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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."]}

Dravidogecko tholpalli sp. nov. (Figs 11 A–D, 13G, 15B; Table 7) Hoplodactylus anamallensis: Boulenger, 1885 Hoplodactylus anamallensis [non Gecko anamallensis Günther, 1875] — Boettger, 1893. Hemidactylus anamallensis: Bauer & Russell, 1995 Hemidactylus anamallensis [non Gecko anamallensis Günther, 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 /ɵɔl/) is an archaic Tamil word for ‘ancient’ and ‘ palli ’ (pronounced /pǝllɪ/) 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.

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 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.

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 douglasadamsi sp. nov. (Figs 9 A���D, 13B, 14C; Table 6) Hoplodactylus anamallensis: Boulenger, 1885 Hoplodactylus anamallensis [non Gecko anamallensis G��nther, 1875]��� Boulenger, 1885 [partim]; Boulenger, 1890 [partim]. Dravidogecko anamallensis: Smith, 1933 Dravidogecko anamallensis [non Gecko anamallensis G��nther, 1875]��� Smith, 1935 [partim]; Murthy, 1993; Sharma, 2002 [partim]. Hemidactylus anamallensis: Bauer & Russell, 1995 Hemidactylus anamallensis [non Gecko anamallensis G��nther, 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., 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 23-27, 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.","Boulenger, G. A. (1890) The Fauna of British India, Including Ceylon and Burma. Reptilia and Batrachia. Taylor & Francis, London, 541 pp., XVIII pls. https: // doi. org / 10.5962 / bhl. title. 57017","Smith, M. A. (1933) Remarks on some Old World geckos. Records of the Indian Museum, 35, 9 - 19.","Smith, M. A. (1935) The fauna of British India including Ceylon and Burma. Reptilia and Amphibia, Vol. II, Sauria. London, Taylor & Francis, London, xiii + 440 pp., 2 folding maps, 1 pl.","Murthy, T. S. N. (1993) An identification key to the reptiles of the Kalakad Wildlife Sanctuary, Tamil Nadu, India. Records of the Zoological Survey of India, 91, 161 - 168.","Sharma, R. C. (2002) The Fauna of India and the Adjacent Countries. Reptilia. Vol. II. Sauria. Zoological Survey of India, Kolkata, 425 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","Johnsingh, A. J. T. (2001) The Kalakad-Mundanthurai Tiger Reserve: A global heritage of biological diversity. Current Science, 80 (3), 378 - 388.","Srinivasulu, C., Srinivasulu, B. & Molur, S. (2014) The Status and Distribution of Reptiles in the Western Ghats, India. Conservation Assessment and Management Plan (CAMP). Wildlife Information Liaison Development Society, Coimbatore, Tamil Nadu, 148 pp.","Ayyanar, M. & Ignacimuthu, S. (2008) Endemic medicinal plants used by tribal people in Tirunelveli Hills, Western Ghats of India. In: Reddy, M. V. (Ed.), Wildlife Biodiversity Conservation. Daya Publishing House, New Delhi, pp. 278 - 285."]}

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).

Dravidogecko meghamalaiensis sp. nov. (Figs 8 A���D, 13D, 14B; Table 6) Hemidactylus anamallensis: Bauer & Russell, 1995 Hemidactylus anamallensis [non Gecko anamallensis G��nther, 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)., 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 21-23, DOI: 10.11646/zootaxa.4688.1.1, http://zenodo.org/record/3514770, {"references":["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","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.","Chandramouli, S. R. & Ganesh, S. R. (2010) Herpetofauna of southern Western Ghats, India-Reinvestigated after decades. Taprobanica, 2 (2), 72 - 85. http: // doi. org / 10.4038 / tapro. v 2 i 2.3145","Bhupathy, S. & Babu, S. (2013) Meghamalai landscape: a biodiversity hotspot. Journal of Threatened Taxa, 5, 4939 - 4944. https: // doi. org / 10.11609 / JoTT. o 3592.4939 - 44","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."]}

Calotes zolaiking sp. nov. Figs 8���12; Tables 3���5. Holotype. Adult female, NCBS-AU152, from Synod Hospital Compound (23��46 19.06 N, 92��43 56.37 E, 1290 m a.s.l.), Durtlang, Aizawl District, Mizoram state, Northeast India, collected by C. Lalrinchhana and Samuel Lalrounga on 10 June 2014. Paratypes. Two subadult females, NCBS-AU153 and ESV 105, an adult female, NCBS-AU154, and a subadult male, NCBS-AU155, collection data same as holotype; subadult male, BNHS 2327, from Hmuifang (23��27 17.93 N, 92��45 07.05 E, 1478 m a.s.l.), Aizawl District, Mizoram state, Northeast India, collected by C. Lalrinchhana and Samuel Lalrounga. Referred specimens. Adult male, NCBS-AU156 collection data same as holotype. Diagnosis. A medium sized Calotes, snout to vent length averaging 64.7 �� 8.17, and maximum to at least 77.0 mm. Body feebly compressed laterally with a weak dorsal crest; scales on top of head highly heterogeneous, keeled, those above orbits are largest, scales surrounding parietal are unequal in size; three spines on each side of head, one above orbit, one above tympanum and one on temporal region; dorsal scales heterogeneous, composed of medium sized, weakly pointed, strongly keeled scales, intermixed with similar but slightly larger scales which are distinct on flanks, upper rows directed backwards and upwards and a few lower rows directed backwards; 49���52 midbody scale rows; a weakly developed fold anterior to forelimb insertion having granular scales; tympanum small; tail rounded; eight to nine supralabials and seven to ten infralabials; lamellae bicarinate, 21���24 on fourth finger and 23���27 on fourth toe. Calotes zolaiking sp. nov can be easily diagnosed from all congeners except Calotes paulus comb. nov. in possessing heterogeneous scales on the dorsum and a weakly developed dorsal crest. Based on dorsal pholidosis, the new species is most similar to Calotes paulus comb. nov., but differs with respect to the following (Calotes paulus comb. nov. given in parentheses): greater number of midbody scales rows, 49���52 (versus 42���46), greater number of caudal vertebrae, 50 (versus 35���45), and fewer caudal vertebrae with transverse processes 12 (versus 14). Description of holotype (NCBS-AU152). Specimen is generally in good condition but slightly dehydrated, tail bent toward left in a sigmoid manner, left manus slightly adpressed with outer three fingers curved, right manus ad- pressed with fingers directed backwards, eyes sunken, orbital spine on left side broken, tips of supra-tympanic spines on both sides curved, all artefacts of preservation. Mensural and meristic data is summarised in Table 3. An adult female, SVL 77.2 mm. Head relatively long (HL/SVL 0.31), broad (HW/HL ratio 0.65), not depressed (HH/HL ratio 0.52), noticeably broader than neck (Fig. 8A). Snout short (SE/HL ratio 0.37), longer than orbit diameter (OD/SE ratio 0.78). Orbit large (OD/HL ratio 0.29); pupil round, eyelids covered with small rounded scales, a single row of scales bordering eyelids slightly elongate, six supraciliaries on each side, elongate, subimbricate, slightly protruding laterally on supraorbital ridge, similar in size, with single anterior most supraciliary smallest. Snout obtusely pointed; rostral wider (2.1 mm) than deep (0.7 mm), contacted laterally by first supralabial, an elongated large prenasal, and two large scales dorsally. Canthus rostralis and supraciliary edge sharp. Nostril circular, laterally positioned and placed at centre of a large, undivided nasal plate (Fig. 8C), which is bordered by seven scales on both sides, including one prenasal, one supranasal, three postnasals, with two subnasals on left side, and one subnasal and second supralabial on right side; separated on both sides from rostral and first supralabial by prenasal. Supralabials roughly rectangular, more or less equal sized, posterior-most being longest, bordered above by two rows of scales starting behind postnasals and extending to posterior border of orbit; upper row distinctly enlarged consisting of equal sized, roughly rectangular scales; scales on lower row are heterogeneous in shape and size, decreasing in size posteriorly. Loreal region concave, scales of loreal region heterogeneous in shape and size, flat or weakly tuberculate. Scales on postorbital and temporal region heterogeneous in shape and size, subimbricate, mostly tuberculate; a row of five much larger longitudinally keeled scales, fourth being largest, extending from posterior midorbital region to just before anterior edge of tympanum (Fig. 8C). Tympanum covered with single scale (Fig. 8C). Canthals enlarged, overlapping, becoming slightly larger along supraciliaries. Scales on dorsal surface of snout and forehead heterogeneous in shape and size, all weakly pointed anteriorly (Fig. 8B), smaller scales weakly tubercular, larger with distinct keels except those on internarial region which are roughly rounded, uniformly large and smooth. Supraorbital scales weakly keeled, increase in size from supraciliaries to inner edges of orbits, of which enlarged scales follow curvature of orbits posterolaterally (Fig. 8B). Two small scale rows divide enlarged scales of inner orbits at narrowest point of frontal. Scales on occipital region heterogeneous in shape and size; smaller scales weakly tubercular, larger ones with distinct keels. Parietal scale roughly triangular, without visible pineal eye (but with distinct depression) (Fig. 8B). Parietal scale bordered laterally by three enlarged, roughly elongated keeled scales, with much smaller scales anteriorly and posteriorly. Single temporal spine on each side, longer than nuchal spines; shorter orbital spine and a small supra-tympanic spine also present on each side (Fig. 8B). Mental subtriangular, approximately as wide as long, bordered posterolaterally by a row of six elongated postmental chin shields on either side, gradually changing shape and size posteriorly to blend with surrounding rows of gular scales (Fig. 8D). First pair of postmental chin shields in contact laterally with first infralabials, remaining shields separated from infralabials by one to two rows of small gular scales. Posterior gular region with considerably enlarged, flat, strongly keeled, pointed, imbricate scales, those on jowls slightly larger in size; anterior gular scales much smaller, rounded, weakly keeled and subimbricate; all gular scales directed posteromedially except a few median rows which are directed posteriorly. No distinct gular pouch present, transverse gular fold absent (Fig. 8D). Ventral scales are like those on posterior gular region, enlarged, flat, strongly keeled, pointed, imbricate, homogenous in shape but heterogeneous in size, generally increasing in size laterally; arranged in regular longitudinal rows that are directed posteriorly, but those on lateral aspect are directed posteromedially (Fig. 9C). Approximately 49 scales around mid-body. Nuchal crest composed of three short, laterally compressed, equal sized spines. Remaining vertebral scales slightly enlarged relative to adjacent rows (Fig. 9A) but possess a more pronounced median keel providing a serrated appearance in profile (Fig. 9B); 33 mid-dorsal scales from first raised nuchal scale to above level of cloaca. Dorsal scales heterogeneous in size and shape; all scales with a moderate median keel, arranged into approximate rows (Fig. 9A & B); keels on those of upper flanks mostly oriented obliquely upward, horizontal on mid-flanks and obliquely downward on lower flanks (Fig. 9B). A distinct shallow oblique fold in front, and curving around anterior forelimb insertion. Scales of forelimbs and ventral hindlimbs form approximate rows, those on dorsal hindlimbs do not form regular rows and are heterogeneous in size. Dorsal scales larger than ventral scales on forelimbs and hindlimbs, all moderately keeled dorsally, very weakly keeled ventrally. Forelimbs moderately long (UAL+LAL+HaL/SVL ratio 0.45); hindlimbs long (ULL+LLL+FoL/SVL ratio 0.66). Digits slender, elongate and ending in a strong, slightly curved claw. Lamellae entire, bicarinate; 21 on fourth finger, 23 on fourth toe. Relative finger lengths: IV(8.5 mm)> III(7.9 mm)> II(4.6 mm)> V(4.1 mm)> I(3.0 mm) and toes IV(11.9 mm)> III(10.5 mm)> V(8.2 mm)> II(7.2 mm)> I(5.5 mm). Tail entire, rounded, nearly twice head-body length (TL/SVL ratio 1.87), not swollen at base (Fig. 8A); uniformly covered with similar sized, strongly keeled, weakly pointed, regularly arranged, posteriorly directed imbricate scales. Coloration (in preservative). Coloration in preservation is more or less similar to live specimens but slightly faded and predominantly light brown. Transverse stripes on body, limbs and tail are darker. Refer to Figures 8 and 9 for further details of coloration and markings. Coloration in life (based on specimens other than the holotype). Coloration in life varies, but is primarily mottled shades of light and darker brown dorsally, with four distinct lighter transverse stripes on body, two on forelimbs and two on hindlimbs; stripes may be primarily green or light brown (Fig. 10). Two thin white longitudinal stripes on upper flanks extend from rear of head to base of tail. Region below orbit and lateral surface of snout white, with narrow white and brown radial stripes extending from eye. Tail with faint light and darker brown bands. Ventral surfaces whitish, throat with irregularly arranged dark mottling or thin stripes. Refer to Figure 10 for further details of coloration and marking on live animals. Variation. Paratypes and referred specimen agree with holotype in general morphology and scalation with following exceptions: NCBS-AQ154 has ten infralabials on both sides, and ESV 105 has seven infralabials on both sides; 35 vertebral scales on BNHS 2327; within paratype series, lamellae on fourth finger varies from 22 to 24, and on fourth toe from 25 to 27. Further variation in mensural and meristic characters within paratype series is documented in Table 3. Hemipenis. Hemipenis single, clavate (divided less than half of length) with length of organ greater than width. Base naked; sulcus spermaticus broad, canal like, shallow at base and deeper at terminal region. Lips of sulcus spermaticus smooth. Calyculate ornamentation present on each lobe. Thick walled smooth calyces form deep oval pits. Fleshy cardioid structure at base of ventral sulcus absent. Apex capitate and divided into four segments (Fig. 11). Hemipenis of Calotes zolaiking sp. nov. matches with 13 out of 14 characters reported for seven other Calotes spp., and 12 out of 14 characters reported for Calotes ceylonensis M��ller, 1887 (Maduwage et al. 2008; Table 4). Osteology. Skull of Calotes zolaiking sp. nov. (NCBS-AU156) is subpentagonal in outline (Fig. 12A): raised in parietal region and slopes steeply towards naris (Fig. 12B). Hyoid apparatus broad with a paired ceratobranchial (CB), CB II shorter than CB I. Maxilla has 14 teeth (including one canine-like tooth [on each side]); three premaxillary teeth; thirteen teeth on either side of mandibles (Fig. 12D & E). Twenty presacral vertebrae excluding atlas and axis and 50 caudal vertebrae (Table 5). Sternum with an oval and elongated central foramen. Ten trunk vertebrae have ribs, and an additional five vertebrae near pelvic girdle have ribs replaced by small transverse processes. Shoulder girdle comprises a broad clavicle; interclavicle long and rod like, suprascapula wedge shaped. Humerus with well-developed proximal and distal ends. Phalangeal formula of manus is 2:3:4:5:3 and of pes is 2:3:4:5:4. Many osteological characters checked for Calotes zolaiking sp. nov. (NCBS-AU156) matched those of Calotes paulus comb. nov., except the former had a higher number of caudal vertebrae (Table 5). Calotes paulus comb. nov. and Calotes zolaiking sp. nov. differ from other congeners in five different osteological characters (Table 5). Calotes paulus comb. nov. and Calotes zolaiking sp. nov. differ from Pseudocalotes in three different osteological characters (Table 5). Within the genera Calotes and Pseudocalotes, there are several variable characters (Table 5). Etymology. The species is named for its occurrence in high elevation regions. The specific epithet is a noun in apposition, derived from the Mizo language and is a portmanteau word for a lizard that inhabits high elevations (Mizo: Zo = highland/cold region, Laiking = agamid lizard). Suggested English name. Mizoram Montane Forest Lizard. Distribution and natural history. Calotes zolaiking sp. nov. is known only from its type and paratype localities, Durtlang (1290 m a.s.l.) and Hmuifang forest (1480 m a.s.l.) in Aizawl District, Mizoram state, Northeast India. These two localities are approximately 36 km apart [straight line distance]. This species is also suspected to be found in other high elevation areas of Mizoram. However, very few herpetological surveys have been carried out in this state, so the full extent of its distribution and preferred elevational range is currently unknown. The holotype was collected from a residential area where both native and non-native trees were present. It was seen perching on a Melaleuca citrina (Curtis) Dum.Cours. (bottlebrush tree), an introduced ornamental species native to Australia. The native trees dominating the locality include Trema orientalis Blume, Callicarpa arborea Roxb., Schima wallichii (DC.) Korth., Castanopsis tribuloides Sm., Albizia chinensis (Osbeck) Merr. and Ficus semicordata Buch. -Ham. ex Sm. The locality where the paratypes were collected is dominated by Rapanea capitellata (Wall) Mez., a Eurya sp. Korth., Quercus spp. L., Elaeocarpus rugosus Roxb., Nyssa javanica (Blume) Koord., a Macropanax sp. Miq., Schima wallichii (DC.) Korth. and Ardisia macrocarpa Wall., with a small patch of grassland (Fig. 13). According to Champion and Seth (1968), these regions fall under their ���Montane Sub-tropical Forest��� category. Most of the natural history observations of Calotes zolaiking sp. nov. were made from Hmuifang where the paratypes were collected. The type series was collected from vegetation at a height of 1.5 ��� 4.5 m above ground level. All individuals of the species were spotted perching on branches, which suggests they are primarily arboreal. In captivity, an adult female (NCBS AU154) laid three large eggs (measuring 14.1 X 8.2 mm, 14.3 X 8.5 mm and 14.5 X 8.5 mm) in July. During fieldwork at the type locality, juveniles were observed in November. Other agamids found in sympatry include Calotes cf. versicolor and Cristidorsa otai (Mahony, 2009). Phylogenetic relationships and morphology. Based on its phylogenetic position within the subfamily Draconinae, Oriocalotes now becomes the second monotypic genus (after Brachysaura: see Deepak et al. 2015) in recent years to be subsumed into Calotes. The poor support in the nuclear dataset is probably due to incongruence in the individual nuclear genes (Appendix 4���6). So, additional nuclear markers need to be analysed to further resolve its position within the Calotes radiation. Our nuclear and mitochondrial trees support a clade encompassing Calotes + ��� Oriocalotes ��� + Psammophilus. Members of this radiation are largely confined to India and harbor multiple ecomorphs; Psammophilus is a predominantly rock dwelling genus with a dorsoventrally compressed body. Calotes minor is completely terrestrial with a relatively short fifth toe (Deepak et al. 2015), whereas all other Calotes (including Calotes paulus comb. nov. and Calotes zolaiking sp. nov.) are predominantly or exclusively arboreal and have a longer fifth toe. Mahony (2010) noted that Oriocalotes shares common external morphological characters with Calotes (elongated spines on the post ocular, temporal and supratympanic regions) and Pseudocalotes (enlarged subocular scale row and heterogeneous dorsal scalation). Mahony (2010) also noted that no Calotes species possessed a noticeably enlarged subocular scale row, however, this character was found to be present or absent in the genus Diploderma Hallowell, 1861, so the plasticity of this character within Calotes (now including Oriocalotes) might not be unusual. The evolutionary relevance of this character is yet to be studied. Hallerman and B��hme (2000) (modified by Deepak et al. 2015) attribute the following characters to the genus Calotes: a broad head (HW/HL ratio 0.58���0.82), groups of spines from eye to tympanum, cheek often swollen in males, relatively long hindlimbs (hindlimb length/SVL ratio 0.62���1.01) and tail (TL/SVL ratio 0.9���3.0). Presence of regular, uniform dorsal scales, thought to be an important character in the diagnosis of the genus Calotes (Smith 1935; Manthey & Denzer 2000), is now demonstrated to have no systematic importance, as observed in the draconine lizards belonging to the genus Bronchocela Kaup, 1827 and Pseudocalotes. Our study reinforces the importance of hemipenial morphology as a generic diagnostic character for agamid lizards. The shape of the hemipenis is largely conserved within draconine genera (Maduwage et al. 2008; Maduwage & Silva 2012), however, the absence of the cardioid structure on the hemipenes of Calotes zolaiking sp. nov. warrants a relook at the usage of this character to diagnose the genus Calotes (Maduwage et al. 2008; Maduwage & Silva 2012; Deepak et al. 2015). The osteological characters suggested by Moody (1980) to diagnose agamid genera will require revision considering the recent changes in agamid taxonomy, e.g. within the distantly related genus, Pseudocalotes, there are multiple osteological characters which are not congruent with Moody���s (1980) diagnoses (Table 5). However, osteological comparisons of additional species in the genus Calotes is required to better understand homologous osteological characters. The genus-level assignment of draconine species has been historically based on synapomorphic characters. Taxonomic stability for most draconine species will only occur following detailed molecular phylogenetic analysis and a thorough reassessment of morphological characters., Published as part of Giri, Ad. B., Chaitanya, R., Mahony, Stephen, Lalrounga, Samuel, Lalrinchhana, C., Das, Abhijit, Sarkar, Vivek, Karanth, Praveen & Deepak, V., 2019, On the systematic status of the genus Oriocalotes G��nther, 1864 (Squamata: Agamidae: Draconinae) with the description of a new species from Mizoram state, Northeast India, pp. 451-484 in Zootaxa 4638 (4) on pages 467-477, DOI: 10.11646/zootaxa.4638.4.1, http://zenodo.org/record/3995524, {"references":["Muller, F. (1887) Funfter Nachtrag zum Katalog der herpetologischen Sammlung des Basler Museums. Verhandlungen der Naturforschenden Gesellschaft in Basel, 8, 249 - 296. https: // doi. org / 10.5962 / bhl. part. 19447","Maduwage, K., Meegaskumbura, M., Silva, A. & Pethiyagoda, R. (2008) Phylogenetic implications of hemipenial morphology in Sri Lankan agamid lizards. Current Science, 95, 838 - 840.","Champion, H. G. & Seth, S. K. (1968) A revised survey of the forest types of India. Government of India Press, Nasik, 404 pp.","Mahony, S. (2009) A new species of Japalura (Reptilia: Agamidae) from northeast India with a discussion of the similar species Japalura sagittifera Smith, 1940 and Japalura planidorsata Jerdon, 1870. Zootaxa, 2212, 41 - 61.","Deepak, V., Vyas, R., Giri, V. B. & Karanth, K. P. (2015) A taxonomic mystery for more than 180 years: the identity and systematic position of Brachysaura minor (Hardwicke & Gray, 1827) Vertebrate Zoology, 65, 371 - 381.","Mahony, S. (2010) Systematic and taxonomic revaluation of four little known Asian agamid species, Calotes kingdonwardi Smith, 1935, Japalura kaulbacki Smith, 1937, Salea kakhienensis Anderson, 1879 and the monotypic genus Mictopholis Smith, 1935 (Reptilia: Agamidae). Zootaxa, 2514 (1), 1 - 23. https: // doi. org / 10.11646 / zootaxa. 2514.1.1","Hallowell, E. (1861) Report upon the Reptilia of the North Pacific Exploring Expedition, under command of Capt. John Rogers, U. S. N. Proceedings of the Academy of Natural Sciences of Philadelphia, 12, 480 - 510.","Smith, M. A. (1935) The Fauna of British India, including Ceylon and Burma. Reptiles and Amphibia. Vol. II. Sauria. Taylor and Francis, London, xiii + 440 pp., pls. 1 + 2 maps.","Manthey, U. & Denzer, W. (2000) Description of a new genus, Hypsicalotes gen. nov. (Sauria: Agamidae) from Mount Kinabalu, North Borneo, with remarks on the generic identity of Gonocephalus schultzewestrumi Urban, 1999. Hamadryad, 25, 13 - 20.","Bahir, M. M. & Maduwage, K. P. (2005) Calotes desilvai, a new species of agamid lizard from Morningside Forest, Sri Lanka. Raffles Bulletin of Zoology, 12, 381 - 392.","Moody, S. M. (1980) Phylogenetic and historical biogeographical relationship of the genera in the family Agamidae (Reptilia: Lacertilia). PhD thesis, The University of Michigan, Ann Arbor, 373 pp.","Maduwage, K. & Silva, A. (2012) Hemipeneal morphology of Sri Lankan dragon lizards (Sauria: Agamidae). Ceylon Journal of Science, Biological Sciences, 41, 111 - 123. https: // doi. org / 10.4038 / cjsbs. v 41 i 2.5381"]}

Calotes zolaiking sp. nov. Figs 8–12; Tables 3–5. Holotype. Adult female, NCBS-AU152, from Synod Hospital Compound (23°46 19.06 N, 92°43 56.37 E, 1290 m a.s.l.), Durtlang, Aizawl District, Mizoram state, Northeast India, collected by C. Lalrinchhana and Samuel Lalrounga on 10 June 2014. Paratypes. Two subadult females, NCBS-AU153 and ESV 105, an adult female, NCBS-AU154, and a subadult male, NCBS-AU155, collection data same as holotype; subadult male, BNHS 2327, from Hmuifang (23°27 17.93 N, 92°45 07.05 E, 1478 m a.s.l.), Aizawl District, Mizoram state, Northeast India, collected by C. Lalrinchhana and Samuel Lalrounga. Referred specimens. Adult male, NCBS-AU156 collection data same as holotype. Diagnosis. A medium sized Calotes, snout to vent length averaging 64.7 ± 8.17, and maximum to at least 77.0 mm. Body feebly compressed laterally with a weak dorsal crest; scales on top of head highly heterogeneous, keeled, those above orbits are largest, scales surrounding parietal are unequal in size; three spines on each side of head, one above orbit, one above tympanum and one on temporal region; dorsal scales heterogeneous, composed of medium sized, weakly pointed, strongly keeled scales, intermixed with similar but slightly larger scales which are distinct on flanks, upper rows directed backwards and upwards and a few lower rows directed backwards; 49–52 midbody scale rows; a weakly developed fold anterior to forelimb insertion having granular scales; tympanum small; tail rounded; eight to nine supralabials and seven to ten infralabials; lamellae bicarinate, 21–24 on fourth finger and 23–27 on fourth toe. Calotes zolaiking sp. nov can be easily diagnosed from all congeners except Calotes paulus comb. nov. in possessing heterogeneous scales on the dorsum and a weakly developed dorsal crest. Based on dorsal pholidosis, the new species is most similar to Calotes paulus comb. nov., but differs with respect to the following (Calotes paulus comb. nov. given in parentheses): greater number of midbody scales rows, 49–52 (versus 42–46), greater number of caudal vertebrae, 50 (versus 35–45), and fewer caudal vertebrae with transverse processes 12 (versus 14). Description of holotype (NCBS-AU152). Specimen is generally in good condition but slightly dehydrated, tail bent toward left in a sigmoid manner, left manus slightly adpressed with outer three fingers curved, right manus ad- pressed with fingers directed backwards, eyes sunken, orbital spine on left side broken, tips of supra-tympanic spines on both sides curved, all artefacts of preservation. Mensural and meristic data is summarised in Table 3. An adult female, SVL 77.2 mm. Head relatively long (HL/SVL 0.31), broad (HW/HL ratio 0.65), not depressed (HH/HL ratio 0.52), noticeably broader than neck (Fig. 8A). Snout short (SE/HL ratio 0.37), longer than orbit diameter (OD/SE ratio 0.78). Orbit large (OD/HL ratio 0.29); pupil round, eyelids covered with small rounded scales, a single row of scales bordering eyelids slightly elongate, six supraciliaries on each side, elongate, subimbricate, slightly protruding laterally on supraorbital ridge, similar in size, with single anterior most supraciliary smallest. Snout obtusely pointed; rostral wider (2.1 mm) than deep (0.7 mm), contacted laterally by first supralabial, an elongated large prenasal, and two large scales dorsally. Canthus rostralis and supraciliary edge sharp. Nostril circular, laterally positioned and placed at centre of a large, undivided nasal plate (Fig. 8C), which is bordered by seven scales on both sides, including one prenasal, one supranasal, three postnasals, with two subnasals on left side, and one subnasal and second supralabial on right side; separated on both sides from rostral and first supralabial by prenasal. Supralabials roughly rectangular, more or less equal sized, posterior-most being longest, bordered above by two rows of scales starting behind postnasals and extending to posterior border of orbit; upper row distinctly enlarged consisting of equal sized, roughly rectangular scales; scales on lower row are heterogeneous in shape and size, decreasing in size posteriorly. Loreal region concave, scales of loreal region heterogeneous in shape and size, flat or weakly tuberculate. Scales on postorbital and temporal region heterogeneous in shape and size, subimbricate, mostly tuberculate; a row of five much larger longitudinally keeled scales, fourth being largest, extending from posterior midorbital region to just before anterior edge of tympanum (Fig. 8C). Tympanum covered with single scale (Fig. 8C). Canthals enlarged, overlapping, becoming slightly larger along supraciliaries. Scales on dorsal surface of snout and forehead heterogeneous in shape and size, all weakly pointed anteriorly (Fig. 8B), smaller scales weakly tubercular, larger with distinct keels except those on internarial region which are roughly rounded, uniformly large and smooth. Supraorbital scales weakly keeled, increase in size from supraciliaries to inner edges of orbits, of which enlarged scales follow curvature of orbits posterolaterally (Fig. 8B). Two small scale rows divide enlarged scales of inner orbits at narrowest point of frontal. Scales on occipital region heterogeneous in shape and size; smaller scales weakly tubercular, larger ones with distinct keels. Parietal scale roughly triangular, without visible pineal eye (but with distinct depression) (Fig. 8B). Parietal scale bordered laterally by three enlarged, roughly elongated keeled scales, with much smaller scales anteriorly and posteriorly. Single temporal spine on each side, longer than nuchal spines; shorter orbital spine and a small supra-tympanic spine also present on each side (Fig. 8B). Mental subtriangular, approximately as wide as long, bordered posterolaterally by a row of six elongated postmental chin shields on either side, gradually changing shape and size posteriorly to blend with surrounding rows of gular scales (Fig. 8D). First pair of postmental chin shields in contact laterally with first infralabials, remaining shields separated from infralabials by one to two rows of small gular scales. Posterior gular region with considerably enlarged, flat, strongly keeled, pointed, imbricate scales, those on jowls slightly larger in size; anterior gular scales much smaller, rounded, weakly keeled and subimbricate; all gular scales directed posteromedially except a few median rows which are directed posteriorly. No distinct gular pouch present, transverse gular fold absent (Fig. 8D). Ventral scales are like those on posterior gular region, enlarged, flat, strongly keeled, pointed, imbricate, homogenous in shape but heterogeneous in size, generally increasing in size laterally; arranged in regular longitudinal rows that are directed posteriorly, but those on lateral aspect are directed posteromedially (Fig. 9C). Approximately 49 scales around mid-body. Nuchal crest composed of three short, laterally compressed, equal sized spines. Remaining vertebral scales slightly enlarged relative to adjacent rows (Fig. 9A) but possess a more pronounced median keel providing a serrated appearance in profile (Fig. 9B); 33 mid-dorsal scales from first raised nuchal scale to above level of cloaca. Dorsal scales heterogeneous in size and shape; all scales with a moderate median keel, arranged into approximate rows (Fig. 9A & B); keels on those of upper flanks mostly oriented obliquely upward, horizontal on mid-flanks and obliquely downward on lower flanks (Fig. 9B). A distinct shallow oblique fold in front, and curving around anterior forelimb insertion. Scales of forelimbs and ventral hindlimbs form approximate rows, those on dorsal hindlimbs do not form regular rows and are heterogeneous in size. Dorsal scales larger than ventral scales on forelimbs and hindlimbs, all moderately keeled dorsally, very weakly keeled ventrally. Forelimbs moderately long (UAL+LAL+HaL/SVL ratio 0.45); hindlimbs long (ULL+LLL+FoL/SVL ratio 0.66). Digits slender, elongate and ending in a strong, slightly curved claw. Lamellae entire, bicarinate; 21 on fourth finger, 23 on fourth toe. Relative finger lengths: IV(8.5 mm)> III(7.9 mm)> II(4.6 mm)> V(4.1 mm)> I(3.0 mm) and toes IV(11.9 mm)> III(10.5 mm)> V(8.2 mm)> II(7.2 mm)> I(5.5 mm). Tail entire, rounded, nearly twice head-body length (TL/SVL ratio 1.87), not swollen at base (Fig. 8A); uniformly covered with similar sized, strongly keeled, weakly pointed, regularly arranged, posteriorly directed imbricate scales. Coloration (in preservative). Coloration in preservation is more or less similar to live specimens but slightly faded and predominantly light brown. Transverse stripes on body, limbs and tail are darker. Refer to Figures 8 and 9 for further details of coloration and markings. Coloration in life (based on specimens other than the holotype). Coloration in life varies, but is primarily mottled shades of light and darker brown dorsally, with four distinct lighter transverse stripes on body, two on forelimbs and two on hindlimbs; stripes may be primarily green or light brown (Fig. 10). Two thin white longitudinal stripes on upper flanks extend from rear of head to base of tail. Region below orbit and lateral surface of snout white, with narrow white and brown radial stripes extending from eye. Tail with faint light and darker brown bands. Ventral surfaces whitish, throat with irregularly arranged dark mottling or thin stripes. Refer to Figure 10 for further details of coloration and marking on live animals. Variation. Paratypes and referred specimen agree with holotype in general morphology and scalation with following exceptions: NCBS-AQ154 has ten infralabials on both sides, and ESV 105 has seven infralabials on both sides; 35 vertebral scales on BNHS 2327; within paratype series, lamellae on fourth finger varies from 22 to 24, and on fourth toe from 25 to 27. Further variation in mensural and meristic characters within paratype series is documented in Table 3. Hemipenis. Hemipenis single, clavate (divided less than half of length) with length of organ greater than width. Base naked; sulcus spermaticus broad, canal like, shallow at base and deeper at terminal region. Lips of sulcus spermaticus smooth. Calyculate ornamentation present on each lobe. Thick walled smooth calyces form deep oval pits. Fleshy cardioid structure at base of ventral sulcus absent. Apex capitate and divided into four segments (Fig. 11). Hemipenis of Calotes zolaiking sp. nov. matches with 13 out of 14 characters reported for seven other Calotes spp., and 12 out of 14 characters reported for Calotes ceylonensis Müller, 1887 (Maduwage et al. 2008; Table 4). Osteology. Skull of Calotes zolaiking sp. nov. (NCBS-AU156) is subpentagonal in outline (Fig. 12A): raised in parietal region and slopes steeply towards naris (Fig. 12B). Hyoid apparatus broad with a paired ceratobranchial (CB), CB II shorter than CB I. Maxilla has 14 teeth (including one canine-like tooth [on each side]); three premaxillary teeth; thirteen teeth on either side of mandibles (Fig. 12D & E). Twenty presacral vertebrae excluding atlas and axis and 50 caudal vertebrae (Table 5). Sternum with an oval and elongated central foramen. Ten trunk vertebrae have ribs, and an additional five vertebrae near pelvic girdle have ribs replaced by small transverse processes. Shoulder girdle comprises a broad clavicle; interclavicle long and rod like, suprascapula wedge shaped. Humerus with well-developed proximal and distal ends. Phalangeal formula of manus is 2:3:4:5:3 and of pes is 2:3:4:5:4. Many osteological characters checked for Calotes zolaiking sp. nov. (NCBS-AU156) matched those of Calotes paulus comb. nov., except the former had a higher number of caudal vertebrae (Table 5). Calotes paulus comb. nov. and Calotes zolaiking sp. nov. differ from other congeners in five different osteological characters (Table 5). Calotes paulus comb. nov. and Calotes zolaiking sp. nov. differ from Pseudocalotes in three different osteological characters (Table 5). Within the genera Calotes and Pseudocalotes, there are several variable characters (Table 5). Etymology. The species is named for its occurrence in high elevation regions. The specific epithet is a noun in apposition, derived from the Mizo language and is a portmanteau word for a lizard that inhabits high elevations (Mizo: Zo = highland/cold region, Laiking = agamid lizard). Suggested English name. Mizoram Montane Forest Lizard. Distribution and natural history. Calotes zolaiking sp. nov. is known only from its type and paratype localities, Durtlang (1290 m a.s.l.) and Hmuifang forest (1480 m a.s.l.) in Aizawl District, Mizoram state, Northeast India. These two localities are approximately 36 km apart [straight line distance]. This species is also suspected to be found in other high elevation areas of Mizoram. However, very few herpetological surveys have been carried out in this state, so the full extent of its distribution and preferred elevational range is currently unknown. The holotype was collected from a residential area where both native and non-native trees were present. It was seen perching on a Melaleuca citrina (Curtis) Dum.Cours. (bottlebrush tree), an introduced ornamental species native to Australia. The native trees dominating the locality include Trema orientalis Blume, Callicarpa arborea Roxb., Schima wallichii (DC.) Korth., Castanopsis tribuloides Sm., Albizia chinensis (Osbeck) Merr. and Ficus semicordata Buch. -Ham. ex Sm. The locality where the paratypes were collected is dominated by Rapanea capitellata (Wall) Mez., a Eurya sp. Korth., Quercus spp. L., Elaeocarpus rugosus Roxb., Nyssa javanica (Blume) Koord., a Macropanax sp. Miq., Schima wallichii (DC.) Korth. and Ardisia macrocarpa Wall., with a small patch of grassland (Fig. 13). According to Champion and Seth (1968), these regions fall under their “Montane Sub-tropical Forest” category. Most of the natural history observations of Calotes zolaiking sp. nov. were made from Hmuifang where the paratypes were collected. The type series was collected from vegetation at a height of 1.5 – 4.5 m above ground level. All individuals of the species were spotted perching on branches, which suggests they are primarily arboreal. In captivity, an adult female (NCBS AU154) laid three large eggs (measuring 14.1 X 8.2 mm, 14.3 X 8.5 mm and 14.5 X 8.5 mm) in July. During fieldwork at the type locality, juveniles were observed in November. Other agamids found in sympatry include Calotes cf. versicolor and Cristidorsa otai (Mahony, 2009). Phylogenetic relationships and morphology. Based on its phylogenetic position within the subfamily Draconinae, Oriocalotes now becomes the second monotypic genus (after Brachysaura: see Deepak et al. 2015) in recent years to be subsumed into Calotes. The poor support in the nuclear dataset is probably due to incongruence in the individual nuclear genes (Appendix 4–6). So, additional nuclear markers need to be analysed to further resolve its position within the Calotes radiation. Our nuclear and mitochondrial trees support a clade encompassing Calotes + “ Oriocalotes ” + Psammophilus. Members of this radiation are largely confined to India and harbor multiple ecomorphs; Psammophilus is a predominantly rock dwelling genus with a dorsoventrally compressed body. Calotes minor is completely terrestrial with a relatively short fifth toe (Deepak et al. 2015), whereas all other Calotes (including Calotes paulus comb. nov. and Calotes zolaiking sp. nov.) are predominantly or exclusively arboreal and have a longer fifth toe. Mahony (2010) noted that Oriocalotes shares common external morphological characters with Calotes (elongated spines on the post ocular, temporal and supratympanic regions) and Pseudocalotes (enlarged subocular scale row and heterogeneous dorsal scalation). Mahony (2010) also noted that no Calotes species possessed a noticeably enlarged subocular scale row, however, this character was found to be present or absent in the genus Diploderma Hallowell, 1861, so the plasticity of this character within Calotes (now including Oriocalotes) might not be unusual. The evolutionary relevance of this character is yet to be studied. Hallerman and Böhme (2000) (modified by Deepak et al. 2015) attribute the following characters to the genus Calotes: a broad head (HW/HL ratio 0.58–0.82), groups of spines from eye to tympanum, cheek often swollen in males, relatively long hindlimbs (hindlimb length/SVL ratio 0.62–1.01) and tail (TL/SVL ratio 0.9–3.0). Presence of regular, uniform dorsal scales, thought to be an important character in the diagnosis of the genus Calotes (Smith 1935; Manthey & Denzer 2000), is now demonstrated to have no systematic importance, as observed in the draconine lizards belonging to the genus Bronchocela Kaup, 1827 and Pseudocalotes. Our study reinforces the importance of hemipenial morphology as a generic diagnostic character for agamid lizards. The shape of the hemipenis is largely conserved within draconine genera (Maduwage et al. 2008; Maduwage & Silva 2012), however, the absence of the cardioid structure on the hemipenes of Calotes zolaiking sp. nov. warrants a relook at the usage of this character to diagnose the genus Calotes (Maduwage et al. 2008; Maduwage & Silva 2012; Deepak et al. 2015). The osteological characters suggested by Moody (1980) to diagnose agamid genera will require revision considering the recent changes in agamid taxonomy, e.g. within the distantly related genus, Pseudocalotes, there are multiple osteological characters which are not congruent with Moody’s (1980) diagnoses (Table 5). However, osteological comparisons of additional species in the genus Calotes is required to better understand homologous osteological characters. The genus-level assignment of draconine species has been historically based on synapomorphic characters. Taxonomic stability for most draconine species will only occur following detailed molecular phylogenetic analysis and a thorough reassessment of morphological characters.

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 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., 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 339-342, DOI: 10.11646/zootaxa.4420.3.2, http://zenodo.org/record/1250911, {"references":["Annandale, N. E. (1906) A new gecko from the Eastern Himalayas. Journal of the Asiatic Society of Bengal, 2 (7), 287 - 288.","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., 1 pl., 2 folding maps.","Ahmed, S. & Dasgupta, G. (1992) Reptilia. In: Ghosh, A. K. (Ed.), Fauna of West Bengal. Part II. (Reptilia, Amphibia, fishes, Hemichordata and archaeozoology). State Fauna Series 3. Zoological Survey of India, Calcutta, pp. 1 - 65."]}

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 ³. 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 ³. (Continueđ)

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 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.

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 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 Tail original, incomplete, rounded, slender, tapering; divided into indistinct caudal segments; two to three rows of enlarged, flat, weakly pointed, smooth, tubercles positioned transversely on first three segments of tail, remaining dorsal caudal scales smooth, rounded, subimbricate, similar in size dorsally but becoming larger on lateral aspect; subcaudal scales larger, smooth, imbricate and heterogeneous in size, no enlarged plates. Three enlarged, smooth, conical post cloacal spurs on each side of tail base. 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. (19̚21 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.

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 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 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., 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 520-524, 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 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., 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 524-528, DOI: 10.11646/zootaxa.4524.5.1, http://zenodo.org/record/2610704

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 Tail original, tip missing, somewhat rounded, slender, tapering, divided into indistinct segments; six or seven rows of enlarged, flat, weakly pointed, feebly keeled, tubercles on base and first segment of tail only, remaining dorsal caudal scales smooth, rounded, subimbricate, similar in size dorsally but becoming larger on lateral aspect; subcaudal scales larger, smooth, imbricate; with no distinct mid-ventral series of paired scales. Five enlarged, smooth post cloacal spurs on each side. 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 ”.

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"]}

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 Tail original with regenerated tip, somewhat rounded, slender, tapering, not divided into segments; about ten rows of enlarged, rounded, feebly keeled tubercles on tail base and 1–4 rows on the first three segments; remaining dorsal caudal scales smooth, roughly rectangular, subimbricate throughout without a single row of tubercles along the posterior edge of the first three segments; subcaudal scales larger, smooth, imbricate; no distinct mid-ventral series; three enlarged smooth post cloacal spurs on each side. 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.

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 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., 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 528-531, DOI: 10.11646/zootaxa.4524.5.1, http://zenodo.org/record/2610704

Sitana gokakensis sp. nov. Fig. 7–9 & 11; Table 3 & 4; Appendix 7 & 8 Holotype. BNHS 2490, adult male (Fig. 7 & 8A) from Gokak plateau, Belagavi district, Karnataka, India (16.18618°N, 74.75952°E), 255 m elevation, collected on 0 8.08.2013 by V. Deepak and Aparna Lajmi. Paratypes. BNHS 2491, adult female (Fig. 9b; Appendix 7b) collected by V. Deepak and Aparna Lajmi; CES 141136, an adult male (Appendix 7a) collected by V. Deepak and Kunal Arekar on 12.5.15; both same collection data as holotype. Diagnosis. A large sized Sitana with a maximum SVL of 53.1 mm, distinguished from its congeners by the following combination of characters: 1) dewlap feebly serrated without bright orange patches in breeding males (vs well serrated with bright orange patches in breeding males in S. ponticeriana, S. visiri, S. marudhamneydhal and S. devakai); 2) dewlap extending beyond forearm insertion (vs not extending in S. sivalensis, S. schleichi and S. fusca); 3) four prominent enlarged non spine like scales bordering occipital region (vs enlarged spine like scales in S. spinaecephalus); 4) dewlap large sized extending up to 55% of trunk (vs up to 44 % of trunk in S. laticeps in southern most population which appears to be an outlier for the species; up to 29% of trunk in remaining population (Fig. 10) and vs up to 42% of trunk in Sitana sp1 (Fig. 10); 5) body size (male & females averaged) relatively large (SVL: mean 48.8 +/- 3.6 SE; range 42.4–53.1; n=14) vs (body size relatively small in S. laticeps SVL: mean 47.8 +/- 3.2 SE; range 40.7– 54.6; n=63 and in S. sp1 SVL: mean 46.2 +/- 3.2 SE; range 38.9–54.1; n=31 and S. spinaecephalus SVL: mean 47.3 +/- 3.7 SE; range 39.5– 56.6; n=25). It has to be noted here that the southernmost population of S. laticeps appears to have some relatively large bodied individuals (SVL: mean 50.1 +/- 4.2 SE; range 45.2–54.6; n=6) but need a larger sample size to further confirm. Description of holotype. The holotype is in good condition; hemipenis partially everted, exposed and seen on both sides when viewed dorsally (Fig. 7a). Tail entire, curved slightly towards the right; loose folds of skin on the dorsum and a small incision of 4mm to extricate tissue, are artefacts of preservation. An adult male, SVL 51.2 mm. Head relatively long (HL/ SVL ratio 0.29), wide (HW/HL ratio 0.70), not depressed (HH/HL ratio 0.51), distinct from neck. Snout short (SE /HL ratio 0.39), longer than diameter of orbit (OD/ SE ratio 0.73), obtusely pointed in profile when viewed dorsally (Fig. 7c); rostral much wider than high, contacted laterally on either side by first supralabial, a prenasal and dorsally by two smaller, keeled scales (Fig. 7e). Canthus rostralis and supraciliary edge, sharp. Nostril roughly circular, laterally positioned and placed in the centre of a large, undivided nasal. Nasal scale bordered by eight scales on either side: one supranasal, three postnasals, one prenasal, the last two scales form a series of enlarged scales bordering the supralabials and a small scale separates them from the first supralabial. Ten supralabials on the right side (11 on left side), first slightly higher than others, broader than long, roughly rectangular, rest more elongate, weakly keeled, bordered above by a row of slightly smaller, rectangular, weakly keeled scales, which start at posterior margin of first supralabial, decreasing in size posteriorly and terminating after the posterior margin of orbit. Twelve infralabials on the right side (13 on left), first slightly smaller, the rest elongate, keeled increasing in size posteriorly. Loreal region concave, with scales of heterogeneous shape and size. Canthals enlarged, overlapping, slightly protruding on supraorbital ridge laterally. Eye large (ED/HL ratio 0.27); pupil rounded, covered under the eyelids; eyelids covered with scales that are heterogeneous in shape and size; larger scales on the upper eyelids, keeled, elongate and bluntly pointed, rest smooth; supraciliaries longer than broad. Orbital scales small but not granular. Scales on postorbital and temporal region, heterogeneous, sub- imbricate, strongly keeled, directed backward and upwards. Tympanum naked. Scales on dorsal surface of snout, forehead, interorbital and occipital region highly heterogeneous in size, shape, mostly elongate, strongly keeled longitudinally (Fig. 7c); scales on snout large in size, those on forehead slightly larger, interorbital region largest; occipital region with much smaller imbricate scales relative to other dorsal head scales; four relatively enlarged scales on a transverse row in the occiput region; nine scales anterior and 14 scales posterior of eyelids in the interorbital region; supraorbital scales along the supraciliary edge elongate, keeled, decreasing in size posteriorly, following curvature of orbit. Parietals larger than surrounding scales, longer than broad, tricarinate, separated from each other by two inter-parietal scales; anterior large, roughly triangular, tricarinate; posterior roughly pentagonal, bicarinate, with distinctly visible pineal eye. Parietals and the posterior inter-parietal part of a series of nine large, strongly keeled scales traversing the forehead. Mental shield narrower than rostral, roughly pentagonal, pointed posteriorly, a pair of roughly hexagonal postmentals, slightly shorter than mental, completely divided by a smaller gular scale (Fig. 7d); scales on the gular region homogenous in shape, those behind mental smooth, increasing in size and carination posteriorly. Dewlap large (DEWL / SVL ratio 0.69), extends posteriorly over 55% of trunk, extending much beyond axilla (Fig. 7b); about nine rows of anterior dewlap scales smaller, elongate, pointed, keeled; remainder of scales much enlarged, keeled, pointed, gradually increasing in size towards margin; single marginal row largest with lanceolate scales. Enlarged scales on dewlap in 23 rows. Nuchal and dorsal crest absent (Fig. 7a). Scales on nuchal region smaller, less than half the size of those on interorbital region, imbricate, strongly keeled. Body slender, 60 rows of scales around midbody; vertebral scale row with 46 scales, vertebral scale row partially paired and alternating. Five enlarged dorsal scale rows, on either side of the vertebral scale row (except in 3 places the five rows on either side are in contact with each other), the 5 dorsal scale rows starts from back of neck until groin, sub equal in size and shape, imbricate, pointed, keeled, directed backwards forming regularly arranged longitudinal rows; scales on flanks heterogeneous in size, shape, smaller than those on back, pointed, keeled, upper rows directed backwards and upwards, lower rows backwards and downwards; ventral scales subimbricate, keeled, homogenous in size and shape, arranged in 101 rows. Fore and hindlimbs relatively slender, tibia short (CL/ SVL ratio 0.33); digits moderately long, ending in strong, elongate, slightly recurved claw; inter-digital webbing absent; subdigital lamellae entire, bi-mucronate, 22 subdigital lamellae on toe IV including claw sheath; relative length of right fingers 3> 4> 2> 5> 1, right toes 4> 3> 2> 1. Fore and hindlimbs covered above and below with regularly arranged, enlarged, pointed, strongly keeled, scales. Tail long (TL/ SVL ratio 2.35), entire, base swollen, uniformly covered with similar sized, keeled, pointed, regularly arranged, backwardly directed imbricate scales; subcaudal scales keeled, weakly pointed near base, becoming pointed posteriorly. Colour of holotype in life. Head darker than body, dorsum of the torso dark brown, darker compared to flank and tail (Fig. 7a). Neck region with blue and dark pink colouration. Iris yellow in colour, tympanum cream coloured. Dorsum with five black blotches the one on neck darker. Flank region brick red colour with mottled dark brown patches. Enlarged scales on the flank and thigh cream in colour. Belly off white in colour with brown speckles on most scales. Forelimbs and hindlimbs dark brown suffused with red on dorsal side, cream patch on the back of thigh extending to the saccaral region and pale on the ventral side. Lower jaws cream mostly, dark blue colouration starts from the mentum (on 7–8 horizontal scale rows (Fig. 7d) and extends along midline of dewlap. Dewlap white, each scale on dewlap with clustered brown speckles. Colour of holotype in preservative. Head dark brown, darker than the body—with scattered blue patches on the neck. Two broken black stripes start below the eye approximately in the middle directed downwards that broaden and end after the forearm insertion. Tympanum cream coloured, slightly lighter than the surrounding scales with small light brown speckles. Dorsum dark brown with two black blotches on the neck; 3 broad, dark brown bars at the tail base and multiple smaller bars found throughout the tail. The five enlarged rows of scales on the dorsum flanking the vertebral scale rows dark brown coloured, a pale brown broken white stripe starting from neck bordering enlarged scale row on the dorsum ends near the tail base, upper 3 /4th of the flanks dark brown with mixed light brown scales, the lower end of flanks paler. Venter mottled with brown on off white scales. Brown bars present on the dorsal side of limbs. Tail with dark brown bands throughout. Enlarged scales on dewlap with dark brown/black spots, mottled, with some scales having dark brown/black spots with a white spot in the center. Throat region on either side of the enlarged scale on dewlap off white with bluish dark gray /black speckles and the dark blue colouration on the throat is prominent 5 scales after the mentum and its surrounding scales, the rest are faint bluish gray. Variation in paratypes. The two paratypes agree with the holotype in overall scalation with some exceptions. CES 141136 has 101 ventral scales and 32 belly ventral scales, 58 mid-body scales, 49 dorsal scales, one less infralabial on each side and one additional supralabial on the left. The female BNHS 2491 agrees with the holotype in overall scalation except that it lacks a dewlap, and has 67 ventral scales, one less infralabial on each side. CES 141136 differs in colouration from the holotype in having a light brown dorsum (an artefact of preservation), four distinct dark brown patches on the dorsum the first one darker. No blue patches on the neck. The stripe below the eye, indistinguishable compared to the surrounding scale, only a short light brown stripe ends near the forearm insertion. Blue line on the dewlap/throat darker than the holotype. Broad bands are present throughout the tail. BNHS 2491 differs in colouration from the holotype in having head colouration similar to body, three rhomboidal/ oval patterns on the dorsum, pale white stripe on the dorsum absent. Head brown with patches of brick red and grey (likely due to shedding scales), no blue patches on the neck. Throat region pale white in colour with dark black spots and speckles. One rhomboidal marking on the dorsum of tail base. Variation in live colouration. Most of the male specimens collected during this study matched with the holotype in live colouration. There were some specimens which were different from the rest which we describe herein. Three out of the ten breeding males were found with blue colouration on the upper eyelids. Seven out of the ten males observed in-situ had mostly dark pink and tinge of blue on the nuchal region. The fifth scale on the enlarged scale rows flanking vertebral scales on the dorsum was bordered with continuous cream coloured/reddish stripe starting from neck till the vent (e.g, see Fig. 9 a,b,d), in some individuals these stripes were broken. The rhomboidal patch on the neck was single in many individuals but the 5 rhomboid patches on the dorsum were separated by a cream line on the vertebral region (e.g, see Fig. 9c). One of the adult males had blue colouration on the tail which became prominent immediately after euthanization (Appendix 8). Hemipenial morphology. Four samples were examined (CES 141136 – 141139). Hemipenis bilobed, relatively small, longer than wide and shallowly forked. Sulcus spermaticus bifurcated and the fork continues onto the apical lobes (Fig. 11a). Sulcal lips raised and papillate, sulcus smooth originating from the side of the base. Apex with small serrated row of calyces and the sulcal region of apex nude (Fig. 11d), medial projection absent. Ornamentation is differentiated and combination of flounces and calyces observed. Papillae present between the apical lobes. Apical regions on the lobes of sulcal side calyces are serrated and continuous; calyces are relatively larger and non-serrated at the base of the lobes (Fig. 11a,b). Calyces are deep regular pits on the asulcal side and become shallow at basal region (Fig. 11b). Ridges between the calyces are thin and show micro-ornamentation which are scalloped. Flounces present, eight to ten in number, all prominent on the asulcal side (Fig. 11B,C). Etymology. The specific epithet is an adjectival toponym and refers to the Gokak plateau of Belagavi district in Karnataka, to which this species is endemic. Suggested common name. Gokak fan-throated lizard. Distribution. Sitana gokakensis sp. nov. is endemic to Gokak plateau in Belagavi district, Karnataka (Fig. 1). The samples collected from north and south outside this plateau (Nipani, Ramdurga, Bagalkot) were of S. laticeps. The Ghataprabha is a rocky river and a potential geographical barrier for this species. Habitat and natural history. Sitana gokakensis sp. nov. is only known from the open rocky habitat in Gokak plateau (Fig. 12). Gokak hills receives 820 mm of average annual rainfall. The habitat is xeric and is dominated by thorny and dwarf succulent species (Malpure et al. 2016). The recently described Euphorbia gokakensis Yadav, Malpure, Chandore, 2016 is endemic to this plateau which is found alongside other succulents such as E. antiquorum L., E. caducifolia Haines and Opuntia elatior Mill. and non-succulents like Mundulea sericea (Willd.) A. Chev. (Malpure et al. 2016). Ophisops sp and Hemidactylus murrayi Gleadow, 1887 are the two sympatric lizards found in this plateau. Breeding males were recorded during May and August months.

Sitana thondalu sp. nov. Fig. 13–14 & 16; Table 3 & 4; Appendix 9 & 10 Holotype. BNHS 2492, adult male (Fig.13, 14a) from East Bank of Nagarjuna Sagar reservoir, Guntur district, Andhra Pradesh, India (16.55557°N, 79.30134°E), 200 m asl, collected on 0 8.08.2015 by V. Deepak, Praveen Karanth, Aniruddha Datta-Roy, Rama Harvey and Aparna Lajmi. Paratypes. CES 141175 adult male (Appendix 9b) and CES 141178 adult female (Appendix 9a) collected on 0 8.08.2015 by V. Deepak, Praveen Karanth, Aniruddha Datta-Roy, Rama Harvey and Aparna Lajmi both same collection data as holotype. Diagnosis. A large sized Sitana with a maximum SVL of 56.3 mm, distinguished from its congeners by the combination of characters: 1) dewlap feebly serrated without bright orange patches (vs well serrated in S. ponticeriana, S. visiri, S. marudhamneydhal, S. devakai with bright orange patches in breeding males), 2) dewlap extending beyond forearm insertion (vs not extending in S. sivalensis and S. fusca), 3) four prominent enlarged non spine like scales bordering occipital region (vs enlarged spine like scales in S. spinaecephalus); 4) relatively higher head HH/HL: mean 0.53 +/- 0.05 SD and HH/HW: mean 0.79 +/-0.06 (vs relatively depressed head HH/HL: mean 0.50 +/- 0.02 SD and HH/HW: mean 0.73 +/- 0.03 SD in S. gokakensis sp. nov.) (Fig. 15). Although these ratios have overlaps S. gokakensis overall have a depressed head compared S. thondalu sp. nov. (Fig. 15). 5) higher number of vertebral scales in females 49–53 (vs lower 45–47 in S. gokakensis sp. nov. however, this needs further verification with a larger sample size). 6) Geographic location confined to Northern Andhra, Guntur District (vs known only from Gokak plateau S. gokakensis sp. nov.). 7) Sitana thondalu sp. nov. has an uncorrected p- distance of 11% in the ND2 gene and 1.5 % in R35 gene, from its sister species Sitana gokakensis sp. nov. (Appendix 3 & 11). Description of holotype: The holotype is in good condition; hemipenis everted, exposed and seen on both sides when viewed dorsally (Fig. 13a). Tail entire, curved towards the right; loose folds of skin on the dorsum are artefacts of preservation. An adult male, SVL 54.25 mm. Head relatively long (HL/ SVL ratio 0.26), wide (HW/HL ratio 0.67), not depressed (HH/HL ratio 0.56), distinct from neck. Snout short (SE /HL ratio 0.38), longer than eye diameter (OD/ SE ratio 0.88), obtusely pointed in profile when viewed dorsally (Fig. 13c); rostral wider than high (RH/ RW ratio 0.55), contacted laterally on either side by first supralabial, a prenasal and dorsally by two large scales. Canthus rostralis and supraciliary edge sharp (Fig. 13e). Nostril roughly circular, laterally positioned and placed roughly in the centre of a large, undivided nasal scale (Fig. 13e). Nasal scale bordered by seven scales on the right and six on the left, one supranasal, two postnasals, one prenasal, the last two of a series of enlarged scales bordering the supralabials, first supralabial only on the right. Ten supralabials on the right side (11 on left side), first higher than others, broader than high, roughly rectangular, rest more elongate, weakly keeled, bordered above by a row of slightly smaller, rectangular, weakly keeled scales, which start at posterior margin of first supralabial, decreasing in size posteriorly and terminating above the eigth (right side). Twelve infralabials on the right side (11 on left), first slightly smaller, the rest elongate, weakly keeled increasing in size posteriorly. Loreal region concave, with scales of heterogeneous shape and size. Canthals enlarged, overlapping, slightly protruding on supraorbital ridge laterally. Loreal region with few weekly keeled scales. Eye large (ED/HL ratio 0.34); pupil rounded, covered under the eyelids; eyelids covered with scales that are heterogeneous in shape and size, predominantly smooth; supraciliaries longer than broad. Orbital scales small but not granular. Scales on postorbital and temporal region heterogeneous (size and shape), sub-imbricate, strongly keeled, directed backward and upwards. Tympanum naked. Canthal scales, and orbit bordered below by a row of fourteen scales that are heterogeneous (size and shape) anteriorly, roughly rectangular under the eye, weekly keeled, starting at the posterior margin of nasal scale and terminating after the posterior margin of the orbit. Scales on dorsal surface of snout, forehead, interorbital and occipital region highly heterogeneous (size and shape), mostly elongate, imbricate, strongly keeled longitudinally (Fig. 13c); scales on snout large, those on forehead slightly larger, interorbital region largest; occipital region with much smaller scales; 11 scales anterior and 13 scales posterior to eyelids in the interorbital region; supraorbital scales along the supraciliary edge elongate, keeled, decreasing in size posteriorly, following curvature of orbit. Parietals larger than surrounding scales, longer than broad, strongly keeled, separated from each other by a series of three smaller scales; single inter-parietal, roughly pentagonal, with no visible pineal eye. Mental shield narrower than rostral, roughly pentagonal, pointed posteriorly, a pair of elongate, curved postmentals, slightly longer than mental, in strong contact with each other (Fig. 13d); scales on the gular region homogenous in shape, those behind mental smooth, increasing in size and carination posteriorly. Dewlap large (DEW/ SVL ratio 0.62), extends posteriorly over 41% of trunk; about seven to eight rows of anterior dewlap scales smaller, elongate, pointed, keeled; remainder of scales much enlarged, keeled, ending obtusely, gradually increasing in size towards margin; single marginal row largest. Enlarged scales on dewlap in 21 rows. Nuchal and dorsal crest absent (Fig. 13a). Scales on nuchal region smaller, less than half the size of those on interorbital region, imbricate, strongly keeled. Body slender (TORW / SVL ratio 0.23), 60 rows of scales around midbody; vertebral scale row with 49 scales, four or five dorsal scale rows on either side of the slightly smaller alternating pair of vertebral scale row, the 4 or 5 dorsal scale rows are larger than those on the neck and the largest enlarged scale on the flank, these scales starts from back of neck until groin, sub equal in size, shape, imbricate, pointed, keeled, directed backwards forming regularly arranged longitudinal rows; those on flanks heterogeneous in size and shape, smaller than those on back, pointed, keeled, upper rows directed backwards and upwards, lower rows backwards and downwards; ventral scales imbricate around the dewlap, subimbricate posteriorly, keeled, homogenous in size and shape, arranged in 94 rows. Fore and hindlimbs relatively slender, tibia short (CL/ SVL ratio 0.32); digits moderately long, ending in strong, elongate, slightly recurved claw; inter-digital webbing absent; subdigital lamellae entire, bi-mucronate, 23 subdigital lamellae on toe IV including claw sheath; relative length of fingers (right) 3> 4> 2> 5> 1, toes (right) 4> 3> 2> 1. Fore and hindlimbs covered above and below with regularly arranged, enlarged, pointed, strongly keeled scales. Tail long (TL/ SVL ratio 2.72), entire, base swollen, uniformly covered with similar sized, keeled, pointed, regularly arranged, backwardly directed imbricate scales; subcaudal scales keeled, weakly pointed near base, becoming pointed posteriorly. Colour of holotype in life. Head visibly darker than body (Fig. 14a). Dorsum of the torso dark brown, darker than flank region and tail. Blue tinge on the upper labials, neck and upper eyelids. Iris yellow, lower jaws pale cream, tympanum cream coloured. Forebody with two prominent cream yellow/orange stripes. First stripe starts from the nasal scale and extends till above the forearm insertion, the second shorter and thinner stripe starts above the tympanum and extends to the dorsum. Confluent with the second thinner stripe are the two prominent dark brown stripes on the dorsum which are discontinous and broken by lighter patches starting from the neck and end before 1/4th of the tail. Enlarged scale rows on the dorsum grayish brown in contrast to flank region which is brick red in colour (Fig. 14a). Some enlarged scales on the flank and thigh cream in colour. Most belly scales were cream yellow, scales adjoining the dewlap with brown speckles. Most scales on belly without speckles. Forelimbs and hindlimbs dark brown on dorsal side and pale on the ventral side; both limbs with dark brown bands on the dorsal side. Dewlap white with most scales having clustered brown speckles and some plain. Tail with broad dark brown bands, darker on the dorsal than the ventral side (Fig. 14a). Colour of holotype in preservative. Head dark brown, darker than body, with a faded Carolina blue patch on the neck and a pale white stripe with brown speckles that starts below the eye approximately in the middle directed downwards which broadens and ends at the anterior edge of the forearm insertion. A dark brown stripe below the off-white stripe on the head starting from the 10th supralabials on the left and 9th supralabial on the right, extends into the neck and ends near the forearm insertion. Tympanum off white with small light brown speckles, slightly lighter than the surrounding scales. A prominent dark brown band between the supraciliaries on top of the head. Dorsum light brown with 5 rhomboidal markings on the back that are variable in size and almost indistinguishable from the dorsal colouration; behind the hind limb insertion is a small rhomboidal marking on the dorsal side of tail base with small dark brown bars found throughout the tail. The 4–5 enlarged rows of scales on the dorsum markings, dark brown coloured. A off white stripe starting from neck bordering the 4th and the 5th lower enlarged scale rows on the dorsum ends near the tail base. Flanks light brown with the ventro-lateral scales paler with dark brown speckles. Venter off white (Fig 13b). Faint brown bars on the dorsal side of limbs. Enlarged scales on dewlap mottled with dark brown/black spots with a pale white spot in the center that is interspersed with dark brown/black spots. Anterior of each dewlap scale pale white without any spots/speckles, the throat region on either side of the enlarged scale on dewlap pale white with the dark blue colouration on the throat visible. Variation in paratypes. The two paratypes agree with the holotype in overall scalation with some exceptions. CES 141175 has 105 ventral scales, 32 belly ventral scales (VENB) and 46 dorsal scales. The female CES 141178 agrees with the holotype in overall scalation except that it lacks a dewlap, and has 68 ventral scales, one additional supralabial on the right and one less supralabial on the left, 53 vertebral scales. CES 141175 differs in colouration from the holotype in having a light brown stripe staring below the eye and ending near the forearm insertion. The thin black stripe near the labium starts from the angle of the jaw instead of the supralabials. The throat region on either side of the enlarged scale on dewlap off white with blue speckles. The first three bands on the tail broader than the rest. CES 141178 differs in colouration from the holotype in having head colouration similar to body, rhomboidal patterns prominent, pale white stripe on the dorsum, absent. Instead, the edges of the enlarged row on the dorsum that is dark brown starting from neck to tail base, forms two stripes on the dorsum. Light brown stripe starting below the eye ends anterior to the tympanum. Stripe near labium faint and brown in colour. Venter mottled with pale brown colouration. Throat region pale white in colour. Bands on the tail broader than the holotype. Variation in live colouration. Some of the male specimens collected during this study matched with the holotype in live colouration but there were some specimens which were different from the holotype. All of the breeding males had blue colouration on the eyelids, upper labials, nasal and the nuchal region (e.g. 14A & B; 16A & B). In two individuals, the blue colouration on the nuchal region extended most of the dorsum most prominent immediately after euthanization (Appendix 10). The gray band marking between the supraciliaries on top of the head, were darker in some males. The last row on the enlarged scale rows on the dorsum were bordered with continous/broken cream coloured or dark brown stripes till the vent. The rhomboidal patch on the neck was relatively darker than the other 4–5 rhomboidal markings which were faint and broken (Fig. 16 a,b). Hemipenial morphology. Hemipenis of seven specimens including the type and the paratype were examined (BNHS 2492, CES 141175, CES 141198, CES 141200, CES 141201, CES 141202 – 141203). Hemipenis bilobed, relatively small, longer than wide and shallowly forked (Fig. 17 a,b). Sulcus spermaticus bifurcated (Fig. 17a). Sulcal lips raised and papillate. Sulcus smooth, originating from the side of the base. Apex with close irregular row of small calyces and the sulcal region of apex nude, medial projection absent (Fig. 17a,d). Ornamentation is differentiated and combination of flounces, calyces observed (Fig. 17a–d). Papillae present between the apical lobes. Calyces are relatively larger at the base of the lobes. Calyces are deep regular pits on the asulcal side and become shallow at basal region (Fig. 17b). Ridges between the calyces are smooth. Eight to eleven flounces present, all of them, prominent on the asulcal side. Etymology. The specific epithet is a noun in apposition, derived from the Telugu word thondalu for an agamid lizard in Andhra Pradesh and Telangana states. Suggested English name. Nagarjuna Sagar fan-throated lizard Distribution. Sitana thondalu sp. nov. is found in Macherala and Nagarjuna Sagar, Guntur District, in Andhra Pradesh state. This species was only recorded from these two localities during this study (Fig. 1). The altitudinal distribution is between 180 and 200 m a.s.l. Habitat and Natural history. Sitana thondalu sp. nov. is found in dry rocky outcrops with boulders (Fig. 18a) and with sparse vegetation predominantly shrubs and few stunted trees (dominated by Mundulea sericea. Macherla receives annual rainfall between 380 to 508 mm (Talukder & Pal, 2007). In the second site Macherla, the habitat is dominated by the invasive Prosopis juliflora (Sw.) DC. (Fig. 18b). Calotes versicolor (Daudin, 1802) and Eutropis nagarjunensis (Sharma, 1969) are the two sympatric diurnal lizards found in this region. Breeding males were recorded during June and August months. useđ in the analyses. ……continued on the next page

Sitana gokakensis sp. nov. Fig. 7���9 & 11; Table 3 & 4; Appendix 7 & 8 Holotype. BNHS 2490, adult male (Fig. 7 & 8A) from Gokak plateau, Belagavi district, Karnataka, India (16.18618��N, 74.75952��E), 255 m elevation, collected on 0 8.08.2013 by V. Deepak and Aparna Lajmi. Paratypes. BNHS 2491, adult female (Fig. 9b; Appendix 7b) collected by V. Deepak and Aparna Lajmi; CES 141136, an adult male (Appendix 7a) collected by V. Deepak and Kunal Arekar on 12.5.15; both same collection data as holotype. Diagnosis. A large sized Sitana with a maximum SVL of 53.1 mm, distinguished from its congeners by the following combination of characters: 1) dewlap feebly serrated without bright orange patches in breeding males (vs well serrated with bright orange patches in breeding males in S. ponticeriana, S. visiri, S. marudhamneydhal and S. devakai); 2) dewlap extending beyond forearm insertion (vs not extending in S. sivalensis, S. schleichi and S. fusca); 3) four prominent enlarged non spine like scales bordering occipital region (vs enlarged spine like scales in S. spinaecephalus); 4) dewlap large sized extending up to 55% of trunk (vs up to 44 % of trunk in S. laticeps in southern most population which appears to be an outlier for the species; up to 29% of trunk in remaining population (Fig. 10) and vs up to 42% of trunk in Sitana sp1 (Fig. 10); 5) body size (male & females averaged) relatively large (SVL: mean 48.8 +/- 3.6 SE; range 42.4���53.1; n=14) vs (body size relatively small in S. laticeps SVL: mean 47.8 +/- 3.2 SE; range 40.7��� 54.6; n=63 and in S. sp1 SVL: mean 46.2 +/- 3.2 SE; range 38.9���54.1; n=31 and S. spinaecephalus SVL: mean 47.3 +/- 3.7 SE; range 39.5��� 56.6; n=25). It has to be noted here that the southernmost population of S. laticeps appears to have some relatively large bodied individuals (SVL: mean 50.1 +/- 4.2 SE; range 45.2���54.6; n=6) but need a larger sample size to further confirm. Description of holotype. The holotype is in good condition; hemipenis partially everted, exposed and seen on both sides when viewed dorsally (Fig. 7a). Tail entire, curved slightly towards the right; loose folds of skin on the dorsum and a small incision of 4mm to extricate tissue, are artefacts of preservation. An adult male, SVL 51.2 mm. Head relatively long (HL/ SVL ratio 0.29), wide (HW/HL ratio 0.70), not depressed (HH/HL ratio 0.51), distinct from neck. Snout short (SE /HL ratio 0.39), longer than diameter of orbit (OD/ SE ratio 0.73), obtusely pointed in profile when viewed dorsally (Fig. 7c); rostral much wider than high, contacted laterally on either side by first supralabial, a prenasal and dorsally by two smaller, keeled scales (Fig. 7e). Canthus rostralis and supraciliary edge, sharp. Nostril roughly circular, laterally positioned and placed in the centre of a large, undivided nasal. Nasal scale bordered by eight scales on either side: one supranasal, three postnasals, one prenasal, the last two scales form a series of enlarged scales bordering the supralabials and a small scale separates them from the first supralabial. Ten supralabials on the right side (11 on left side), first slightly higher than others, broader than long, roughly rectangular, rest more elongate, weakly keeled, bordered above by a row of slightly smaller, rectangular, weakly keeled scales, which start at posterior margin of first supralabial, decreasing in size posteriorly and terminating after the posterior margin of orbit. Twelve infralabials on the right side (13 on left), first slightly smaller, the rest elongate, keeled increasing in size posteriorly. Loreal region concave, with scales of heterogeneous shape and size. Canthals enlarged, overlapping, slightly protruding on supraorbital ridge laterally. Eye large (ED/HL ratio 0.27); pupil rounded, covered under the eyelids; eyelids covered with scales that are heterogeneous in shape and size; larger scales on the upper eyelids, keeled, elongate and bluntly pointed, rest smooth; supraciliaries longer than broad. Orbital scales small but not granular. Scales on postorbital and temporal region, heterogeneous, sub- imbricate, strongly keeled, directed backward and upwards. Tympanum naked. Scales on dorsal surface of snout, forehead, interorbital and occipital region highly heterogeneous in size, shape, mostly elongate, strongly keeled longitudinally (Fig. 7c); scales on snout large in size, those on forehead slightly larger, interorbital region largest; occipital region with much smaller imbricate scales relative to other dorsal head scales; four relatively enlarged scales on a transverse row in the occiput region; nine scales anterior and 14 scales posterior of eyelids in the interorbital region; supraorbital scales along the supraciliary edge elongate, keeled, decreasing in size posteriorly, following curvature of orbit. Parietals larger than surrounding scales, longer than broad, tricarinate, separated from each other by two inter-parietal scales; anterior large, roughly triangular, tricarinate; posterior roughly pentagonal, bicarinate, with distinctly visible pineal eye. Parietals and the posterior inter-parietal part of a series of nine large, strongly keeled scales traversing the forehead. Mental shield narrower than rostral, roughly pentagonal, pointed posteriorly, a pair of roughly hexagonal postmentals, slightly shorter than mental, completely divided by a smaller gular scale (Fig. 7d); scales on the gular region homogenous in shape, those behind mental smooth, increasing in size and carination posteriorly. Dewlap large (DEWL / SVL ratio 0.69), extends posteriorly over 55% of trunk, extending much beyond axilla (Fig. 7b); about nine rows of anterior dewlap scales smaller, elongate, pointed, keeled; remainder of scales much enlarged, keeled, pointed, gradually increasing in size towards margin; single marginal row largest with lanceolate scales. Enlarged scales on dewlap in 23 rows. Nuchal and dorsal crest absent (Fig. 7a). Scales on nuchal region smaller, less than half the size of those on interorbital region, imbricate, strongly keeled. Body slender, 60 rows of scales around midbody; vertebral scale row with 46 scales, vertebral scale row partially paired and alternating. Five enlarged dorsal scale rows, on either side of the vertebral scale row (except in 3 places the five rows on either side are in contact with each other), the 5 dorsal scale rows starts from back of neck until groin, sub equal in size and shape, imbricate, pointed, keeled, directed backwards forming regularly arranged longitudinal rows; scales on flanks heterogeneous in size, shape, smaller than those on back, pointed, keeled, upper rows directed backwards and upwards, lower rows backwards and downwards; ventral scales subimbricate, keeled, homogenous in size and shape, arranged in 101 rows. Fore and hindlimbs relatively slender, tibia short (CL/ SVL ratio 0.33); digits moderately long, ending in strong, elongate, slightly recurved claw; inter-digital webbing absent; subdigital lamellae entire, bi-mucronate, 22 subdigital lamellae on toe IV including claw sheath; relative length of right fingers 3> 4> 2> 5> 1, right toes 4> 3> 2> 1. Fore and hindlimbs covered above and below with regularly arranged, enlarged, pointed, strongly keeled, scales. Tail long (TL/ SVL ratio 2.35), entire, base swollen, uniformly covered with similar sized, keeled, pointed, regularly arranged, backwardly directed imbricate scales; subcaudal scales keeled, weakly pointed near base, becoming pointed posteriorly. Colour of holotype in life. Head darker than body, dorsum of the torso dark brown, darker compared to flank and tail (Fig. 7a). Neck region with blue and dark pink colouration. Iris yellow in colour, tympanum cream coloured. Dorsum with five black blotches the one on neck darker. Flank region brick red colour with mottled dark brown patches. Enlarged scales on the flank and thigh cream in colour. Belly off white in colour with brown speckles on most scales. Forelimbs and hindlimbs dark brown suffused with red on dorsal side, cream patch on the back of thigh extending to the saccaral region and pale on the ventral side. Lower jaws cream mostly, dark blue colouration starts from the mentum (on 7���8 horizontal scale rows (Fig. 7d) and extends along midline of dewlap. Dewlap white, each scale on dewlap with clustered brown speckles. Colour of holotype in preservative. Head dark brown, darker than the body���with scattered blue patches on the neck. Two broken black stripes start below the eye approximately in the middle directed downwards that broaden and end after the forearm insertion. Tympanum cream coloured, slightly lighter than the surrounding scales with small light brown speckles. Dorsum dark brown with two black blotches on the neck; 3 broad, dark brown bars at the tail base and multiple smaller bars found throughout the tail. The five enlarged rows of scales on the dorsum flanking the vertebral scale rows dark brown coloured, a pale brown broken white stripe starting from neck bordering enlarged scale row on the dorsum ends near the tail base, upper 3 /4th of the flanks dark brown with mixed light brown scales, the lower end of flanks paler. Venter mottled with brown on off white scales. Brown bars present on the dorsal side of limbs. Tail with dark brown bands throughout. Enlarged scales on dewlap with dark brown/black spots, mottled, with some scales having dark brown/black spots with a white spot in the center. Throat region on either side of the enlarged scale on dewlap off white with bluish dark gray /black speckles and the dark blue colouration on the throat is prominent 5 scales after the mentum and its surrounding scales, the rest are faint bluish gray. Variation in paratypes. The two paratypes agree with the holotype in overall scalation with some exceptions. CES 141136 has 101 ventral scales and 32 belly ventral scales, 58 mid-body scales, 49 dorsal scales, one less infralabial on each side and one additional supralabial on the left. The female BNHS 2491 agrees with the holotype in overall scalation except that it lacks a dewlap, and has 67 ventral scales, one less infralabial on each side. CES 141136 differs in colouration from the holotype in having a light brown dorsum (an artefact of preservation), four distinct dark brown patches on the dorsum the first one darker. No blue patches on the neck. The stripe below the eye, indistinguishable compared to the surrounding scale, only a short light brown stripe ends near the forearm insertion. Blue line on the dewlap/throat darker than the holotype. Broad bands are present throughout the tail. BNHS 2491 differs in colouration from the holotype in having head colouration similar to body, three rhomboidal/ oval patterns on the dorsum, pale white stripe on the dorsum absent. Head brown with patches of brick red and grey (likely due to shedding scales), no blue patches on the neck. Throat region pale white in colour with dark black spots and speckles. One rhomboidal marking on the dorsum of tail base. Variation in live colouration. Most of the male specimens collected during this study matched with the holotype in live colouration. There were some specimens which were different from the rest which we describe herein. Three out of the ten breeding males were found with blue colouration on the upper eyelids. Seven out of the ten males observed in-situ had mostly dark pink and tinge of blue on the nuchal region. The fifth scale on the enlarged scale rows flanking vertebral scales on the dorsum was bordered with continuous cream coloured/reddish stripe starting from neck till the vent (e.g, see Fig. 9 a,b,d), in some individuals these stripes were broken. The rhomboidal patch on the neck was single in many individuals but the 5 rhomboid patches on the dorsum were separated by a cream line on the vertebral region (e.g, see Fig. 9c). One of the adult males had blue colouration on the tail which became prominent immediately after euthanization (Appendix 8). Hemipenial morphology. Four samples were examined (CES 141136 ��� 141139). Hemipenis bilobed, relatively small, longer than wide and shallowly forked. Sulcus spermaticus bifurcated and the fork continues onto the apical lobes (Fig. 11a). Sulcal lips raised and papillate, sulcus smooth originating from the side of the base. Apex with small serrated row of calyces and the sulcal region of apex nude (Fig. 11d), medial projection absent. Ornamentation is differentiated and combination of flounces and calyces observed. Papillae present between the apical lobes. Apical regions on the lobes of sulcal side calyces are serrated and continuous; calyces are relatively larger and non-serrated at the base of the lobes (Fig. 11a,b). Calyces are deep regular pits on the asulcal side and become shallow at basal region (Fig. 11b). Ridges between the calyces are thin and show micro-ornamentation which are scalloped. Flounces present, eight to ten in number, all prominent on the asulcal side (Fig. 11B,C). Etymology. The specific epithet is an adjectival toponym and refers to the Gokak plateau of Belagavi district in Karnataka, to which this species is endemic. Suggested common name. Gokak fan-throated lizard. Distribution. Sitana gokakensis sp. nov. is endemic to Gokak plateau in Belagavi district, Karnataka (Fig. 1). The samples collected from north and south outside this plateau (Nipani, Ramdurga, Bagalkot) were of S. laticeps. The Ghataprabha is a rocky river and a potential geographical barrier for this species. Habitat and natural history. Sitana gokakensis sp. nov. is only known from the open rocky habitat in Gokak plateau (Fig. 12). Gokak hills receives 820 mm of average annual rainfall. The habitat is xeric and is dominated by thorny and dwarf succulent species (Malpure et al. 2016). The recently described Euphorbia gokakensis Yadav, Malpure, Chandore, 2016 is endemic to this plateau which is found alongside other succulents such as E. antiquorum L., E. caducifolia Haines and Opuntia elatior Mill. and non-succulents like Mundulea sericea (Willd.) A. Chev. (Malpure et al. 2016). Ophisops sp and Hemidactylus murrayi Gleadow, 1887 are the two sympatric lizards found in this plateau. Breeding males were recorded during May and August months., Published as part of Deepak, V., Khandekar, Akshay, Chaitanya, R. & Karanth, Praveen, 2018, Descriptions of two new endemic and cryptic species of Sitana Cuvier, 1829 from peninsular India, pp. 327-365 in Zootaxa 4434 (2) on pages 336-343, DOI: 10.11646/zootaxa.4434.2.5, http://zenodo.org/record/1290584, {"references":["Malpure, N. V., Chandore, A. N. & Yadav, S. R., 2016. Euphorbia gokakensis sp. nov. (Euphorbiaceae) from sandstone formations in Karnataka, India. Nordic Journal of Botany 34, 380 - 383. https: // doi. org / 10.1111 / njb. 00997"]}

Sitana thondalu sp. nov. Fig. 13���14 & 16; Table 3 & 4; Appendix 9 & 10 Holotype. BNHS 2492, adult male (Fig.13, 14a) from East Bank of Nagarjuna Sagar reservoir, Guntur district, Andhra Pradesh, India (16.55557��N, 79.30134��E), 200 m asl, collected on 0 8.08.2015 by V. Deepak, Praveen Karanth, Aniruddha Datta-Roy, Rama Harvey and Aparna Lajmi. Paratypes. CES 141175 adult male (Appendix 9b) and CES 141178 adult female (Appendix 9a) collected on 0 8.08.2015 by V. Deepak, Praveen Karanth, Aniruddha Datta-Roy, Rama Harvey and Aparna Lajmi both same collection data as holotype. Diagnosis. A large sized Sitana with a maximum SVL of 56.3 mm, distinguished from its congeners by the combination of characters: 1) dewlap feebly serrated without bright orange patches (vs well serrated in S. ponticeriana, S. visiri, S. marudhamneydhal, S. devakai with bright orange patches in breeding males), 2) dewlap extending beyond forearm insertion (vs not extending in S. sivalensis and S. fusca), 3) four prominent enlarged non spine like scales bordering occipital region (vs enlarged spine like scales in S. spinaecephalus); 4) relatively higher head HH/HL: mean 0.53 +/- 0.05 SD and HH/HW: mean 0.79 +/-0.06 (vs relatively depressed head HH/HL: mean 0.50 +/- 0.02 SD and HH/HW: mean 0.73 +/- 0.03 SD in S. gokakensis sp. nov.) (Fig. 15). Although these ratios have overlaps S. gokakensis overall have a depressed head compared S. thondalu sp. nov. (Fig. 15). 5) higher number of vertebral scales in females 49���53 (vs lower 45���47 in S. gokakensis sp. nov. however, this needs further verification with a larger sample size). 6) Geographic location confined to Northern Andhra, Guntur District (vs known only from Gokak plateau S. gokakensis sp. nov.). 7) Sitana thondalu sp. nov. has an uncorrected p- distance of 11% in the ND2 gene and 1.5 % in R35 gene, from its sister species Sitana gokakensis sp. nov. (Appendix 3 & 11). Description of holotype: The holotype is in good condition; hemipenis everted, exposed and seen on both sides when viewed dorsally (Fig. 13a). Tail entire, curved towards the right; loose folds of skin on the dorsum are artefacts of preservation. An adult male, SVL 54.25 mm. Head relatively long (HL/ SVL ratio 0.26), wide (HW/HL ratio 0.67), not depressed (HH/HL ratio 0.56), distinct from neck. Snout short (SE /HL ratio 0.38), longer than eye diameter (OD/ SE ratio 0.88), obtusely pointed in profile when viewed dorsally (Fig. 13c); rostral wider than high (RH/ RW ratio 0.55), contacted laterally on either side by first supralabial, a prenasal and dorsally by two large scales. Canthus rostralis and supraciliary edge sharp (Fig. 13e). Nostril roughly circular, laterally positioned and placed roughly in the centre of a large, undivided nasal scale (Fig. 13e). Nasal scale bordered by seven scales on the right and six on the left, one supranasal, two postnasals, one prenasal, the last two of a series of enlarged scales bordering the supralabials, first supralabial only on the right. Ten supralabials on the right side (11 on left side), first higher than others, broader than high, roughly rectangular, rest more elongate, weakly keeled, bordered above by a row of slightly smaller, rectangular, weakly keeled scales, which start at posterior margin of first supralabial, decreasing in size posteriorly and terminating above the eigth (right side). Twelve infralabials on the right side (11 on left), first slightly smaller, the rest elongate, weakly keeled increasing in size posteriorly. Loreal region concave, with scales of heterogeneous shape and size. Canthals enlarged, overlapping, slightly protruding on supraorbital ridge laterally. Loreal region with few weekly keeled scales. Eye large (ED/HL ratio 0.34); pupil rounded, covered under the eyelids; eyelids covered with scales that are heterogeneous in shape and size, predominantly smooth; supraciliaries longer than broad. Orbital scales small but not granular. Scales on postorbital and temporal region heterogeneous (size and shape), sub-imbricate, strongly keeled, directed backward and upwards. Tympanum naked. Canthal scales, and orbit bordered below by a row of fourteen scales that are heterogeneous (size and shape) anteriorly, roughly rectangular under the eye, weekly keeled, starting at the posterior margin of nasal scale and terminating after the posterior margin of the orbit. Scales on dorsal surface of snout, forehead, interorbital and occipital region highly heterogeneous (size and shape), mostly elongate, imbricate, strongly keeled longitudinally (Fig. 13c); scales on snout large, those on forehead slightly larger, interorbital region largest; occipital region with much smaller scales; 11 scales anterior and 13 scales posterior to eyelids in the interorbital region; supraorbital scales along the supraciliary edge elongate, keeled, decreasing in size posteriorly, following curvature of orbit. Parietals larger than surrounding scales, longer than broad, strongly keeled, separated from each other by a series of three smaller scales; single inter-parietal, roughly pentagonal, with no visible pineal eye. Mental shield narrower than rostral, roughly pentagonal, pointed posteriorly, a pair of elongate, curved postmentals, slightly longer than mental, in strong contact with each other (Fig. 13d); scales on the gular region homogenous in shape, those behind mental smooth, increasing in size and carination posteriorly. Dewlap large (DEW/ SVL ratio 0.62), extends posteriorly over 41% of trunk; about seven to eight rows of anterior dewlap scales smaller, elongate, pointed, keeled; remainder of scales much enlarged, keeled, ending obtusely, gradually increasing in size towards margin; single marginal row largest. Enlarged scales on dewlap in 21 rows. Nuchal and dorsal crest absent (Fig. 13a). Scales on nuchal region smaller, less than half the size of those on interorbital region, imbricate, strongly keeled. Body slender (TORW / SVL ratio 0.23), 60 rows of scales around midbody; vertebral scale row with 49 scales, four or five dorsal scale rows on either side of the slightly smaller alternating pair of vertebral scale row, the 4 or 5 dorsal scale rows are larger than those on the neck and the largest enlarged scale on the flank, these scales starts from back of neck until groin, sub equal in size, shape, imbricate, pointed, keeled, directed backwards forming regularly arranged longitudinal rows; those on flanks heterogeneous in size and shape, smaller than those on back, pointed, keeled, upper rows directed backwards and upwards, lower rows backwards and downwards; ventral scales imbricate around the dewlap, subimbricate posteriorly, keeled, homogenous in size and shape, arranged in 94 rows. Fore and hindlimbs relatively slender, tibia short (CL/ SVL ratio 0.32); digits moderately long, ending in strong, elongate, slightly recurved claw; inter-digital webbing absent; subdigital lamellae entire, bi-mucronate, 23 subdigital lamellae on toe IV including claw sheath; relative length of fingers (right) 3> 4> 2> 5> 1, toes (right) 4> 3> 2> 1. Fore and hindlimbs covered above and below with regularly arranged, enlarged, pointed, strongly keeled scales. Tail long (TL/ SVL ratio 2.72), entire, base swollen, uniformly covered with similar sized, keeled, pointed, regularly arranged, backwardly directed imbricate scales; subcaudal scales keeled, weakly pointed near base, becoming pointed posteriorly. Colour of holotype in life. Head visibly darker than body (Fig. 14a). Dorsum of the torso dark brown, darker than flank region and tail. Blue tinge on the upper labials, neck and upper eyelids. Iris yellow, lower jaws pale cream, tympanum cream coloured. Forebody with two prominent cream yellow/orange stripes. First stripe starts from the nasal scale and extends till above the forearm insertion, the second shorter and thinner stripe starts above the tympanum and extends to the dorsum. Confluent with the second thinner stripe are the two prominent dark brown stripes on the dorsum which are discontinous and broken by lighter patches starting from the neck and end before 1/4th of the tail. Enlarged scale rows on the dorsum grayish brown in contrast to flank region which is brick red in colour (Fig. 14a). Some enlarged scales on the flank and thigh cream in colour. Most belly scales were cream yellow, scales adjoining the dewlap with brown speckles. Most scales on belly without speckles. Forelimbs and hindlimbs dark brown on dorsal side and pale on the ventral side; both limbs with dark brown bands on the dorsal side. Dewlap white with most scales having clustered brown speckles and some plain. Tail with broad dark brown bands, darker on the dorsal than the ventral side (Fig. 14a). Colour of holotype in preservative. Head dark brown, darker than body, with a faded Carolina blue patch on the neck and a pale white stripe with brown speckles that starts below the eye approximately in the middle directed downwards which broadens and ends at the anterior edge of the forearm insertion. A dark brown stripe below the off-white stripe on the head starting from the 10th supralabials on the left and 9th supralabial on the right, extends into the neck and ends near the forearm insertion. Tympanum off white with small light brown speckles, slightly lighter than the surrounding scales. A prominent dark brown band between the supraciliaries on top of the head. Dorsum light brown with 5 rhomboidal markings on the back that are variable in size and almost indistinguishable from the dorsal colouration; behind the hind limb insertion is a small rhomboidal marking on the dorsal side of tail base with small dark brown bars found throughout the tail. The 4���5 enlarged rows of scales on the dorsum markings, dark brown coloured. A off white stripe starting from neck bordering the 4th and the 5th lower enlarged scale rows on the dorsum ends near the tail base. Flanks light brown with the ventro-lateral scales paler with dark brown speckles. Venter off white (Fig 13b). Faint brown bars on the dorsal side of limbs. Enlarged scales on dewlap mottled with dark brown/black spots with a pale white spot in the center that is interspersed with dark brown/black spots. Anterior of each dewlap scale pale white without any spots/speckles, the throat region on either side of the enlarged scale on dewlap pale white with the dark blue colouration on the throat visible. Variation in paratypes. The two paratypes agree with the holotype in overall scalation with some exceptions. CES 141175 has 105 ventral scales, 32 belly ventral scales (VENB) and 46 dorsal scales. The female CES 141178 agrees with the holotype in overall scalation except that it lacks a dewlap, and has 68 ventral scales, one additional supralabial on the right and one less supralabial on the left, 53 vertebral scales. CES 141175 differs in colouration from the holotype in having a light brown stripe staring below the eye and ending near the forearm insertion. The thin black stripe near the labium starts from the angle of the jaw instead of the supralabials. The throat region on either side of the enlarged scale on dewlap off white with blue speckles. The first three bands on the tail broader than the rest. CES 141178 differs in colouration from the holotype in having head colouration similar to body, rhomboidal patterns prominent, pale white stripe on the dorsum, absent. Instead, the edges of the enlarged row on the dorsum that is dark brown starting from neck to tail base, forms two stripes on the dorsum. Light brown stripe starting below the eye ends anterior to the tympanum. Stripe near labium faint and brown in colour. Venter mottled with pale brown colouration. Throat region pale white in colour. Bands on the tail broader than the holotype. Variation in live colouration. Some of the male specimens collected during this study matched with the holotype in live colouration but there were some specimens which were different from the holotype. All of the breeding males had blue colouration on the eyelids, upper labials, nasal and the nuchal region (e.g. 14A & B; 16A & B). In two individuals, the blue colouration on the nuchal region extended most of the dorsum most prominent immediately after euthanization (Appendix 10). The gray band marking between the supraciliaries on top of the head, were darker in some males. The last row on the enlarged scale rows on the dorsum were bordered with continous/broken cream coloured or dark brown stripes till the vent. The rhomboidal patch on the neck was relatively darker than the other 4���5 rhomboidal markings which were faint and broken (Fig. 16 a,b). Hemipenial morphology. Hemipenis of seven specimens including the type and the paratype were examined (BNHS 2492, CES 141175, CES 141198, CES 141200, CES 141201, CES 141202 ��� 141203). Hemipenis bilobed, relatively small, longer than wide and shallowly forked (Fig. 17 a,b). Sulcus spermaticus bifurcated (Fig. 17a). Sulcal lips raised and papillate. Sulcus smooth, originating from the side of the base. Apex with close irregular row of small calyces and the sulcal region of apex nude, medial projection absent (Fig. 17a,d). Ornamentation is differentiated and combination of flounces, calyces observed (Fig. 17a���d). Papillae present between the apical lobes. Calyces are relatively larger at the base of the lobes. Calyces are deep regular pits on the asulcal side and become shallow at basal region (Fig. 17b). Ridges between the calyces are smooth. Eight to eleven flounces present, all of them, prominent on the asulcal side. Etymology. The specific epithet is a noun in apposition, derived from the Telugu word thondalu for an agamid lizard in Andhra Pradesh and Telangana states. Suggested English name. Nagarjuna Sagar fan-throated lizard Distribution. Sitana thondalu sp. nov. is found in Macherala and Nagarjuna Sagar, Guntur District, in Andhra Pradesh state. This species was only recorded from these two localities during this study (Fig. 1). The altitudinal distribution is between 180 and 200 m a.s.l. Habitat and Natural history. Sitana thondalu sp. nov. is found in dry rocky outcrops with boulders (Fig. 18a) and with sparse vegetation predominantly shrubs and few stunted trees (dominated by Mundulea sericea. Macherla receives annual rainfall between 380 to 508 mm (Talukder & Pal, 2007). In the second site Macherla, the habitat is dominated by the invasive Prosopis juliflora (Sw.) DC. (Fig. 18b). Calotes versicolor (Daudin, 1802) and Eutropis nagarjunensis (Sharma, 1969) are the two sympatric diurnal lizards found in this region. Breeding males were recorded during June and August months. useđ in the analyses. ������continued on the next page, Published as part of Deepak, V., Khandekar, Akshay, Chaitanya, R. & Karanth, Praveen, 2018, Descriptions of two new endemic and cryptic species of Sitana Cuvier, 1829 from peninsular India, pp. 327-365 in Zootaxa 4434 (2) on pages 343-351, DOI: 10.11646/zootaxa.4434.2.5, http://zenodo.org/record/1290584, {"references":["Talukder, B & Pal, T. K. (2007) Animal remains front Nagarjunakonda archaeological site (Andhra Pradesh) and their relevance to the ancient civilization. Records of the Zoological Survey of India, Occasional Paper, 264, 1 - 92.","Sharma, R. C. (1969) Two new izards of the genera Mabuya Fitzinger and Riopa Gray (Scincidae) from India. Bulletin of Systematic Zoology, Calcutta, 1, 71 - 75."]}

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 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 Dum��ril & 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 Dum��ril & Bibron, and H. flaviviridis R��ppel 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 Dum��ril & 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 v��n��m), 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., Published as part of Chaitanya, R., Lajmi, Aparna & Giri, Varad B., 2018, A new cryptic, rupicolous species of Hemidactylus Oken, 1817 (Squamata: Gekkonidae) from Meghamalai, Tamil Nadu, India, pp. 49-70 in Zootaxa 4374 (1) on pages 51-58, DOI: 10.11646/zootaxa.4374.1.3, http://zenodo.org/record/1152206, {"references":["Scherz, M. D., Daza, J. D., Kohler, J, Vences, M. & Glaw, F. (2017) Off the scale: a new species of fish-scale gecko (Squamata: Gekkonidae: Geckolepis) with exceptionally large scales. PeerJ, 5, e 2955 https: // doi. org / 10.7717 / peerj. 2955"]}

Sitana marudhamneydhal sp. nov. Fig. 6, Fig.7; Table 3 Holotype. NCBS AQ 460, an adult male (Fig. 6) from Kallidaikurichi, Tirunelveli District, Tamil Nadu, India (8��39���54��� N; 77��27���15��� E), 68 m elevation, collected on 12 November 2014 by V. Deepak and R. Chaitanya. Paratypes CES 141124, an adult male from Kallidaikurichi, Tirunelveli District, Tamil Nadu, India (8��39���54��� N; 77��27���15��� E), 68 m elevation, collected on 12 November 2014 by V. Deepak and R. Chaitanya. BNHS 2313 an adult female from Kanyakumari, Kanyakumari District, Tamil Nadu, India (8��4���45��� N 77��31���55��� E), 68 m elevation, collected on 10 January 2014 by V. Deepak Diagnosis. A large-sized Sitana with a maximum SVL of 55 mm, distinguished from all congeners by the following combination of characters: 1) dewlap extending beyond forearm insertion (vs not extending in S. sivalensis complex including S. kalesari); 2) highly serrated dewlap (vs faintly serrated in S. spinaecephalus and S. laticeps); 3) large dewlap extending over up to 64% of trunk vs S. ponticeriana (46%), S. bahiri (29%), S. devakai (33%) and S. visiri (56%), 4. Large body size (average SVL) 51 mm vs S. bahiri (Rameshwaram population 42.1 mm; Sri Lankan population 34 mm), S. devakai 35 mm, S. visiri 50 mm and S. ponticeriana 40.5 mm, 5. Fewer dorsal scales (39���40) vs S. visiri (41���50). Description of holotype. The holotype is in good condition, hemipenis partially everted, exposed and seen on both sides when viewed dorsally. Tail entire, curved towards right side. An adult male, SVL 55 mm. Head relatively long (HL/SVL ratio 0.27), wide (HW/HL ratio 0.63), not depressed (HH/HL ratio 0.56), distinct from neck. Snout short (SE/HL ratio 0.38), longer than eye diameter (OD/SE ratio 0.77), obtusely pointed in profile when viewed dorsally; rostral much wider than high, contacted laterally on either side by first supralabial and a large prenasal, dorsally by three smaller scales. Canthus rostralis and supraciliary edge sharp. Nostril elliptical, laterally positioned and placed in the centre of a large, undivided, pentagonal nasal plate. Nasal plate bordered by six scales on each side, two postnasals, one supranasal, one prenasal (separating it from rostral), first supralabial and one more enlarged scale, separating the nasal from the second supralabial. Seven supralabials on the right side (eight on left side), first twice as large as others, broader than long, roughly rectangular, strongly keeled, bordered above by a row of slightly smaller, rectangular, keeled scales, which start at posterior margin of first supralabial and terminate above the seventh. Loreal region concave, with scales of heterogeneous shape and size. Canthals enlarged, overlapping, keeled, slightly protruding on supraorbital ridge laterally. Eye large (ED/HL ratio 0.20); pupil rounded, eyelids covered with scales that are heterogeneous in shape and size, larger ones on the upper eyelid keeled; supraciliaries longer than broad. Orbital scales small but not granular. Scales on postorbital and temporal region homogenous, imbricate, strongly keeled, and directed backward and upwards. Tympanum naked. Canthal scales 4 on both sides. Orbit bordered below by a row of nine roughly rectangular, elongate, weekly keeled scales starting from posterior margin of nasal scale and terminating after the posterior margin of the orbit. Scales on dorsal surface of snout, forehead, interorbital and occipital region heterogeneous in size, shape, mostly elongate, imbricate, strongly keeled longitudinally; those on snout smaller, rhomboidal, those on forehead large, much elongate; supraorbital scales along the supraciliary edge elongate, keeled, decreasing in size posteriorly, following curvature of orbit; occipital region with slightly smaller, less elongate, imbricate and keeled scales. Parietals larger than surrounding scales, longer than broad, strongly keeled, completely divided by two smaller inter-parietal scales; anterior inter-parietal bicarinate, posterior elliptical, keeled, without visible pineal eye. Mental shield narrower than rostral, roughly pentagonal, pointed posteriorly; scales on the gular region keeled. Dewlap large (DEWL /TrL ratio 0.61), extends posteriorly over 60% of trunk, with posterior scales extending much beyond axilla, about seven to eight rows of anterior dewlap scales (colour is bluish in life & blackish in the specimen) smaller, elongate, pointed, keeled, remainder of scales much enlarged, keeled, lanceolate, bluntly pointed, gradually increasing in size towards margin; single marginal row largest with sharply pointed scales. Enlarged scales on dewlap in 21 rows. Nuchal and dorsal crest absent. Scales on nuchal region smaller, less than half the size of those on interorbital region, imbricate, strongly keeled. Body slender, 55 rows of scales around midbody; 10 to 12 dorsal rows including the vertebral scale row, starting from back of neck until groin, homogenous in size, shape, slightly larger than those on neck, imbricate, pointed, keeled, directed backwards forming regularly arranged longitudinal rows; those on flanks heterogeneous in size, shape, smaller than those on back, pointed, keeled, irregularly scattered with larger, pointed, keeled scales; scales of upper rows on flanks directed backwards and upwards, lower rows backwards and downwards; ventral scales subimbricate, keeled, homogenous in size and shape, arranged in 80 rows; no precloacal or femoral pores. Fore and hindlimbs relatively slender, tibia short (CL/ SVL ratio 0.33); digits moderately long, ending in strong, elongate, slightly recurved claw; inter-digital webbing absent; subdigital lamellae entire, tri-mucronate, 23 subdigital lamellae on toe IV; relative length of fingers 3> 4> 2> 5> 1, toes 4> 3> 2> 1. Fore and hindlimbs covered above and below with regularly arranged, enlarged, pointed, strongly keeled, scales. Tail long (TL/SVL ratio 2.7), entire, base swollen, uniformly covered with similar sized, keeled, pointed, regularly arranged, backwardly directed imbricate scales; subcaudal scales keeled, rounded near base, becoming pointed posteriorly; no enlarged subcaudal row. Colour of holotype in life: Dorsum light to dark brown with a black patch on the neck, three brown rhomboidal markings on the trunk, relatively lighter than the one on the neck. A black ���V��� shaped marking between the eyes. Limbs and tail with dark brown or black bands of variable width. A cream-coloured streak begins behind the nostril, extending below the eye and into flank region. Belly pale white (iridescent) in colour. Both forelimbs, hindlimbs, and tail have dark brown bars of variable width. Dewlap with dark blue colouration from the mental extending up to the anterior half margin of the dewlap, bright orange patches or spots present on the plain iridescent white dewlap; bright yellow patch on either side of dewlap starting approximately in the middle of lower jaw. Colour of holotype in preservative: A dark roughly ���V��� shaped band across the interorbital region of head. Dorsum dark brown with two much darker blotches on either side of neck. Vertebral scale row much lighter than dorsolateral scales. Flank much darker with light brown markings mostly on enlarged scales. Dark brown bars on limbs and tail, faintly visible. The dewlap colouration is faint. Variation. The paratypes agree with the holotype in overall scalation except MBS (CES 141124 has 55 scale rows vs 52), VS (BNHS 2313 has 39 scales vs 40) and LAM 4 (BNHS 2313 has 22 lamellae vs 23). Morphometric data for the paratypes are summarized in Table 3. Morphological characters of females mostly agree with males except for the absence of a dewlap, lack of a swollen tail base, and a narrower head. In the type series, the nasal scale always touches the first supralabial. However, this character is not seen in other referred specimens (Appendix 1), where the nasal scale is separated from the first supralabial by the first scale in a row of enlarged scales above the supralabials. Etymology. The species is named for its occurrence in the grassy plains as well as the seashore in Tamil Nadu, India. The specific epithet is derived from a combination of two ancient Tamil words (Tamil: Marudham = cultivable grasslands, Neydhal = land by the seashore). Hemipenial morphology. One specimen from Kallidaikurichi (CES 141196) was examined. The hemipenis is bilobed, divided almost half its length (Fig.8 A), relatively small, as wide as long, and shallowly forked. Base naked. Sulcus spermaticus single (Fig. 8 B). Lips smooth. Calyces present on each lobe. Proximal half of sulcal and asulcal side with small deep polygonal calyces and apex unornamented. Ornamentation includes a combination of flounces and calyces. Papillae are present between the apical lobes. Ridges between the calyces are thin and show scalloped micro-ornamentation. Eight flounces present on the asulcal side (Fig. 8 A) and 10���11 flounces are present on the sulcal side (Fig. 8 B). Natural history. The new species Sitana marudhamneydhal sp.nov. is found in coastal sand dunes, grassland habitats, open areas dominated with Prosopis juliflora and plains in the southern Tamil Nadu (Fig. 9). It is so far recorded only south of the Thamirabarani river in Tamil Nadu. Other diurnal lizard species found in these habitats include Eutropis carinata, Eutropis bibronii, and Calotes versicolor. Breeding males were observed during the month of September and October. Hatchlings were recorded during the month of January., Published as part of Deepak, V., Khandekar, Akshay, Varma, Sandeep & Chaitanya, R., 2016, Description of a new species of Sitana Cuvier, 1829 from southern India, pp. 167-182 in Zootaxa 4139 (2) on pages 171-178, DOI: 10.11646/zootaxa.4139.2.2, http://zenodo.org/record/261330