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Additional file 1: Figure S1. Maximum likelihood(ML) phylogeny of Himalayan langur for mitochondrial cytochrome b (Cyt-b) gene.Numbers at the node indicate ML bootstrap (MLBS) values. Node support values of > 75 are shown. Node support values only for major clades are shown.

Additional file 2: Figure S2. Divergence time tree for the mitochondrial cytochrome b (Cyt-b) dataset constructed in BEASTv2.6. The bars at the nodes indicate the 95% credible interval. Details of the collapsed IHR node can be found in Additional file 4: Table S3. IHR = Indian HimalayanRegion; WMP = Western metapopulation; EMP = Eastern metapopulation.

Additional file 4: Table S1. Samples used for molecular phylogenetic analysis. Samples that are marked with † were sequenced in this study. Samples with the same accession numbers are identical sequences. Table S2. Sequences used for divergence dating analysis in BEAST v2.6.6. Sequences that are marked with † were sequenced in this study. Sequences from sr. no. 1–66 constitutes the collapsed clade IHR in Additional file 2: Fig. S2. Table S3. Model selection for Maxent analysis—the table shows AUC values for different models. AUC values in bold shows the features and RM values selected.

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.

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."]}

Island systems from around the world have provided fascinating opportunities for studies pertaining to various evolutionary processes. One recurring feature of isolated islands is the presence of endemic radiations. In this regard, the Indian subcontinent is an interesting entity given it has been an island during much of its history following separation from Madagascar and currently is isolated from much of Eurasia by the Himalayas in the north and the Indian Ocean in the south. Not surprisingly, recent molecular studies on a number of endemic taxa from India have reported endemic radiations. These studies suggest that the uniqueness of Indian biota is not just due to its diverse origin, but also due to evolution in isolation. The isolation of India has generated some peculiarities typically seen on oceanic islands. However, these patterns might be confined to, groups with low dispersal ability.

Sphenomorphus apalpebratus sp. nov. (Figures 1–4) Holotype. CES 10 / 830 (adult, female), from Mawphlang (25.44563329 °N, 91.7428503 °E, alt. 1,815 m asl; datum WGS 84), East Khasi Hills District, Meghalaya State, north-east India (Fig. 6). 17 October 2010 (Aniruddha Datta- Roy, Ishan Agarwal, Ronald K. Lyngdoh Tron, N. P. I. Das and Tarun Khichi, collectors). Paratypes. CES 10 / 831 –CES 10 / 833 (unsexed juveniles). Collection data as for holotype. Diagnosis. We allocate the lizard specimens collected from Mawphlang to the genus Sphenomorphus for showing the following characters: parietals meet behind interparietals; median preanals overlap outer preanals; and iris as dark as pupil, considered apomorphies of the Sphenomorphus group (Greer 1979), in addition to the absence of supranasals; five digits on fore- and hindlimbs, limbs well developed, and body elongated, but non-vermiform. Further, the new species can be differentiated from congeners from India and mainland south-east Asia in showing the following characters: inner preanals overlapping the outer ones, small body size (SVL to 42.0 mm); midbody scale rows 27–28; longitudinal scale rows between parietals and base of tail 62–64; subdigital lamellae of toe IV 8– 9; supraoculars five; supralabials 5–6; infralabials 4–5; subcaudals 92; and dorsum golden brown, except at dorsal margin of lateral line, which is lighter, with four faintly spotted lines, two along each side of vertebral row of scales, extending to tail base. Etymology. Latin for lacking eyelids, a distinctive morphological character in the new species. Nomenclatural Notes. The name of the genus Sphenomorphus Fitzinger 1843 (type species: Gongylus [Lygosoma] melanopogon Duméril & Bibron 1839) was derived from Greek (meaning wedge-shaped, an allusion to the shape of the head) and originally coined as masculine. However, a number of species, some of which were transferred from the catchall lygosomatine genus Lygosoma (of neuter gender), have name terminations not matching the gender of their current generic allocation, in recent lists (e.g., Uetz 2012). These include S. crassa Inger et al. 2001, S. mimikanum (Boulenger 1914) and S. nigrolineata (Boulenger 1897). Following Article 30.1. 3 of the International Code of Zoological Nomenclature (International Commission of Zoological Nomenclature 1999), the termination of these species nomina requires emendation. Description of holotype. Based on an adult female, SVL 42.0 mm, TL 65.0 mm; snout blunt (IN:IO ratio 0.35), not projecting beyond lower jaws; nostril laterally oriented; oval, situated closer to snout-tip than to orbit; eye-nostril distance 1.44 mm, eye-snout distance 2.32 mm (E-N:E-S ratio 0.62); head long, much longer than wide (HL 6 mm, HW 4.68 mm (HL:HW ratio 1.29); head shape slightly flattened, HD 4 mm (HL:HD ratio 1.5); rostral width (1.28 mm), broad, not projecting onto snout; posterior border of rostral curved; frontonasal trapezoidal, wider than long; frontal elongated, arrow-shaped, wider anteriorly; frontoparietals separate; prefrontals small, widely separated; interparietal single, smaller than frontonasal; parietals contact posterior to interparietal, half length of interparietal; parietal eye spot absent; five supraoculars; four supraciliaries; nostrils located in midnasal; postnasal absent; supranasals absent; loreals two, trapezoidal; anterior loreal higher than posterior; two presuboculars, separating supralabial III from orbit; six supralabials (supralabial IV contacting orbit); three postsuboculars; three postoculars; two pretemporals; three temporals, the upper secondary temporal largest; upper secondary temporals overlapped by lower; temporals larger than lateral body scales; one pair of nuchals; five infralabials; one scale separates second pair of enlarged chin shields; three scales separate third pair of chin shields; enlarged chin shields contact infralabials; auricular opening scaleless, its location indicated by a shallow depression; eyes relatively small; pupil not discernable in preserved specimen; no moveable eyelids (Fig. 2); upper palpebrals 11; lower palpebrals 12; tongue short; undivided anteriorly, tip obtuse, not pointed; teeth relatively small and somewhat blunt. Body slender, BW 6.36 mm (BW:SVL ratio 0.15); head slightly distinct from neck and body; 64 longitudinal scale rows from parietal to above level of anterior margin of hind limb; 64 scales dorsally; 60 ventrals, counted from first postgular to last scale before preanals; body scales smooth, subcycloid; median rows not enlarged, as wide as adjacent scales; 28 transverse scale rows at midbody; 92 subcaudals; abdominal scales larger than throat scales and marginally larger than pectoral scales; median ventral scales enlarged relative to scales on flanks; median and lateral preanals enlarged, larger than adjacent ventrals (Fig. 4); median pair of preanals overlap lateral preanals and are smaller than lateral ones; tail rounded, relatively long, longer than snout-vent length (TL:SVL ratio 65.42); tail tip acute; tail base wider than rest of tail; tail gradually tapering to a point; median row of subcaudals enlarged. Visceral fat bodies present in abdomen. Limbs short and pentadactyle, digits short and clawed; lamellae smooth, enlarged; adpressed limbs fail to meet; lamellae under finger I- 2; II- 3; III- 5; IV- 5; V- 4; lamellae under toes I- 3; II- 6; III- 8; IV- 9; V- 5. Coloration (in preservative). Lateral side of head with brown-black flecks; radiating broken lines from tympanic region to axilla; broken line, in the form of spots, extending from nostril to preocular region, continuing after orbit to almost two-thirds along the length of tail. On flanks, dark stripe fuses with its partner, dorsum olive brown (Medium Spring bud, colour no. C 9 DC 87), with four faint lines, two on each side of the vertebral row of scales. Lines start at row of scales just below single pair of nuchals, and continue to tail base, where they become diffused. Dorsal surface of tail marginally darker than rest of dorsum; tail tip (regenerated) slightly lighter than rest of tail; flanks speckled with brown-black below lateral line; venter anterior to preanal scales lacking dark pigment, gulars cream coloured; pectoral to preanal region bright yellow; venter of tail grey, spotted with brown-black speckles to tip, darker apically. Tail venter with lighter pigments than dorsal side. Both fore- and hind limbs heavily spotted on dorsal side and lightly spotted ventrally. Lamellae chocolate brown in colour. Coloration (in life). Lateral side of head freckled with patches of brown-black spots. A few radiating spotted lines extend from tympanic region to insertion of forelimb. A spotted line on lateral side of head from nostril to preocular, continuing beyond orbit to nearly two-thirds along the length of tail. Upper and lower palpebral series lemon yellow in colour, except at anterior and posterior corners of eye. Iris and pupil not clearly discernible. Head scales edged with black spots. Dorsum golden brown (Gamboge colour no. E 49 B0F), except at dorsal margin of lateral line, which is lighter. Four faintly spotted lines on dorsum, two along each side of vertebral row of scales, that continue to tail base; dorsum of tail darker than that of trunk at midbody; sides of head from snout to auricular depression, to below orbit cream coloured, gradually becoming pink around midbody between posterior margin of forelimb to anterior margin of hindlimb; yellowish near tail base. Flanks freckled with brownish-black longitudinal spots that are denser towards hindlimb than immediately behind forelimb. Dorsal surface of forelimb and hindlimb heavily spotted and appear dark. Gulars to 17 th ventral scale cream-coloured and devoid of dark spots. Starting from 18 th ventral row of scales, colour abruptly becomes bright lemon yellow until preanal scales or ventral margin of tail base. Yellow spots more dense in midregion of ventral scales, gradually becoming non-pigmented at scale boundary. Ventral side of forelimbs less spotted and pale pink in colour, ventral side of hindlimbs lemon yellow, with marginally more brownish-black spots. Toe lamellae of both limbs dark, unspotted. Preanal region, a light shade of pink, gradually fading to white and turning cobalt blue towards tail tip; venter of tail with brownish-black flecks, becoming denser apically. Measurements (in millimeters; holotype with variation shown in paratype series, all juveniles, in parentheses). SVL 42.00 (range 26.76–27.50, mean 26.90); HL 10.00 (range 4.32–5.10 range, mean 4.73); HW 5.50 (range 4.10–4.94, mean 4.65); HD 4.00 (range 2.98 – 2.98, mean 2.98); BW 6.36 (range 4.10–4.60, mean 4.37); TBL 3.38 (range 2.24–2.46, mean 2.35); ED 1.46 (range 1.00– 1.16, mean 1.05); IN 1.24 (range 1.00–1.00, mean 1.00); E– S 2.32 (range 1.42–1.80, mean 1.59); E– N 1.44 (range 1.00– 1.20, mean 1.11); N– S 0.70 (range 0.40–0.48, mean 0.44); A–G 25.82 (range 14.50–15.58, mean 15.03); and TL 65.00 (tail tips of paratypes clipped for DNA extractions). Squamation (holotype with variation shown in paratype series, all juveniles, in parentheses). Transverse scale rows at midbody 28 (range 27–28, mean 27.33); longitudinal scale rows 64 (range 62–64, mean 63); ventral scale rows 60 (range 56–58, mean 57); supralabials 6 (range 5 – 5, mean 5); infralabials 5 (range 4 – 4, mean 4); subcaudals 92 (tail of paratypes clipped for DNA extraction); and lamellae under toe IV 9 (range 8–9, mean 8.33)., Published as part of Datta-Roy, Aniruddha, Das, Indraneil, Bauer, Aaron M., Lyngdoh Tron, Ronald K. & Karanth, Praveen, 2013, Lizard Wears Shades. A Spectacled Sphenomorphus (Squamata: Scincidae), from the Sacred Forests of Mawphlang, Meghalaya, North-east India, pp. 257-276 in Zootaxa 3701 (2) on pages 259-264, DOI: 10.11646/zootaxa.3701.2.7, http://zenodo.org/record/218482

A new species of lygosomatine scincid lizard is described from the sacred forests of Mawphlang, in Meghalaya, northeastern India. Sphenomorphus apalpebratus sp. nov. possesses a spectacle or brille, an unusual feature within the Scincidae, and a first for the paraphyletic genus Sphenomorphus. The new species is compared with other members of the genus to which it is here assigned, as well as to members of the lygosomatine genera Lipinia and Scincella from mainland India, the Andaman and Nicobar Islands, and south-east Asia, to which it also bears resemblance. The new taxon is diagnosable in exhibiting the following combination of characters: small body size (SVL to 42.0 mm); moveable eyelids absent; auricular opening scaleless, situated in a shallow depression; dorsal scales show a line of demarcation along posterior edge of ventral pes; midbody scale rows 27-28; longitudinal scale rows between parietals and base of tail 62-64; lamellae under toe IV 8-9; supraoculars five; supralabials 5-6; infralabials 4-5; subcaudals 92; and dorsum golden brown, except at dorsal margin of lateral line, which is lighter, with four faintly spotted lines, two along each side of vertebral row of scales, that extend to tail base. The new species differs from its congeners in the lack of moveable eyelids, a character shared with several distantly related scincid genera.

Numerous morphology-based classification schemes have been proposed for langurs and leaf monkeys of South Asia but there is very little agreement between them. An incorrect classification scheme when used as a basis for biogeographic studies can support erroneous hypotheses. Further, lack of taxonomic resolution will also confound conservation efforts, given that conservation biologists use traditional morphology-based-classification schemes to prioritize species for conservation. Here, I have revisited recent molecular phylogenetic studies done on langurs and leaf monkeys of South Asia. Results from these studies are in turn used to derive a rational and scientific basis for prioritizing species for conservation. Molecular data support the classification of langurs of the Indian subcontinent-Hanuman, Nilgiri and purple-faced langurs-in the genus Semnopithecus, whereas Phayre's leaf monkey along with other Southeast Asian leaf monkeys form another distinct clade (Trachypithecus). The phylogenetic position of capped and golden langurs remains unresolved. Molecular data suggest that they are closely related to each other but this group might have evolved through past hybridization between Semnopithecus and Trachypithecus. Additionally, genetic data also support the splitting of the so-called Hanuman langurs into at least three species. The scores for taxonomic uniqueness of langurs and leaf monkeys of South Asia were revised using this molecular phylogeny-based classification. According to the revised scores, Phayres leaf monkey and golden langur are priority species for conservation followed by capped and Nilgiri langurs.

A recent phylogenetic study of langurs and leaf monkeys of South Asia suggested a reticulate evolution of capped and golden leaf monkeys through ancient hybridization between Semnopithecus and Trachypithecus. To test this hybridization scenario, I analysed nuclear copies of the mitochondrial cytochrome b gene (numts) from capped, golden and Phayre's leaf monkeys. These numts were aligned with mitochondrial cytochrome b sequences of various species belonging to the genera Semnopithecus and Trachypithecus. In the phylogenetic tree derived from this alignment, the numts fell into three distinct clades (A, B and C) suggesting three independent integration events. Clade A was basal to Semnopithecus, and clades B and C were basal to Trachypithecus. Among the numts in clades A and C were sequences derived from species not represented in their respective sister mitochondrial groups. This unusual placement of certain numts is taken as additional support for the hybridization scenario. Based on the molecular dating of these integration events, hybridization is estimated to have occurred around 7.1 to 3.4 million years ago. Capped and golden leaf monkeys might have to be assigned to a new genus to reconcile their unique evolutionary history. Additionally, northeast India appears to be a `hot spot' for lineages that might have evolved through reticulate evolution.

Many hypotheses have been proposed to explain the presence of Gondwanan lineages in tropical Asia including, the out-of-India hypothesis, the Eurasian route hypothesis, long-distance transoceanic dispersal and the boreotropical hypothesis. Recent molecular studies support an out-of-India Gondwanan origin of some tropical Asian taxa. Specifically these studies suggest that ancient Gondwanan lineages survived on peninsular India in spite of dramatic climatic changes and Late Cretaceous volcanism. Under this model, Gondwanan lineages dispersed into Asia when peninsular India collided with Asian plate. These Gondwanan lineages represent the remnants of unique ancient biota that require urgent attention from conservationists if we are to preserve the overall evolutionary history of Indian biota.