in shape and size throughout dorsum; an indistinct ventrolateral fold on either side comprises a single row of small conical tubercles; tubercles extend anteriorly onto eyelids and frontal region; tubercles on nape smaller than those of dorsum; 21 or 22 dorsal tubercles across mid-dorsum; 40 paravertebral tubercles; ventral scales much larger than dorsals, smooth, cycloid, subimbricate; slightly smaller in size under thighs; a row of enlarged subimbricate scales along posterior border of precloacal scales; 42 ventral scales between ventrolateral folds; gular region with much smaller granular scales throughout, except two rows bordering infralabials originating from posterior margin of outer postmentals, which are larger, flat and juxtaposed. A continuous series of eight enlarged precloacal scales, each with a shallow depression, no precloacal or femoral pores, no precloacal groove. Fore and hind limbs slender; forearm (FL/SVL 0.15) and tibia (CL/SVL 0.18) short; digits short, strongly inflected at each joint, all bearing robust recurved claws that are only slightly longer than their respective claw sheaths; four subdigital lamellae in basal series and eight in distal series with one non-lamellar granule at inflection (digit IV, right manus); six basal and 10 distal lamellae with one non-lamellar granule at inflection (digit IV, right pes); inter-digital webbing absent; relative length of digits: I Tail original with tip removed, somewhat rounded, slender, tapering, divided into indistinct segments; about ten rows of enlarged keeled sub-trihedral tubercles scattered on tail base; 2–5 conical tubercles at end of first to third segments; remaining dorsal caudal scales smooth, rounded, subimbricate, similar in size dorsally and on lateral aspect; subcaudal scales larger than dorsal, smooth, imbricate; no distinct mid-ventral series of enlarged subcaudals; three enlarged smooth post cloacal spurs on each side. Colouration in life. Not documented for holotype. Colouration in preservative (Figs. 16 & 17). Dorsal ground colour of head, body, limbs and tail brown; top of head with indistinct darker markings; labials darker than head dorsum with about one light brown blotch per labial; a discontinuous light postorbital streak extends to above ear opening; neck with two longitudinal discontinuous dark streaks; dorsum and dorsolateral aspect with about eight rows of four dark paravertebral blotches; limbs with light grey and tan spots; seven dark and light caudal bands, dark bands twice as wide as light bands; rest of ventral surfaces off-white, immaculate. Variation (Fig 9). Compared to the holotype, BNHS 2231 has much thicker dark dorsal markings, regenerated tail without spots, an inverted light chevron shaped marking on occiput flanked on either side by a similar coloured spot; postorbital streaks interrupted but meet on base of head; dark and light bands on tail of approximately equal width in some types. The male paratypes have a continuous series of 8–10 precloacal pores (Table 4), except BNHS 2233 which has four pores on each side separated by a median unpored scale. Distribution and Natural History. Cyrtodactylus montanus sp. nov. is known only from its type locality, in the Jampui Hills. We collected the type series from rocky roadside cuttings on a mountain-top in approximately one-half of an hour of searching after dark. This high elevation region has a small patch of montane evergreen forest, as opposed to deciduous forest at lower elevations where C. tripuraensis is distributed. A couple of additional populations were observed but not collected in other parts of the Jampui Hills. These populations are provisionally referred to this species due to overall morphological similarity with the type series. One female was found near the peak (~ 23.8098°N, 92.2610°E, ~ 920 m asl.) of Betalongchhip (or Thaidawr: highest peak in Tripura state) approximately 1 km south of Phuldungsei. The individual was located during dusk emerging from a tree hole at approximately 1.5 m above ground level within a forest patch. Two additional populations photographed but not collected were observed ~ 20 km north of the type locality near Vanghmun (~ 24.0055°N, 92.2817°E, ~ 600 m asl.). Eight adult individuals (three males and five females) were observed on an old brick wall after dusk on consecutive nights during the months of June to mid August, after which this population appeared to have dispersed from the wall, as none were observed on the wall during several visits in late August. Nearby the old brick wall was a large dense bamboo brake. Adults spent the day concealed within holes in the wall, emerging only after dusk where they were observed in the open, typically spaced> 1.5 m apart. This population had considerable variation in dorsal markings. Elsewhere near Vanghmun, a male and female were observed on a steep rocky roadside embankment about 1 h after night-fall. Three females observed during late August were gravid with a large pair of eggs (visible through the skin). Comparisons. Cyrtodactylus montanus sp. nov. is a member of the mountain clade and is sister to C. sp. Mizoram from which it differs by 10.1% uncorrected ND2 p- distance, separated from other members of the clade by p- distance of 19.0–21.5%. Cyrtodactylus montanus sp. nov. can be diagnosed by the presence of 8–10 precloacal pores in a single series (or separated medially by a single scale) in males only from C. gansi (16–29 PcFP) and Cyrtodactylus jaintiaensis sp. nov. (11–12 PcP on both males and females), also distinguished from the latter by its slightly smaller size (SVL up to 78.2 mm vs. up to 96.2 mm); and by the number of dorsal tubercle rows (21–23) from C. brevidactylus (27 DTR) and C. chrysopylos (16 DTR). Cyrtodactylus montanus sp. nov. is distinguished from Cyrtodactylus nagalandensis sp. nov. after its description. Major diagnostic characters for the new species and regional congeners are summarized in Table 3.

Cyrtodactylus jaintiaensis sp. nov. Figs. 9 and 12–14. Holotype. Adult female (BNHS 2248, field number CES09/1228), collected from the vicinity of Tyrshi Falls near Jowai town, West Jaintia Hills district, Meghalaya state, India (25.46956°N, 92.18313°E, 1350 m asl.) by Tarun Khichi, Aniruddha Datta-Roy and Ishan Agarwal on 15 November 2010. Paratypes. Adult males (BNHS 2246 & BNHS 2247), bear the same collection data as holotype. Etymology. The specific epithet is a toponym named after the type locality of the new species in the Jaintia Hills, West Jaintia Hills district, Meghalaya. Diagnosis. Cyrtodactylus jaintiaensis sp. nov. can be distinguished from all congeners by its large body size (SVL to at least 96.2 mm); eight or nine supralabials; nine or 10 infralabials; 19 or 20 longitudinal rows of rounded, conical, feebly keeled dorsal tubercles; tubercles extending to third tail segment; 30–34 paravertebral tubercles; 40–42 ventral scales between ventrolateral folds; no precloacal groove; 11 or 12 precloacal pores on males and females, no femoral pores; a row of enlarged scales posterior to precloacal pore-bearing scales, slightly larger than pore-bearing scales; 15–18 total subdigital lamellae beneath toe IV of pes; subcaudal scalation of original tail without enlarged subcaudals; dorsal pattern of indistinct light and dark blotches; tail with alternating dark and lighter bands. Description of holotype. Holotype in good preservation condition: tail tip removed for tissue voucher, constriction just anterior to hind limb insertion due to overzealous tag tying. Adult female, SVL 96.2 mm. Head long (HL/SVL 0.26), slightly wide (HW/HL 0.69), somewhat depressed (HD/HW 0.57), distinct from neck; loreal region somewhat inflated, interorbital region flat, canthus rostralis not prominent; snout short (SE/HL 0.42), twice as long as orbital diameter (ED/SE 0.54); scales on snout and canthus rostralis granular, juxtaposed, homogenous in shape with scales on snout larger; scales on interorbital region, forehead and occipital region smaller, granular, juxtaposed; forehead and occipital region interspersed with larger tubercles that are rounded, smooth and twice the size of adjacent granules, enlarged tubercles sparse on forehead, denser on occipital region. Eye small (ED/HL 0.23); pupil vertical with crenulated margins; supraciliaries similar in size and shape to adjacent granules, weakly pointed; dorsal surface of eyelids covered with small granular scales, row of enlarged tubercles bordering supraciliaries absent. Ear opening oval, obliquely oriented, large; eye to ear distance larger than diameter of eye (ED/EE 0.71). Rostral wider (3.6 mm) than deep (2.0 mm), partially divided dorsally by a weakly developed rostral groove; single much enlarged, rounded supranasals on either side, in contact with each other; rostral in contact with supralabial I, nostrils and supranasals; nostrils oval, laterally oriented, posterior half covered by an indistinct nasal pad, each in broad contact with rostral and surrounded by supranasal, supralabial I, and five smaller, granular postnasal scales; five rows of granular scales between mid-orbit and supralabials; mental wider (2.8 mm) than deep (2.2 mm), two well-developed postmentals on either side, inner pair in broad contact (1.6 mm) with each other behind mental, twice size (2.3 mm) of and separating outer pair (1.1 mm); each inner postmental bordered by mental, infralabial I, outer postmental and three gular scales; outer postmentals bordered by inner postmental, infralabials I and II, a large, roughly hexagonal scale posteriorly and three or four slightly enlarged gular scales; seven supralabials on each side at midorbital position; supralabials to angle of jaw, eight on right side and nine on left side, bordered by a row of flat, somewhat elongated scales slightly larger than their adjacent granules; infralabials, 10 on left side, infralabials II–VI bordered by two rows of enlarged scales, anteriormost largest. Body slender (BW/TRL 0.42), short (TRL/SVL 0.48); dorsal scales heterogeneous, mostly rounded granules, intermixed with irregularly arranged, enlarged tubercles, tubercles rounded, feebly keeled, flatter and more keeled towards vertebral region, more pointed toward flanks; indistinct ventrolateral fold on either side covered by a single row of enlarged smooth scales interspersed with much smaller granules; tubercles extend from occipital region to anterior half of remaining portion of tail; tubercles on nape smaller than those of dorsum; 20 dorsal tubercles across mid-dorsum; 34 paravertebral tubercles; ventral scales much larger than dorsals, smooth, cycloid, subimbricate; slightly smaller under thighs; two rows of enlarged subimbricate scales along posterior border of precloacal pores; 40 ventral scales between ventrolateral folds; gular region with much smaller granular scales throughout, except two rows which are twice as large, flat and juxtaposed, and run from posterior margin of outer postmentals, bordering infralabials; 12 minute precloacal pores in a continuous series; femoral pores absent; no precloacal groove. Fore and hind limbs slender; forearm (FL/SVL 0.14) and tibia (CL/SVL 0.16) short; digits short, strongly inflected at each joint, all bearing robust recurved claws that are slightly longer than their claw sheath; five subdigital lamellae in basal series and 11 in distal series (digit IV, right manus); seven basal and 10 distal lamellae with three non-lamellar granules at inflection (digit IV, right pes); slight inter-digital webbing between toes I–IV, absent between fingers; relative length of digits: I Tail tip absent, two thirds original, somewhat rounded, slender, tapering, divided into indistinct segments; seven rows of enlarged, rounded, feebly keeled tubercles scattered on tail base; four to six rounded, flattened tubercles at end of first to third segments; remaining dorsal caudal scales smooth, rounded, subimbricate, similar in size dorsally getting larger on lateral aspect; subcaudal scales larger, smooth, imbricate; no distinct median series; three enlarged smooth post cloacal spurs on each side. Colouration in life (for paratype, BNHS 2246, Fig 12). Dorsal ground colour of head, body, limbs and tail grey-brown, strongly mottled with tan, yellow and white; top of head strongly mottled toward occiput; labials lighter than head dorsum and with some yellow streaks; a light yellow narrow postorbital streak that does not extend as far as ear opening; neck and back with black light grey spots forming a reticulate pattern of thick markings, numerous light and dark brown tubercles on dorsum; limbs with light tan spots; ten dark and nine light caudal bands on complete original tail, dark bands more than four times as wide as light bands; rest of ventral surfaces immaculate; iris green-grey with dark reticulations, pupil bordered by orange-red. Colouration in preservative (Figs. 13 & 14). Dorsal ground colour of head, body, limbs and tail light-brown; top of head strongly mottled with fine dark markings; labials similar to top of head bearing scattered darker spots; a light narrow postorbital streak does not extend to ear opening; nape and dorsum bear black and light-grey spots, numerous light and dark brown tubercles on dorsum; limbs with slightly darker reticulate markings; four dark and three light bands on original portion of tail, darker bands wider than light, regenerated portion colouration and markings similar to limbs; rest of ventral surfaces dirty white. Variation. The two adult male paratypes have 11 or 12 distinct precloacal pores and a distinct hemipenal bulge, and are similar in colour pattern to the holotype except there are approximately eight dark and nine light caudal bands on BNHS 2247, and ten dark and nine light caudal bands on BNHS 2246. Variation in scale counts and morphometrics is summarized in Table 4. Distribution and Natural History. The species is only known from the vicinity of Tyrshi Falls, in the Jaintia Hills. The geckos were spotted at night by eye-shine on vertical rock faces. The landscape is made up of patches of forest in an agricultural matrix. Comparisons. Cyrtodactylus jaintiaensis sp. nov. is a member of the mountain clade and is the poorly supported sister lineage to C. montanus sp. nov. and C. sp. Mizoram, and is separated from members of the clade by pairwise uncorrected genetic distance of approximately 18.7–21.4%. Cyrtodactylus jaintiaensis sp. nov. can be diagnosed by its large body size (SVL up to 96 mm), and the presence of 11 or 12 precloacal pores in a single series in males from C. chrysopylos (SVL up to 79 mm, enlarged pore-bearing post-precloacal scales in males), and C. gansi (SVL up to 63 mm, 16–29 PcFP). Cyrtodactylus jaintiaensis sp. nov. can be diagnosed by its reticulate dorsal pattern from C. brevidactylus (3–dark dorsal blotches between nape and sacrum). The differentiation of C. jaintiaensis sp. nov. from C. montanus sp. nov., and C. nagalandensis sp. nov. is detailed following their descriptions. Major diagnostic characters for the new species and regional congeners are summarized in Table 3.

Cyrtodactylus nagalandensis sp. nov. Figs. 9 and 18–20. Holotype. Adult female (BNHS 2253, field number CES09/1233) collected from near Khonoma village, Kohima district, Nagaland state, India (25.65818°N, 94.02142°E, 1450 m asl.) by Ishan Agarwal on 23 October 2009. Paratype. Adult female (BNHS 2254) bears the same collection information as holotype. Etymology. This is the first endemic gecko from Nagaland, and the specific epithet is a toponym for the state. Diagnosis. Cyrtodactylus nagalandensis sp. nov. can be distinguished from all congeners by its moderate body size (SVL to at least 72.0 mm); 10–12 supralabials; 8–10 infralabials; 16–18 longitudinal rows of rounded, conical, feebly keeled dorsal tubercles; tubercles extending posteriorly to third tail segment; 35–37 paravertebral tubercles; 34 or 35 ventral scales between ventrolateral folds; no precloacal groove; six depressions in precloacal series but no precloacal or femoral pores on females, male condition unknown; 16 total subdigital lamellae beneath toe IV of pes; subcaudal scalation of original tail with paired enlarged subcaudals not forming median plates; dorsal pattern of indistinct light and dark blotches forming longitudinal markings on neck and forebody; tail with alternating dark and lighter bands. Description of holotype. Holotype in good preservation condition: tail tip removed for tissue voucher, tail curved towards left, with folds of skin along either side of ventrolateral aspect; constriction around midbody due to overzealous tag tying, toes on left and right pes recurved; all artifacts of preservation. Adult female, SVL 72.0 mm. Head long (HL/SVL 0.30), slightly wide (HW/HL 0.70), dorsoventrally depressed (HD/HW 0.59), distinct from neck; loreal region inflated, interorbital region flat, canthus rostralis not prominent; snout short (SE/HL 0.40), twice as long as orbital diameter (ED/SE 0.50); scales on snout and canthus rostralis granular, juxtaposed, homogenous in shape, with scales on snout being slightly larger; scales on interorbital region, forehead and occipital region slightly smaller, granular, juxtaposed; forehead and occipital region interspersed with larger tubercles that are rounded, smooth and two to four times the size of adjacent granules; enlarged tubercles smallest on anterior and central parietal regions, increasing in size laterally and posteriorly. Eye small (ED/HL 0.20); pupil vertical with crenulated margins; supraciliaries roughly triangular, pointed, decreasing in size towards posterior and anterior ends of orbit, largest about one-third the way from anterior edge of orbit; dorsal surface of eyelid primarily covered with small granular scales except a single row of granular scales bordering anterior supraciliaries are enlarged relative to adjacent granular scales. Ear opening oval, obliquely oriented; eye to ear distance almost twice diameter of eye (ED/EE 0.60). Rostral wider (2.7 mm) than deep (1.8 mm), partially divided dorsally by a weakly developed rostral groove; single enlarged rounded supranasals on either side, separated by a smaller, roughly hexagonal internasal scale which is about three times size of granules on snout; rostral in contact with supralabials I, nostrils, supranasals and a single internasal; nostrils oval, laterally oriented, posterior half covered by a conspicuous nasal pad, each in broad contact with rostral and surrounded by supranasal, supralabial I, and four smaller granular postnasal scales; one to three rows of granular scales separate orbit from supralabials; mental wider (3.1 mm) than long (1.8 mm), triangular, two pairs of well-developed postmentals on either side, inner pair in broad contact (0.9 mm) with each other posteriorly behind mental, inner pair larger (1.8 mm) than and separating outer pair (1.2); each inner postmental bordered by mental, infralabial I, outer postmental and a row of four or five gular scales of similar size to other gulars; outer postmentals each bordered by inner postmental, infralabials I and II, a large, roughly hexagonal scale posteriorly and four or five gular scales; eight supralabials to midorbital position and 10 to angle of jaw on each side, bordered by a row of flat, somewhat elongated scales slightly larger than their adjacent granules; infralabials, 10 on right side and eight on left side, infralabials II to V bordered by a row of enlarged scales, anteriormost largest. Body slender (BW/TRL 0.4), short (TRL/SVL 0.50); dorsal scales heterogeneous with mostly rounded granules, intermixed with irregularly arranged enlarged tubercles, rounded, conical, feebly keeled, decreasing in size towards vertebral region; tubercles extend anteriorly to interorbital region; tubercles on nape smaller than those of dorsum; dorsal tubercles in 16 or 17 longitudinal rows at mid-dorsum; 35 paravertebral tubercles; a distinct ventrolateral fold on either side covered by a series of slightly enlarged, flat, granular scales; ventral scales much larger than dorsals, smooth, cycloid, subimbricate; a row of enlarged subimbricate scales along posterior border of precloacal pores; 36 ventral scales between ventrolateral folds; gular region with much smaller, granular scales throughout except for one or two rows originating from posterior margin of outer postmentals bordering infralabials, which are larger, flat and juxtaposed. A continuous series of six enlarged precloacal scales each with a shallow depression, precloacal and femoral pores absent, no precloacal groove. Fore and hind limbs slender; forearm (FL/SVL 0.20) and tibia (CL/SVL 0.20) short; digits short, strongly inflected at each joint, all bearing robust recurved claws that are slightly longer than their associated claw sheath; five subdigital lamellae in basal series and seven in distal series with four non-lamellar granules at inflection (digit IV, right manus); five basal and 11 distal lamellae with one non-lamellar granule at inflection (digit IV, right pes); slight inter-digital webbing between toes I–III, absent between fingers; relative length of digits: I Tail original with tip removed, somewhat rounded, slender, tapering, divided into indistinct segments; eight rows of enlarged, sub-trihedral, strongly keeled tubercles on tail base; approximately four rounded, flattened tubercles form a transverse row at posterior border of each of first three segments; remaining dorsal caudal scales smooth, rounded, subimbricate, similar in size dorsally getting slightly larger on lateral aspect; subcaudal scales larger, smooth, imbricate; a distinct mid-ventral series comprising of two large scales; enlarged smooth post cloacal spurs, three on left side and two on right side. Colouration in life (Fig. 18). Dorsal ground colour of head, body, limbs and tail buff-brown; top of head with numerous dark spots; labials strongly suffused with yellow; a dark postorbital streak that extends beyond ear opening; neck and back with black spots that form longitudinal markings on neck and forebody; limbs with indistinct darker bars and spots; 11 dark and 10 light caudal bands, dark bands more than twice as wide as light bands; rest of ventral surfaces immaculate; iris green-grey with dark reticulations, pupil bordered by orange-red Colouration in preservative (Figs. 19 & 20). Dorsal ground colour of head, body, limbs and tail buff-brown; top of head with numerous dark spots; labials suffused with lighter markings; a dark postorbital streak that extends beyond ear opening; neck and back with black spots that form longitudinal markings on neck and forebody; limbs with indistinct darker bars and spots; 11 dark and 10 light caudal bands, dark bands more than twice as wide as light bands; rest of ventral surfaces immaculate except some mottling toward flanks. Variation (Fig 9). Female paratype (BNHS 2254) similar to holotype in general colouration but with less distinct longitudinal markings on neck; 10 dark bands on tail. Refer to Table 4 for further variation. Distribution and Natural History. Cyrtodactylus nagalandensis sp. nov. is known only from the vicinity of Khonoma village, in the Naga Hills. The type series was collected from a road cutting after dark. We only saw two specimens of this species in two nights of searching. The surrounding forest habitats were mainly secondary forest. Comparisons. Cyrtodactylus nagalandensis sp. nov. is a member of the mountain clade and is poorly supported as sister to the other Indian species in this clade, with uncorrected p- distance between the new species and other members of the clade 18.0–21.7%. Cyrtodactylus nagalandensis sp. nov. can be diagnosed by the number of ventral scales (36 or 37 MVSR) from C. brevidactylus (45 MVSR), and Cyrtodactylus jaintiaensis sp. nov. (40–42 MVSR); by having fewer dorsal tubercle rows (16–18) from C. gansi (20–25 DTR) and Cyrtodactylus montanus sp. nov. (21–23 DTR); by its irregular dorsal colour pattern from C. chrysopylos (dorsal pattern of alternating brown and white bands). Major diagnostic characters for the new species and regional congeners are summarized in Table 3.

Cyrtodactylus septentrionalis sp. nov. Figs. 9���11. Holotype. Adult female (BNHS 1990), collected from near Abhayapuri town, Bongaigaon district, Assam state, India (26.345417��N, 90.677875��E, 40 m asl.) by Varad Giri and Kedar Bhide on 13 November 2009. Paratype. Adult male (BNHS 1989), bears the same collection details as holotype. Etymology. The specific epithet is a nominative, masculine, singular, Latin adjective meaning ���northern���, as this species is the only known member of the lowland clade (Agarwal et al. 2014) that is found north of the Brahmaputra River, the other five known species of the clade are found south of the Brahmaputra River. Diagnosis. Cyrtodactylus septentrionalis sp. nov. can be distinguished from all congeners by its moderate body size (SVL to at least 65.2 mm); nine or 10 supralabials; nine infralabials; 23 or 24 longitudinal rows of bluntly conical, feebly keeled dorsal tubercles; tubercles extend onto first three tail segments; 38���42 paravertebral tubercles; 35���38 ventral scales between ventrolateral folds; no precloacal groove; 14 fine precloacal pores and no femoral pores on males; 15���20 total subdigital lamellae beneath toe IV of pes; subcaudal scalation of original tail without enlarged plates; dorsal colour pattern of 6���9 rows of paired dark blotches, tail with alternating dark and lighter bands. Description of holotype. Holotype generally in good preservation condition: trunk curved to left, folds of skin along either side of ventrolateral aspect, constriction just anterior to hind limb insertion due to overzealous tag tying. Adult female, SVL 65.2 mm. Head long (HL/SVL 0.30), slightly wide (HW/HL 0.70), not depressed (HD/ HW 0.63), distinct from neck; loreal region somewhat inflated, interorbital region flat, canthus rostralis not prominent; snout short (SE/HL 0.50), twice as long as orbital diameter (ED/SE 0.50); scales on snout and canthus rostralis granular, juxtaposed, homogenous in shape with scales on dorsum of snout being slightly larger; scales on interorbital region, forehead and occipital region slightly smaller, granular, juxtaposed; forehead sparsely and occipital region densely interspersed with larger tubercles that are rounded, smooth, and two times the size of adjacent granules. Eye small (ED/HL 0.20); pupil vertical with crenulated margins; supraciliaries flat, smooth, roughly triangular, decreasing in size towards posterior and anterior ends of orbit, largest about one-third the way from anterior edge of orbit; a single row of enlarged tubercles bordering supraciliaries present on dorsal surface of upper eyelid. Ear opening oval, obliquely oriented, large. Eye to ear distance slightly larger than diameter of eye (ED/EE 0.70). Rostral wider (2.7 mm) than deep (1.7 mm), divided dorsally by a weakly developed rostral groove; single much enlarged rounded supranasal on either side, separated by two rows of three granular scales; rostral in contact with supralabials I, nostrils, supranasals and three scales separating supranasals; nostrils oval, laterally oriented, posterior half covered by a conspicuous nasal pad, each nostril in broad contact with rostral and surrounded by internasal, supralabial I, and four smaller, granular postnasal scales; three rows of granular scales separate orbit from supralabials; mental wider (2.3 mm) than deep (2.0 mm) triangular, two well-developed pairs of postmentals on either side, inner pair in broad contact (1.2 mm) behind mental, twice the size (2.1 mm) of and separating outer pair (1.0 mm); each inner postmental bordered by mental, infralabial I, outer postmental and three gular scales, outermost of which is largest; outer postmentals bordered by inner postmentals, infralabials I and II, a large, roughly hexagonal scale posteriorly and four gular scales; seven supralabials to midorbital position on each side; supralabials to angle of jaw, nine on right side and 10 on left side, bordered by a row of flat, somewhat elongated scales slightly larger than adjacent granular scales; nine infralabials on each side, infralabials II���V bordered by one or two rows of enlarged scales, anteriormost largest. Body slender (BW/TRL 0.4), short (TRL/SVL 0.40); dorsal scales heterogeneous with mostly rounded granules intermixed with irregularly arranged, enlarged tubercles that are rounded, slightly conical, feebly keeled; a distinct ventrolateral fold on either side with enlarged, rounded, flat tubercles interspersed by one or two slightly enlarged, flat, juxtaposed scales; tubercles extend from forehead posteriorly onto base of tail; tubercles on nape smaller than those of dorsum; 23 or 24 dorsal tubercles across mid-dorsum; 38 paravertebral tubercles; ventral scales much larger than dorsals, smooth, cycloid, juxtaposed; slightly smaller in size under thighs; a row of slightly enlarged, flat, subimbricate scales along posterior border of precloacal scales; 36���38 ventral scales between ventrolateral folds; gular region with small granular scales throughout, except for one or two rows originating from posterior margin of outer postmentals, bordering infralabials, which are larger, flat, juxtaposed. A continuous series of 14 indistinct depressions in precloacal scales, no pores, no precloacal groove. Fore and hind limbs slender; forearm (FL/SVL 0.20) and tibia (CL/SVL 0.20) short; digits short, strongly inflected at each joint, all bearing robust recurved claws that are slightly longer than claw sheath; six subdigital lamellae in basal series and nine in distal series with three non-lamellar granules at inflection (digit IV, right manus); five basal and 10 distal lamellae with four non-lamellar granules at inflection (digit IV, right pes); slight inter-digital webbing between toes I���III, absent between fingers; relative length of digits: I Colouration in life. Not documented. Colouration in preservative (Figs. 10 & 11). Dorsal ground colour of body, head, limbs and tail light brown; top of head scattered with few dark brown markings including longitudinal streak extending from occiput to nape; labials pale cream with fine black stippling; dark postorbital streak flanked by lighter markings that extend to ear openings, dark preorbital streak present extending to nostrils; neck with three longitudinal dark markings; dorsal pattern of six or seven pairs of dark elongated blotches from fore limb insertion to tail base; ventral surfaces immaculate; limbs with dark blotches; original portion of tail with eight dark and light markings that circle tail; regenerated portion of tail light brown, with a few dark spots. Variation (Fig. 9). BNHS 1989 is a male with 13 fine precloacal pores and nine pairs of dark dorsal markings; six dark and light markings on incomplete but original tail; broadly agreeing in scalation with the holotype (Table 4). Distribution and Natural History. Cyrtodactylus septentrionalis sp. nov. is only known from the vicinity of Abhayapuri town, close to the base of Bamun Gaon Pahar (a small hill). Nine individuals of this species were observed during a single night of search. They were found on open rocks along a small stream running through degraded forest near a human settlement. Comparisons. Cyrtodactylus septentrionalis sp. nov. is a member of the lowland clade and differs from other members of the clade by 11.3���18.6 % uncorrected genetic distance. Cyrtodactylus septentrionalis sp. nov. can be diagnosed by the presence of 14 precloacal pores and the absence of femoral pores on males from C. ayeyarwadyensis (10���28 PcP���PcFP), Cyrtodactylus guwahatiensis sp. nov. (26 PcFP in discontinuous series), Cyrtodactylus kazirangaensis sp. nov. (10 or 11 PcP), C. khasiensis (10���12 PcP), and C. tripuraensis (29���37 PcFP). Cyrtodactylus septentrionalis sp. nov. further differs from C. khasiensis in having more dorsal tubercle rows (23 or 24 vs. 19���23). Major diagnostic characters for the new species and regional congeners are summarized in Table 3., Published as part of Agarwal, Ishan, Mahony, Stephen, Giri, Varad B., Chaitanya, R. & Bauer, Aaron M., 2018, Six new Cyrtodactylus (Squamata: Gekkonidae) from northeast India, pp. 501-535 in Zootaxa 4524 (5) on pages 514-517, DOI: 10.11646/zootaxa.4524.5.1, http://zenodo.org/record/2610704, {"references":["Agarwal, I., Bauer, A. M., Jackman, T. R. & Karanth, K. P. (2014) Insights into Himalayan biogeography from geckos: a molecular phylogeny of Cyrtodactylus (Squamata: Gekkonidae). Molecular Phylogenetics and Evolution, 80, 145 - 155. https: // doi. org / 10.1016 / j. ympev. 2014.07.018"]}

Agarwal, Ishan, Khandekar, Akshay, Ramakrishnan, Uma, Vyas, Raju, Giri, Varad B. (2018): Two new species of the Ophisops microlepis (Squamata: Lacertidae) complex from northwestern India with a key to Indian Ophisops. Journal of Natural History 52 (13-16): 819-847, DOI: 10.1080/00222933.2018.1436203, URL: http://dx.doi.org/10.1080/00222933.2018.1436203

Hemidactylus vanam sp.nov. Figs. 1–10 Hemidactylus maculatus Bhupathy & Sathishkumar 2012 Holotype. NCBS-AU158, adult male; collected en-route to the High Wavy Mountains, Tamil Nadu, India (9.7808°N; 77.4427°E, 610 m asl) on 21 November 2016. Collected by R. Chaitanya, Akshay Khandekar and Vanasundara Pandian. Paratypes. NCBS-AU159, sub-adult female, NCBS-AU160, NCBS-AU161, adult females; ESV106, ESV107, adult males, ESV108, adult female; BNHS 2328 sub-adult female, BNHS 2329 adult female; ZSI/ WGRC /IR/V.NO 2634, adult male, ZSI/ WGRC /IR/V.NO 2635, adult female. Collection data same as holotype. ZSI/ WGRC /IR/V.NO 2635, collected near Suruli falls, Theni district (9.6563° N; 77.3061° E, 430 m asl). Diagnosis. A large sized Hemidactylus, SVL averaging 88.24 ± 16 mm to a maximum of at least 112.2 mm (n=9). Dorsal pholidosis heterogeneous, composed of roughly circular, striated, granular scales intermixed with much enlarged, fairly regularly arranged longitudinal rows of 17–19 strongly keeled, striated tubercles that are heterogeneous in shape and size, extending from occipital region to tail (Fig. 1); enlarged tubercles on the two most medial parasagittal rows are small, flat, strongly keeled and rounded gradually increasing in size and becoming conical towards flanks, last two to three rows on flanks are smaller and strongly conical. Two well-developed pairs of postmentals, inner pair much larger than the outer and in broad contact with each other behind the mental. Ventrolateral folds distinct. About 34–40 scale rows across the venter. All digits with enlarged scansors, lamellae in straight transverse series, all divided except the apical and a few basal that are undivided, 10–12 (manus and pes) lamellae beneath fourth digit and 9–10 (manus) and 8–9 (pes) beneath first digit. Femoral pores on each side 17– 22, separated by 10–11 poreless scales in males. Dorsal coloration pale-brown with transversely arranged saddleshaped markings running from the occiput to the sacrum that are of a darker shade. The large size (up to 112.2 mm SVL) of Hemidactylus vanam sp. nov. can be used to easily diagnose it from most other congeners from the Indian subcontinent and Southeast Asia, including H. garnotii Duméril & Bibron, H. platyurus (Schneider), H. aquilonius McMahan & Zug, H. scabriceps (Annandale), H. imbricatus Bauer, Giri, Greenbaum, Jackman, Dharne & Shouche, H. gracilis Blanford, H. reticulatus Beddome, H. albofasciatus Grandison & Soman, H. sataraensis Giri & Bauer, H. brookii Gray, H. gujaratensis Giri, Bauer, Vyas & Patil, H. frenatus Schlegel, H. persicus Anderson, H. robustus Heyden, H. parvimaculatus Deraniyagala, H. treutleri Mahony, H. gleadowi Murray, H. kushmorensis Murray, H. murrayi Gleadow, H. chipkali Mirza & Raju, H. triedrus (Daudin), H. subtriedrus Jerdon, H. lankae Deraniyagala, H. depressus Gray, H. pieresii Kelaart, H. leschenaultii Duméril & Bibron, and H. flaviviridis Rüppel all of which reach maximum sizes of up to 90 mm SVL. Hemidactylus vanam sp. nov. shares its large adult size (adult SVL upto 112.2 mm) only with H. giganteus Stoliczka, H. aaronbaueri Giri, H. yajurvedi Murthy, Bauer, Agarwal, Lajmi & Giri, H. hemchandrai Dandge & Tiple, H. prashadi Smith, H. hunae Deraniyagala, H. graniticolus Agarwal, Giri & Bauer, H. maculatus Duméril & Bibron, H. acanthopholis, H. kangerensis Mirza, Bhosale & Patil and H. sushilduttai Giri, Bauer, Mohapatra, Srinivasulu & Agarwal. Hemidactylus giganteus differs from H. vanam sp. nov. in the complete absence of enlarged dorsal tubercles while H. aaronbaueri, H. yajurvedi and H. hemachandrai have slightly enlarged, rounded, weakly-keeled tubercles at midbody. Hemidactylus prashadi, H. hunae, H. graniticolus H. kangerensis and H. sushilduttai differ from the new species in dorsal pholidosis, number of femoral pores and the number of poreless scales separating them (differing character states indicated in parenthesis): Hemidactylus prashadi (weakly keeled, sub-trihedral, slightly enlarged dorsal tubercles arranged in 14–16 rows; 17–20 femoral pores separated by 3 poreless scales), H. hunae (keeled, sub-trihedral, slightly enlarged dorsal tubercles arranged in 16– 20 rows; 22–24 femoral pores separated by 3–6 poreless scales), H. graniticolus (keeled, sub-trihedral, slightly enlarged dorsal tubercles arranged in 16–18 rows; 23–28 femoral pores separated by 1–3 poreless scales), H. kangerensis (keeled, trihedral, enlarged dorsal tubercles arranged in 18–20 rows; 18–21 femoral pores separated by four poreless scales), H. sushilduttai (keeled, trihedral, enlarged dorsal tubercles arranged in 16–17 rows; 19–24 femoral pores separated by four poreless scales. Based on size, dorsal pholidosis and general colouration, Hemidactylus vanam sp. nov. is most similar to H. maculatus and H. acanthopholis but can be diagnosed on the basis of the following combination of characters: dorsal pholidosis, femoral pores and the number of poreless scales separating them (differing character states indicated in parenthesis): H. maculatus (enlarged tubercles trihedral throughout, parasagittal rows slightly smaller; 16–19 femoral pores separated by 5–9 poreless scales ), H. acanthopholis (enlarged tubercles trihedral, parasagittal rows slightly smaller, last few rows on flanks, slightly conical; 19–21 femoral pores separated by 13–14 poreless scales) (Fig. 6). Additionally, H. vanam sp. nov. can be diagnosed from H. maculatus and H. acanthopholis by the presence of a dense aggregation of enlarged, strongly conical, striated tubercles on dorsal aspect of thigh (enlarged tubercles on dorsal aspect of thigh sparse in H. maculatus and H. acanthopholis) (Fig. 3). Description. The holotype is in good condition except for loss of skin on a small triangular patch on the neck, body is dorsoventrally flattened, ventrolateral fold is more pronounced on the left, and second toe on right pes curved upwards—all artefacts of preservation. Tail is regenerated and slightly curved towards the left. Hemipenes everted, exposed and seen on both sides when viewed dorsally (Fig. 1). Adult male, SVL 108.8 mm. Head short (HL/SVL ratio 0.27), slightly elongate (HW/HL ratio 0.76), not strongly depressed (HH/HL ratio 0.46), distinct from neck (Fig. 4). Loreal region slightly inflated, canthus rostralis indistinct (Fig. 4B). Snout short (SE/HL ratio 0.40); longer than eye diameter (OD/SE ratio 0.61); scales on snout, canthus rostralis, forehead and inter-orbital region heterogenous, mostly granular and rounded; scales on the snout and canthus rostralis much larger than those on occipital, forehead and inter-orbital regions (Fig. 4A). Eye small (OD/HL ratio 0.24); pupil vertical with crenulated margins; supraciliaries small, mucronate, gradually increasing in size towards front of the orbit, those at the anterior end of orbit larger. Ear opening roughly elliptical (greatest diameter 3.2 mm); eye to ear distance slightly greater than diameter of eye (EE/OD ratio 1.8). Rostral wider than deep (RL/RW ratio 0.65), incompletely divided dorsally by a weakly developed rostral groove; two internasals, enlarged and separated by a single scale of similar size; one supranasal on each side, much smaller than internasals; three diminutive postnasals on each side; rostral in contact with nostril, supralabial I, internasals and the small scale separating the internasals; nostrils small (0.8 mm), oval shaped; nasal surrounded by supranasal, internasal, rostral, supralabial I and three postnasals on either side; 6–7 rows of scales separate orbit from supralabials. Mental triangular; two well-developed postmentals, the inner pair slightly shorter (4.2 mm) than the mental (4.3 mm), and in strong contact with each other (1.6 mm) behind mental, outer pair smaller (2.4 mm) than the inner pair and separated from each other by inner pair (Fig. 4C). Inner postmental bordered by mental, infralabial I and II, outer postmental and five small gular scales each; outer postmental bordered by infralabial II and III (barely touching), inner postmental, and 6 (right)—7(left) gular scales increasing in size laterally, outer-most of which is much enlarged and continues as a single row of enlarged scales below the infralabials. Infralabials bordered by a single row of enlarged scales; 2 to 3 rows of scales below infralabials IV to VIII are enlarged and weakly imbricate. Supralabials (to midorbital position) 8 (right)—9 (left); supralabials (to angle of jaw) 13 (right)—13 (left); infralabials (to angle of jaw) 10 (right)—10 (left). Body relatively stout, not elongate (TRL/SVL ratio 0.41), ventrolateral folds indistinct. Dorsal pholidosis heterogeneous, composed of roughly circular, striated, granular scales intermixed with enlarged, fairly regularly arranged longitudinal rows of 17–18 strongly keeled, striated tubercles, extending from occipital region to tail that are heterogeneous in shape and size; enlarged tubercles on the two most medial parasagittal rows smaller, flatter and more rounded, increasing in size and becoming conical at the flanks; the last two rows on flanks slightly larger than the medial parasagittal rows and strongly conical; each enlarged tubercle surrounded by a rosette of 13–16 small granules with 1–5 granules between two longitudinally adjacent enlarged tubercles; enlarged tubercles on nape, shoulder, smaller and pointed, those on occipital, temporal region still smaller. Ventral scales larger than dorsal granular scales, smooth, imbricate, slightly larger on precloacal and femoral region than on chest and abdominal region; midbody scale rows across belly 36–39; gular region with much smaller, juxtaposed scales, anterior gular scales being slightly larger than the rest. Scales on palm and sole smooth; on palm sub-imbricate, rounded and those on sole imbricate and weakly pointed; scales on dorsal aspect of upper arm much larger than granules on dorsum, sub-imbricate and weakly keeled; dorsal aspect of forearm with smaller, striated, granular scales, intermixed with a few enlarged, striated, conical tubercles; scales on dorsal part of thigh and shank circular, striated, intermixed with enlarged, striated, conical tubercles, which are larger on thigh compared to shank; anterior aspect of thigh with flatter tubercles; posterior aspect with a congregation of enlarged, strongly conical, striated tubercles. Fore and hind limbs relatively short, stout; forearm short (FL/SVL ratio 0.10); tibia short (CL/SVL ratio 0.20); digits moderately short, strongly clawed; all digits of manus and digits I–IV of pes indistinctly webbed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad, half or more than half as long as associated toepad; scansors beneath each toe in a straight transverse series, divided except for distal and three to four basal scansors on digit I and one or two in other digits: 10-11-11-11-10 (left manus), 10-11-11-11-11 (right manus) (Fig. 5A), 9-11-11-11-10 (left pes), 9-11-11-12-12 (right pes) (Fig. 5 B). Relative length of digits (measurements in mm in parentheses): III (7.1)> IV (6.8) = V (6.8)> II (6.7)> I (5.8) (left manus); II (7.4)> III (7.3)> V (6.9)> IV (7.2)> I (5.8) (left pes). Tail with first segment intact, rest regenerated, depressed, flat beneath, verticillate with a distinct median furrow; length of the partially regenerated tail less than snout-vent length (TL/SVL ratio 0.76); first segment of tail covered above with small granules that are heterogeneous in shape and size, juxtaposed, striated, interspersed with 11 enlarged, rounded, strongly keeled, striated, posteriorly pointed and flattened tubercles; subcaudal scales on first tail segment larger, imbricate, weakly pointed. Nineteen femoral pores on the left and 21on the right separated medially by a diastema of 10 poreless scales (Fig. 6). Colouration (in preservative). Dorsum uniformly grayish-brown interspersed with slightly darker blotches in the snout, forehead and mid-vertebral regions (Fig. 1). Darker blotches on dorsal aspect of forelimb and hindlimb. Mid-vertebral tubercle rows mostly dark brown, those on flanks a lighter shade of brown. Crown of head with scattered vague dark-brown markings. Anterior supralabials dark-brown, getting paler towards the posterior. Infralabials uniformly mid-brown. Limbs mid-brown, with vague irregular dark markings that are more distinct on the forelimbs than the hind limbs, alternating pale and dark interspaces distally, especially on metapodial segments. First segment of original tail grayish-brown, interspersed with vague darker markings on the tubercles. Regenerated portion of tail, a lighter shade of gray interspersed with inconsistent darker markings. Venter cream with scattered patches of diffusely pigmented scales on head. A darker patch of scales in the chest region which is brownish-red spreads onto the forearms; palms and soles grayish. Ventral surface of tail pale, with scattered midbrown speckling throughout. Colouration (in life) (based on photographs of an uncollected topotype) Dorsal markings are much more evident in life (Fig. 7). Dorsum pale-brown with no distinct vertebral stripe. Head dorsum pale-brown with a mottling of white and darker-brown spots in the forehead region. Snout light brown, with a scattering of white blotches. A coffee coloured, roughly L-shaped stripe runs from the post orbital region to the forelimb insertion. Four saddle-shaped markings running from the occiput to the sacrum are a darker shade of brown. Interspaces between saddles are similar in colouration to the rest of the dorsum. Most tubercles on the saddle-shaped markings, of a much darker shade of brown than the rest of the body. Tail distinctly banded, with alternating light and dark bands similar to trunk dorsum; darker bands slightly wider than pale bands on tail. Mottled pattern on limb bases with distinct banding distally. Iris marbled, light olive-green, suffused with prominent rust-brown venation; pupil black with crenulated margins that are a lighter shade of olive-green. Etymology. The specific epithet is a noun in apposition honouring Vanam (pronounced vʌnʌm), a nongovernmental organization based in Theni District, Tamil Nadu, India, for carrying out exemplary conservation work in the region. Their unwavering support has been vitally important to our work in the Meghamalai Wildlife Sanctuary. Suggested Common Name. Meghamalai Rock Gecko Variation and additional information from type series. Mensural data for the type series and additional material is given in Table 1. There are seven females and four males, ranging in size from 61.4 mm to at least 112.2 mm. Males have a series of 17–22 femoral pores separated medially by 10–11 poreless scales (ESV106—20/20 pores on left/right side separated by 11 poreless scales, ZSI/ WGRC /IR/V.NO 2634—20/22 pores on left/right side separated by 11 poreless scales, ESV107—17/21 pores on left/right side separated by 11 poreless scales). All paratypes resemble the holotype in most of the morphological characters except as follows: Tail entire in paratypes ESV107 and NCBS-AU 159, broken or regenerated in all other paratypes; enlarged dorsal tubercles in 17–19 rows; range of supralabials is from 11–15 (9–11 below eye) and infralabials from 10–14. The scales across belly range from 34–40 in paratypes. Phylogenetic relationships. Hemidactylus vanam sp. nov. is nested within the H. prashadi group of the Indian radiation. It is sister to the recently described species H. acanthopholis (Fig. 8) with a high posterior probability and bootstrap value. H. vanam sp. nov. is genetically distinct from H. acanthopholis and H. maculatus, its two closest congeners, with an uncorrected p-distance of 0.19 each from the cytb dataset. Distribution and habitat. The following southern tropical forest types are identified in Meghamalai as per the working plan of the Tamil Nadu Forest Department: west coast semi-evergreen forests, moist mixed deciduous forests, secondary moist mixed deciduous forests, dry mixed deciduous forests, carnatic umbrella thorn forests and dry deciduous scrub forests (Bhupathy & Babu 2013). The type-series of Hemidactylus vanam sp. nov. was collected en route to the High Wavy Mountains (Fig. 10) and from near Suruli falls both within the Meghamalai Wildlife Sanctuary, where the habitat chiefly constitutes southern tropical dry mixed deciduous forests with progressions of large charnockite rock formations abutting them (Palanivelu et al. 1998). These habitats are at an altitude of 400–600 m asl and receive an average annual rainfall of 1500 mm (Bhupathy et al. 2009). Similar habitats are seen in the Vellimalai range within the Meghamalai Wildlife Sanctuary and in many parts of the Srivilliputtur Grizzled Squirrel Wildlife Sanctuary, where these geckos could also possibly occur. Natural history. Hemidactylus vanam sp. nov. appears to be predominantly rupicolous (Fig. 9) and is occasionally found on the higher branches of trees that surround rocky boulders. These strictly nocturnal geckos are seen occupying rock crevices during the day when they are easily disturbed by human presence. They seem less perturbed during the night when they actively forage on rock boulders. Juveniles and adults are syntopic. Most individuals observed had a regenerated tail. They also seem to readily slough their scales when captured, a defence strategy employed by other geckos and an exaggerated version of which was more recently described in Geckolepis megalepis from northern Madagascar (Scherz et al. 2017). Hemidactylus vanam sp. nov. is found in sympatry with Psammophilus cf. dorsalis, Cnemaspis sp. and Hemidactylus cf. triedrus. Conservation concerns. The landscape at the type locality is highly affected by anthropogenic factors. In some parts, large portions of these rock faces, the preferred niche for these geckos, have been carved out in an effort to widen the approach road to the High Wavys hill station, which is a popular tourist destination. The first author (RC) observes that these geckos were commonly seen in their habitat in November 2014, and that sightings of the same population had reduced in November 2016.

Hemidactylus sushilduttai sp. nov. Figs. 1���7 Hemidactylus maculatus (non Dum��ril & Bibron, 1836) Smith 1935; McCann 1945: 435 Hemidactylus maculatus maculatus (non Dum��ril & Bibron, 1836) Javed et al., 2010 Hemidactylus cf. maculatus Mirza & Sanap 2014 Holotype. NCBS-AU157, adult male; Simhachalam, Visakhapatnam District, Andhra Pradesh, India (17.767�� N 83.248�� E), collected by Aparna Lajmi, Aniruddha Datta-Roy and V. Deepak, 0 1 April 2014. Paratypes. ESV 109, adult male, near Lambasingi, Visakhapatnam District, Andhra Pradesh, India (17.798�� N, 82.502�� E, 750 masl), collected by Aniruddha Datta-Roy, V. Deepak, Ishan Agarwal and Prudhviraj, 0 6 October 2014; ESV 110, adult female, Maredumilli, East Godavari District, Andhra Pradesh (17.443�� N 81.753�� E), same collectors as holotype, 29 March 2014; ESV 111, subadult male, ESV 112 and ESV 113, adult females, same collection data as holotype. NCBS-AU160, adult female, near Ananthagiri, Visakhapatnam District, Andhra Pradesh, India (18.255�� N, 82.991�� E, 1170 masl), 18 September 2013, collected by Aniruddha Datta-Roy, Ishan Agarwal and Tarun Khichi. Diagnosis. A large sized Hemidactylus, snout-vent averaging 91.9 �� 13.3 mm (n=6) and up to at least 105.0 mm. Dorsal pholidosis heterogeneous, composed of granular scales intermixed with 16���17 fairly regularly arranged longitudinal rows of enlarged, strongly keeled, trihedral tubercles at midbody. First supralabial in contact with nasal; two well-developed pairs of postmentals, the inner pair slightly larger than the outer pair and in contact behind the mental. Ventrolateral folds indistinct, 30���33 scale rows across venter. All digits with enlarged scansors, 11���12 (manus) and 11���13 (pes) divided lamellae beneath fourth digit and 9���11 (manus) and 9���11 (pes) beneath first digit; 21���24 femoral pores on each side separated by four poreless scales in males. Original tail depressed, oval in transverse section with a median dorsal furrow; scales on the dorsal aspect of tail heterogenous, slightly larger than granular scales on dorsum, weakly imbricate, intermixed with a longitudinal series of six to eight, enlarged, strongly keeled and pointed tubercles. Dorsal coloration of transversely arranged, pale grey to ashy markings on a pale mustard-brown background. Hemidactylus sushilduttai sp. nov. can be easily distinguished from most congeners from India, Sri Lanka and Pakistan based on its heterogenous dorsal pholidosis that consists of small granules intermixed with 16���17 rows of fairly regularly arranged, longitudinal rows of distinct, pointed trihedral tubercles at midbody versus H. aquilonius Zug & McMahan, H. garnotii Dum��ril & Bibron and H. platyurus (Schneider), which all have homogenous dorsal pholidosis of small granules without enlarged tubercles; H. imbricatus Bauer, Giri, Greenbaum, Jackman, Dharne & Shouche and H. scabriceps (Annandale), which have homogenous dorsal pholidosis with imbricate scales and no enlarged tubercles; H. albofasciatus Grandison & Soman, H. gracilis Blanford, H. reticulatus Beddome and H. sataraensis Giri & Bauer which have heterogenous dorsal pholidosis with irregularly arranged indistinct tubercles; H. frenatus Dum��ril & Bibron, H. leschenaultii Dum��ril & Bibron, and H. flaviviridis R��ppel, which either lack enlarged tubercles or have small rounded tubercles mainly on the flanks; and H. gujaratensis Giri, Bauer, Vyas & Patil which has 12���16 rows of irregularly arranged, flattened to weakly conical dorsal tubercles. The large size of Hemidactylus sushilduttai sp. nov. (up to 105.0 mm SVL) easily distinguishes it from the smaller sized, tuberculate congeners H. brookii Gray, H. chipkali Mirza & Raju, H. depressus Gray, H. gleadowi Murray, H. kushmorensis Murray, H. lankae Deraniyagala, H. murrayi Gleadow 1887, H. parvimaculatus Deraniyagala, H. persicus Anderson, H. pieresii Kelaart, H. robustus Heyden, H. tenkatei Lidth de Jeude, H. treutleri Mahony, H. triedrus (Daudin), and H. turcicus (Linnaeus), all of which reach maximum sizes of 90 mm SVL or less. A number of congeners in India and Sri Lanka approach or exceed maximum sizes of 100 mm, including H. aaronbaueri Giri 2008, H. acanthopholis, H. giganteus Stoliczka, H. graniticolus, H. hemchandrai Dandge & Tiple, H. hunae Deraniyagala, H. maculatus Dum��ril & Bibron, H. prashadi Smith, and H. yajurvedi Murthy, Bauer, Agarwal, Lajmi & Giri. Hemidactylus sushilduttai sp. nov. can be diagnosed from all large-bodied Indian and Sri Lankan congeners by the number and shape of enlarged dorsal tubercles (16���18 rows of fairly regularly arranged, longitudinal rows of distinct trihedral tubercles at midbody) and number and arrangement of femoral pores (20���23 femoral pores separated by 3���6 poreless scales) (opposing character states indicated parenthetically): H. giganteus (complete absence of enlarged dorsal tubercles), H. yajurvedi and H. hemchandrai (10���15 rows of irregularly arranged, slightly larger, rounded, weakly-keeled tubercles at midbody), H. prashadi (14���16 rows of enlarged subtrihedral tubercles and 17���20 femoral pores on each side separated by three poreless scales), H. hunae (16���20 relatively regular rows of keeled, subtrihedral tubercles and 22���24 femoral pores on each side with a gap of 3���6 scales), H. graniticolus (16���18 relatively regular rows of subtrihedral, weakly keeled, striated tubercles and 23���28 femoral pores on each side separated by 1���3 poreless scales), H. maculatus (20 relatively regular longitudinal rows of large trihedral tubercles and 15���19 femoral pores on each side with a gap of 5���6 poreless scales), H. acanthopholis (18���20 relatively regular longitudinal rows of trihedral, moderately keeled, striated tubercles and 19���21 femoral pores separated by 13���14 poreless scales). Description of the holotype. The holotype is generally in good condition with some exceptions related to preservation (Fig. 1). The head is slightly bent to the left and the tail is fixed in a curl toward the right. There is an 11 mm long incision in the sternal region (for tissue collection), a ventrolateral fold of skin on the right side running from neck to mid-body and a ventrolateral fold on the left from behind the eye up to the neck. There is a slight constriction near the groin caused by a field tag and the venter is flattened as an artifact of preservation. Head short (HL/SVL ratio 0.29), slightly elongate (HW/HL ratio 0.75), not strongly depressed (HH/HL ratio 0.45), distinct from neck (Fig. 2 A). Loreal region slightly inflated, canthus rostralis not prominent. Snout short (SE/HL ratio 0.40); more or similar to eye diameter (OD/SE ratio 0.50); scales on snout, canthus rostralis granular and pointed, 2���3 times larger than those on interorbital and occipital region, occipital region has intermixed enlarged, keeled, pointed tubercles, which are 3���4 larger than adjacent granules. Eye small (OD/HL ratio 0.20); pupil vertical with crenulated margins; supraciliaries small, pointed, those at the anterior end of orbit slightly larger, posterior half with smaller spinose scales. Ear opening oval (greatest height 3.2 mm); eye to ear distance slightly greater than diameter of eye (EE/OD ratio 1.5). Rostral wider (3.9 mm) than deep (2.9 mm), incompletely divided dorsally by weakly developed rostral groove; internasals separated by a single small scale, one supranasal and one postnasal on each side, all subequal; rostral in contact with nostril, supralabial I, internasals and scale separating the internasals; nostrils large (1.1 mm), subcircular, each surrounded by supranasal, internasal, rostral, supralabial I and postnasal; 4���5 rows of scales separate orbit from supralabials, single row of enlarged elongate scales bordering supralabials (Fig. 2 B). Mental triangular, broader (4.3 mm) than high (4 mm), two pairs of postmentals, inner pair slightly shorter (3.3 mm) than mental and in extensive contact with each other (1.5 mm) behind mental (Fig. 2 C); outer post mental small (1.8 mm), medially divided. Inner postmentals bordered by mental, infralabial I and II, outer postmentals and 12 gular scales; outer postmentals bordered by inner postmental, infralabial II and 7���8 much enlarged gular scales on either side, of which outer row continues as a single row of enlarged scales below infralabials. About 2���7 rows of scales below infralabials III to VIII are enlarged and weakly imbricate. Supralabials (to midorbital position) 8; supralabials (to angle of jaw) 11; infralabials (to angle of jaw) 8 (right)���9 (left). Body relatively stout, not elongate (TRL/SVL ratio 0.39), with indistinct ventrolateral folds without denticulate scales. Dorsal pholidosis heterogeneous, composed of conical, granular scales intermixed with enlarged, relatively regularly arranged, longitudinal rows of 16���17 trihedral, strongly keeled tubercles at midbody (Fig. 3), extending from occiput onto the tail, each enlarged tubercle roughly 4���7 times longer than adjacent granules, surrounded by 10���18 small granules, 2���4 granules separate adjacent enlarged tubercles; enlarged tubercles similar in size except those on most medial parasagittal rows less than half the size of adjacent tubercle; shape of enlarged tubercles on back homogenous except those on flanks conical; enlarged tubercles on nape slightly smaller and more conical than those of back; tubercles on occipital and temporal region still smaller, strongly pointed. Ventral scales larger than dorsal granules, smooth, imbricate, slightly larger on femoral and largest on precloacal region; midbody scale rows across belly 33; gular region with granular scales that are smaller than ventrals. Femoral pores 23 on left thigh and 21 on right thigh, with left and right series separated by a diastema of four un-pored scales (Fig. 4). Scales on the palm and sole smooth, granular, rounded; dorsal aspect of manus and pes heterogenous, granular with densely packed enlarged tubercles, upper arm with a few granular scales which are larger than granules on dorsum intermixed with much larger tubercles; dorsal aspect of forearm predominantly bearing much larger, flattened, strongly keeled tubercles; scales on elbow are strongly keeled and conical, slightly smaller than the enlarged tubercles on the upper arm; those on dorsal aspect of thigh heterogeneous with a few granular scales which are similar to those on dorsum, intermixed with much larger trihedral tubercles which are slightly larger than enlarged dorsal tubercles; tubercles on the back of the thigh much smaller, conical and sparsely spaced; large trihedral tubercles on dorsal aspect of shank slightly smaller than those on dorsum of thigh, intermixed with a few granular scales; scales on knee are strongly weakly keeled and conical, ~ 2 times larger than adjacent granules. Fore- and hind limbs relatively short, stout; forearm short (FL/SVL ratio 0.14); tibia short (CL/SVL ratio 0.15); digits moderately short, strongly clawed; all digits of manus and digits I���IV of pes indistinctly webbed; terminal phalanx of all digits curved, arising angularly from distal portion of expanded lamellar pad or scansor, half or more than half as long as associated toepad; lamellae beneath each toe in straight transverse series, divided except for three to four basal scansors on digit I and single distal on all digits and some single basal ones too; scansors from proximal most at least twice diameter of palmar scales to distalmost single scansor: 10-11-11-12-11 (right manus; Fig. 5 A), 9-11-12-11-11 (right pes; Fig. 5 B). Relative length of digits (measurements in mm in parentheses): IV (8.5)> III (8.1)> V (8)> II (7.5)> I (7) (right manus); III (9.5)> IV (9.4)> V (8.6)> II (8.5)> I (6.5) (right pes). Tail depressed, flat beneath, verticillate, with well-defined median furrow; length of the original tail slightly longer than snout-vent length (TL/SVL ratio 1.24); scales on the tail subimbricate, slightly conical and keeled, larger than granules on dorsum, with a series of 6-8 much enlarged, strongly pointed and keeled conical tubercles, lateral row on both sides slightly elongated and weakly keeled (Fig. 6 A); ventral scales enlarged, imbricate, median row (subcaudal plates) covering almost entire base of the tail with two rows of larger pointed, smooth, imbricate scales on lateral aspects (Fig. 6 B). Coloration (in preservative). Overall specimen in preservative appears pale with indistinct transverse markings on dorsum and tail. Dorsum grayish- brown with darker speckling (darker small dots) on granular scales and enlarged tubercles throughout, a slightly darker vertebral stripe and a series of still darker, transverse markings, defining the edges of somewhat paler bands (weakly evident on occiput, one across shoulders, one between fore- and hindlimb insertions, and one on sacrum) (Fig. 1) Crown of head similar to dorsum, except slightly darker pigmentation on snout. A thin, dark brown stripe extending from behind nostril to front of orbit, bordered above with lighter stripe, continuing behind eye and on to temporal region. Infralabials and posterior supralabials slightly pale, anterior supralabials with scattered dark pigment. Limbs similar in colour to dorsum with slightly dark pigmentation on dorsal aspects of digits. Tail similar colour to dorsum but becoming paler posteriorly. Venter cream with dark speckling on all scales throughout; palms and soles grayish. Ventrolateral surfaces of tail darkly pigmented, with scattered speckling throughout, more evident on proximal portions and distal portions of midventral scutes. Colouration (in life). Dorsum with a series of broad transverse bands, one across shoulder, two between fore and hind limb insertions, and one over anterior sacral region; dorsal ground colour russet with paler transverse bands, borders of light bands slightly wavy or scalloped, with narrow seal brown, almost black edging (Fig. 7). Dark edging lacking and division between background coloration and light bands indistinct on flanks. Dorsum of head tawny, without any mottling. Incomplete, very thin greyish occipital band, discontinuously associated with whitish stripe on side of head; supraciliaries yellowish. Basal regions of limbs mottled tawny and grayish, with elbows and knees predominantly light and distal portions of limbs slightly darker; limbs without any prominent markings or bands. Tail banded, with predominant colours as on body dorsum except that pale bands tend towards a lighter ashy colour. Six pale bands on tail, all narrower than adjacent russet bands. Iris a deep golden color. Juvenile coloration similar to that of the adult, although darker areas are chocolate brown, light bands with yellowish border, limbs are slightly paler, and pale tubercles on the flanks, limbs, head and tail-base are maize yellow rather than dull whitish as in adults. Pale markings distal to the pygal portion of the tail are white, with all but the basal most very bright white. Darker bands on the tail grade from dark brown basally to nearly black at the tip. A total of nine pale bands on the original tail. Etymology. The specific epithet is a patronym honoring Sushil Kumar Dutta for his contributions to research on Indian amphibians and reptiles, as well as for his central role in encouraging many young herpetologists through talks, workshops, and the dedicated School in Herpetology. The name is particularly apt as the new species is endemic to the Eastern Ghats, the region in which much of S.K. Dutta���s herpetological research has been. Suggested common name. Dutta���s Mahendragiri Gecko/ Hemidactyl Variation and additional information from type series. Mensural data for the type series is given in Table 1. The three males and three females range from 55.0 mm to 105.0 mm SVL. All paratypes resemble the holotype in most of the morphological characters except as follows: Range of supralabials is from 9���13 (8���9 below eye) and infralabials from 9���10. The scales across belly range from 30���33 in the paratypes. Range of subdigital lamellae on first digit 9���10 and on fourth digit 1 1���13. continued. Distribution and Natural History. The type series of Hemidactylus sushilduttai sp. nov. is from the eastern slopes of the northern Mahendragiri Range and scattered coastal hills (Fig. 8). This species was found in degraded deciduous forests on the coastal hills, and in degraded and intact deciduous forest and coffee plantations at the Mahendragiri Hills; in association with either large boulders or large trees. The species is known from below 250 m close to the coast at Simhachalam to above 1,100 m at Araku. These large, nocturnal geckos are scansorial, found on large boulders and large trees, as well as culverts and occasionally buildings. The skin is fragile and is easily torn. The species is relatively common where it occurs, with most nights of survey effort in areas with large rocks or trees resulting in at least one sighting, and up to more than 10. Sympatric geckos at the type locality include Hemidactylus parvimaculatus Deraniyagala and Cyrtodactylus cf. nebulosus (Beddome). Molecular data. A partial cytochrome b sequence of topotypical Hemidactylus sushilduttai sp. nov. (ESV 112 GenBank MF668228) is 15.9���21.8 % divergent from published sequences of members of the prashadi clade (Table 2). Importantly, the new species is>16% divergent from Hemidactylus maculatus, H. graniticolus and H. prashadi; other large-bodied, tuberculate members of the H. prashadi group. 1 2 3 4 5 1 Hemidactylus sushilduttai sp. nov. ESV 112 (MF668228) 2 Hemidactylus aaronbaueri (HM595640) 20.2 3 Hemidactylus graniticolus (HM595663) 18.8 22.3 4 Hemidactylus maculatus (HM559607) 15.8 17.3 15.5 5 Hemidactylus prashadi (HM595668) 18.1 23.9 15.7 15.4 6 Hemidactylus triedrus (HM559616) 15.6 20.4 17.4 13.1 15.5, Published as part of Giri, Varad B., Bauer, Aaron M., Mohapatra, Pratyush P., Srinivasulu, Chelmala & Agarwal, Ishan, 2017, A new species of large-bodied, tuberculate Hemidactylus Oken (Squamata: Gekkonidae) from the Eastern Ghats, India, pp. 331-345 in Zootaxa 4347 (2) on pages 332-341, DOI: 10.11646/zootaxa.4347.2.8, http://zenodo.org/record/1045733, {"references":["Dumeril, A. M. C. & Bibron, G. (1836) Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Vol. 3. Libr. Encyclopedique Roret, Paris, 528 pp.","Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma. Reptilia and Amphibia. Vol. II. Sauria. Taylor and Francis, London, xiii + 440 pp., 2 folding maps, 1 pl.","McCann, C. (1945) Reptiles and amphibians of Vizagapatam and neighbouring Ghats. Journal of the Bombay Natural History Society, 45, 435 - 436.","Mirza, Z. & Sanap, R. (2014) New cryptic species of gecko of the genus Hemidactylus Oken, 1817 (Reptilia: Gekkonidae) from Southern India. Taprobanica, 6, 12 - 20.","Gleadow, F. (1887) Description of a new lizard from the Dangs. Journal of the Bombay Natural History Society, 2, 49 - 51."]}