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Microauris gen. nov. (Fig. 8b) Type species. Calotes aurantolabium (Krishnan, 2008) Etymology. The genus epithet is derived by adding the word ‘Micro’ as a prefix to the Latin word ‘auris’ meaning ear, referring to the extremely small tympanum of this genus. Diagnosis. Microauris gen.nov. differs from all other members of draconinae lizards from the Indian subcontinent in having the smallest tympanum. Microauris gen.nov. differs from Mantheyus phuwuanensis by the absence of femoral pores (Manthey & Nabhitabhata, 1991; Ananjeva & Stuart 2001); from the members of the genus Bronchocela by the absence of a fold of skin from angle of jaw to shoulder (Hallermann & Böhme 2000). Microauris gen. nov. can be easily diagnosed from the genera Otocryptis, Sarada and Sitana by the presence of a well-developed fifth toe (Smith, 1935; Deepak et al, 2016); from Cophotis, Ceratophora, Lyriocephalus, Ptyctolaemus, Phoxophrys, Japalura otai, J. planidorsata, J. sagittifera and Otocryptis by the presence of external tympanum (Boulenger, 1885; Smith, 1935; Inger, 1 960; Pethiyagoda & Manamendra-Arachchi 1998; Schulte II et al. 2004; Bahir & Silva 2005; Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006). Microauris gen. nov. can be diagnosed from other members of genus Japalura by the absence of heterogenous dorsal scales and short and thick nuchal scales; from Salea by the absence of enlarged plate like scales between the eye and tympanum (Smith, 1 935). Microauris gen. nov. can be diagnosed from the members of Psammophilus and Monilesaurus by the absence of a well developed antehumeral fold. Microauris gen. nov. can be diagnosed from Complictus nigrigularis, Hypsicalotes kinabaulensis, Malayadracon robinsonii, Oriocalotes, Pseudocophotis and Pseudocalotes by the absence of enlarged row of suborbital scales (Smith, 1935; Taylor 1963; Ota & Hikida, 1991; Manthey & Denzer 1992; Inger & Steubing 1994; Ota & Hikida, 1996; Hallermann & Böhme 2000; Manthey & Denzer, 2000; Hallermann & McGuire 2001; Leong 2001; Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006; Ananjeva et al. 2007; Hallermann & Böhme 2007; Das & Lakim 2008; Hallermann et al. 2010; Mahony 2010; Harvey et al. 2014; Denzer et al. 2015; Grismer LL et al. 2016; Harvey et al. 2017). Microauris gen.nov. can be distinguished from the members of the genus Calotes (C. bachae, C. bhutanensis, C. calotes, C. ceylonensis, C. chincollium, C. desilvai, C. emma, C. grandisquamis, C. hutunwini, C. irawadi, C. jerdoni, C. liocephalus, C. liolepis, C. versicolor, C. manamendrai, C. maria, C. medgoensis, C. minor, C. mystaceus, C. nemoricola, C. nigrilabris, C. nigriplicatus and C. pethiyagodai) and Lophocalotes (Günther, 1872) in having the smallest tympanum (Hardwicke & Gray, 1827; Duméril and Bibron, 1837; Gray, 1845; Jerdon, 1854; Günther, 1864; Günther, 1870, Günther, 1872, Günther, 1875; Boulenger, 1885; Biswas, 1975; Zhao & Li, 1984; Hallerman, 2000; Vindum et al. 2003; Hallerman et al. 2004; Bahir & Maduwage, 2005; Zug et al. 2006; Manthey, 2008; Krishnan, 2008; Hartmann et al. 2013; Amarasinghe et al. 2014a, b; Deepak et al. 2015). Suggested English. Small-eared dragon Holotype. BNHS 1436, an adult female collected from Kalakkad Mundanthurai Tiger Reserve, Tamil Nadu, India; by N. M. Ishwar in 1997. Referred specimens. CESL 104, an adult female collected from a branch in the canopy of a shola forest tree near Chemunji peak, Peppara Wildlife Sanctuary, Kerala, India; by SPP, SPV, MVP, VTR and ADR in 18th August 2010. Taxonomic comments. Microauris aurantolabium comb. nov. was described as Calotes aurantolabium in 2008 based on a single specimen (Krishnan 2008). Prior to this, the specimen was wrongly identified and reported as the rediscovery of Calotes andamanensis. (Ishwar and Das, 1998). During our fieldwork in the Agasthyamalai hills of Kerala in August 2010, we found a single female specimen sleeping on a branch of a tree in a shola forest. There have been no other reports of this species thereafter and thus all the known specimens are females. There is a clear gap in the knowledge of the morphology of this rare lizard considering that male lizards are known to be better representatives for morphological characterization because of the presence of secondary sexual characters. In spite of this, the two specimens share some common characters, which are unique to this lizard. Based on the presence of unique morphological characters along with the phylogenetic position of this species in the subfamily Draconinae, we designate this species as a new monotypic genus. Diagnosis. A medium sized light green agamid lizard with a distinct orange streak from above the lip scales till behind the jaw; a moderately broad head with a pointed snout; very small tympanum relative to the orbit diameter (TymD/OrbD = 0.14–0.19); acutely keeled scales over the body and throat; scales on head small, sub-triangular (vs. uniform shield like in members of genus Calotes); 3rd and 4th toe almost equal in length; dorsal and lateral body scales oriented backwards and downwards; nuchal crest indistinct, poorly developed; supratympanic and postorbital spines absent and a long and slender tail. Description of the Holotype. BNHS 1436; a medium sized agamid, adult female (SVL- 68.5 mm) Morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately compressed. Head moderate (HL/SVL ratio 0.28), broad (HW/HL ratio 0.56), maximum height less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield, which is separated from rostral by a single scale; mental shield almost as wide as the rostral; two postmentals; first pair touching each other at the base; genials partially keeled; gular sac small, composed of partially keeled scales, slightly smaller than genials; scales on the snout smooth, unkeeled; scales on top of head heterogenous, keeled, small, subtriangular; supraorbital scales keeled; orbit diameter 95% of distance between anterior border of orbit and snout tip; tympanum very small, exposed, its greatest diameter 11% of horizontal diameter of orbit; slightly keeled scales between tympanum and orbit; posterior region of jaws partially swollen; supralabials 10/10; infralabials 11/11. Nuchal crest poorly developed, composed of 13 small conical spines; the remaining vertebral scales slightly enlarged relative to adjacent rows and possess a more pronounced median keel; 54 longitudinal scale rows around midbody; scales on dorsum keeled; lateral scales smaller than dorsal, keeled, oriented postero-ventrally; ventrals slightly larger then dorsals, genials and gular scales strongly keeled. Limbs slender and covered with strongly keeled scales, larger than laterals; scales under thighs keeled; length of hindlimb ca. 64% SVL; relative lengths of toes 4=3>5>2>1; fourth toe and third toe subequal; 20 subdigital lamellae under fourth finger; 21 subdigital lamellae under fourth toe; tail slender, long; scales on dorsal and ventral surface of tail with sharp keels, mucronate, slightly larger than laterals; tail length 117 mm. Colouration. In life (CESL 104): dorsum and head uniform, light green overall with a distinct reddish orange streak from above the third lip scale covering 2–3 scale rows below the tympanum, ending just at the shoulder. A distinct pale whitish grey band, edged with black speckles from the base of tail till underside of knee. Mid-dorsal scales edged anteriorly with black, which is visible when the body is inflated; gular scales lighter, yellowish green; ventrally brighter, uniform green with a few rows of pale whitish scales towards the lateral side. This lizard was observed to quickly change its body color to dark blackish-brown on disturbance but the orange streak and band on thigh remained unchanged. In preservative: dorsum and head uniform, overall pale blue with brownish patches above the eyes and neck continuing on the mid-dorsal line, broader towards the tail; tail pale brown; orange lip streak paler turning whitish towards the neck; laterally pale blue with anteriorly black edged scales; ventrally pale whitish grey overall; limbs edged with brown blotches, broader towards the tips. Distribution. Currently, this lizard is known only from two sites in the high elevation tropical evergreen forests of Agasthyamalai hills. The second locality near Chemunji peak in Peppara wildlife sanctuary is approximately 50 kms northeast of the type locality near Kakachi in KMTR (See Appendix 1 for details). Both these sites are separated by almost contiguous shola grassland mosaic on undulating hills. Ecology and natural history. Nothing much is known about the ecology and natural history of this species. Ishwar and Das (1998) reported sightings of 3 individuals, all gravid females in the month of August. Two of these lizards laid 4 elliptical eggs each. This suggests that monsoon might be a breeding season for this lizard. Ishwar and Das (1998) also mentioned that this is a canopy dwelling species. The fact that it was only found again 12 years after its discovery and that too from a fairly well explored habitat suggests that this might be a rare lizard or it may occur high in the canopy of rain forests and thus escape detection.

Calotes Cuvier, 1817 The genus Calotes Cuvier, 1817 with the type species Lacerta calotes Linnaeus, 1758 belongs to the subfamily Draconinae of the family Agamidae, containing 26 species. The geographical distribution of this genus is largely restricted to South Asia and Indo-china, except for one isolated species Calotes nigriplicatus Hallermann, 2000 found in Ambon Island in Indonesia. The greatest diversity of the genus is in the Western Ghats and Sri Lanka. Four species of Calotes (C. grandisquamis, C. nemoricola, Calotes cf. versicolor , Calotes calotes) are known from the Western Ghats of which two are endemic to this region. Diagnosis. Medium to large sized agamids with a relatively broader head; dorsal and lateral scales uniform, arranged in rows of backwardly pointed scales; dorsal and nuchal crest more or less developed; groups or rows of spines from eye to above tympanum; often an antehumeral skin fold or pit present covered with small scales; gular sac usually present; tympanum naked; tail long and slender, usually swollen and rounded at the base in males; no preanal or femoral pores. Distribution. Lizards belonging to the Genus Calotes are widely distributed across Iran, Afghanistan, Bhutan, Bangladesh, Maldives, Nepal, India, Pakistan and Sri Lanka as well as southern China; parts of Myanmar, Thailand, Laos, Cambodia, Vietnam, Peninsular Malaysia and parts of Indonesia. Moreover, Calotes versicolor has been introduced into Kenya, Borneo, Sulawesi, Seychelles, Mauritius, Oman and Florida (USA) (Das et al. 2008; Manthey, 2008; Uetz and Ho��ek, 2016). Taxonomic groups. Smith (1935) separated members of the genus Calotes into four distinct groups, I. C. cristatella group, II. C. microlepis group, III. C. versicolor group, and IV. C. liocephalus group, each composed of multiple species and another group composed of two dwarf species based on their morphological similarities. Later, Moody (1980) separated the genus into four different genera, namely Bronchocela Kaup, 1827 (group I of Smith), Dendragama Doria, 1888, Pseudocalotes Fitzinger, 1843 (group II of Smith) and Calotes (group III, IV and the two dwarf species of Smith). Most of the Calotes species occurring in the Western Ghats belong to the Calotes versicolor group of Smith (1935) except Calotes rouxii and Calotes ellioti, which belong to the dwarf group. Here, we transfer these two species to the new genus Monilesaurus gen. nov. In the present study, we follow Smith���s species groups for taxonomic accounts and morphological comparisons. Since Smith, there have been various new additions to the genus Calotes and to the Calotes versicolor group. In this study, we compare each individual species only with members of their own group. Calotes versicolor group. Members of the C. versicolor group are characterized by a medium to large adult body size; body more or less compressed, dorsal and lateral scales large, uniform, smooth or feebly keeled and oriented postero-dorsally; antehumeral fold absent; dorsal scales larger than the ventrals except in C. calotes where it is equal to or slightly smaller than the ventrals; nuchal and dorsal crest well developed, continuous, at least till behind the shoulders; postorbital spine absent except in C. emma Gray, 1845; supratympanic spines in form of paired patches or as a single longitudinal series of spines (Fig. 4 & 5). For taxonomic comparisons, we add the recently designated C. minor to Smith���s C. versicolor group (See Deepak et al. 2015). Content. Calotes bachae Hartmann, Geissler, Poyarkov, Ihlow, Galoyan, R��dder & B��hme, 2013; C. bhutanensis Biswas, 1975; C. calotes; C. ceylonensis M��ller, 1887; C. chincollium Vindum, 2003; C. desilvai Bahir & Maduwage, 2005; C. emma; C. grandisquamis, C. hutunwini Zug & Vindum, 2006; C. irawadi Zug, Grown, Schulte & Vindum, 2006; C. jerdoni G��nther, 1870; C. liocephalus G��nther, 1872; C. liolepis Boulenger, 1885; C. versicolor, C. manamendrai Amarasinghe & Karunarathna, 2014; C. maria Gray, 1845; C. medgoensis Zhao & Li, 1984; C. minor, C. mystaceus Dum��ril & Bibron, 1837; C. nemoricola, C. nigrilabris Peters, 1860; C. nigriplicatus Hallermann, 2000 and C. pethiyagodai Amarasinghe, Karunarathna, Hallermann, Fujinuma, Grillitsch & Campbell, 2014. Diagnosis. All the species belonging to this group can be differentiated from the other groups of Calotes based on the combination of the following characters: From members of the genus Monilesaurus gen.nov., they can be easily distinguished morphologically based on a larger adult size (vs. smaller); dorsal and lateral scales rows directed posterodorsally, much larger in size (vs. posteroventrally, smaller in size); dorsal and lateral scales larger than ventrals, except in C. calotes (vs. not larger than ventrals) (Fig. 4���5 vs 6���7) and absence of antehumeral fold (vs. presence of a well developed antehumeral fold) (Fig. 10). From members of the genus Psammophilus, group they can be distinguished by the presence of compressed body shape (vs. depressed) (Fig. 4���5 vs 8a); absence of an antehumeral fold (vs. antehumeral fold present) (Fig. 10f vs 10g); lateral scales much larger (vs. smaller); presence of well developed nuchal crest (vs. reduced) and lesser number of midbody scale rows, less than 60 (vs. scale rows higher in number, more than 80). From Microauris gen. nov. by having a relatively large tympanum (Fig. 9a vs 9b); scales on head large uniform shield like (vs. small, sub-triangular) (Fig. 10b vs 10d)., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 414-418, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674, {"references":["Manthey, U. (2008) Agamid Lizards of Southern Asia, Draconinae 1. Edition Chimaira, Frankfurt, 160 pp.","Uetz, P. & Hosek, J. (2016) The Reptile Database. Available from: http: // reptile-database. reptarium. cz (accessed 6 May 2016)","Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Fol. II. Sauria. Taylor and Francis, London, 440 pp.","Moody, S. M. (1980) Phylogenetic and historical biogeographical relationships of the genera in the family Agamidae (Reptilia; Lacertilia). Unpublished thesis, The University of Michigan, Ann Arbor, Michigan, 373 pp.","Gray, E. (1845) Catalogue of the specimens of lizards in the collection British museum. Trustees of die British Museum / Edward Newman, London, 289 pp.","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) Fertebrate Zoology, 65 (3), 371 - 381.","Hartmann, T., Geissler, P., Nikolay, A. J., Ihlow, P. F., Galoyan, E. A., Rodder, D. & Bohme, W. (2013) A new species of the genus Calotes Cuvier, 1817 (Squamata: Agamidae) from southern Vietnam. Zootaxa, 3599 (3), 246 - 260. https: // doi. org / 10.11646 / zootaxa. 3599.3.3","Biswas, S. (1975) Reptilia from Bhutan with description of a new species of Calotes Rafinesque. Journal of Bombay Natural History Society, 72, 774 - 777.","Vindum, J. V., Win, H., Thin, T., Lwin, K. S., Shein, A. K. & Tun, H. (2003) A new Calotes (Squamata: Agamidae) from the Indo-Burman range of western Myanmar (Burma). Proceedings of the California Academy of Science, 54 (1), 1 - 16.","Bahir, M. M. & Silva, A. (2005) Otocryptis nigristigma, a new species of agamid lizard from Sri Lanka. In: Yeo, C. J., Ng, P. K. L. & Pethiyagoda, R. (Eds.), Contributions to biodiversity exploration and research in Sri Lanka. The Raffles Bulletin of Zoology, 12 (Supplement), 393 - 406.","Zug, G. R., Brown, H. H., Schulte, J. A. & Vindum, J. V. (2006) Systematics of the Garden Lizards, Calotes versicolor Group (Reptilia, Squamata, Agamidae), in Myanmar: Central Dry Zone Populations. Proceedings of the California Academy of Sciences, 57 (2), 35 - 68.","Gunther, A. (1870) Descriptions of a new Indian lizard of the genus Calotes. Proceedings of the Zoological Society of London, 778 - 779.","Gunther, A. (1872) Descriptions of some Ceylonese Reptiles and Batrachians. Annals Magazine of natural History, 4, 85 - 88. https: // doi. org / 10.1080 / 002229372011951778","Boulenger, G. A. (1885) Catalogue of lizards in the British museum. Fol. 1. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae, Second edition. British Museum (Natural History), London, 436 pp.","Zhao, E. & Li, S. (1984) A new species of Calotes (Lacertilia: Agamidae) from Xizang (Tibet) [in Chinese]. Acta Herpetologica Sinica, 3 (4), 77 - 78.","Dumril, A. M. C. & Bibron, G. (1837) Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Fol. 4. Librairie Encyclopedique Roret, Paris, 570 pp.","Amarasinghe, A. A. T., Karunarathna, D. M. S. S., Hallermann, J., Fujinuma, J., Grillitsch, H. & Campbell, P. D. (2014 b) A new species of the genus Calotes (Squamata: Agamidae) from high elevations of the Knuckles Massif of Sri Lanka. Zootaxa, 3785 (1), 059 - 078. http: // doi. org / 10.11646 / zootaxa. 3785.1.5"]}

Monilesaurus gen. nov. Type species. Calotes rouxii (Duméril & Bibron, 1837) Content. Monilesaurus ellioti comb. nov., Monilesaurus montanus gen. et sp. nov., Monilesaurus rouxii comb. nov. and Monilesaurus acanthocephalus gen. et sp. nov. Etymology. The genus epithet is derived by adding the word ‘Monile’ meaning necklace in Latin referring to the distinct neck fold in this genus and the Greek word sauros meaning lizard which is latinized here as saurus. Diagnosis. Monilesaurus gen. nov. can be easily diagnosed from all members of draconinae lizards from the Indian subcontinent except Psammophilus in having an antehumeral fold, which mostly extends below the dewlap forming a fused fold on the shoulder and the neck. It differs from Psammophilus in having a dorso-laterally compressed body and lower number of scales on the mid-body. Supratympanic spines are present, in the form of two separated spines vs clusters in Calotes and Psammophilus. From Microauris gen. nov. by having a relatively large tympanum (Fig. 9c vs. 9b). Scales on head large uniform shield like (vs small, sub-triangular) (Fig. 10a vs 10d). Monilesaurus gen. nov. can be easily diagnosed from the genera Otocryptis, Sarada Deepak, Karanth & Giri, 2016 and Sitana Cuvier, 1829 by the presence of a well-developed fifth toe (Smith, 1935; Deepak et al. 2016). Monilesaurus gen. nov. differs from Mantheyus phuwuanensis by the absence of femoral pores (Manthey & Nabhitabhata, 1991; Ananjeva & Stuart 2001); from the members of the genus Bronchocela by the presence of postorbital and supratympanic spines (Hallermann & Böhme 2000); from Cophotis, Ceratophora , Lyriocephalus, Ptyctolaemus Peters, 1864, Phoxophrys Hubrecht, 1881 by the presence of external tympanum (Boulenger, 1885; Smith, 1935; Inger, 1960; Pethiyagoda & Manamendra-Arachchi 1998; Schulte II et al. 2004; Bahir & Silva 2005; Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006). Monilesaurus gen. nov. can be diagnosed from Japalura, Gray 1853 by the absence of heterogenous dorsal scales and short and thick nuchal scales; from Salea Gray, 1845 (S. anamallayana and S. horsfieldii) by the presence of small regular lateral scales and the absence of enlarged plate like scales between the eye and tympanum (Smith, 1935); from Complictus nigrigularis (Ota & Hikida, 1991), Hypsicalotes kinabaulensis (de Grijs, 1937), Malayadracon robinsonii (Boulenger, 1908), Oriocalotes (Günther, 1864) Pseudocophotis (Manthey & Grossmann, 1997) and Pseudocalotes by the absence of enlarged row of suborbital scales (Smith, 1935; Taylor 1963; Ota & Hikida, 1991; Manthey & Denzer 1992; Inger & Steubing 1994; Ota & Hikida, 1996; Hallermann & Böhme 2000; Manthey & Denzer, 2000; Hallermann & McGuire 2001; Leong 2001; Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006; Ananjeva et al. 2007; Hallermann & Böhme 2007; Das & Lakim 2008; Hallermann et al. 2010; Mahony 2010; Harvey et al. 2014; Denzer et al. 2015; Grismer LL et al. 2016; Harvey et al. 2017). Taxonomic comments. Monilesaurus rouxii comb. nov. was described as Calotes rouxii (Duméril & Bibron, 1837). The name bearing type is a male specimen housed at MNHN. The type locality of this species is named as “Indes orientales”. Günther (1864) described another similar looking species from southern India, Calotes ellioti. Based on their size similarity Smith (1935) called them dwarf species. Here, we formally transfer these two extant species to this new genus. We also describe two new species belonging to this genus based on similar morphological characters and their phylogenetic position.

Monilesaurus gen. nov. Type species. Calotes rouxii (Duméril & Bibron, 1837) Content. Monilesaurus ellioti comb. nov., Monilesaurus montanus gen. et sp. nov., Monilesaurus rouxii comb. nov. and Monilesaurus acanthocephalus gen. et sp. nov. Etymology. The genus epithet is derived by adding the word ‘Monile’ meaning necklace in Latin referring to the distinct neck fold in this genus and the Greek word sauros meaning lizard which is latinized here as saurus. Diagnosis. Monilesaurus gen. nov. can be easily diagnosed from all members of draconinae lizards from the Indian subcontinent except Psammophilus in having an antehumeral fold, which mostly extends below the dewlap forming a fused fold on the shoulder and the neck. It differs from Psammophilus in having a dorso-laterally compressed body and lower number of scales on the mid-body. Supratympanic spines are present, in the form of two separated spines vs clusters in Calotes and Psammophilus. From Microauris gen. nov. by having a relatively large tympanum (Fig. 9c vs. 9b). Scales on head large uniform shield like (vs small, sub-triangular) (Fig. 10a vs 10d). Monilesaurus gen. nov. can be easily diagnosed from the genera Otocryptis, Sarada Deepak, Karanth & Giri, 2016 and Sitana Cuvier, 1829 by the presence of a well-developed fifth toe (Smith, 1935; Deepak et al. 2016). Monilesaurus gen. nov. differs from Mantheyus phuwuanensis by the absence of femoral pores (Manthey & Nabhitabhata, 1991; Ananjeva & Stuart 2001); from the members of the genus Bronchocela by the presence of postorbital and supratympanic spines (Hallermann & Böhme 2000); from Cophotis, Ceratophora , Lyriocephalus, Ptyctolaemus Peters, 1864, Phoxophrys Hubrecht, 1881 by the presence of external tympanum (Boulenger, 1885; Smith, 1935; Inger, 1960; Pethiyagoda & Manamendra-Arachchi 1998; Schulte II et al. 2004; Bahir & Silva 2005; Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006). Monilesaurus gen. nov. can be diagnosed from Japalura, Gray 1853 by the absence of heterogenous dorsal scales and short and thick nuchal scales; from Salea Gray, 1845 (S. anamallayana and S. horsfieldii) by the presence of small regular lateral scales and the absence of enlarged plate like scales between the eye and tympanum (Smith, 1935); from Complictus nigrigularis (Ota & Hikida, 1991), Hypsicalotes kinabaulensis (de Grijs, 1937), Malayadracon robinsonii (Boulenger, 1908), Oriocalotes (Günther, 1864) Pseudocophotis (Manthey & Grossmann, 1997) and Pseudocalotes by the absence of enlarged row of suborbital scales (Smith, 1935; Taylor 1963; Ota & Hikida, 1991; Manthey & Denzer 1992; Inger & Steubing 1994; Ota & Hikida, 1996; Hallermann & Böhme 2000; Manthey & Denzer, 2000; Hallermann & McGuire 2001; Leong 2001; Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006; Ananjeva et al. 2007; Hallermann & Böhme 2007; Das & Lakim 2008; Hallermann et al. 2010; Mahony 2010; Harvey et al. 2014; Denzer et al. 2015; Grismer LL et al. 2016; Harvey et al. 2017). Taxonomic comments. Monilesaurus rouxii comb. nov. was described as Calotes rouxii (Duméril & Bibron, 1837). The name bearing type is a male specimen housed at MNHN. The type locality of this species is named as “Indes orientales”. Günther (1864) described another similar looking species from southern India, Calotes ellioti. Based on their size similarity Smith (1935) called them dwarf species. Here, we formally transfer these two extant species to this new genus. We also describe two new species belonging to this genus based on similar morphological characters and their phylogenetic position., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 426-427, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674, {"references":["Dumril, A. M. C. & Bibron, G. (1837) Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Fol. 4. Librairie Encyclopedique Roret, Paris, 570 pp.","Deepak, V., Giri, V. B., Asif, M., Dutta, S. K., Vyas, R., Zambre, A. M., Bhosale, H. & Karanth, K. P. (2016) Systematics and phylogeny of Sitana (Reptilia: Agamidae) of Peninsular India, with description of one new genus and five new species. Contributions to Zoology, 85, 67 - 111.","Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Fol. II. Sauria. Taylor and Francis, London, 440 pp.","Manthey, U. & Nabhitabhata, J. (1991) Eine neue Agame, Ptyctolaemus phuwuanensis sp. n. (Sauria: Agamidae), aus Nordost- Thailand. Sauria, 13, 3 - 6.","Ananjeva, N. B. & Stuart, B. L. (2001) The agamid lizard Ptyctolaemus phuwuanensis Manthey and Nabhitabhata, 1991 from Thailand and Laos represents a new genus. Russian Journal of Herpetology, 8 (3), 165 - 170.","Hallermann, J. & Bohme, W. (2000) A review of the genus Pseudocalotes (Squamata: Agamidae), with description of a new species from West Malaysia. Amphibia-Reptilia, 21 (2), 193 - 210. https: // doi. org / 10.1163 / 156853800507372","Boulenger, G. A. (1885) Catalogue of lizards in the British museum. Fol. 1. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae, Second edition. British Museum (Natural History), London, 436 pp.","Inger, R. F. (1960) A review of the agamid lizards of the genus Phoxophrys Hubrecht. Copeia, 3, 221 - 225. https: // doi. org / 10.2307 / 1439661","Pethiyagoda, R. & Manamendra-Arachchi, K. (1998) A revision of the endemic Sri Lankan agamid lizard genus Ceratophora Gray, 1835, with description of two new species. Journal of South Asian natural History, 3 (1), 1 - 50.","Schulte II, J. A., Vindum, J. V., Win, H., Thin, T., Lwin, K. S. & Shein, A. K. (2004) Phylogenetic relationships of the genus Ptyctolaemus (Squamata: Agamidae), with a description of a new species from Chin Hills of Western Myanmar. Proceedings of the California Academy of Sciences, 55, 222 - 247.","Bahir, M. M. & Silva, A. (2005) Otocryptis nigristigma, a new species of agamid lizard from Sri Lanka. In: Yeo, C. J., Ng, P. K. L. & Pethiyagoda, R. (Eds.), Contributions to biodiversity exploration and research in Sri Lanka. The Raffles Bulletin of Zoology, 12 (Supplement), 393 - 406.","Manamendra-Arachchi, K., de Silva, A. & Amarasinghe, T. (2006) Description of a second species of Cophotis (Reptilia: Agamidae) from the highlands of Sri Lanka. Lyriocephalus, 6, 1 - 8.","Samarawickrama, V. A. M. P. K., Ranawana, K. B., Rajapaksha, D. R. N. S., Ananjeva, N. B., Orlov, N. L., Ranasinghe, J. M. A. S. & Samarawickrama, V. A. P. (2006) A new species of the genus Cophotis (Squamata: Agamidae) from Sri Lanka. Russian Journal of Herpetology, 13, 207 - 214.","Gray, E. (1845) Catalogue of the specimens of lizards in the collection British museum. Trustees of die British Museum / Edward Newman, London, 289 pp.","Ota, H. & Hikida, T. (1991) Taxonomic review of the lizards of the genus Calotes Cuvier, 1817 (Agamidae Squamata) from Sabah, Malaysia. Tropical Zoology, 4 (2), 179 - 192. https: // doi. org / 10.1080 / 03946975.1991.10539488","Gunther, A. (1864) The Reptiles of British India. Taylor & Francis, London, 452 pp.","Denzer, W., Gunther, R. & Manthey, U. (1997) Annotated type catalogue of the Agamid lizards (Reptilia: Squamata: Agamidae) in the Museum fur Naturkunde der Humbolt-Universitat zu Berlin (former Zoological Museum Berlin). Mitteilungen aus dem Zoologischen Museum in Berlin, 73 (2), 309 - 332. https: // doi. org / 10.1002 / mmnz. 19970730209","Taylor, E. H. (1963) The lizards of Thailand. University of Kansas Science Bulletin, 44, 687 - 1077.","Manthey, U. & Denzer, W. (1992) The true Angle-head agamas of the genus Gonocephalus Kaup (Sauria: Agamidae) IV. Gonocephalus mjoebergi Smith, 1925 and Gonocephalus robinsonii Boulenger, 1908. Sauria, 1, 15 - 19.","Inger, R. F. & Stuebing, R. B. (1994) First record of the lizard genus Pseudocalotes (Lacertilia: Agamidae) in Borneo, with description of a new species. Raffles Bulletin of Zoology, 42 (4), 961 - 965.","Ota, H. & Hikida, T. (1996) The second specimen of Calotes kinabaluensis de Grijs (Squamata: Agamidae) from Sabah, Malaysia, with comments on the taxonomic status of the species. Journal of Herpetology, 30 (2), 288 - 291. https: // doi. org / 10.2307 / 1565529","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.","Hallermann, J. & McGuire, J. A. (2001) A new species of Pseudocalotes (Squamata: Agamidae) from Bukit Larut, West Malaysia. Herpetologica, 57 (3), 255 - 265.","Leong, T. M. (2001) The first female topotype of the female agamid Pseudocalotes flavigula (Smith, 1924) from Peninsular Malaysia, with description of colouration in life. Hamadryad, 26, 336 - 338.","Ananjeva, N. B., Orlov, N. L., Nguyen, Q. T. & Nazarov, R. A. (2007) A new species of Pseudocophotis (Agamidae: Acrodonta: Lacertilia: Reptilia) from central Vietnam. Russian Journal of Herpetology, 14 (2), 153 - 160.","Hallermann, J. & Bohme, W. (2007) On the validity of two recently described species of the endemic Sri Lankan genus Cophotis Peters, 1861 (Squamata: Agamidae) from the Knuckles Forest Range. Salamandra, 43, 187 - 190.","Das, I. & Lakim, M. (2008) A second specimen of Complicitus nigrigularis (Ota & Hikida, 1991). Hamadryad, 32 (1), 82 - 84.","Hallermann, J., Truong, N. Q., Orlov, N. & Ananjeva, N. (2010) A new species of the genus Pseudocalotes (Squamata: Agamidae) from Vietnam. Russian Journal of Herpetology, 17 (1), 31 - 40.","Mahony, S. (2010) Systematic and taxomonic 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 - 23.","Harvey, M. B., Hamidy, A., Kurniawan, N., Shaney, K. & Smith, E. N. (2014) Three new species of Pseudocalotes (Squamata: Agamidae) from southern Sumatra, Indonesia. Zootaxa, 3841 (2), 211 - 238. https: // doi. org / 10.11646 / zootaxa. 3841.2.3","Denzer, W., Manthey, U., Mahlow, K. & Bohme W. (2015) The systematic status of Gonocephalus robinsonii Boulenger, 1908 (Squamata: Agamidae: Draconinae). Zootaxa, 4039 (1), 129 - 144. https: // doi. org / 10.11646 / zootaxa. 4039.1.5","Harvey, M. B., Shaney, K., Hamidy, A., Kurniawan, N. & Smith, E. N. (2017) A new species of Pseudocalotes (Squamata: Agamidae) from the Bukit Barisan Range of Sumatra with an Estimation of its phylogeny. Zootaxa, 4276 (2), 215 - 232. https: // doi. org / 10.11646 / zootaxa. 4276.2.4"]}

Psammophilus Fitzinger, 1843 Type species: Agama dorsalis (Gray, 1845) Content: Psammophilus dorsalis, Psammophilus blanfordanus (Stoliczka, 1871) Etymology: None provided but probably from Latin “Psammo” meaning sand and “Philus” meaning loving. Diagnosis. The genus Psammophilus can be distinguished from the genera Cophotis, Ceratophora, Lyriocephalus, Ptyctolaemus, Phoxophrys, Japalura otai Mahony 2009, J. planidorsata Jerdon, 1870, J. sagittifera Smith, 1940 and Otocryptis by the presence of an external tympanum (Boulenger, 1885; Smith, 1935; Inger, 1960; Pethiyagoda & Manamendra-Arachchi 1998; Schulte II et al. 2004; Bahir & Silva 2005; Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006); from other members of genus Japalura by the absence of heterogenous dorsal scales and short and thick nuchal scales. The genus Psammophilus can be diagnosed from all other species of draconinae lizards from the Indian subcontinent except Calotes minor, in having a dorso-ventrally compressed body. Psammophilus differs from Calotes minor in having small body scales and higher number of around the body scales 80–150 (vs 48–60). Psammophilus are characterized by a medium to large adult body size (SVL mm to mm); body dorso-ventrally compressed; presence of an antehumeral fold; supratympanic spines present, reduced in size; dorsal and lateral scales rows slightly irregular; nuchal and dorsal crest small, reduced; dorsal and lateral scales small, keeled, scale rows directed postero-dorsally. Psammophilus can be distinguished from its sister genus Monilesaurus gen. nov. and Calotes in having a dorso-ventrally compressed body (vs dorso-laterally compressed) and higher number of scales on the mid-body scales (more than 80 vs less than 65) and reduced nuchal crest (vs. well developed); and from Microauris gen. nov. by having a relatively large tympanum (Fig. 9d vs 9b). Scales on head large uniform shield like (vs small, sub-triangular) (Fig. 10c vs 10d). Supratympanic spines are present, in the form of two separated spines vs clusters in Calotes. Psammophilus are sexually dimorphic, adult males are larger than females and have enlarged cheeks. Psammophilus are also sexually dichromatic; during breeding season males acquire bright yellow and orange colors on the dorsum while females remain dull to dark brown with or without orange/ yellow spots. Taxonomic comments. The two species Psammophilus dorsalis and Psammophilus blanfordanus were earlier placed in the genus Agama (Daudin, 1802) and the now obsolete genus Charasia (Gray, 1845). Smith (1935) transferred the two species Charasia dorsalis and Charasia blanfordanus to the genus Psammophilus. Most likely due to their dorso-ventrally compressed body, Psammophilus was placed among the Agaminae but molecular data suggest that they are nested well within the subfamily Draconinae. Both P. dorsalis and P. blanfordanus have been recorded in the Western Ghats (Smith, 1935). Stoliczka (1871) provided a brief description of P. blanfordanus without mentioning a type locality and later provided a detailed description (Stoliczka, 1872) based on a large number of samples collected from central India (Udaypore west of Chotanagpur, West of Raipore, West of Ranchi) which are currently recognized as syntypes in ZSI with additional syntypes in NMW and ZMB (Das et al. 1998). Stoliczka (1872) also mentions that P. blanfordanus is not uncommon in Parisnath (Parasnath) hill. Psammophilus dorsalis was described by Gray (1831). The key character to diagnose this species is number of scales around the body, which overlap with each other (Boulenger, 1885; Smith, 1935). Boulenger (1885) used a combination of specimens from Southern India (Nilgiris, Malabar, Madras and a few specimens with locality “ India ”) to describe P. dorsalis. To describe P. blanfordanus, Boulenger (1885) used specimens from Godavari valley and “Jeypore” (Northern Andhra) and Ranchi (now in Jharkhand state). One single specimen collected from Parasnath, Jharkhand that is relatively close to the type locality (Central India) of P. blanfordanus had 1–3% genetic divergence in the 16S gene from P. dorsalis samples used in this study. We were unable to find P. blanfordanus from the Western Ghats during our fieldwork. Given the amount of morphological variation among the observed individuals and wide distribution of this group, we would suggest a thorough evaluation of this species group from the entire peninsula with detailed sampling. In the present work, we only describe P. dorsalis as we were unable to confirm the presence of P. blanfordanus in the Western Ghats. Members. Psammophilus dorsalis and Psammophilus blanfordanus., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 437-438, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674, {"references":["Gray, E. (1845) Catalogue of the specimens of lizards in the collection British museum. Trustees of die British Museum / Edward Newman, London, 289 pp.","Stoliczka, F. (1871) Notes on new or little known Indian lizards. Proceedings of the Asiatic Society of Bengal, 9, 192 - 195.","Boulenger, G. A. (1885) Catalogue of lizards in the British museum. Fol. 1. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae, Second edition. British Museum (Natural History), London, 436 pp.","Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Fol. II. Sauria. Taylor and Francis, London, 440 pp.","Inger, R. F. (1960) A review of the agamid lizards of the genus Phoxophrys Hubrecht. Copeia, 3, 221 - 225. https: // doi. org / 10.2307 / 1439661","Pethiyagoda, R. & Manamendra-Arachchi, K. (1998) A revision of the endemic Sri Lankan agamid lizard genus Ceratophora Gray, 1835, with description of two new species. Journal of South Asian natural History, 3 (1), 1 - 50.","Schulte II, J. A., Vindum, J. V., Win, H., Thin, T., Lwin, K. S. & Shein, A. K. (2004) Phylogenetic relationships of the genus Ptyctolaemus (Squamata: Agamidae), with a description of a new species from Chin Hills of Western Myanmar. Proceedings of the California Academy of Sciences, 55, 222 - 247.","Bahir, M. M. & Silva, A. (2005) Otocryptis nigristigma, a new species of agamid lizard from Sri Lanka. In: Yeo, C. J., Ng, P. K. L. & Pethiyagoda, R. (Eds.), Contributions to biodiversity exploration and research in Sri Lanka. The Raffles Bulletin of Zoology, 12 (Supplement), 393 - 406.","Manamendra-Arachchi, K., de Silva, A. & Amarasinghe, T. (2006) Description of a second species of Cophotis (Reptilia: Agamidae) from the highlands of Sri Lanka. Lyriocephalus, 6, 1 - 8.","Samarawickrama, V. A. M. P. K., Ranawana, K. B., Rajapaksha, D. R. N. S., Ananjeva, N. B., Orlov, N. L., Ranasinghe, J. M. A. S. & Samarawickrama, V. A. P. (2006) A new species of the genus Cophotis (Squamata: Agamidae) from Sri Lanka. Russian Journal of Herpetology, 13, 207 - 214.","Stoliczka, F. (1872) Notes on various new or little known Indian lizards. Proceedings of the Asiatic Society of Bengal, 41, 86 - 135."]}

Monilesaurus ellioti (Günther, 1864) comb. nov. Calotes rouxii— (not of Dum. & Bibr., 1837), Jerdon, 1853. J. Asiat. Soc. Beng. (2) xxii, 1853: 471 Calotes ellioti — Günther, 1864. Rept. Brit. Ind. 1864: 142. Bronchocela indica —Theobald, 1876. Cat. Rept. Brit. Ind. 1876: 105. Calotes elliotti — Smith, 1935. Fauna of British India, ii, 1935: 207. Original description. Günther, A. 1864. The Reptiles of British India. London (Taylor & Francis), xxvii, 452 pp. Taxonomic comments. Monilesaurus ellioti was described, based on Jerdon’s observation of C. rouxii like animals from “Malabar” in his list of Reptiles of Southern India, 1853 and a collection of drawings of this species in the possession of W. Elliot. Historically, southern Karnataka and northern Kerala have been referred to as Malabar. Boulenger (1885) gave a short description of this species based on the collections made by Col. R.H. Beddome from Anamalai hills, Sivagiri Ghat, Tirunelveli and Malabar. As there are no type specimens associated with this species, we hereby designate a neotype for this species. The selection of the neotype is roughly congruent with the original type locality of this species “Malabar” as mentioned by Jerdon (1853). Murthy (1978) described a subspecies Calotes ellioti amarambalamensis the type locality of which is Amarambalam, Nilambur. The holotype of this species is now at ZSI Madras (ZSI Madras 159) (Das et al. 1998). We were unable to trace the specimen in ZSI collections despite repeated attempts. For now, the specimen may be considered as lost. Hence, we compare another specimen collected from the same locality (BNHS 1033), a female individual whose characters match well with other M. ellioti and the description provided by Murthy. Also, the locality is within the distribution range of M. ellioti and contiguous with the type locality. Therefore, we consider Calotes ellioti amarambalamensis as a junior synonym of M. ellioti (Günther, 1864). ZSI Madras 159 is wrongly mentioned as a type specimen of Calotes ellioti (Uetz & Hošek, 2016). Neotype. CESL 0 45, adult male collected from Chembra reserve forest, Kerala by SPP, MVP and SPV on 10th June 2010. Other material examined. ZSI 4325 gravid female & ZSI 4328 male collected from Southern India; CESL 0 42 adult male collected from Manikunjamalai, Wayanad reserve forest, Kerala; CESL 0 45 adult male collected from Chembra reserve forest, Kerala; CESL 0 47 adult female collected from Vaithiri, Wayanad reserve forest, Kerala; CESL 0 57, CESL 0 59 and CESL 0 61 adult females and CESL 0 58 and CESL 0 60 adult males collected from Vallerimala, Wayanad reserve forest, Kerala; CESL 0 77, adult female collected from Arlam Wildlife Sanctuary, Kerala; CESL 0 33 adult male collected from Bonacord, Peppara Wildlife Sanctuary, Kerala; CESL 162, adult male collected from Sholyar, Vazhachal reserve forest, Kerala; CESL 204, adult female collected from Pamba, Periyar tiger reserve, Kerala and CESL 369 adult male collected from Peppara, Cardemom hills, Kerala and BNHS 1033, adult female collected from Nilambur reserve forest, Kerala. Details of collection locality, specimen voucher and GenBank accession number in Appendix 1. Diagnosis and comparison. A small sized Monilesaurus (SVL up to 73.8 mm) characterized by the posteroventral orientation of lateral scales; antehumeral fold well developed, extending across the throat; 52–58 midbody scale rows; nuchal crest composed of 3–4 long, well developed spines; two separated supratympanic spines; a long, distinct isolated postorbital spine; dorsal and lateral scales keeled, ventral scales strongly keeled; paired postmentals, first pair separated by a single scale; 24–28 subdigital lamellae under fourth finger, 26–34 subdigital lamellae under fourth toe; 9–10 supralabials and 8–9 infralabials; olive brown above with angular darker cross bars on dorsum, a white spot below the eye. Morphologically M. ellioti is superficially similar to M. montanus gen. et sp. nov., M. rouxii comb. nov.; and M. acanthocephalus gen. et sp. nov., but can be distinguished by a combination of the following characters: 52–58 midbody scale rows (vs. 46–52 in M. montanus gen. et sp. nov., 62–64 in M. acanthocephalus gen. et sp. nov., and 52–56 in M. rouxii) presence of a long, distinct isolated spine in the posterior corner of orbit (vs. absent in M. rouxii; very small, indistinct tubercle like in M. montanus gen. et sp. nov., and much longer in M. acanthocephalus gen. et sp. nov.); 3–4 long nuchal spines (vs. 3–6 small nuchal spines in M. montanus gen. et sp. nov., 6 much longer nuchal spines in M. acanthocephalus gen. et sp. nov., 7–8 smaller nuchal spines in C. rouxii); longer, prominent isolated spine on the back of head and above tympanum (vs. much smaller in M. montanus gen. et sp. nov., and M. rouxii) and presence of a white spot below the eye (vs. absent in M. rouxii; in the form of a band in M. montanus gen. et sp. nov.) and smaller body size: adult SVL 59.4–73.8 mm, n=9 (vs. C. montanus gen. et sp. nov., adult SVL 61–83.4 mm, n=8; and M. acanthocephalus gen. et sp. nov. adult SVL 68.9–72.6 mm, n=3). Description. Based on CESL 0 45. A medium sized adult male (SVL- 63.7 mm), Morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately compressed. Head moderately large (HL/SVL ratio 0.29), elongate (HW/HL ratio 0.63), maximum height less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield, which is separated from rostral by a single scale; mental shield narrower than rostral; two postmentals, first pair in contact with each other; genials keeled; gular scales strongly keeled, slightly smaller than genials; scales on top of snout smooth except median row, which is keeled; scales on top of head heterogenous in size and shape, keeled; supraorbital scales keeled; canthus-rostralis and supraciliary edge sharp; a spine at the posterior corner of the orbit; two separated long spines on posterior end of head, the anterior slightly longer, midway between nuchal crest and tympanum, posterior above tympanum; orbit diameter 70% of distance between anterior border of orbit and snout tip; tympanum exposed, its greatest diameter 54% horizontal diameter of orbit; enlarged keeled scale between tympanum and orbit; posterior region of jaws swollen; supralabials 10/10; infralabials 9/9. Nuchal crest well developed, composed of four primary, long conical spines, the first being the smallest and the third longest; the remaining vertebral scales slightly enlarged relative to adjacent rows and possess a more pronounced median keel forming a serrated ridge like the dorsal crest which continues till the tail base; 54 longitudinal scale rows around midbody; scales on dorsum keeled, oriented postero-dorsally, while lateral ones oriented postero-ventrally; lateral scales smaller than dorsal, keeled; ventrals strongly keeled, irregular, slightly smaller than dorsals but of similar size as laterals, genials and gular scales; a strong, oblique antehumeral fold, nearly extending across the throat. Limbs slender and covered with strongly keeled scales, larger than laterals, forming parallel longitudinal rows; scales under thighs weakly keeled; length of hindlimb ca. 87% SVL; relative length of fingers 4>3>2>5>1; relative lengths of toes 4>3>5>2>1; fourth toe longer than fifth finger; 23 subdigital lamellae under fourth finger; 27 subdigital lamellae under fourth toe; subdigital lamellae with sharp keels, bicarinate; tail slender, swollen at the base; scales on dorsal and ventral surface of tail with sharp keels, larger than laterals; tail length 172 mm. Colouration. In life: dorsum and head blackish-brown with irregular lighter patches on the back; a light brown band from above the shoulder till the dorsal crest forming a ‘v’ shape; head laterally dark blackish with a white spot below the posterior end of eye; stripe from above nostril to anterior margin of orbit extending till the tympanum from the posterior margin of orbit in the form of a black band; tympanum pale grey, lip scales white; a blackish triangular patch behind the tympanum continuing to the antehumeral fold; ventral uniformly lighter, pale grey; gular pouch with a reddish stripe extending beyond the throat; tail with alternating dark and light blotches forming irregular bands towards the end. Representative image showing live colouration (Fig. 6b). In preservative: colouration pattern mostly similar to that in life, except overall paler; bands on the head dull greyish brown. Variation and secondary sexual characteristics. The other specimens examined agree with the CESL 0 45 in general morphology and scalation except for some differences that are summarised in Appendix 2 & 3. All the examined female specimens (CESL 0 47, CESL 0 58 and CESL 061) have much smaller nuchal spines compared to the males, reduced postero-orbital spine; lack a dorsal crest and gular sac; overall colouration olive-brown, lighter head and vertebral region, a dark band along the side of the head to the neck and presence of a black antehumeral fold. Genetic distance. M. ellioti comb. nov. shows 1–2% intraspecific genetic divergence in the 16S gene; 4–7% interspecific genetic divergence from M. rouxii comb. nov.; 5–7% interspecific genetic divergence from M. acanthocephalus gen. et sp. nov and 4–8% interspecific genetic divergence from M. montanus gen. et sp. nov. (Appendix 5). Distribution. Monilesaurus ellioti comb. nov. is endemic to the Western Ghats and is distributed across the low and medium elevation forests (up to 1000 m asl) of southern parts of central Western Ghats (Coorg plateau and south) and the southern Western Ghats. This is one of the most common evergreen forest dwelling agamid lizards in this region. During this study, M. ellioti comb.nov. was recorded in various sites in the central and southern Western Ghats (See Fig. 3 & Appendix 1 for details). Ecology and natural history. Monilesaurus ellioti is a diurnal lizard, semi-arboreal to arboreal in habit, and so far, has been recorded mostly in semi-evergreen and evergreen forests. Individuals were mostly seen perching on shrubs, branches and actively moving on tree trunks. In some instances, it has also been observed in coffee plantations surrounded by evergreen forests. In some sites like Parambikulam, Vazhachal reserve forest and Brahmagiri hills, it was observed to occur syntopically in the same habitat as M. rouxii comb. nov., but more abundant within dense forests, while M. rouxii comb. nov. is generally restricted to forest edges. In some instances, gravid females were recorded during pre-and mid monsoon (June–August), which hints that monsoon might be a breeding season for this species.

Monilesaurus rouxii (Dum��ril & Bibron, 1837) comb.nov. Calotes rouxii ���Dum��ril & Bibron, 1837. Erp. Gen, iv, 1837: 407. Calotes ellioti ���(not of G��nther) Stoliczka, 1872. J. Asiat. Soc. Beng. (2) xli, 1872: 113. Calotes rouxii ��� Smith, 1935. Fauna of British India, ii, 1935: 206. Syntypes. MNHN���MNHN-RA-0.6894 & MNHN-RA-1994.1857 collected from ��� Indes Orientales ��� and deposited in the National Museum of Natural History (France). Original description. Dum��ril, A. M. C. and G. Bibron. 1837. Erp��tologie G��n��rale ou Histoire Naturelle Complete des Reptiles. Vol. 4. Libr. Encyclop��dique Roret, Paris, 570 pp. Taxonomic comments. The exact type locality of M. rouxii comb.nov. is unknown, as the localities for the syntypes in MNHN and in the original publication are given as ���Indes Orientales��� or India. The only precise locality of a specimen in the catalogue of British Museum is given as ���Matheran, Bombay Presidency��� (Boulenger 1885, 1890); Smith (1935) gives the range of M. rouxii comb. nov. as ���Bombay Presidency (Matheran, Khandala, Kanara, Jog); Travancore���. Of these, other than ���Travencore��� all the other localities are from the northern and central Western Ghats. Material examined. CESL 129, adult male collected from Matheran, Maharashtra; CESL 523, adult male collected from Brahmagiri Wildlife Sanctuary, Karnataka; CESL 554, adult male collected from Pushpagiri Wildlife Sanctuary, Karnataka; CESL 669, adult female collected from Bondla Wildlife Sanctuary, Goa; CESL 834, adult female collected from Narsimparvata, Kudremukh National Park, Karnataka, CESL 678, adult male collected from Madhei wildlife sanctuary, Goa; CESL 875, juvenile collected from Radhanagri, Maharashtra; CESL 0 95, juvenile, collected from Agumbe, Karnataka; 0 72, adult male collected from Wayanad wildlife sanctuary, Kerala; CESL 123 and CESL 153 adult males collected from Vazhachal, Kerala; CESL 215, adult male collected from near Parambikulam wildlife sanctuary, Kerala and CESL 581, adult male collected from near Pooyamkutty, Kerala. MNHN-RA-0.6894 (photographs only). Details of collection locality, specimen voucher and GenBank accession number in Appendix 1. Diagnosis and comparisons. A small sized Monilesaurus (SVL up to 74.8 mm) characterized by the posteroventral orientation of lateral scales; antehumeral fold small, triangular spines; two separated small supratympanic spines; dorsal and lateral scales keeled, ventral scales strongly keeled; paired postmentals, first pair in contact or separated by a single scale; 18���21 subdigital lamellae under fourth finger, 24���29 subdigital lamellae under fourth toe; 9���10 supralabials and 8���9 infralabials; olive-brown to above, antehumeral fold black, top of head often darker than dorsum, body often speckled with dark and light blotches, prominent in juveniles and sub-adults. Morphologically, M. rouxii comb.nov. is superficially similar to M. montanus gen. et sp. nov., M. ellioti comb. nov.; and M. acanthocephalus gen. et sp. nov., but can be distinguished by a combination of the following characters: 52���56 midbody scale rows (vs. 46���52 in M. montanus gen. et sp. nov., 62���64 in M. acanthocephalus gen. et sp. nov., and 52���58 in M. ellioti comb. nov.) spine in the posterior corner of orbit absent (vs. very small, indistinct tubercle like in M. montanus gen. et sp. nov., long, distinct in M. ellioti comb.nov. and much longer in M. acanthocephalus gen. et sp. nov.); 7���8 small nuchal spines (vs. 3���6 small nuchal spines in M. montanus gen. et sp. nov., 6 much longer nuchal spines in C. acanthocephalus gen. et sp. nov., 3���4 long nuchal spines in M. ellioti comb.nov.); small isolated spines on the back of head and above tympanum (vs. longer, prominent spines in M. ellioti comb. nov. and M. acanthocephalus gen. et sp. nov.) white spot below the eye absent (vs. present in M. ellioti comb.nov. and M. acanthocephalus gen. et sp. nov.; in the form of a band in M. montanus gen. et sp. nov.) and smaller body size: adult SVL 51.4���74.8 mm, n=9 (vs. C. montanus gen. et sp. nov., adult SVL 61���83.4 mm, n=8; and M. acanthocephalus gen. et sp. nov. adult SVL 68.9���72.6 mm, n=3). Description. Based on CESL 129. A medium sized adult male (SVL- 74.79 mm). Morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately compressed. Head moderately large (HL/SVL ratio 0.29), elongate (HW/HL ratio 0.73), maximum height less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield, which is separated from rostral by a single scale; mental shield narrower than rostral; two postmentals; first pair in contact with each other; genials keeled; gular scales strongly keeled, slightly smaller than genials; scales on top of snout smooth except median row, which is keeled; scales on top of head heterogenous in size and shape, keeled; supraorbital scales keeled; canthus-rostralis and supraciliary edge sharp; two separated spines on posterior end of head, the anterior slightly longer, midway between nuchal crest and tympanum, posterior above tympanum; orbit diameter 75% of distance between anterior border of orbit and snout tip; tympanum exposed, its greatest diameter 58% horizontal diameter of orbit; enlarged keeled scale between tympanum and orbit; posterior region of jaws swollen; supralabials 10/10; infralabials 9/9. Nuchal crest well developed, composed of eight primary, broadly conical spines, the first and last smaller than the rest; the remaining vertebral scales slightly enlarged relative to adjacent rows and possessing a more pronounced median keel forming a serrated ridge like the dorsal crest which continues till the tail base; 56 longitudinal scale rows around midbody; scales on dorsum keeled, oriented postero-dorsally, while lateral ones oriented postero-ventrally; lateral scales smaller than dorsal, keeled; ventrals strongly keeled, irregular, slightly smaller than dorsals but of similar size as laterals, genials and gular scales; a strong, oblique antehumeral fold, nearly extending across the throat. Limbs slender and covered with strongly keeled scales, larger than laterals, forming parallel longitudinal rows; scales under thighs weakly keeled; length of hindlimb ca. 82% SVL; relative length of fingers 4>3>2>5>1; relative lengths of toes 4>3>5>2>1; fourth toe longer than fifth finger; 20 subdigital lamellae under fourth finger; 24 subdigital lamellae under fourth toe; subdigital lamellae with sharp keels, bicarinate; tail slender, swollen at the base; scales on dorsal and ventral surface of tail with sharp keels, mucronate, slightly larger than laterals; tail length 118 mm (tail incomplete, broken at the tip). Colouration. In life: dorsum uniform blackish-brown; head bright reddish-orange, from snout tip to slightly behind mid vertebral; laterally a blackish-brown stripe from above nostril to anterior margin of orbit extending till the tympanum from the posterior margin of orbit in the form of black band; a bright reddish-orange stripe from snout covering labials till the anterior margin of tympanum, continuing backwards from posterior margin and ending abruptly in the antehumeral fold; tympanum pale grey; ventral uniformly black; gular pouch with a small orange stripe in the median row. Representative image showing live colouration (Fig. 6a). In preservative: colouration pattern mostly similar to that in life, except overall paler; bands on the head dull greyish brown. Variation and secondary sexual characteristics. The other specimens examined agree with CESL 129 in general morphology and scalation except for some differences that are summarised in Appendix 2 & 3. Both the examined female specimens (CESL 669 and CESL 834) have much smaller nuchal spines and lack a dorsal crest and gular sac; overall colouration olive-brown, lighter head and vertebral region, a dark band along the side of the head to the neck and the presence of a black antehumeral fold. Genetic distance. M. rouxii comb. nov. shows 1% intraspecific genetic divergence in the 16S gene; 4���7% interspecific genetic divergence from M. ellioti comb. nov.; 6% genetic divergence from M. acanthocephalus gen. et sp. nov. and 7���8% interspecific genetic divergence from M. montanus gen. et sp. nov. (Appendix. 5). Distribution. Monilesaurus rouxii comb. nov. is distributed across the low and medium elevation forests (up to 1000 m asl) of the Western Ghats and has also been reported from parts of the southern Eastern Ghats (Daniels & Ishwar 1994). This is one of the most common forest dwelling agamid lizards in the northern and central Western Ghats. During this study, M. rouxii comb. nov. was recorded in various sites across the Western Ghats (See Fig. 3 & Appendix 1 for details). Ecology and natural history. Monilesaurus rouxii is a diurnal lizard, semi-arboreal to arboreal in habit, and has so far been recorded mostly in deciduous, secondary and semi-evergreen forests. Individuals were mostly seen perching on branches and actively moving on tree trunks. In some instances, it has also been observed in forest fragments and plantations. In some sites, they occur syntopically in the same habitat as C. versicolor, but tend to prefer higher and thicker perches than C. versicolor, and were found to be more abundant than C. versicolor inside forests. In many instances, gravid females were recorded during pre and mid monsoon (June���August), which suggests that monsoon might be a breeding season for this species., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 427-429, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674, {"references":["Dumril, A. M. C. & Bibron, G. (1837) Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Fol. 4. Librairie Encyclopedique Roret, Paris, 570 pp.","Gunther, A. (1872) Descriptions of some Ceylonese Reptiles and Batrachians. Annals Magazine of natural History, 4, 85 - 88. https: // doi. org / 10.1080 / 002229372011951778","Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Fol. II. Sauria. Taylor and Francis, London, 440 pp.","Boulenger, G. A. (1885) Catalogue of lizards in the British museum. Fol. 1. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae, Second edition. British Museum (Natural History), London, 436 pp.","Boulenger, G. A. (1890) The Fauna of British India, including Ceylon and Burma: Reptilia and Batrachia. Taylor and Francis, London, 541 pp.","Daniels, R. J. R. & Ishwar, N. M. (1994) Rarity and the herpetofauna of the Southern Eastern Ghats, India. Cobra, 16, 2 - 14."]}

Calotes versicolor (Daudin, 1802) Original description. Daudin, 1802. Hist. Nat., Rept. iii, 1802: 395, xliv. Taxonomic comments. Calotes versicolor was described in 1802 by Daudin, based on specimens at MNHN. There is no mention of a precise locality or even a type specimen. Later Smith (Smith 1935) and other workers designated the type-locality to populations from Pondicherry on the east coast of India, Chennai (Madras), and Kolkata (Calcutta) (Zug et al. 2006). The specimen MNHN 2548, collected from Pondicherry by Leschenault, cannot be a syntype of C. versicolor Daudin 1802, because Leschenault visited Pondicherry only after 1816 (Amarasinghe et al. 2009). Amarasinghe et al (2009) clarified that MNHN 2548 collected from Pondicherry by Leschenault is the syntype of Agama tiedemanni Kuhl, 1820. Gowande et al (2016) synonymized Calotes tiedemanni and designated a neotype from Pondicherry without any comparison with the syntype of Calotes tiedemanni. This neotype designation was recently invalidated (Chaitanya et al. 2017). Currently, Calotes versicolor is the most widely distributed species of the genus Calotes. Many researchers have mentioned the high level of morphological variation in this species across different populations and some have even considered it to be a species complex (Zug et al. 2006). Recently two new species were described from dry zone populations of C. versicolor from Myanmar based on DNA sequence and morphological analysis (Zug et al. 2006). In our phylogenetic analyses, the southern Western Ghats population forms a distant cluster within the C. versicolor clade (Fig. 1). The studied specimens also show 0–5% genetic divergence across their distribution. This shows that the Indian population might also be a species complex (see Gowande, 2016). A phylogeographic study with thorough sampling across its range and a rigorous comparison of the available type materials for the existing species and subspecies is necessary to resolve the complexity in this species. Since we did not study any historical type materials of Calotes versicolor or the synonyms, we consider the populations in this study as Calotes cf. versicolor. The diagnosis is based on specimens collected from the Southern Western Ghats only and used to separate Calotes cf. versicolor from other distinct species. Diagnosis and comparison. A medium to large sized Calotes (SVL up to 99.4 mm) characterized by the posterodorsal orientation of lateral scales; antehumeral fold absent; 38-44 midbody scale rows; nuchal and dorsal crest well developed, continuous, extending till the start of tail in males; two well separated supratympanic spines; dorsal and lateral scales large, more or less distinctly keeled, ventral scales strongly keeled, mucronate; paired postmentals, first pair separated by 1–2 median scales; 15–22 subdigital lamellae under fourth finger, 21–25 subdigital lamellae under fourth toe; 10–12 supralabials and 9–11 infralabials; uniform, light brown or greyish above, juveniles and females with darker cross bars or blotches on lateral side, often with paired lighter dorsolateral stripes, gular pouch brick red and overall body reddish in breeding males. From other members of Smith’s C. versicolor group (C. nemoricola, C. grandisquamis, C. calotes, C. minor, C. maria, C. jerdoni, C. emma and C. mystaceus) C. cf. versicolor can be distinguished by a combination of the following characters: smaller body size: adult SVL 73.8–99.4 mm, n=9 (vs. C. nemoricola, adult SVL 112–134.8 mm, n=7, n=5; and C. grandisquamis, adult SVL 110.0– 136.5 mm, n=5); 38–44 midbody scale rows (vs. 27–35 in C. grandisquamis; 30–35 in C. calotes, n=3; 36–43 in C. nemoricola; 58–63 in C. maria; 45–57 in C. jerdoni; 49– 65 in C. emma; 48–58 in C. mystaceus and 48–60 in C. minor); crescent-shaped patch of granular scales in front of the shoulder absent (vs. present in C. nemoricola, C. grandisquamis, C. emma, C. mystaceus and C. jerdoni); nuchal and dorsal crest well developed, composed of almost equal spines (vs. nuchal spines much longer, dorsal crest reduced in C. nemoricola and C. maria; nuchal spines much longer than dorsal spines in C. grandisquamis, C. calotes and C. emma); two well separated supratympanic spines (vs. row of 3–4 compressed supratympanic spines in C. nemoricola and C. grandisquamis; 8–9 compressed spines in C. calotes; two parallel rows of compressed supratympanic scales in C. maria and C. jerdoni; single well developed postorbital spine in C. emma). Calotes cf. versicolor can be distinguished from its other Sri Lankan congeners based on posterodorsal orientation of lateral scales and absence of oblique fold or pit in front of the shoulder. Description. Based on specimen CESL 182. A large sized adult male (SVL 99.4 mm), morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately compressed. Head moderate (HL/ SVL ratio 0.22), slightly elongate (HW/HL ratio 0.95), maximum height slightly less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield which is separated from rostral by two scales; mental shield narrower than rostral; two postmentals, first pair separated from each other by two small scales; genials keeled; gular sac small, composed of strongly keeled scales, slightly smaller than genials; scales on top of snout smooth except median row, which is keeled; scales on top of head heterogenous in size and shape, keeled; supraorbital scales keeled; canthus-rostralis and supraciliary edge sharp; two separated spines on posterior end of head, the anterior much longer, closer to the nuchal crest, posterior above tympanum; orbit diameter 87% of distance between anterior border of orbit and snout tip; tympanum exposed, its greatest diameter 42% of horizontal diameter of orbit; slightly keeled, scales between tympanum and orbit smooth, slightly enlarged; posterior region of jaws swollen; supralabials 11/10; infralabials 10/10. Nuchal and dorsal crest well developed, continuous; nuchal crest composed of 11 long, conical spines, the first being the smallest; dorsal crest composed of 33 curved spines till above the vent, slightly smaller than the nuchal spines; 36 longitudinal scale rows around midbody; dorsal and lateral scales sub-triangular, keeled, oriented postero-dorsally; ventrals strongly keeled, mucronate, smaller than dorsal and lateral scales. Limbs slender and covered with strongly keeled scales, similar to laterals, forming parallel longitudinal rows; scales under thighs weakly keeled; length of hindlimb ca. 75 % SVL; relative length of fingers 4>3>2>5>1, third and fourth finger almost equal; relative lengths of toes 4>3>5>2>1; fourth toe much longer than fifth finger; 21 subdigital lamellae under fourth finger; 25 subdigital lamellae under fourth toe; subdigital lamellae with sharp keels, bicarinate; slender, swollen at the base; dorsal and ventral tail scales mucronate, strongly keeled; tail broken, incomplete; tail length 90 mm. Colouration. In life: dorsum and head uniform, greyish-brown with irregular darker patches towards the flank; each lateral scale with a darker blackish spot towards the base, followed by lighter grey edges; head laterally paler, whitish towards the cheeks; a small, thin darker stripe from posterior corner of the eye till end of orbit; legs with irregular dark crossbars; a black coloured patch above the shoulder, near throat; tympanum pale grey with many small black spots, ventral uniformly lighter, pale grey; tail with alternating dark and light bands. Representative image showing live colouration (Fig. 4b). In preservative: colouration mostly similar to that in life, except overall paler. Variation and secondary sexual characteristics. Meristic and morphometric data of four adult male and four adult female specimens are given in Appendix 2 & 3. The examined specimens agree with each other in general morphology and scalation. All the examined female specimens (CESL 0 48, CESL 163, CESL 306 and BNHM 374) have much smaller nuchal and dorsal spines compared to the males and lack a gular sac. Distribution. Calotes cf. versicolor is common in the Western Ghats at elevations between 70 m to 1250 m M.S.L. (See Appendix 1 for details). Ecology and natural history. Calotes cf. versicolor is a diurnal lizard, semi-arboreal to arboreal in habit, and so far, has been recorded mostly in scrub, deciduous forests and plantations. Individuals were mostly seen perching on shrubs, hedges, gardens and along compound walls of houses in villages along forests. C. cf. versicolor has often been found within plantations and prefers open patches to forested areas. During this study, it was never recorded from primary evergreen forests. In some sites, they might occur syntopically in the same habitat as M. rouxii comb. nov. but tend to be restricted to forest edges., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 421-423, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674, {"references":["Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Fol. II. Sauria. Taylor and Francis, London, 440 pp.","Zug, G. R., Brown, H. H., Schulte, J. A. & Vindum, J. V. (2006) Systematics of the Garden Lizards, Calotes versicolor Group (Reptilia, Squamata, Agamidae), in Myanmar: Central Dry Zone Populations. Proceedings of the California Academy of Sciences, 57 (2), 35 - 68.","Amarasinghe, A. A. T., Manthey, U., Stockli, E., Ineich, I., Kullander, S. O., Tiedemann, F., McCarthy, C. & Gabadage, D. E. (2009) The original descriptions and figures of Sri Lankan Agamid lizards (Squamata: Agamidae) of the 18 th and 19 th centuries. Taprobanica, 1 (1), 2 - 15. https: // doi. org / 10.4038 / tapro. v 1 i 1.2771","Chaitanya, R., Giri, V. B. & Deepak, V. (2017) On the recent designation of a neotype for the taxon Calotes versicolor (Daudin, 1802) (Reptilia: Agamidae). Zootaxa, 4317 (3), 585 - 587. https: // doi. org / 10.11646 / zootaxa. 4317.3.10"]}

Calotes nemoricola Jerdon, 1853 Calotes nemoricola��� Jerdon, 1853. J. Asiat. Soc. Beng. xxii, 1853: 471. Original description. Jerdon, 1853. Journal of the Asiatic Society of Bengal, xxii, 1853: 471. Taxonomic comments. Calotes nemoricola was described based on specimens collected from ���Foot of ���Coonoor Ghat, Nilgherries��� (���Coonoor���, Nilgiri hills, Tamil Nadu, India). The type collected by Jerdon is a much mutilated male specimen (Smith 1935) and is housed in the collection of ZSIK (Das et al. 1998). Another male topotype is housed in the collection of BNHS (BNHS 373) (Smith 1935). Holotype: ZSI 6560. The specimen is in poor condition; the posterior part of the body along with the hindlimbs and tail is broken. Material examined. CESL 0 38 adult male collected from Bonaccord, Peppara Wildlife Sanctuary, Kerala; CESL 406 adult male collected from Kanyakumari Wildlife Sanctuary, Tamil Nadu; CESL 545 adult male collected from Makuta, Brahmagiri Wildlife Sanctuary, Karnataka; CESL 555 adult male collected from Bidahalli, Pushpagiri Wildlife Sanctuary, Karnataka; BNHS 1778 adult male collected from Kudremukh National Park, Karnataka; BNHS 373 adult male collected from ��� Coonoor ��� Ghat, Nilgiri hills, Tamil Nadu; BMNH 74.4.29.224 male, 74.4.29.225 female both collected from Malabar and ZSI 6560 male collected from the foothills of ��� Coonoor Ghat. Details of collection locality, specimen voucher and GenBank accession number in Appendix 1. Diagnosis and comparison. A large sized Calotes (SVL up to 134.8 mm) characterized by the posterodorsal orientation of lateral scales; antehumeral fold absent; 36���43 midbody scale rows; nuchal crest well developed composed of long spines, dorsal crest reduced, almost indistinct behind the midbody; row of 3���4 compressed supratympanic spines; postorbital spine absent; a crescent-shaped patch of granular scales in front of the shoulder present; dorsal and lateral scales large, mostly smooth, weakly keeled towards the flanks, ventral scales strongly keeled, mucronate; paired postmentals, first pair separated by 1���2 median scales; 21���23 subdigital lamellae under fourth finger, 25���31 subdigital lamellae under fourth toe; 10���12 supralabials and 10���11 infralabials; green to brownish above, uniform, often with a dark streak from eye to above tympanum, ventral white to pale grey. C. nemoricola can be distinguished from members of Smith���s C. versicolor group by a combination of the following characters: larger body size: mean adult SVL 125.8 mm, n=7 (vs. C. versicolor, adult SVL 73.8���99.4 mm, n=9 and C. calotes, adult SVL 73.6���96.8 mm, n=3); 36���43 midbody scale rows (30���35 in C. calotes, n=3, 38��� 44 in C. versicolor; 58���63 in C. maria; 45���57 in C. jerdoni; 49���65 in C. emma; 48���58 in C. mystaceus and 48���60 in C. minor); nuchal crest well developed with longer spines, dorsal crest reduced (vs. nuchal and dorsal crest well developed, composed of almost equal spines in C. versicolor and C. mystaceus); row of 3���4 compressed supratympanic spines (vs. two well separated supratympanic spines in C. versicolor, 8���9 compressed spines in C. calotes, two parallel rows of compressed supratympanic scales in C. maria and C. jerdoni; single well developed postorbital spine in C. emma); presence of a crescent-shaped patch of granular scales in front of the shoulder (vs. absent in C. versicolor, C. calotes and C. maria) and lateral scales much larger than ventrals (vs. almost equal to the ventrals in C. calotes). Morphologically, it is mostly similar to the closely related C. grandisquamis but can be distinguished based on higher number of midbody scale rows (36���43 vs. 27���35); smaller scales between eye and tympanum, much smaller than tympanum (vs. much larger scales, almost equal to the tympanum); scales on the cheek smaller, weakly keeled (vs. larger, smooth); gular scales keeled, smaller than mental (vs. gular scales smooth, equal to or slightly larger than the mental) and dorsal crest reduced, not extending beyond mid body (dorsal crest well developed, till above the base of tail). Even though these two large bodied species show the above-mentioned morphological differences, there has been some confusion in their correct identification in the recent past. To clarify this, a detailed comparison to distinguish these two species was provided by Ganesh & Chandramouli (2013). Description. Based on specimen CESL 555. A large sized adult male (SVL- 120.4 mm), morphometric and meristic data are summarised in Appendix 2 & 3. General habitus compressed. Head moderate (HL/SVL ratio 0.25), elongate (HW/HL ratio 0.75), maximum height slightly less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield which is separated from rostral by two scales; mental shield narrower than rostral; two postmentals, first pair separated from each other by two small scales; genials keeled; gular sac small, composed of keeled scales, strongly keeled towards the medial row, slightly smaller than genials; scales on top of snout smooth except median row, which is keeled; scales on top of head heterogenous in shape and size, keeled towards the edges; supraorbital scales keeled; canthus-rostralis and supraciliary edge sharp; 4 supratympanic spines in a row, compressed; orbit diameter 69% of distance between anterior border of orbit and snout tip; tympanum exposed, its greatest diameter 46% of horizontal diameter of orbit; partially keeled, scales between tympanum and orbit smooth, not enlarged; posterior region of jaws distinctly swollen; supralabials 11/11; infralabials 11/11. Nuchal crest well developed, continuous with the much reduced dorsal crest; dorsal crest reaching slightly behind midbody; nuchal crest composed of 16 long, conical spines, the first being the smallest and fourth the longest; longest nuchal spine 65% of the orbit; dorsal spines less than half the nuchal spines in length; 41 longitudinal scale rows around midbody; dorsal and lateral scales sub-triangular, smooth, oriented posterodorsally; a crescent-shaped patch of small, granular scales present in front of the shoulder. Limbs moderate and covered with weakly keeled scales, stronger towards the end forming parallel longitudinal rows; scales under thighs weakly keeled; length of hindlimb ca. 71 % SVL; relative length of fingers 4>3>2>5>1, third and fourth finger almost equal; relative lengths of toes 4>3>5>2>1; fourth toe much longer than fifth finger; 21 subdigital lamellae under fourth finger; 27 subdigital lamellae under fourth toe; subdigital lamellae keeled, bicarinate; slender, swollen at the base; scales on dorsal and ventral surface of tail keeled, ventral tail scales mucronate; tail length 268 mm. Colouration. In life: dorsum and head uniform, light green above with slightly brownish lateral scales; a thick milky white band from behind the head till end of nuchal crest extending across the neck; head laterally brighter green, whitish below the cheeks; pupil black surrounded by a brick red iris; lip scales green; legs uniform darker shade of green; tympanum pale bluish green; ventral uniformly white; tail greenish at the start turning dull brownish towards the end. This individual changed colour at the slightest disturbance and turned dark brown overall with a dark olive head and a prominent white band behind the head. Representative image showing live colouration (Fig. 5b). In preservative: colouration mostly similar to that in life, except overall paler. Variation and secondary sexual characteristics. Meristic and morphometric data of the examined specimens are given in Appendix 2 & 3. The examined specimens agree with each other in general morphology and scalation. The only female specimen we examined (BMNH 74.4.29.255) had no cheek pouches or a swollen tail base. Distribution. Calotes nemoricola is endemic to the Western Ghats and is distributed in the low to mid elevation evergreen forests of the southern part of central Western Ghats (Agumbe and south) and the southern Western Ghats (Inger et al. 1984; Naniwadekar & Deepak 2008). During the present study, this species was recorded from various sites in the hills of central and southern Western Ghats (See Appendix 1 for details). Ecology and natural history. Calotes nemoricola is a diurnal lizard, mostly arboreal in habit, and so far, has been recorded mostly from primary evergreen forests. Individuals were mostly seen perched on high branches and actively moving on tree trunks inside forests. In a few instances, it was also found sleeping on lower branches at night. This species has mostly been recorded from low elevations (up to 600 m msl) in the southern Western Ghats but has sometimes been recorded from high elevation evergreen forests in the central Western Ghats (around 1000 m msl). It might overlap in distribution with the other large green agamid C. grandisquamis at its higher elevation range extremes given their preference of similar microhabitat. During this study, they were not recorded to be syntopic at any site., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 425-426, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674, {"references":["Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Fol. II. Sauria. Taylor and Francis, London, 440 pp.","Ganesh, S. R. & Chandramouli, S. R. (2013) Identification of two similar Indian Agamid lizards Calotes nemoricola Jerdon, 1853 and C. grandisquamis Gunther, 1875. Russian Journal of Herpetology, 20, 33 - 35.","Inger, R. F., Shaffer, H. B, Koshy, M. & Bakde, R. (1984) A report on a collection of amphibians and reptiles from the Ponmudi, Kerala, South India. Journal of Bombay Natural History Society, 81, 551 - 570.","Naniwadekar, R. & Deepak, V. (2008) New distribution record for Calotes nemoricola Jerdon, 1853 from the Kudremukh hills, Karnataka, India. Journal of Bombay Natural History Society, 105 (1), 99."]}

Microauris gen. nov. (Fig. 8b) Type species. Calotes aurantolabium (Krishnan, 2008) Etymology. The genus epithet is derived by adding the word ‘Micro’ as a prefix to the Latin word ‘auris’ meaning ear, referring to the extremely small tympanum of this genus. Diagnosis. Microauris gen.nov. differs from all other members of draconinae lizards from the Indian subcontinent in having the smallest tympanum. Microauris gen.nov. differs from Mantheyus phuwuanensis by the absence of femoral pores (Manthey & Nabhitabhata, 1991; Ananjeva & Stuart 2001); from the members of the genus Bronchocela by the absence of a fold of skin from angle of jaw to shoulder (Hallermann & Böhme 2000). Microauris gen. nov. can be easily diagnosed from the genera Otocryptis, Sarada and Sitana by the presence of a well-developed fifth toe (Smith, 1935; Deepak et al, 2016); from Cophotis, Ceratophora, Lyriocephalus, Ptyctolaemus, Phoxophrys, Japalura otai, J. planidorsata, J. sagittifera and Otocryptis by the presence of external tympanum (Boulenger, 1885; Smith, 1935; Inger, 1 960; Pethiyagoda & Manamendra-Arachchi 1998; Schulte II et al. 2004; Bahir & Silva 2005; Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006). Microauris gen. nov. can be diagnosed from other members of genus Japalura by the absence of heterogenous dorsal scales and short and thick nuchal scales; from Salea by the absence of enlarged plate like scales between the eye and tympanum (Smith, 1 935). Microauris gen. nov. can be diagnosed from the members of Psammophilus and Monilesaurus by the absence of a well developed antehumeral fold. Microauris gen. nov. can be diagnosed from Complictus nigrigularis, Hypsicalotes kinabaulensis, Malayadracon robinsonii, Oriocalotes, Pseudocophotis and Pseudocalotes by the absence of enlarged row of suborbital scales (Smith, 1935; Taylor 1963; Ota & Hikida, 1991; Manthey & Denzer 1992; Inger & Steubing 1994; Ota & Hikida, 1996; Hallermann & Böhme 2000; Manthey & Denzer, 2000; Hallermann & McGuire 2001; Leong 2001; Manamendra-Arachchi et al. 2006; Samarawickrama et al. 2006; Ananjeva et al. 2007; Hallermann & Böhme 2007; Das & Lakim 2008; Hallermann et al. 2010; Mahony 2010; Harvey et al. 2014; Denzer et al. 2015; Grismer LL et al. 2016; Harvey et al. 2017). Microauris gen.nov. can be distinguished from the members of the genus Calotes (C. bachae, C. bhutanensis, C. calotes, C. ceylonensis, C. chincollium, C. desilvai, C. emma, C. grandisquamis, C. hutunwini, C. irawadi, C. jerdoni, C. liocephalus, C. liolepis, C. versicolor, C. manamendrai, C. maria, C. medgoensis, C. minor, C. mystaceus, C. nemoricola, C. nigrilabris, C. nigriplicatus and C. pethiyagodai) and Lophocalotes (Günther, 1872) in having the smallest tympanum (Hardwicke & Gray, 1827; Duméril and Bibron, 1837; Gray, 1845; Jerdon, 1854; Günther, 1864; Günther, 1870, Günther, 1872, Günther, 1875; Boulenger, 1885; Biswas, 1975; Zhao & Li, 1984; Hallerman, 2000; Vindum et al. 2003; Hallerman et al. 2004; Bahir & Maduwage, 2005; Zug et al. 2006; Manthey, 2008; Krishnan, 2008; Hartmann et al. 2013; Amarasinghe et al. 2014a, b; Deepak et al. 2015). Suggested English. Small-eared dragon Holotype. BNHS 1436, an adult female collected from Kalakkad Mundanthurai Tiger Reserve, Tamil Nadu, India; by N. M. Ishwar in 1997. Referred specimens. CESL 104, an adult female collected from a branch in the canopy of a shola forest tree near Chemunji peak, Peppara Wildlife Sanctuary, Kerala, India; by SPP, SPV, MVP, VTR and ADR in 18th August 2010. Taxonomic comments. Microauris aurantolabium comb. nov. was described as Calotes aurantolabium in 2008 based on a single specimen (Krishnan 2008). Prior to this, the specimen was wrongly identified and reported as the rediscovery of Calotes andamanensis. (Ishwar and Das, 1998). During our fieldwork in the Agasthyamalai hills of Kerala in August 2010, we found a single female specimen sleeping on a branch of a tree in a shola forest. There have been no other reports of this species thereafter and thus all the known specimens are females. There is a clear gap in the knowledge of the morphology of this rare lizard considering that male lizards are known to be better representatives for morphological characterization because of the presence of secondary sexual characters. In spite of this, the two specimens share some common characters, which are unique to this lizard. Based on the presence of unique morphological characters along with the phylogenetic position of this species in the subfamily Draconinae, we designate this species as a new monotypic genus. Diagnosis. A medium sized light green agamid lizard with a distinct orange streak from above the lip scales till behind the jaw; a moderately broad head with a pointed snout; very small tympanum relative to the orbit diameter (TymD/OrbD = 0.14–0.19); acutely keeled scales over the body and throat; scales on head small, sub-triangular (vs. uniform shield like in members of genus Calotes); 3rd and 4th toe almost equal in length; dorsal and lateral body scales oriented backwards and downwards; nuchal crest indistinct, poorly developed; supratympanic and postorbital spines absent and a long and slender tail. Description of the Holotype. BNHS 1436; a medium sized agamid, adult female (SVL- 68.5 mm) Morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately compressed. Head moderate (HL/SVL ratio 0.28), broad (HW/HL ratio 0.56), maximum height less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield, which is separated from rostral by a single scale; mental shield almost as wide as the rostral; two postmentals; first pair touching each other at the base; genials partially keeled; gular sac small, composed of partially keeled scales, slightly smaller than genials; scales on the snout smooth, unkeeled; scales on top of head heterogenous, keeled, small, subtriangular; supraorbital scales keeled; orbit diameter 95% of distance between anterior border of orbit and snout tip; tympanum very small, exposed, its greatest diameter 11% of horizontal diameter of orbit; slightly keeled scales between tympanum and orbit; posterior region of jaws partially swollen; supralabials 10/10; infralabials 11/11. Nuchal crest poorly developed, composed of 13 small conical spines; the remaining vertebral scales slightly enlarged relative to adjacent rows and possess a more pronounced median keel; 54 longitudinal scale rows around midbody; scales on dorsum keeled; lateral scales smaller than dorsal, keeled, oriented postero-ventrally; ventrals slightly larger then dorsals, genials and gular scales strongly keeled. Limbs slender and covered with strongly keeled scales, larger than laterals; scales under thighs keeled; length of hindlimb ca. 64% SVL; relative lengths of toes 4=3>5>2>1; fourth toe and third toe subequal; 20 subdigital lamellae under fourth finger; 21 subdigital lamellae under fourth toe; tail slender, long; scales on dorsal and ventral surface of tail with sharp keels, mucronate, slightly larger than laterals; tail length 117 mm. Colouration. In life (CESL 104): dorsum and head uniform, light green overall with a distinct reddish orange streak from above the third lip scale covering 2–3 scale rows below the tympanum, ending just at the shoulder. A distinct pale whitish grey band, edged with black speckles from the base of tail till underside of knee. Mid-dorsal scales edged anteriorly with black, which is visible when the body is inflated; gular scales lighter, yellowish green; ventrally brighter, uniform green with a few rows of pale whitish scales towards the lateral side. This lizard was observed to quickly change its body color to dark blackish-brown on disturbance but the orange streak and band on thigh remained unchanged. In preservative: dorsum and head uniform, overall pale blue with brownish patches above the eyes and neck continuing on the mid-dorsal line, broader towards the tail; tail pale brown; orange lip streak paler turning whitish towards the neck; laterally pale blue with anteriorly black edged scales; ventrally pale whitish grey overall; limbs edged with brown blotches, broader towards the tips. Distribution. Currently, this lizard is known only from two sites in the high elevation tropical evergreen forests of Agasthyamalai hills. The second locality near Chemunji peak in Peppara wildlife sanctuary is approximately 50 kms northeast of the type locality near Kakachi in KMTR (See Appendix 1 for details). Both these sites are separated by almost contiguous shola grassland mosaic on undulating hills. Ecology and natural history. Nothing much is known about the ecology and natural history of this species. Ishwar and Das (1998) reported sightings of 3 individuals, all gravid females in the month of August. Two of these lizards laid 4 elliptical eggs each. This suggests that monsoon might be a breeding season for this lizard. Ishwar and Das (1998) also mentioned that this is a canopy dwelling species. The fact that it was only found again 12 years after its discovery and that too from a fairly well explored habitat suggests that this might be a rare lizard or it may occur high in the canopy of rain forests and thus escape detection., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 440-441, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674, {"references":["Krishnan, S. (2008) New Species of Calotes (Reptilia: Squamata: Agamidae) from the Southern Western Ghats, India. Journal of Herpetology, 42 (3), 530 - 535. https: // doi. org / 10.1670 / 06 - 198.1","Manthey, U. & Nabhitabhata, J. (1991) Eine neue Agame, Ptyctolaemus phuwuanensis sp. n. (Sauria: Agamidae), aus Nordost- Thailand. Sauria, 13, 3 - 6.","Ananjeva, N. B. & Stuart, B. L. (2001) The agamid lizard Ptyctolaemus phuwuanensis Manthey and Nabhitabhata, 1991 from Thailand and Laos represents a new genus. Russian Journal of Herpetology, 8 (3), 165 - 170.","Hallermann, J. & Bohme, W. (2000) A review of the genus Pseudocalotes (Squamata: Agamidae), with description of a new species from West Malaysia. Amphibia-Reptilia, 21 (2), 193 - 210. https: // doi. org / 10.1163 / 156853800507372","Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Fol. II. Sauria. Taylor and Francis, London, 440 pp.","Deepak, V., Giri, V. B., Asif, M., Dutta, S. K., Vyas, R., Zambre, A. M., Bhosale, H. & Karanth, K. P. (2016) Systematics and phylogeny of Sitana (Reptilia: Agamidae) of Peninsular India, with description of one new genus and five new species. Contributions to Zoology, 85, 67 - 111.","Boulenger, G. A. (1885) Catalogue of lizards in the British museum. Fol. 1. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae, Second edition. British Museum (Natural History), London, 436 pp.","Pethiyagoda, R. & Manamendra-Arachchi, K. (1998) A revision of the endemic Sri Lankan agamid lizard genus Ceratophora Gray, 1835, with description of two new species. Journal of South Asian natural History, 3 (1), 1 - 50.","Schulte II, J. A., Vindum, J. V., Win, H., Thin, T., Lwin, K. S. & Shein, A. K. (2004) Phylogenetic relationships of the genus Ptyctolaemus (Squamata: Agamidae), with a description of a new species from Chin Hills of Western Myanmar. Proceedings of the California Academy of Sciences, 55, 222 - 247.","Bahir, M. M. & Silva, A. (2005) Otocryptis nigristigma, a new species of agamid lizard from Sri Lanka. In: Yeo, C. J., Ng, P. K. L. & Pethiyagoda, R. (Eds.), Contributions to biodiversity exploration and research in Sri Lanka. The Raffles Bulletin of Zoology, 12 (Supplement), 393 - 406.","Manamendra-Arachchi, K., de Silva, A. & Amarasinghe, T. (2006) Description of a second species of Cophotis (Reptilia: Agamidae) from the highlands of Sri Lanka. Lyriocephalus, 6, 1 - 8.","Samarawickrama, V. A. M. P. K., Ranawana, K. B., Rajapaksha, D. R. N. S., Ananjeva, N. B., Orlov, N. L., Ranasinghe, J. M. A. S. & Samarawickrama, V. A. P. (2006) A new species of the genus Cophotis (Squamata: Agamidae) from Sri Lanka. Russian Journal of Herpetology, 13, 207 - 214.","Taylor, E. H. (1963) The lizards of Thailand. University of Kansas Science Bulletin, 44, 687 - 1077.","Ota, H. & Hikida, T. (1991) Taxonomic review of the lizards of the genus Calotes Cuvier, 1817 (Agamidae Squamata) from Sabah, Malaysia. Tropical Zoology, 4 (2), 179 - 192. https: // doi. org / 10.1080 / 03946975.1991.10539488","Manthey, U. & Denzer, W. (1992) The true Angle-head agamas of the genus Gonocephalus Kaup (Sauria: Agamidae) IV. Gonocephalus mjoebergi Smith, 1925 and Gonocephalus robinsonii Boulenger, 1908. Sauria, 1, 15 - 19.","Inger, R. F. & Stuebing, R. B. (1994) First record of the lizard genus Pseudocalotes (Lacertilia: Agamidae) in Borneo, with description of a new species. Raffles Bulletin of Zoology, 42 (4), 961 - 965.","Ota, H. & Hikida, T. (1996) The second specimen of Calotes kinabaluensis de Grijs (Squamata: Agamidae) from Sabah, Malaysia, with comments on the taxonomic status of the species. Journal of Herpetology, 30 (2), 288 - 291. https: // doi. org / 10.2307 / 1565529","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.","Hallermann, J. & McGuire, J. A. (2001) A new species of Pseudocalotes (Squamata: Agamidae) from Bukit Larut, West Malaysia. Herpetologica, 57 (3), 255 - 265.","Leong, T. M. (2001) The first female topotype of the female agamid Pseudocalotes flavigula (Smith, 1924) from Peninsular Malaysia, with description of colouration in life. Hamadryad, 26, 336 - 338.","Ananjeva, N. B., Orlov, N. L., Nguyen, Q. T. & Nazarov, R. A. (2007) A new species of Pseudocophotis (Agamidae: Acrodonta: Lacertilia: Reptilia) from central Vietnam. Russian Journal of Herpetology, 14 (2), 153 - 160.","Hallermann, J. & Bohme, W. (2007) On the validity of two recently described species of the endemic Sri Lankan genus Cophotis Peters, 1861 (Squamata: Agamidae) from the Knuckles Forest Range. Salamandra, 43, 187 - 190.","Das, I. & Lakim, M. (2008) A second specimen of Complicitus nigrigularis (Ota & Hikida, 1991). Hamadryad, 32 (1), 82 - 84.","Hallermann, J., Truong, N. Q., Orlov, N. & Ananjeva, N. (2010) A new species of the genus Pseudocalotes (Squamata: Agamidae) from Vietnam. Russian Journal of Herpetology, 17 (1), 31 - 40.","Mahony, S. (2010) Systematic and taxomonic 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 - 23.","Harvey, M. B., Hamidy, A., Kurniawan, N., Shaney, K. & Smith, E. N. (2014) Three new species of Pseudocalotes (Squamata: Agamidae) from southern Sumatra, Indonesia. Zootaxa, 3841 (2), 211 - 238. https: // doi. org / 10.11646 / zootaxa. 3841.2.3","Denzer, W., Manthey, U., Mahlow, K. & Bohme W. (2015) The systematic status of Gonocephalus robinsonii Boulenger, 1908 (Squamata: Agamidae: Draconinae). Zootaxa, 4039 (1), 129 - 144. https: // doi. org / 10.11646 / zootaxa. 4039.1.5","Harvey, M. B., Shaney, K., Hamidy, A., Kurniawan, N. & Smith, E. N. (2017) A new species of Pseudocalotes (Squamata: Agamidae) from the Bukit Barisan Range of Sumatra with an Estimation of its phylogeny. Zootaxa, 4276 (2), 215 - 232. https: // doi. org / 10.11646 / zootaxa. 4276.2.4","Gunther, A. (1872) Descriptions of some Ceylonese Reptiles and Batrachians. Annals Magazine of natural History, 4, 85 - 88. https: // doi. org / 10.1080 / 002229372011951778","Hardwicke, T. & Gray, J. E. (1827) A synopsis of the species of saurian reptiles, collected in India by Major-General Hardwicke. Zoological Journal, 3, 213 - 229.","Dumril, A. M. C. & Bibron, G. (1837) Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Fol. 4. Librairie Encyclopedique Roret, Paris, 570 pp.","Gray, E. (1845) Catalogue of the specimens of lizards in the collection British museum. Trustees of die British Museum / Edward Newman, London, 289 pp.","Jerdon, T. C. (1854) Catalogue of the Reptiles inhabiting the Peninsula of India. Part 1. Journal of Asiatic Society of Bengal, 22, 462 - 479.","Gunther, A. (1864) The Reptiles of British India. Taylor & Francis, London, 452 pp.","Gunther, A. (1870) Descriptions of a new Indian lizard of the genus Calotes. Proceedings of the Zoological Society of London, 778 - 779.","Gunther, A. (1875) Second report on collections of Indian Reptiles obtained by the British Museum. Proceedings of the Zoological Society of London, 1875, 224 - 234.","Biswas, S. (1975) Reptilia from Bhutan with description of a new species of Calotes Rafinesque. Journal of Bombay Natural History Society, 72, 774 - 777.","Zhao, E. & Li, S. (1984) A new species of Calotes (Lacertilia: Agamidae) from Xizang (Tibet) [in Chinese]. Acta Herpetologica Sinica, 3 (4), 77 - 78.","Vindum, J. V., Win, H., Thin, T., Lwin, K. S., Shein, A. K. & Tun, H. (2003) A new Calotes (Squamata: Agamidae) from the Indo-Burman range of western Myanmar (Burma). Proceedings of the California Academy of Science, 54 (1), 1 - 16.","Zug, G. R., Brown, H. H., Schulte, J. A. & Vindum, J. V. (2006) Systematics of the Garden Lizards, Calotes versicolor Group (Reptilia, Squamata, Agamidae), in Myanmar: Central Dry Zone Populations. Proceedings of the California Academy of Sciences, 57 (2), 35 - 68.","Manthey, U. (2008) Agamid Lizards of Southern Asia, Draconinae 1. Edition Chimaira, Frankfurt, 160 pp.","Hartmann, T., Geissler, P., Nikolay, A. J., Ihlow, P. F., Galoyan, E. A., Rodder, D. & Bohme, W. (2013) A new species of the genus Calotes Cuvier, 1817 (Squamata: Agamidae) from southern Vietnam. Zootaxa, 3599 (3), 246 - 260. https: // doi. org / 10.11646 / zootaxa. 3599.3.3","Amarasinghe, A. A. T., Karunarathna, D. M. S. S. & Fujinuma, J. (2014 a) A new Calotes species from Sri Lanka with a redescription of Calotes liolepis Boulenger, 1885. Herpetologica, 70 (3), 323 - 338. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00087","Amarasinghe, A. A. T., Karunarathna, D. M. S. S., Hallermann, J., Fujinuma, J., Grillitsch, H. & Campbell, P. D. (2014 b) A new species of the genus Calotes (Squamata: Agamidae) from high elevations of the Knuckles Massif of Sri Lanka. Zootaxa, 3785 (1), 059 - 078. http: // doi. org / 10.11646 / zootaxa. 3785.1.5","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) Fertebrate Zoology, 65 (3), 371 - 381.","Ishwar, N. M. & Das, I. (1998) Rediscovery of Calotes andamanensis Boulenger 1891 and a reassessment of the type locality. Journal of Bombay Natural History Society, 95, 513 - 514."]}

Monilesaurus acanthocephalus gen. et. sp. nov. (Fig. 7a) Etymology. The species epithet is derived by combining the Greek word ‘acanthos’, meaning spine or thorn, and ‘kephale’ latinized as ‘cephalus’ meaning head; referring to the long posterorbital and supratympanic spines. Holotype. BNHS 2409, an adult male (Fig. 11) collected from a disturbed habitat, adjoining evergreen forest—tea garden edge in Upper Manalar (Fig. 12), Periyar tiger reserve, Megamalai (934'35. 81"N, 7720'11. 43"E; 1562 m a.s.l) by SPV on 8th April 2009. Paratypes. CESL 0 0 1 adult male and CESL 112 juvenile male collected by SPV on 8th April 2009; BNHS 2410 adult male collected by SPP on 4th September 2011 from Upper Manalar, Periyar tiger reserve, Megamalai (9°34'18. 25"N, 77°20'5. 02"E, 1547 m). Diagnosis and comparison. A medium sized agamid (SVL = 72.6 mm) characterized by backward and downward orientation of lateral body scales; antehumeral fold present, throat fold present; 62–64 midbody scale rows; nuchal crest composed of 6 long, well developed spines; dorsal crest developed, in the form of a serrated ridge; two long, separated supratympanic spines; a long, well developed postorbital spine; dorsal and lateral scales keeled, ventral scales strongly keeled; paired postmentals, first pair separated by a 1 or 2 scales; 22–24 subdigital lamellae under fourth finger, 27–31 subdigital lamellae under fourth toe; 9 supralabials and 8 infralabials; reddish brown above with alternating dark and light crossbars on the dorsum, two white spots below the eye. Monilesaurus acanthocephalus gen. et sp. nov. can be can be distinguished from its congeners by a combination of the following characters: 62–64 midbody scale rows (vs. 46–52 in M. montanus gen. et sp. nov., 52–58 in M. ellioti and 52–56 in M. rouxii); presence of much longer, distinct isolated spine in the posterior corner of orbit (vs. absent in M. rouxii; very small, indistinct tubercle like in M. montanus gen. et sp. nov., and smaller in M. ellioti); 6 very long nuchal spines (vs. 3–6 small nuchal spines in M. montanus gen. et sp. nov., 3–4 long nuchal spines in M. ellioti, 7–8 smaller nuchal spines in C. rouxii); longer, prominent isolated spine on the back of head and above tympanum (vs. much smaller in M. montanus gen. et sp. nov., and M. rouxii) and presence of a white spot below the eye (vs. absent in M. rouxii; in the form of a band in M. montanus gen. et sp. nov.). Description of holotype. BNHS 2409, a medium sized adult male (SVL: 72.6 mm), morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately compressed. Head moderately large (HL/SVL ratio: 0.29), broad (HW/HL ratio: 0.65), maximum height less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield, which is separated from rostral by a single scale; mental shield narrower than rostral; two postmentals; first pair in contact with each other; genials keeled; gular scales strongly keeled, slightly smaller than genials; scales on top of snout smooth except median row, which is keeled; scales on top of head heterogenous in size and shape, keeled; supraorbital scales keeled; canthus-rostralis and supraciliary edge sharp; a long, well developed, thick spine at the posterior corner of the orbit, ca. 54% the orbit in length; two separated spines on posterior end of head, the anterior much longer, between the nuchal crest and tympanum, posterior above tympanum; orbit diameter 56% of distance between anterior border of orbit and snout tip; tympanum exposed, its greatest diameter 49% horizontal diameter of orbit; enlarged keeled scale between tympanum and orbit; posterior region of jaws swollen; supralabials 9/8; infralabials 8/8. Nuchal crest well developed, composed of six primary, long conical spines, the first being the smallest and the fourth longest; the remaining vertebral scales subtriangular, pointed, much larger than adjacent rows of scales, and possessing a strong, median keel forming an elevated, serrated ridge like the dorsal crest which continues till the tail base; 62 longitudinal scale rows around midbody; scales on dorsum keeled, oriented postero-dorsally, while lateral ones oriented postero-ventrally; lateral scales smaller than dorsal, keeled; ventrals strongly keeled, irregular, slightly smaller than dorsals but of similar size as laterals, genials and gular scales; a strong, oblique antehumeral fold, extending across the throat. Limbs long, slender and covered with strongly keeled scales, larger than laterals, forming parallel longitudinal rows; scales under thighs keeled; length of hindlimb ca. 79 % SVL; relative length of fingers 4>3>5>2>1; relative lengths of toes 4>3>5>2>1; fourth toe longer than fifth finger; 22 subdigital lamellae under fourth finger; 27 subdigital lamellae under fourth toe; subdigital lamellae with sharp keels, bicarinate; tail slender, swollen at the base; scales on dorsal and ventral surface of tail with sharply keeled, mucronate, larger than laterals; tail length 215 mm. Colouration. In life: dorsum olive red with a light brown head, irregular alternating red and black blotches on the mid dorsum, the black blotches extend to the lateral side in the form of alternate ‘v’ shaped bands; 4–5 light green blotches on the lateral side followed by thin whitish stripes from behind shoulder till start of tail; two light broken dorsolateral stripes present; head laterally light olive with three dark black bands from posterior part of eye till tympanum and two white spots below the eye; tympanum off-white with light green edge, lip scales light grey; a large blackish triangular patch behind the tympanum continuing to the antehumeral fold; ventral uniformly white; gular pouch light reddish; tail with alternating spaced dark blotches forming irregular bands towards the end. In preservative: dorsum and head uniform light brown, back banded with five ‘U’ shaped black bands from neck to start of tail; the black bands extend midway laterally forming an alternating pale brown and black pattern; tail banded with alternating thick brown and grey blotches; head laterally pale brown with yellowish white lip scales, tympanum pale grey; ventrally uniform pale whitish yellow with darker blotches below the limbs. Variation and secondary sexual characteristics. Morphometric and meristic data for the type specimens is presented in Appendix 2 & 3. The adult paratypes are all males and range from 68.9–72 mm in SVL. The paratypes agree with the holotype (CESL 002) in general morphology and scalation except for the following characters: 62– 64 longitudinal scale rows around midbody; 22–24 subdigital lamellae under fourth finger, 27–31 subdigital lamellae under fourth toe; infralabials 9 on the left in CESL 0 0 1 and 9 on the right in CESL 410; 1 st pair separated by a single scale in CESL 410. Dorsal and nuchal crest, spines above the orbit and tympanum not developed in the juvenile paratype CESL 112. Genetic distance. M. acanthocephalus gen. et sp. nov. shows 4–6% interspecific genetic divergence in the 16S gene from M. rouxii comb. nov.; 5–6 % interspecific genetic divergence from M. ellioti comb. nov. and 2–3 % interspecific genetic divergence from M. montanus gen. et sp. nov. (Appendix 5). Distribution. Monilesaurus acanthocephalus gen. et sp. nov. is currently known only from high elevations (above 1500 m asl) of Megamalai hills of southern Western Ghats (See Fig. 3 & Appendix 1 for details). Ecology and natural history. Monilesaurus acanthocephalus gen. et sp. nov. is a diurnal lizard, semiarboreal to arboreal in habit, and has, so far, been recorded from high elevation evergreen forests and along foresttea garden edges (Fig. 12). Individuals were seen perching on shrubs, branches and on tree trunks. One of the type specimens (CESL 410) was found sleeping on a tree branch in a forest patch.

Monilesaurus acanthocephalus gen. et. sp. nov. (Fig. 7a) Etymology. The species epithet is derived by combining the Greek word ���acanthos���, meaning spine or thorn, and ���kephale��� latinized as ���cephalus��� meaning head; referring to the long posterorbital and supratympanic spines. Holotype. BNHS 2409, an adult male (Fig. 11) collected from a disturbed habitat, adjoining evergreen forest���tea garden edge in Upper Manalar (Fig. 12), Periyar tiger reserve, Megamalai (934'35. 81"N, 7720'11. 43"E; 1562 m a.s.l) by SPV on 8th April 2009. Paratypes. CESL 0 0 1 adult male and CESL 112 juvenile male collected by SPV on 8th April 2009; BNHS 2410 adult male collected by SPP on 4th September 2011 from Upper Manalar, Periyar tiger reserve, Megamalai (9��34'18. 25"N, 77��20'5. 02"E, 1547 m). Diagnosis and comparison. A medium sized agamid (SVL = 72.6 mm) characterized by backward and downward orientation of lateral body scales; antehumeral fold present, throat fold present; 62���64 midbody scale rows; nuchal crest composed of 6 long, well developed spines; dorsal crest developed, in the form of a serrated ridge; two long, separated supratympanic spines; a long, well developed postorbital spine; dorsal and lateral scales keeled, ventral scales strongly keeled; paired postmentals, first pair separated by a 1 or 2 scales; 22���24 subdigital lamellae under fourth finger, 27���31 subdigital lamellae under fourth toe; 9 supralabials and 8 infralabials; reddish brown above with alternating dark and light crossbars on the dorsum, two white spots below the eye. Monilesaurus acanthocephalus gen. et sp. nov. can be can be distinguished from its congeners by a combination of the following characters: 62���64 midbody scale rows (vs. 46���52 in M. montanus gen. et sp. nov., 52���58 in M. ellioti and 52���56 in M. rouxii); presence of much longer, distinct isolated spine in the posterior corner of orbit (vs. absent in M. rouxii; very small, indistinct tubercle like in M. montanus gen. et sp. nov., and smaller in M. ellioti); 6 very long nuchal spines (vs. 3���6 small nuchal spines in M. montanus gen. et sp. nov., 3���4 long nuchal spines in M. ellioti, 7���8 smaller nuchal spines in C. rouxii); longer, prominent isolated spine on the back of head and above tympanum (vs. much smaller in M. montanus gen. et sp. nov., and M. rouxii) and presence of a white spot below the eye (vs. absent in M. rouxii; in the form of a band in M. montanus gen. et sp. nov.). Description of holotype. BNHS 2409, a medium sized adult male (SVL: 72.6 mm), morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately compressed. Head moderately large (HL/SVL ratio: 0.29), broad (HW/HL ratio: 0.65), maximum height less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield, which is separated from rostral by a single scale; mental shield narrower than rostral; two postmentals; first pair in contact with each other; genials keeled; gular scales strongly keeled, slightly smaller than genials; scales on top of snout smooth except median row, which is keeled; scales on top of head heterogenous in size and shape, keeled; supraorbital scales keeled; canthus-rostralis and supraciliary edge sharp; a long, well developed, thick spine at the posterior corner of the orbit, ca. 54% the orbit in length; two separated spines on posterior end of head, the anterior much longer, between the nuchal crest and tympanum, posterior above tympanum; orbit diameter 56% of distance between anterior border of orbit and snout tip; tympanum exposed, its greatest diameter 49% horizontal diameter of orbit; enlarged keeled scale between tympanum and orbit; posterior region of jaws swollen; supralabials 9/8; infralabials 8/8. Nuchal crest well developed, composed of six primary, long conical spines, the first being the smallest and the fourth longest; the remaining vertebral scales subtriangular, pointed, much larger than adjacent rows of scales, and possessing a strong, median keel forming an elevated, serrated ridge like the dorsal crest which continues till the tail base; 62 longitudinal scale rows around midbody; scales on dorsum keeled, oriented postero-dorsally, while lateral ones oriented postero-ventrally; lateral scales smaller than dorsal, keeled; ventrals strongly keeled, irregular, slightly smaller than dorsals but of similar size as laterals, genials and gular scales; a strong, oblique antehumeral fold, extending across the throat. Limbs long, slender and covered with strongly keeled scales, larger than laterals, forming parallel longitudinal rows; scales under thighs keeled; length of hindlimb ca. 79 % SVL; relative length of fingers 4>3>5>2>1; relative lengths of toes 4>3>5>2>1; fourth toe longer than fifth finger; 22 subdigital lamellae under fourth finger; 27 subdigital lamellae under fourth toe; subdigital lamellae with sharp keels, bicarinate; tail slender, swollen at the base; scales on dorsal and ventral surface of tail with sharply keeled, mucronate, larger than laterals; tail length 215 mm. Colouration. In life: dorsum olive red with a light brown head, irregular alternating red and black blotches on the mid dorsum, the black blotches extend to the lateral side in the form of alternate ���v��� shaped bands; 4���5 light green blotches on the lateral side followed by thin whitish stripes from behind shoulder till start of tail; two light broken dorsolateral stripes present; head laterally light olive with three dark black bands from posterior part of eye till tympanum and two white spots below the eye; tympanum off-white with light green edge, lip scales light grey; a large blackish triangular patch behind the tympanum continuing to the antehumeral fold; ventral uniformly white; gular pouch light reddish; tail with alternating spaced dark blotches forming irregular bands towards the end. In preservative: dorsum and head uniform light brown, back banded with five ���U��� shaped black bands from neck to start of tail; the black bands extend midway laterally forming an alternating pale brown and black pattern; tail banded with alternating thick brown and grey blotches; head laterally pale brown with yellowish white lip scales, tympanum pale grey; ventrally uniform pale whitish yellow with darker blotches below the limbs. Variation and secondary sexual characteristics. Morphometric and meristic data for the type specimens is presented in Appendix 2 & 3. The adult paratypes are all males and range from 68.9���72 mm in SVL. The paratypes agree with the holotype (CESL 002) in general morphology and scalation except for the following characters: 62��� 64 longitudinal scale rows around midbody; 22���24 subdigital lamellae under fourth finger, 27���31 subdigital lamellae under fourth toe; infralabials 9 on the left in CESL 0 0 1 and 9 on the right in CESL 410; 1 st pair separated by a single scale in CESL 410. Dorsal and nuchal crest, spines above the orbit and tympanum not developed in the juvenile paratype CESL 112. Genetic distance. M. acanthocephalus gen. et sp. nov. shows 4���6% interspecific genetic divergence in the 16S gene from M. rouxii comb. nov.; 5���6 % interspecific genetic divergence from M. ellioti comb. nov. and 2���3 % interspecific genetic divergence from M. montanus gen. et sp. nov. (Appendix 5). Distribution. Monilesaurus acanthocephalus gen. et sp. nov. is currently known only from high elevations (above 1500 m asl) of Megamalai hills of southern Western Ghats (See Fig. 3 & Appendix 1 for details). Ecology and natural history. Monilesaurus acanthocephalus gen. et sp. nov. is a diurnal lizard, semiarboreal to arboreal in habit, and has, so far, been recorded from high elevation evergreen forests and along foresttea garden edges (Fig. 12). Individuals were seen perching on shrubs, branches and on tree trunks. One of the type specimens (CESL 410) was found sleeping on a tree branch in a forest patch., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 431-434, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674

Psammophilus dorsalis (Gray, 1831) (Fig. 8a) Agama dorsalis –Gray, 1831. In Griffith, E & E. Pidgeon’s Anim. King. ix, 1851: 56 Charasia dorsalis – Gray, 1845. Cat. Liz. Brit. Mus. 1845: 246. Charasia dorsalis — Boulenger, 1885. Cat. Liz. Brit. Mus. 1845: 450. Psammophilus dorsalis — Smith, 1935. Fauna of British India, ii, 1935: 209. Original description. Gray, J. E. 1831. A synopsis of the species of Class Reptilia. In: Griffith, E & E. Pidgeon: The animal kingdom arranged in conformity with its organisation by the Baron Cuvier with additional descriptions of all the species hither named, and of many before noticed [V Whittaker, Treacher and Co., London: 481 + 110 pp. [1830] Taxonomic comments. Gray (1831) failed to mention which specimens he used to describe P. dorsalis. Currently, there are 20 specimens labelled as P. dorsalis at the Natural History Museum. Boulenger (1885) used eight specimens at NHM to describe Charasia dorsalis collected from Madras, Nilgiris, Malabar and India and labelled these collections as being from Southern India. There was no precise type locality given in the original description but most of the localities mentioned in older publications suggest collections from different parts of southern India, especially hilly regions (Boulenger 1890; Smith 1935). The chronological collections of the specimens from NHM need to be verified to choose a lectotype from the specimens at NHM. Since there is very little genetic variation (1–2% in 16S mtDNA) in the samples collected from the Western Ghats of Peninsular India, we provide an expanded description of this species based on the largest male collected during this study and compare with other specimens collected during the study. Topotypic material. CESL 273, adult male collected from a boulder in grassland in Kotavasal, Achankovil reserve forest, Kerala by SPP and MVP on 28th May 2011. Other material examined. CESL 326 adult male collected from Kolakumbai, Nilgiri hills, Tamil Nadu; CESL 0 64 adult female collected from sunrise valley, Wayanad reserve forest, Kerala and CESL 181 adult female collected from near Rishi Valley school, Andhra Pradesh; details of collection locality, specimen voucher and GenBank accession number in Appendix 1. Diagnosis and comparison. A medium to large sized Psammophilus characterized by the posterodorsal orientation of lateral scales; antehumeral fold well developed; 94–148 midbody scale rows (vs 115–140 in Boulenger, 1885; 115–150 Smith, 1935); ventral scales 112–152; nuchal crest reduced, composed of 15–18 very small, triangular spines, continuous with almost equal to or even smaller dorsal crest which appears like slightly raised denticulation; dorsal crest absent in females and juveniles; two separated, very small supratympanic spines; dorsal and lateral scales keeled, ventral scales strongly keeled; paired postmentals, first pair separated by 1–2 scales; 24–28 subdigital lamellae under fourth finger, 26–34 subdigital lamellae under fourth toe; 11–13 supralabials and 10–12 infralabials; adult males with uniform greyish brown to black body, dorsum lighter; indistinct lighter blotches on the sides, lip scales lighter grey or off-white; females and juveniles with a pair of dorsolateral stripes from neck to above tail, dorsum and lateral speckled with lighter spots. Morphologically, P. dorsalis is similar to P. blanfordanus, the only other species known from this group, but can be distinguished by a combination of the following characters: 100–140 midbody scale rows (vs 80–100); three samples collected from eastern India close to the type locality of P. blanfordanus also had lower number 84–94 (80–100 Boulenger 1885 & Smith, 1935). Midbody scales row counts clearly overlap with P. dorsalis. Furthermore, all the meristic characters we checked for few specimens overlap (Appendix 3). There is, therefore a need for more thorough examination of morphological and morphometric characters with a larger sample size to diagnose P. dorsalis from P. blanfordanus. Description. Based on CESL 273, a large sized adult male (SVL- 107.7 mm), morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately depressed. Head moderately large (HL/SVL 0.25), not elongated (HW/HL 1.10), maximum width much more than maximum height; snout rounded; rostral broader than high; nostrils in single nasal shield, which is separated from the rostral by two scales; mental shield broader than rostral; two postmentals, first pair separated by two small scales; genials smooth; gular scales keeled, smaller than genials; scales on top of snout feebly keeled towards the edges; scales on top of head heterogenous in size and shape, keeled; supraorbital scales keeled; canthus-rostralis and supraciliary edge sharp; two separated small, triangular spines on posterior end of head, the anterior midway between nuchal crest and tympanum, posterior above tympanum; orbit diameter 65% of distance between anterior border of orbit and snout tip; tympanum exposed, its greatest diameter 39% horizontal diameter of orbit; few slightly enlarged, smooth scales between tympanum and orbit; posterior region of jaws swollen; supralabials 13/13; infralabials 12/12. Nuchal crest developed, composed of 15 primary, small, triangular spines, the first being the smallest and the eight longest; nuchal crest continuous with a reduced dorsal crest present in the form of elevated serrated ridge, composed of vertebral scales with a more pronounced median keel continuing till above the vent, on the tail in the form of enlarged strongly keeled median scale; 104 longitudinal scale rows around midbody; dorsal and lateral scales keeled, oriented postero-dorsally; ventrals smooth, irregular, as large as the dorsals, slightly larger than the genial and gular scales; an oblique antehumeral fold present, not extending across the throat. Limbs strong, robust, covered with uniform keeled scales, slightly larger than laterals forming parallel longitudinal rows; scales under thighs weakly keeled; length of hindlimb 71% SVL; relative length of fingers 4>3>2>5>1; relative lengths of toes 4>3>5>2>1; fourth toe slightly smaller than fifth finger; 15 subdigital lamellae under fourth finger; 19 subdigital lamellae under fourth toe; subdigital lamellae with sharp keels, bicarinate; tail thick, gradually tapering, swollen at the base; scales on dorsal and ventral surface of tail with sharp keels, larger than laterals; median scales on the dorsal surface of tail enlarged, with a pronounced median keel; tail length 147 mm (incomplete, broken at the tip). Colouration. In life: head and body dull blackish-grey with irregular lighter patches near the flank; a longitudinal thick, lighter buff band from behind the head till above the tail; head laterally darker with indistinct reddish brown speckles behind the orbit, a broken reddish stripe from above the nostril till behind the posterior part of orbit covering the supraorbital region; another pale reddish stipe from the nostril till behind the cheek covering the lip scales and lower margin of tympanum; tympanum pale grey; ventral uniformly lighter, pale grey; gular and genial scales lighter, dirty white; tail with a thick lighter band just at the start, then uniform dark grey followed by indistinct lighter bands, bands more prominent towards the tip. In preservative: colouration pattern mostly similar to that in life, except overall paler; bands on lateral side of head distinct. Variation and secondary sexual characteristics. The other specimens examined agree with the CESL 273 in general morphology and scalation except for some differences that are summarised in Appendix 2 & 3. The examined female specimen (CESL 064) has slightly smaller nuchal spines compared to the males, dorsal crest almost nonexistent; dorsal and lateral scales feebly keeled; tail base not swollen; overall colouration olive-brown, speckled with irregular lighter blotches, 3–4 triangular darker patches on the mid dorsum from neck to the start of tail and series of whitish spots forming broken parallel dorso-lateral stripes. Distribution. Psammophilus dorsalis is distributed all across the low and medium elevation rocky habitats of central and southern Western Ghats. This species has often been observed on huge rock boulders and sheath rocks in and around forests but seem to be more common in open, drier habitats. During this study, P. dorsalis was recorded in various sites from the southern Western Ghats (See Appendix 1 for details). It is also known from drier habitats throughout peninsular India. Ecology and natural history. Psammophilus dorsalis is a diurnal lizard, chiefly rupicolous in habit, and so far, has been recorded exclusively in and around rocky habitats. Individuals were mostly observed basking on boulders and rock cliffs in the mornings. In some instances, it has also been observed on isolated rocks inside tea estates and plantations. It is a very shy and active species and escapes inside the gaps and crevices among boulders at the slightest hint of an approaching threat. In the breeding season, the adult males of this species develop a thick bright red to vermilion coloured band from the head extending till behind the lower back and the entire remaining body turns black. Adult males have been observed only during summer and pre-monsoon while gravid females were found in the early monsoon seasons., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 438-440, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674, {"references":["Gray, E. (1845) Catalogue of the specimens of lizards in the collection British museum. Trustees of die British Museum / Edward Newman, London, 289 pp.","Boulenger, G. A. (1885) Catalogue of lizards in the British museum. Fol. 1. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae, Second edition. British Museum (Natural History), London, 436 pp.","Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Fol. II. Sauria. Taylor and Francis, London, 440 pp.","Boulenger, G. A. (1890) The Fauna of British India, including Ceylon and Burma: Reptilia and Batrachia. Taylor and Francis, London, 541 pp."]}

Calotes calotes (Linnaeus, 1758) Lacerta calotes ���Linnaeus, 1758. Syst. Nat. 10th ed. 1758: 207. Iguana calotes ���Laurenti 1768. Syn. Rept. 1768: 49. Agama calotes ���Daudin 1802. Hist. Nat. Rept. iii.1802: 361. Calotes calotes ���L��nnberg 1896. Bih. Svensk. Vet. Akad. Stockholm, xxii. 1896:15. Original description. Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiae. 10th Edition, 1758: 207. Taxonomic comments. Calotes calotes was described based on specimens collected from ���Zeylona��� (Sri Lanka). This species is currently known to commonly occur in the wet zones of Sri Lanka and dry zones of southern India. Smith (1935) reported this species from the Nicobar Islands but this was not included in later works (Harikrishnan et al. 2010). The original description by Linnaeus was a two-line description, which was translated by Amarasinghe et al. (2009). Our analysis shows 1% genetic variance (in the 16S gene) between the Sri Lankan and the Peninsular Indian population of Calotes calotes. This suggests that these species moved between the dry zones in Peninsular India and Sri Lanka. Material examined. CESL 0 37 adult female collected from Peppara Wildlife Sanctuary, Kerala; CESL 374 adult male collected from Marayoor, Kerala and BNHS 378 adult male collected Chatikulam, Sri Lanka. Details of collection locality, specimen voucher and GenBank accession number in Appendix 1. Diagnosis and comparison. A medium to large sized Calotes characterized by the posterodorsal orientation of lateral scales; antehumeral pit indistinct; 30���35 midbody scale rows; nuchal and dorsal crest continuous, nuchal crest composed of long lanceolate spines; dorsal spines smaller, gradually diminishing in size, extending beyond the midbody; a row of 8���9 compressed supratympanic spines, divided into two groups; dorsal and lateral scales large, smooth or feebly keeled, ventral scales strongly keeled, mucronate, equal to or slightly larger than laterals; paired postmentals, first pair separated by 1 to 2 median scales; 21���24 subdigital lamellae under fourth finger, 27��� 29 subdigital lamellae under fourth toe; 10���12 supralabials and 9���11 infralabials; uniform, bright green above, with 5���6 thin white transverse stripes, often continuing on the tail as indistinct bands; a bright red horizontal stripe across the orbit; posterior part of the tail dull brown; ventrally pale green; juveniles bright green, white transverse stripes indistinct. From other members of Smith���s C. versicolor group (C. versicolor, C. nemoricola, C. grandisquamis, C. minor, C. maria, C. jerdoni, C. emma and C. mystaceus) C. calotes can be distinguished by a combination of the following characters: smaller body size: adult SVL 73.6���96.8 mm, n=3 (vs. C. nemoricola, adult SVL 112���134.8 mm, n=7; and C. grandisquamis, adult SVL 110.0��� 136.5 mm, n=5); 30���35 midbody scale rows (vs. 38���45 in C. versicolor, n=9, 36��� 43 in C. nemoricola; 58���63 in C. maria; 45���57 in C. jerdoni; 49���65 in C. emma; 48���58 in C. mystaceus and 48���60 in C. minor); ventral scales equal to or slightly larger than laterals (vs. ventral scales smaller than laterals in C. nemoricola, C. grandisquamis, C. versicolor, C. minor, C. maria, C. jerdoni, C. emma and C. mystaceus); crescent-shaped patch of granular scales in front of the shoulder absent (vs. present in C. nemoricola, C. grandisquamis, C. jerdoni, C. emma and C. mystaceus); and a row of 8���9 compressed supratympanic spines (vs. row of 3���4 compressed supratympanic spines in C. nemoricola and C. grandisquamis; two well separated supratympanic spines in C. versicolor; two parallel rows of compressed supratympanic scales in C. maria and C. jerdoni; single well developed postorbital spine in C. emma). Description. Based on specimen CESL 374. A large sized adult male (SVL- 84.7 mm) collected from a shrub along the road near Marayoor, Kerala. Morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately compressed. Head moderate (HL/SVL ratio 0.23), elongate (HW/HL ratio 0.73), maximum height slightly less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield which is separated from rostral by three scales; mental shield broader than rostral; two postmentals, first pair separated from each other by two small scales; genials smooth; gular sac small, indistinct, composed of strongly keeled scales, slightly smaller than genials; scales on top of snout smooth; scales on top of head heterogenous in size and shape, smooth; canthus-rostralis and supraciliary edge sharp; a row of 8 compressed, spines divided into two groups above the tympanum, the one closer to the nuchal crest longer; orbit diameter 86% of distance between anterior border of orbit and snout tip; tympanum exposed, its greatest diameter 49% of horizontal diameter of orbit; partially keeled, scales between tympanum and orbit smooth, some enlarged, a few with partially keeled edges; posterior region of jaws swollen; supralabials 10/11; infralabials 10/10. Nuchal crest well developed, dorsal crest slightly reduced, continuous; nuchal crest composed of 12 long, conical spines, the first being the smallest; dorsal crest composed of much smaller spines reaching slightly behind midbody; 30 longitudinal scale rows around midbody; dorsal and lateral scales sub-triangular, smooth, ones on the lower back feebly keeled, oriented postero-dorsally; ventrals strongly keeled, mucronate, almost equal to the dorsal and lateral scales. Limbs slender and covered with keeled scales forming parallel longitudinal rows; scales under thighs weakly keeled; length of hindlimb ca. 101 % SVL; relative length of fingers 4>3>5>2>1, second and fifth finger almost equal; relative lengths of toes 4>3>5>2>1; fourth toe much longer than fifth finger; 21 subdigital lamellae under fourth finger; 27 subdigital lamellae under fourth toe; subdigital lamellae with sharp keels, bicarinate; slender, swollen at the base; dorsal and ventral tail scales keeled, ventral tail scales mucronate, strongly keeled; tail length 268 mm. Colouration. In life: dorsum and head uniform, bright green with 5 thin, white transverse stripes, continuing on the tail as indistinct bands; head bright green laterally, lighter towards the cheeks; nuchal spines brownish, a brick red horizontal stripe from posterior corner of the eye till end of orbit, darker behind the eye; few blackish scales below the eye; gular scales lighter, yellowish green; posterior half of the tail dull greyish brown with alternating white bands; tympanum pale grey with greenish edge, ventrally lighter, uniform pale green. Representative image showing live colouration (Fig. 4a). In preservative: colouration mostly similar to that in life, except overall paler. Variation and secondary sexual characteristics. Meristic and morphometric data of two adult males and one adult female are given in Appendix 2 & 3. The examined specimens agree with each other in general morphology and scalation. The examined female specimen (CESL 037) is slightly larger in size and has much smaller nuchal spines, compared to the males, almost equal to the dorsal spines. Distribution. Calotes calotes is distributed in the low and medium elevation dry forests (up to 1000 m asl) of the southern Western Ghats, Eastern Ghats and across Sri Lanka. In the present study, C. calotes was recorded from various sites in the low to mid elevation dry and deciduous forests of the southern Western Ghats (See Appendix 1 for details). This species is common in and around open scrub, secondary and degraded forest patches. Ecology and natural history. Calotes calotes is a diurnal lizard, semi-arboreal to arboreal in habit, and so far, has been recorded mostly in scrub, secondary deciduous forests and plantations. Individuals were mostly seen perching on shrubs, bushes and on trees near edges of forests. In some instances, it has also been observed near villages and within plantations. During this study, it was never recorded from primary evergreen forests. In some sites, they might occur syntopically in the same habitat as C. versicolor., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 419-421, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674, {"references":["Smith, M. A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Fol. II. Sauria. Taylor and Francis, London, 440 pp.","Harikrishnan, S., Vasudevan, K. & Choudhury, B. C. (2010) A review of herpetofaunal descriptions and studies from Andaman and Nicobar Islands, with an updated checklist. In: Ramakrishna, Raghunathan, C. & Sivaperuman, C. (Eds.), Recent Trends in Biodiversity of Andaman and Nicobar Islands. Zoological Survey of India, Kolkata, pp 387 - 398.","Amarasinghe, A. A. T., Manthey, U., Stockli, E., Ineich, I., Kullander, S. O., Tiedemann, F., McCarthy, C. & Gabadage, D. E. (2009) The original descriptions and figures of Sri Lankan Agamid lizards (Squamata: Agamidae) of the 18 th and 19 th centuries. Taprobanica, 1 (1), 2 - 15. https: // doi. org / 10.4038 / tapro. v 1 i 1.2771"]}

Monilesaurus montanus gen. et. sp. nov. (Fig. 7b) Etymology. The species epithet is derived from the word ���montane��� referring to the restricted distribution of this species to high elevation forests (> 1500 m a.s.l). Holotype. BNHS 2411, adult male (Fig. 13), collected at Kudremukh National Park (Fig. 14), Karnataka (1307'54" N, 07516'39" E; 1534.9 m a.s.l); by SPP, SPV and KPD on 28th September 2011. Paratypes: BNHS 2412, adult male and BNHS 2413, adult female collected from Kudremukh National Park, Karnataka by SPV and ADR on 21st August 2009; CESL 131, adult female collected from Siruvani reserve forest, Kerala by SPP and MPV on 0 6th July 2010; CESL 133, adult female collected from Walakkad, Silent Valley National Park, Kerala by SPP and MPV on 23rd November 2010; CESL 330 adult female and CESL 331 an adult male collected from Naduvattam, Tamil Nadu by SPP on 23rd June 2011 and CESL 529 adult male collected from Narimala, Brahmagiri Wildlife Sanctuary, Karnataka by SPP and SRC on 0 9th March 2012. Diagnosis and comparison. A medium sized Monilesaurus with a maximum SVL of 83.4 mm, arboreal species characterized by the backward and downward orientation of lateral body scales; presence of antehumeral fold, throat fold not prominent as the antehumeral fold; 46���52 midbody scale rows; nuchal crest composed of 3���6 small spines; two small separated supratympanic spines; a very small tubercle like postorbital spine barely distinguishable from the surrounding head scales; dorsal and lateral scales feebly keeled, stronger towards ventrals, ventral scales strongly keeled; scales on ventral thigh region feebly keeled; paired postmentals, first pair separated by a single scale; 21���24 subdigital lamellae under fourth finger, 25���30 subdigital lamellae under fourth toe; 9���10 supralabials and 8���9 infralabials; greenish brown above with darker dorsum, a white band below the eye extending till end of jaw. Monilesaurus montanus gen. et sp. nov. can be distinguished from its sister species based on the combination of following characters: larger body size: adult SVL 61���83.4 mm, n=8 (vs. M. ellioti comb. nov. adult SVL 59.4��� 73.8 mm, n=9; M. rouxii comb. nov. adult SVL 51.4���74.8 mm, n=9); lower number of midbody scale rows 46���52 (vs. 62���64 in Monilesaurus acanthocephalus gen. et sp. nov., 52���58 in M. ellioti comb. nov., 52���56 in M. rouxii comb. nov.), presence of a very small, indistinct tubercle like, isolated spine in the posterior corner of orbit (vs. absent in M. rouxii comb. nov., long, distinct isolated spine in M. ellioti comb. nov. and M. acanthocephalus gen. et sp. nov.), 3���6 small nuchal spines (vs. 7���8 small nuchal spines in M. rouxii comb. nov., 6 much longer nuchal spines in C. acanthocephalus gen. et sp. nov., 3���4 long nuchal spines in M. ellioti comb. nov.); small isolated spine on the back of head and above tympanum (vs. longer, prominent spines in M. ellioti comb. nov. and M. acanthocephalus gen. et sp. nov.) presence of white band below the eye (vs. none in M. rouxii comb. nov.; in the form of a spot in M. ellioti comb. nov. and M. acanthocephalus gen. et sp. nov.). Description of holotype. BNHS 2411, a medium sized agamid, adult male (SVL- 78.2 mm). Morphometric and meristic data are summarised in Appendix 2 & 3. General habitus moderately compressed. Head moderately large (HL/SVL ratio 0.31), broad (HW/HL ratio 0.64), maximum height less than maximum width; snout pointed; rostral broader than high; nostrils in single nasal shield, which is separated from rostral by a single scale; mental shield narrower than rostral; two postmentals; first pair separated from each other by a single, small scale; genials keeled; gular sac small, composed of strongly keeled scales, slightly smaller than genials; scales on top of snout smooth except median row, which is keeled; scales on top of head heterogenous in size and shape, keeled; supraorbital scales keeled; canthus-rostralis and supraciliary edge sharp; a very small tubercle like spine at the posterior corner of the orbit; two separated small spines on posterior end of head, the anterior slightly longer, midway between nuchal crest and tympanum, posterior above tympanum; orbit diameter 59% of distance between anterior border of orbit and snout tip; tympanum exposed, its greatest diameter 48% of horizontal diameter of orbit; slightly keeled, enlarged scales between tympanum and orbit; posterior region of jaws swollen; supralabials 9/9; infralabials 8/8. Nuchal crest well developed, composed of three primary, conical spines, the first being the shortest, the third longest; the remaining vertebral scales slightly enlarged relative to adjacent rows and possess a median keel forming a slightly elevated ridge like dorsal crest which continues till the tail base; 50 longitudinal scale rows around midbody; 40 scales on the mid dorsum; scales on dorsum feebly keeled, oriented backwards, lateral scales smaller than dorsal, keeled, oriented backwards and downwards; ventrals strongly keeled, irregular, slightly smaller than dorsals but of similar size as laterals, genials and gular scales; an oblique antehumeral fold present, weakly developed, feebly extending into a throat fold which is not prominent. Limbs slender and covered with strongly keeled scales, larger than laterals, forming parallel longitudinal rows; scales under thighs weakly keeled; length of hindlimb ca. 71% SVL; relative length of fingers 4>3>2>5>1; relative lengths of toes 4>3>5>2>1; fourth toe much longer than fifth finger; 23 subdigital lamellae under fourth finger; 28 subdigital lamellae under fourth toe; subdigital lamellae with sharp keels, bicarinate; slender, swollen at the base; scales on dorsal and ventral surface of tail with sharp keels, mucronate; tail length 190 mm. Colouration. In life: dorsum and head yellowish-green with irregular, alternating light and dark brown patches on the back; a dark brownish band from above the shoulder till the dorsal crest forming a ���v��� shape; head laterally greenish with a whitish band below the posterior end of eye to end of jaw; broken dark striations from above nostril to anterior margin of orbit extending till the tympanum from the posterior margin of orbit in the form of black band; tympanum pale green, lip scales white; thick blackish triangular patch behind the tympanum continuing to the antehumeral fold; an indistinct thin stripe above the labials from the nostril, ending into a black spot anterior to the tympanum; ventral uniformly lighter, pale grey; gular pouch white, irregular dark striations on the sides; tail with alternating dark and light blotches forming irregular bands towards the end. In preservative: dorsum and head buff to light brown with irregular, alternating grey and darker brown patches on the back; tail banded with alternating thick brown and grey blotches; laterally paler with darker brown blotches extending down from the dorsum; lead laterally speckled with grey and brown with a whitish yellow below the posterior end of eye to end of jaw; tympanum pale white, lip scales whitish yellow; ventrally uniform pale grey, lighter below the limbs. Variation and secondary sexual characteristics. Morphometric and meristic data for the type specimens is presented in Appendix 2 & 3. Adult male paratypes range from 61���78.8mm in SVL, whereas adult female paratypes range from 68.4���83.4mm in SVL. The paratypes agree with the holotype (BNHS 2411) in general morphology and scalation except for the following characters: 46���52 longitudinal scale rows around midbody; 21��� 24 subdigital lamellae under fourth finger, 25���30 subdigital lamellae under fourth toe. Supralabials 8 on the right in CESL 124; infralabials 8 on the right in CESL 133 and 8 on the left in CESL 529. All the examined female paratypes (CESL 126, CESL 131, CESL 133 and CESL 331) have slightly smaller nuchal spines compared to the males; lack a dorsal crest and gular sac. Genetic distance. M. montanus gen. et sp. nov. shows 0���1% intraspecific genetic divergence in the 16S gene; 7���8% interspecific genetic divergence from M. rouxii comb. nov.; 4���8% interspecific genetic divergence from M. ellioti comb. nov. and 2���3 % interspecific genetic divergence from M. acanthocephalus gen. et sp. nov. (Appendix 5). Distribution. Monilesaurus montanus gen. et sp. nov., is endemic to the Western Ghats and distributed across the high elevation evergreen forests (above 1250 m asl) of central Western Ghats. During this study, M. montanus gen. et sp. nov. was recorded from montane forests of the following hill ranges: Kudremukh, Brahmagiri, Nilgiri and Elivalmalai (See Fig. 3 & Appendix 1 for details). Ecology and natural history. Monilesaurus montanus gen. et sp. nov., is a diurnal lizard, semi-arboreal to arboreal in habit and so far, has been recorded from montane shola forests (Fig. 14). Individuals were mostly found at night, sleeping on branches of stunted trees within sholas or actively moving on tree trunks. Only in one instance, a female specimen (CESL 133) was found in evergreen forest at a slightly lower elevation (ca. 1250 m asl). In some sites, the lower elevational limits of their distributional range might overlap with the range of M. ellioti comb. nov., but these two species were not observed to be syntopic in any of the sites. A gravid female was recorded in the month of July in Siruvani reserve forest, which hints that pre-monsoon might be a breeding season for this species. No other congeners were found to be syntopically distributed with M. montanus gen. et sp. nov., Published as part of Pal, Saunak, Vijayakumar, S. P., Shanker, Kartik, Jayarajan, Aditi & Deepak, V., 2018, A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species, pp. 401-450 in Zootaxa 4482 (3) on pages 435-437, DOI: 10.11646/zootaxa.4482.3.1, http://zenodo.org/record/1440674