Two New Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from the Shevaroy Massif, Tamil Nadu, India, with a Preliminary ND2 Phylogeny of Indian Cnemaspis

Zootaxa 4609 (1): 068–100 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2019 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4609.1.3 http://zoobank.org/urn:lsid:zoobank.org:pub:FF36B5CC-0ECE-4F16-9002-CA04F9352A05

Two new Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from the Shevaroy massif, Tamil Nadu, India, with a preliminary ND2 phylogeny of Indian Cnemaspis

AKSHAY KHANDEKAR, NIKHIL GAITONDE & ISHAN AGARWAL* National Centre for Biological Sciences, Tata Institute of Fundamental Research, Bengaluru, 560065, India *Corresponding author. E-mail: [email protected]

Abstract

We present a preliminary ND2 phylogeny of South Asian Cnemaspis, recovering a number of deeply divergent clades within Indian Cnemaspis, endemic to the southern and northern Western Ghats besides the Mysore Plateau and hills of Tamil Nadu. There are a number of unnamed lineages that are >5% divergent on ND2 across the phylogeny, including three from the gracilis clade on an elevation gradient (800–1400 m asl.) around Yercaud in the Shevaroy massif, Salem district, Tamil Nadu. We describe two of these as new species— Cnemaspis shevaroyensis sp. nov. and Cnemaspis thackerayi sp. nov. are both allied to Cnemaspis gracilis and can be diagnosed from all other Indian Cnemaspis by the absence of spine-like scales on flank, heterogeneous dorsal pholidosis, presence of femoral and precloacal pores, tail with enlarged, strongly keeled, conical tubercles forming whorls, a median row of enlarged and smooth sub-caudals. They differ from C. gracilis and each other in body size, the number of tubercles around midbody, the number of tubercles in paravertebral rows, the number of femoral and precloacal pores, the number of poreless scales in-between precloacal pores and between femoral and precloacal pores, and subtle colour pattern differences; besides uncorrected mitochondrial sequence divergence (7.9–16.6 %). We also provide a description of Cnemaspis yercaudensis from its type locality and an additional locality. The discovery of two endemic species and a third unnamed divergent lineage from an isolated massif in peninsular India outside the Western Ghats indicate that many other such understudied hill ranges may harbour high endemic biodiversity.

Key words: Cryptic species, Dwarf geckos, Gekkonidae, mountains, peninsular India, taxonomy, Tamil Nadu

Introduction

The gekkonid genus Cnemaspis as currently defined is polyphyletic, including 140 species of round-pupiled geckos patchily distributed across Africa and Asia (Gamble et al. 2012; Grismer et al. 2014; Cyriac et al. 2018; Sayyed et al. 2018; Uetz et al. 2018; Khandekar 2019). The three clades within South Asian Cnemaspis that are not closely related to each other include Southeast Asian Cnemaspis (Cnemaspis sensu stricto), African Cnemaspis and South Asian Cnemaspis (Gamble et al. 2012, 2015; Grismer et al., 2014). South Asian Cnemaspis have a disjunct distribution including India, Sri Lanka, and at least some offshore islands in southeast Asia (Bauer et al. 2007; Agarwal et al. 2017). The main centres of diversity within South Asian Cnemaspis are Sri Lanka (24 species) and peninsular India (34 species), with a single species known from northeast India, and three from southeast Asia (Grismer et al. 2014; Agarwal et al. 2017; Sayyed et al. 2018; Uetz et al. 2018; Khandekar 2019). Most species from peninsular India are from the Western Ghats (29 species), and though the first Cnemaspis outside the Western Ghats in peninsular India was discovered 165 years ago (Jerdon 1853), only five more species have been described since, all in the last two decades (Das & Bauer 2000; Srinivasulu et al. 2015; Khandekar 2019). These five species are disjunctly distributed across a vast area of peninsular India, spanning ~4° latitude and ~3° longitude. Cnemaspis adii Srinivasulu, Kumar & Srinivasulu is from rocky habitats around Hampi on the northern Mysore Plateau in Karnataka and C. mysoriensis (Jerdon) is known from around Bangalore on the southern edge of the Mysore

68 Accepted by A. Bauer: 12 Mar. 2019; published: 22 May 2019 Plateau; while the other three species are from the hills of Tamil Nadu at the edge and just off the Mysore Plateau— C. agarwali Khandekar from a small isolated hillock C. otai Das & Bauer from Vellore Fort, and C. yercaudensis Das & Bauer is from Yercaud in the Shevaroy massif (Shevaroy Hills or Shevaroys or Shevaroyan, the anglicized form of Serverayan) (Fig 1).

FIGURE. 1. Relief map showing distribution of four divergent Cnemaspis lineages in the Shevaroy massif and Kollimalai. Inset map of India shows location of study area (black rectangle).

The Shevaroys are an isolated massif, about 150 km east of the Nilgiris in the Western Ghats and 50 km north of Kollimalai (Kolli Hills), contiguous to the east with the Chitteri and Kalrayan hills. Elevations range from <350 m asl. to ~450 m asl. at the base on the southern and northern aspects up to a maximum of 1623 m asl., with a large plateau ~1000 m asl. and above. The affinities of the Shevaroys are with the Western Ghats (Raheem et al. 2014) or more broadly the hills of southern India (Agarwal et al. 2019), and at least two reptiles are endemic to the massif — the gecko Hemiphyllodactylus aurantiacus (Beddome) and the uropeltid snake Uropeltis shorttii (Beddome) (Ganesh et al. 2014). Cnemaspis yercaudensis was the only Cnemaspis reported from the Shevaroys, described based on a single sub-adult male specimen collected from Yercaud town (Das & Bauer 2000). Subsequently, Ganesh and Arumugam (2016) reported that C. yercaudensis is distributed at high elevations above 1350 m asl. from the Shevaroys and Kollimalai. During recent herpetological explorations in Salem and Namakkal districts in north-western Tamil Nadu, India, we collected a number of Cnemaspis, including C. yercaudensis from its type locality and Kollimalai, and three deeply divergent lineages from a steep elevation gradient (800–1400 m asl.) within about 10 km straight-line distance in the Shevaroys. In this paper, we describe two new species of Cnemaspis from the Shevaroys based on morphological differences and present mitochondrial sequence divergence to show they are distinct lineages. We also provide sequences on distribution, natural history, morphological variation, and genetic data of the poorly known C. yercaudensis. This comprehensive comparative dataset on three species of Cnemaspis from Yercaud in the Shevaroys will be valuable for further systematic studies on this group.

Materials and methods

Taxon sampling. Specimens were caught by hand and euthanized using halothane. Tail tips/liver tissues of at least three individuals per species were collected as tissue samples in molecular grade ethanol and subsequently stored at

TWO NEW CNEMASPIS FROM TAMIL NADU, INDIA Zootaxa 4609 (1) © 2019 Magnolia Press · 69 –200C for genetic analysis. Specimens were fixed in 4% formaldehyde for ~24 hours, washed in water and transferred to 70% ethanol for long-term storage. Specimens have been deposited at the National Centre for Biological Sciences (NCBS), Bengaluru, and Bombay Natural History Society (BNHS), Mumbai. Morphological and meristic data. A total of 13 specimens of the new species and seven specimens of C. yercaudensis were used to collect morphological data, besides representative Indian Cnemaspis spp. (Appendix 1). Counts and measurements were taken under a Leica S6E stereo dissecting microscope and on the right side of the body, where possible. Colour pattern was recorded from photographs taken in life. The following measurements (after Grismer et al. 2014; Khandekar 2019) were taken with a Digimax digital vernier calliper (to the nearest 0.1 mm): snout vent length (SVL, from tip of snout to vent); axilla to groin length (AGL, from posterior margin of forelimb insertion to anterior margin of hindlimb insertion); body width (BW, maximum body width); forearm length (FL, from elbow to distal end of wrist); crus length (CL, from knee to heel); tail length (TL, from vent to tip of tail); tail width (TW, measured at widest point of tail); head length (HL, distance between retroarticular process of jaw and snout-tip); head width (HW, maximum width of head); head depth (HD, maximum head depth at occiput); eye diameter (ED, greatest horizontal diameter of eye); eye to nares distance (EN, distance between anterior margin of eye and posterior edge of nostril); eye to snout distance (ES, distance between anterior margin of eye and tip of snout); eye to ear distance (EE, distance from anterior edge of ear opening to posterior margin of eye); ear length (EL, maximum length of ear opening); internarial distance (IN, distance between nares); interorbital distance (IO, shortest distance between left and right supraciliary scale rows). Meristic data recorded for all specimens were number of supralabials (SL), and infralabials (IL), from rostral and mental, respectively, to posterior-most enlarged scale at angle of the jaw; supralabials at midorbital position (SL M), and infralabials at midorbital position (IL M), from rostral and mental, respectively, to below the middle of the orbit; dorsal tubercle rows (DTR, number of longitudinal rows of enlarged tubercles around the body counted at mid-body), paravertebral tubercles (PVT, number of enlarged tubercles between limb insertions counted in a straight line immediately left of the vertebral column), ventral scales (VS, counted from posterior of mental to anterior border of cloaca); mid-body scale rows across the belly (MVSR, counted at midbody ventral between the lowest rows of dorsal scales); femoral pores (FP); precloacal pores (PP), the number of poreless scales between precloacal pores (SBPP), and the number of poreless scales between femoral and precloacal pores (SB FP&PP); transverse subdigital lamellae, counted from the base of the digits to the claw and including the claw sheath on finger 1 (LamF1), finger 4 (LamF4), toe 1 (LamT1), toe 4 (LamT4), and toe 5 (LamT5). Additional character states evaluated (after Grismer et al. 2014) were presence or absence of a row of enlarged, widely spaced tubercles and spine like scales between the limb insertions along the ventrolateral edge of the body (flank); the general arrangement (i.e., random or more or less linear) of the dorsal body tubercles; orientation and shape of femoral and precloacal pores; the degree and arrangement of body and tail tuberculation; the relative size and morphology of subcaudal scales, subtibial scales; and the number of postcloacal tubercles on each side of the tail base. Longitudinal rows of caudal tubercles on the non-regenerated portion of the tail are quite variable between species and useful in differentiating several taxa. As per Grismer et al. (2014), up to five pairs of the following rows may be present in varying combinations: caudal furrow; dorsolateral row; lateral row, and the ventrolateral row, usually restricted to the anterior 25% of the tail when present were examined. Twenty-six colour pattern characteristics were evaluated and are described in Table 6 (following Grismer et al. 2014 in part). Comparative data on described species of Cnemaspis from peninsular India was taken from Smith (1935), Das & Sengupta (2000), Das & Bauer (2000), Bauer (2002), Mukherjee et al. 2005; Manamendra-Arachchi et al. (2007), Giri et al. (2009a & 2009b), Ganesh et al. 2011; Cyriac & Umesh (2013 & 2014), Mirza et al. (2014), Srinivasulu et al. (2015), Sayyed et al. (2016, 2018), and Cyriac et al. (2018). Molecular data. Total genomic DNA was extracted from tail/ liver tissues using the Qiagen DNeasy extraction kit for Cnemaspis gracilis (Beddome), C. yercaudensis, the new species, and representative Cnemaspis from across peninsular India (including topotypical material for most species; Table 1). The primers MetF1 and H5934 (Macey et al. 1997) were used to amplify the ND2 gene, and MetF1 was used for sequencing. Both PCR and sequencing were carried out by Medauxin and Chromgene (Bangalore, India). We added published South Asian Cnemaspis sequences and used African and Southeast Asian Cnemaspis as outgroups (Table 1). We used MEGA 5.2.2 (Tamura et al. 2011) to align sequences and calculate uncorrected % pairwise sequence divergence (Table 2). The GTR + G model was applied on the codon partitioned ND2 data in RAxML HPC 7.4.2 (Stamatakis 2006) implemented through the raxmlGUI 1.3 (Silvestro and Michalak 2012), and a maximum likelihood (ML) phylogeny was constructed with 10 independent ML runs and support assessed with 1000 rapid bootstraps.

70 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. eld series; GeneBank Accession number MK792466 MK792467 MK792498 MK792499 KY037997 MK792480 MK792481 MK792494 MK792495 MK792496 MK792497 KY037998 KY037999 MK792491 MK792475 MK792476 MK792477 MK792478 MK792479 MK792465 MK792470 MK792461 MK792462 MK792463 one private collection; MVZ, collection; Museum private one G, Varad series; Giri field WHT, Wildlife series; ADS, Anslem de Silva fi Silva de Anslem series; ADS, continued on the next page next page on the continued … ws: AA, Rohan Pethiyagoda field Rohan AA, ws: i District, KWC Campus batore District, Valparai li District, KWC Campus Satara District, Kaas plateau

Locality Salem District, Nadu, Tamil India, Sankari Salem District, Nadu, Tamil India, Sankari Satara District, Mahabaleshwar Maharashtra, India, Satara District, Mahabaleshwar Maharashtra, India, Sri Lanka, Anuradhapura District, Ritigala District, Sindhudurg Amboli Maharashtra, India, District, Sindhudurg Amboli Maharashtra, India, District, Sindhudurg Amboli Maharashtra, India, District, Sindhudurg Amboli Maharashtra, India, Goa, Goa North India, District, Mhadei Kolhapur Maharashtra, District, TilariIndia, Nagar District, Nuwara Eliya Sri Lanka, Hakgala District, Nuwara Eliya Hakgala Sri Lanka, near Maharashtra, India, District, Uttara Kannada Maharashtra, GundIndia, District, Uttara Kannada Ganeshgudi Maharashtra, India, Sangl Maharashtra, India, Sang Maharashtra, India, Goa District, Goa, South CanaconaIndia, Kerala, PalakkadIndia, District, river Chittur near Nadu, Coim Tamil India, Salem District, Nadu, Tamil India, Yercaud Salem Nadu, Tamil District,India, Yercaud Salem District, Nadu, Tamil India, Yercaud series; CES, Centre for Ecological Sciecnes, Bangalore; JB,Jon Bo ological Sciences, Bangalore;ological Biswas SB, Sayantan field series; V

Voucher 107 AK 108 AK 429 AK 432 AK AMB7447 VG 393 VG 394 VG 389 VG 354 518 AK 517 AK AMB7495 AMB7507 439 AK VG 385 VG 388 VG 398 VG 399 360 AK CES G385 135 AK 208 AK 209 AK 212 AK sequences this used in study. Museum abbreviations are as follo Cnemaspis flaviventralis flaviventralis gracilis gracilis gracilis cf. cf. cf. cf. cf. cf. List of

TABLE 1. series; M. Bauer field AMB, KhandekarAK, Aaron field Akshay (Berkeley); VertebrateNCBS, Nationalof Zoology Centre for Bi TrustHeritage (Sri Lanka). Species agarwali Cnemaspis agarwali Cnemaspis Cnemaspis ajijae Cnemaspis ajijae Cnemaspis alwisi Cnemaspis amboliensis Cnemaspis amboliensis Cnemaspis flaviventralis Cnemaspis flaviventralis Cnemaspis Cnemaspis Cnemaspis gemunu Cnemaspis gemunu Cnemaspis girii goaensis Cnemaspis goaensis Cnemaspis goaensis Cnemaspis goaensis Cnemaspis goaensis Cnemaspis Cnemaspis gracilis Cnemaspis gracilis Cnemaspis Cnemaspis Cnemaspis

TWO NEW CNEMASPIS FROM TAMIL NADU, INDIA Zootaxa 4609 (1) © 2019 Magnolia Press · 71 GeneBank Accession number MK792464 MK792503 KY037970 KY037971 MK792500 MK792501 KY037974 KY037976 MK792482 MK792483 MK792484 MK792485 MK792486 MK792492 MK792493 KY037977 MK792474 KY037980 KY038001 KY038005 KY038007 7448 AMB KY037983 KY038008 MK792468 MK792469 continued on the next page next page on the continued … Urban District, IISc Campus

Locality Salem District, Nadu, Tamil India, Yercaud Hassan Karnataka, India, District, Sakleshpur Matale District,Sri Lanka, Rattota, Gammaduwa Kandy District,Sri Lanka, Gannoruwa District, Sindhudurg Amboli Maharashtra, India, Kolhapur Maharashtra, District, DajipurIndia, Moneragala District,Sri Lanka, Rathataakanda (Buttala) District, Nuwara Eliya Sri Lanka, Bandarawela Kolhapur Maharashtra, District, DajipurIndia, Kolhapur Maharashtra, District, DajipurIndia, Kolhapur Maharashtra, District, DajipurIndia, Kolhapur Maharashtra, District, DajipurIndia, Kolhapur Maharashtra, District, DajipurIndia, Raigad Maharashtra, District,India, Phansad Ratnagiri Maharashtra, India, District, Guhagar near Kecematan Enggano, Enggano, Pulau Sumatra, Malakoni Bangalore Karnataka, India, District, Nuwara Eliya LabookellieSri Lanka, Matale District,Sri Lanka, Rattota, Gammaduwa Moneragala District,Sri Lanka, Kukulagoda Matale District,Sri Lanka, Rattota, Gammaduwa Sri Lanka, Anuradahapura District, Mihintale District, Nuwara Eliya Sri Lanka, Upcot tea estate Kandy District, Sri Lanka, Gannnoruwa Salem District, Nadu, Tamil India, Yercaud Salem District, Nadu, Tamil India, Yercaud AK 204) AK (AK 205) (AK

Voucher 213 AK unvouchered AA82 AA57 unvouchered Unvouchered AMB7431 AA05 362 AK 380 AK 381 AK AKDAJ unvouchered 398 AK 389 AK MVZ239314 unvouchered 7494 AMB 81AA 58A AA80 7448 AMB 7505 AMB AA47 NCBS-BH674 ( NCBS-BH675

nov. nov.

sp. sp.

gracilis heteropholis mahabali mahabali evaroyensis evaroyensis evaroyensis cf. cf. cf. cf. cf. (continued) (continued)

TABLE 1. Species Cnemaspis Cnemaspis Cnemaspis kallima Cnemaspis kandiana kolhapurensis Cnemaspis kolhapurensis Cnemaspis Cnemaspis kumarasinghei Cnemaspis latha Cnemaspis limayei Cnemaspis limayei Cnemaspis limayei Cnemaspis limayei Cnemaspis limayei Cnemaspis Cnemaspis Cnemaspis modigliani Cnemaspis mysoriensis Cnemaspis pava Cnemaspis phillipsi podihuna Cnemaspis Cnemaspis punctata Cnemaspis retigalensis Cnemaspis samanalensis Cnemaspis scalpensis Cnemaspis sh Cnemaspis sh

72 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. GeneBank Accession number KY037984 MK792471 KY037986 MK792472 MK792473 MK792487 MK792488 MK792489 MK792490 MK792502 KY037991 KY037989 KY037990 KY037993 KY037995 KY038006 KY038009 KY038011 KY038012 KY038013 KY037991 Satara District, Humbarli Satara District, Humbarli

Locality Galle District, HiyareSri Lanka, reserve forest Nadu, Salem Tamil District,India, Yercaud Kandy District,Sri Lanka, Pussellawa Namakkal Nadu, Tamil District, KollimalaiIndia, Salem Nadu, Tamil District,India, Yercaud Kolhapur Maharashtra, District, India, Amba Kolhapur Maharashtra, District, India, Amba Maharashtra, India, Maharashtra, India, KeralaIndia, pet trade Karnataka, India, Matara District, Naotunna (near Sri Lanka, Thalalla) HambantotaSri Lanka, District, Sandagala Budulla District,Sri Lanka, Haputale Kodagu Karnataka, District,India, Kumarahalli Sri Lanka, Anuradahapura District, Mihintale Moneragala District,Sri Lanka, Serawa Moneragala District,Sri Lanka, Maligathenna MoneragalaSri Lanka, District, estate Kumaradola Kerala, ThrissurIndia, District, Falls Athirappilly Sri Lanka, Ratnapura District, Masimbula, Godakawela

Voucher AA88 CES G143 AA83 280 AK CES G133 VG 407 VG 408 470 AK CES G349 unvouchered JB239 AA17 WHT 7331 AA87 SB048 AMB7449 7418 AMB 47A AA60 SB151 AMB7508

nov.

sp.

sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. (continued) (continued)

TABLE 1. Species Cnemaspis silvula Cnemaspis thackerayi Cnemaspis upendrai Cnemaspis yercaudensis Cnemaspis yercaudensis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis Cnemaspis

Phylogenetic relationships

The monophyly of South Asian Cnemaspis is well supported, with a deep basal split (~29.8–33.9% sequence divergence) separating the C. kolhapurensis Giri, Bauer & Gaikwad + C. heteropholis clades from the rest of South Asian Cnemaspis (Fig. 1), similar to the results of Sayyed et al. (2018). Within other South Asian Cnemaspis, there is strong support for relationships only at the base of the phylogeny and for individual broad clades (Fig. 2); with a basal split separating the podihuna clade + a single sampled species from Kerala (SB151) from the girii, goaensis, gracilis, kandiana + single sampled species from Karnataka (SB048) and mysoriensis clades. The podihuna and kandiana clades have the same composition as that recovered by Agarwal et al. (2017); the girii clade is distributed in the northern Western Ghats and includes C. ajijae Sayyed, Pyron & Dileepkumar, C. flaviventralis Sayyed, Pyron & Dahanukar, C. girii Mirza, Pal, Bhosale & Sanap, C. limayei Sayyed, Pyron & Dileepkumar, and a lineage allied to C. mahabali Sayyed, Pyron & Dileepkumar, besides three unnamed divergent lineages; the goaensis clade includes C. amboliensis Sayyed, Pyron & Dileepkumar besides C. goaensis Sharma and topotypical C. indraneildasii Bauer (VG 385) that is identical to C. goaensis; the mysoriensis clade includes C. mysoriensis and C. yercaudensis; and the gracilis clade include C. agarwali, C. gracilis, C. sp. JB239 and three unnamed lineages from Yercaud, which are 8.0–16.6 % divergent from C. agarwali and C. gracilis (Table 2). We describe two of the divergent lineages from Yercaud as new species, C. shevaroyensis sp. nov. and C. thackerayi sp. nov.

Cnemaspis sp. AMB7508 Cnemaspis latha AA05 Cnemaspis sp. AA87 Cnemaspis ingerorum WHT 7331 Cnemaspis silvula AA88 Cnemaspis kumarsinghei AMB7431 Cnemaspis kallima AA82 kandiana clade Cnemaspis kandiana AA57 Cnemaspis retigalensis AMB7448 Cnemaspis pava AMB7494 Cnemaspis upendrai AA83 Cnemaspis samanalensis AMB7505 Cnemaspis sp. AA17 Cnemaspis modigliani MVZ239314 Cnemaspis sp. SB048 Cnemaspis cf. gracilis AK 213 Cnemaspis cf. gracilis AK 209 Cnemaspis cf. gracilis AK 208 Cnemaspis cf. gracilis AK 212 AK 107 Cnemaspis agarwali clade Cnemaspis agarwali AK 108 gracilis Cnemaspis sp. JB239 Cnemaspis shevaroyensis sp. nov. AK204 Cnemaspis shevaroyensis sp. nov. AK205 Cnemaspis gracilis CESG385 Cnemaspis gracilis AK135 Cnemaspis thackerayi sp. nov. Cnemaspis yercaudensis Cnemaspis yercaudensis AK 280 mysoriensis Cnemaspis mysoriensis clade Cnemaspis goaensis VG388 Cnemaspis goaensis VG385 Cnemaspis goaensis VG399 goaensis clade Cnemaspis goaensis AK360 Cnemaspis goaensis VG398 Cnemaspis amboliensis VG 393 Cnemaspis amboliensis VG 394 Cnemaspis limayei AK380 Cnemaspis limayei DV40.5 Cnemaspis limayei AK381 Cnemaspis limayei AK362 Cnemaspis limayei AKDAJ Cnemaspis sp. VG408 Cnemaspis sp. VG407 Cnemaspis sp. AK470 sp. G349 Cnemaspis girii clade Cnemaspis girii AK439 Cnemaspis cf. mahabali AK398 Cnemaspis cf. mahabali AK389 Cnemaspis flaviventralis VG389 Cnemaspis flaviventralis VG354 Cnemaspis cf. flaviventralis AK518 Cnemaspis cf. flaviventralis AK517 Cnemaspis ajijae AK429 Cnemaspis ajijae AK432 Cnemaspis sp. 47A Cnemaspis nilgala AMB7418 Cnemaspis punctata AA80 Cnemaspis alwisi AMB 7447 sp. AA60 Cnemaspis clade Cnemaspis cf. podihuna AMB 7449 podihuna Cnemaspis podihuna 58A Cnemaspis phillipsi AA81 Cnemaspis scalpensis AA47 Cnemaspis gemunu AMB 7495 Cnemaspis gemunu AMB 7507 Cnemaspis sp. SB151 Amboli Cnemaspis kolhapurensis kolhapurensis clade Cnemaspis kolhapurensis DV40.4 DAJ sp. Kerala Cnemaspis heteropholis clade Cnemaspis cf. heteropholis

0.1

FIGURE 2. Maximum likelihood tree of South Asian Cnemaspis based on ND2. Solid circles at nodes indicate bootstrap support >90% and hollow circles 80–90%. Major clades are marked with a vertical line, outgroups not shown.

74 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. TABLE 2. Pairwise uncorrected ND2 sequence divergence between members of the Cnemaspis gracilis and Cnemaspis mysoriensis clades of Indian Cnemaspis, numbers in bold along diagonal represent intraspecific diversity. Species 1234567 1 Cnemaspis agarwali 0.0 2 Cnemaspis gracilis 8.9 0.0 3 Cnemaspis cf. gracilis 11.0 13.7 0.8 4 Cnemaspis mysoriensis 25.3 24.4 27.2 - 5 Cnemaspis shevaroyensis sp. nov. 8.0 10.3 13.3 25.9 0.7 6 Cnemaspis thackerayi sp. nov. 13.3 12.9 16.6 23.7 13.6 - 7 Cnemaspis yercaudensis 19.9 20.5 21.9 21.2 21.2 20.6 0.0

Systematics Cnemaspis yercaudensis Das & Bauer 2000 Yercaud Day Gecko Figs. 6 A, B; Table 3.

Cnemaspis yercaudensis Das & Bauer, 2000: 7 (1), 17–28

Holotype. AMB 5748, sub-adult male, from Yercaud Town in the Shevaroys (11°.48’ N 78°.14’ E; ca. 1515 m asl.), Salem District, Tamil Nadu, south-western India, collected by Aaron M. Bauer and Indraneil Das, 13 January 1998. Additional material. NCBS-BH678, adult male, NCBS-BH677, sub-adult female, from near Grange resort, Yercaud town, in the Shevaroys (11.775° N 78.219° E; ca. 1388 m asl.), Salem district, Tamil Nadu state, India, collected by A. Khandekar, I. Agarwal and N. Gaitonde on 12 September 2018. NCBS-BH679, BNHS 2533, adult males, NCBS-BH680, BNHS 2532, BNHS 2534, adult females, from near Nallathambi resort, Kollimalai, (11.286° N, 78.337° E; ca. 1146 m asl.), Namakkal district, Tamil Nadu state, India, collected by A. Khandekar, I. Agarwal and N. Gaitonde on 16 September 2018. Diagnosis and comparison with Indian congeners: A small-sized Cnemaspis, snout to vent length less than 30 mm. Dorsal pholidosis heterogeneous; weakly keeled, granular scales in the vertebral and paravertebral region with a few scattered enlarged keeled tubercles, intermixed with about three rows of large, strongly keeled, conical scales on the flanks, five or six rows of dorsal scales, spine-like scales absent on flank. Ventral scales smooth, subimbricate, 18–20 scales across belly, 90–105 longitudinal scales from mental to cloaca. Subdigital scansors smooth, entire, unnotched; lamellae under digit IV of pes 15–19. Males with three femoral pores on each thigh, separated on either side by five or six poreless scales from two continuous precloacal pores. Tail with enlarged, strongly keeled, conical scales forming whorls; a median row of sub-caudals smooth, slightly enlarged. Dorsum grayish-brown with light vertebral stripe from occiput to tail base, throat yellow, unpatterned; original tail bluish- gray in males, regenerated tail light brown. Cnemaspis yercaudensis can be distinguished from all other Indian congeners on the basis of the following differing or non-overlapping characters: spine-like scales absent on flanks (versus spine-like scales present on flanks in C. amboliensis, C. assamensis Das & Sengupta, C. flaviventralis, C. goaensis, C. jerdonii (Theobald), C. littoralis (Jerdon), C. monticola Manamendra-Arachchi, Batuwita & Pethiyagoda and C. nilagirica Manamendra- Arachchi, Batuwita & Pethiyagoda); scales on dorsal aspect of trunk heterogeneous (versus scales on dorsal aspect of trunk homogeneous in C. adii, C. assamensis, C. australis Manamendra-Arachchi, Batuwita & Pethiyagoda, C. boiei (Gray), C. indica (Gray), C. jerdonii, C. kolhapurensis, C. littoralis, C. nilagirica and C. sisparensis (Theobald) ); tail with median row of sub-caudal scales smooth and slightly enlarged (versus median row of subcaudal scales smooth and not enlarged in C. ajijae, C. flaviventralis, C. girii, C. limayei; C. monticola, C. australis with keeled sub-caudals); absence of keeled scales on the venter or gular regions (versus keeled scales on the venter or gular region in C. beddomei (Theobald) and C. goaensis); males with three femoral pores on each thigh, separated on either side by five or six poreless scales from two continuous precloacal pores (versus precloacal pores absent, femoral pores present in C. ajijae, C. anaikattiensis, C. flaviventralis, C. girii, C. indica,

BNHS 2534 BNHS

BNHS 2533 BNHS ……continued on the next page

Abbreviations are listed in Materials and Methods. = tail/ incomplete* digit Cnemaspis yercaudensis Cnemaspis

Measurements (mm) and datameristic for the type series of Sex TABLE 3. Sex SVL TL TW LAL CL AGL BH F M BW F M F M HL HW HD ED EE L/R ES L/R 30 29.8 23.1 27.7 28.4 30 EN 3.5* L/R 28.7 29 3.8* IN 10.4* 3.6* 24.2 20.6 L/R 3 2.8 2.4 4 3 3 IO 3 2.7 M 3.7 3.1 3.5 3.8 4 4.7 3.5 EL M 4.5 3.8 4.8 4.6 4.4 13.5 SL 11.8 9.5 IL 10.7 11.4 13.5 12.3 7.1 3.8 3 1.8 2.1 3.1 4.1 3 SL 5.8 3.6 5.5 5.7 7.3 6.4 IL 7 6.8 6 5 PVT 5 L/R 6.9 6.6 6.9 6.7 4.4 5.2 5.1 4.9 5.1 RBS 6/6 3.1 1.3 2.3 2.8 2.9 3 0.8 6/6 RVS 2.9 1.4 3 5/5 0.7 6/6 6/6 7/6 7/6 2.7 1.2 1.4 1.3 1.3 VS 5/5 0.7 0.9 0.8 0.8 2.5 5/5 5/5 6/5 6/5 3.5 7/8 1.9 2.5 2.4 LamF1 2.5 2.6 3.4 7/7 2.9 2.8 3.5 7/6 3.2 3.4 3.2 7/77/7 8/8 7/7 2.7 7/7 2.1 2.7 2.6 2.7 2.8 7/7 7/6 7/7 7/7 7/7 8/8 2.3 8/8 0.3 2.3 1.7 2 8/8 8/8 7/7 8/8 0.5 2.2 2.3 2.2 0.3 0.3 0.5 0.4 0.4 0 0 6 0 0 6 18 6 5 6 18 19 18 20 18 18 90 95 102 105 95 93 97 or missing, /= data not available. Specimen No. Specimen 2532 NCBS-BH678 NCBS-BH677BNHS NCBS-BH679 NCBS-BH680

BNHS 2534 BNHS

BNHS 2533

(Continued) (Continued) . 3 Ventral scales keeled (1) or smooth (0) (0) 0 0 smooth 0 0 0 0 / 0 0 0 0 0 (0) or heterogeneous tubercles (1) 0 (0) L/R 0 L/R TABLE or 0 0 0 postcloacal L/R L/R 1 LamT1L/R keeled 1 LamT4 1 1 PP&FP absent L/R LamT5 (1) of PP SBPP scales SB 16/16 or FP 16/16 15/15 homogeneous 8/8 15/15 16/16 17/16 17/17 19/18 No. 14/14 8/8 15/14 16/15 16/15 15/14 16/14 8/8 7/7 7/7 8/8 scales Anteriorenlarged (1) extra-brillar fringe not or enlarged (0) 0 0 6/6 0 Ventral 5/5 0 1 (0) 0 smooth or (1) keeled scales Gular (1) 2 pholidosis keeled Pectoral scales (1) smooth (0) or 0 0 2 0 separated (0) or (1) continuous Precloacal pores 3/3 (1)Precloacal or pores elongate round (0) 0 3/3 1 pores elongate (1)femoral round (0) or 3/3 0 present Dorsal 0 (0)Dorsal keeled ortubercles keeled (1) not 0 0 0 linearly random arranged (1) (0)Tubercles more or Spine-like present on (0) or(1) tubercles flanks absent 1 - scales 0 present(0) flanks Tubercles on (1) lower absent or (0) 0 present caudal orLateral (0) (1) furrows absent - Caudal inor(1) tubercles not furrow lateral (0) Subcaudals(0) (1) keeled smooth or 1 1 - 1 0 subtibial keeled Singleof row subcaudals median (1) (0) or smooth 0 1 0 1 Caudal encircle tail or tubercles (1) not (0) (1)row Enlarged ornot subcaudal (0) median scale 1 presentscales Enlarged (1) femoral absent or (0) 1 1 1 0 1 1 0 Shield-like 0 1 1 0 1 Subtibial keeled (1) smooth (0) or scales 1 1 1 1 1 0 - 1 0 0 0 1 1 1 1 0 1 - 1 1 / 1 / - 0 - 0 0 1 / 1 1 1 0 1 - / 1 1 0 1 0 0 / - 0 / 1 1 1 / 1 1 - 0 1 0 1 1 / 0 / 1 0 1 1 / 1 1 1 1 0 - 1 1 / 0 1 / 1 / 1 / 1 0 1 1 0 1 1 1 1 0 1 1 1 1 Specimen No. Specimen 2532 NCBS-BH678 NCBS-BH677BNHS NCBS-BH679 NCBS-BH680

TWO NEW CNEMASPIS FROM TAMIL NADU, INDIA Zootaxa 4609 (1) © 2019 Magnolia Press · 77 C. jerdonii, C. kottiyoorensis Cyriac & Umesh, C. limayei, C. littoralis, C. mahabali, C. sisparensis, C. heteropholis, C. wynadensis (Beddome); only precloacal pores present in C. anamudiensis Cyriac, Johny, Umesh, & Palot, C. beddomei, C. maculicollis Cyriac, Johny, Umesh, & Palot, C. nairi Inger, Marx & Koshy, C. ornata (Beddome); both femoral and precloacal pores absent in C. boiei, C. assamensis; three femoral and four precloacal pores present in C. otai; two femoral and two precloacal pores present in C. adii; 4–6 femoral and two precloacal pores on each side separated by a single poreless scales in C. agarwali; four or five femoral and three precloacal pores in C. australis; 2–4 femoral and three precloacal pores in C. goaensis; 3–5 femoral pores and two precloacal pores separated by two or three poreless scales C. mysoriensis; a continuous series of 26–28 precloacal-femoral pores in C. kolhapurensis). Cnemaspis yercaudensis is 19.9–21.9% divergent in ND2 sequences from all sampled congeners (Table 2). Diagnoses against the new species are provided after their descriptions. Variation and additional information from additional material examined. Mensural data for the type series is given in Table 3. There are four male and three female specimens ranging in SVL from 23.1 mm to 30.0 mm. All paratypes resemble the holotype except as follows: the number of lamellae on digit I of the manus ranges from seven or eight and on digit IV from 12–14; on digit IV of the pes from 15–19 and on digit V from 14–16; the number of supralabials ranges from seven to eight to the angle of jaw; infralabials between six and seven; five or six dorsal tubercle rows. Ventral scale counts in longitudinal and transverse series vary from 90–105 and 18–20, respectively; males have five or six poreless scales between precloacal and femoral pores. The complete tail is slightly shorter than body (TL/SVL ratio 0.87–1.00). The three males in our collection match the three female specimens in overall colouration, except the tail is bluish-grey in males and brown in females. Distribution and Natural history. Cnemaspis yercaudensis is so far known from around the type locality, Yercaud town, in the Shevaroy massif, Salem district across an elevation range of 1350–1500 m asl. and in Kolli Hills between 1140–1350 m asl. At Yercaud, we collected individuals of C. yercaudensis after dusk (~ 1830h) on roadside walls and observed a single individual during an overcast day on a low wall. At Kolli Hills, we collected a total of five individuals of C. yercaudensis, out of which four were found at night along roadside walls approximately at a height of 1m above ground, and a single inactive individual under a rock during the daytime. Sympatric lizards at localities in Yercaud where we observed the species include Hemidactylus frenatus Duméril & Bibron, Hemidactylus parvimaculatus Deraniyagala, Hemiphyllodactylus aurantiacus (Beddome), Cnemaspis thackerayi sp. nov., Eutropis carinata (Schneider), Eutropis macularia (Blyth), Lygosoma albopunctata (Gray), and Calotes versicolor (Daudin). Sympatric lizards in Kollimalai include Hemidactylus frenatus, Hemidactylus parvimaculatus, Hemiphyllodactylus kolliensis Agarwal, Khandekar, Giri, Ramakrishnan & Karanth Cnemaspis sp., Eutropis carinata, Eutropis macularia, Eutropis sp., Lygosoma albopunctata and Calotes versicolor.

Cnemaspis thackerayi sp. nov. Figs. 3, 4, 5, 6 C&D, 7 A&B; Table 4.

Holotype. NCBS-BH670, adult male, from near Grange resort, Yercaud town, in the Shevaroys (11.775° N 78.219° E; ca. 1388 m asl.), Salem district, Tamil Nadu state, India, collected by A. Khandekar and T. Thackeray on 12 September 2018. Paratypes. NCBS-BH671, BNHS 2527, adult males, NCBS-BH672, NCBS-BH673, BNHS 2526, BNHS 2528, adult females, same data as holotype except collected by A. Khandekar, I. Agarwal and N. Gaitonde on 18 September 2018. Etymology. The specific epithet is a patronym in recognition of the contributions to natural history and systematic zoology of Mr. Tejas Thackeray, who also collected the holotype of this species. Suggested Common Name. Thackeray’s dwarf gecko Diagnosis and comparison with Indian congeners: A medium-sized Cnemaspis, snout to vent length less than 41 mm. Dorsal pholidosis heterogeneous, weakly keeled, granular scales intermixed with large, strongly keeled, conical tubercles, 11–14 rows of dorsal tubercles, 12–14 tubercles in paravertebral rows; spine-like scales absent on flank. Ventral scales smooth, subimbricate, 22–25 scales across belly, 105–122 longitudinal scales from mental to cloaca. Subdigital scansors smooth, entire, unnotched; lamellae under digit IV of pes 21–24. Males with 5–9 femoral pores on each thigh, separated on either side by 1–6 poreless scales from five or six precloacal pores; precloacal pores separated medially by one or two poreless scales. Tail with enlarged, strongly keeled, conical

78 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. tubercles forming whorls; a median row of subcaudals smooth, enlarged. Dorsum with 4–6 light grey vertebral blotches forms an indistinct vertebral stripe extending from neck to tail base; a single dorsal ocellus between forelimb insertions, throat off-white with black speckles; original tail in males with 8–9 alternating black and whitish-grey bands, regenerated tail brown. Cnemaspis thackerayi sp. nov. can be distinguished from all other Indian congeners on the basis of the following differing or non-overlapping characters: spine-like scales absent on flank (versus spine-like scales present on flank in C. amboliensis, C. assamensis, C. flaviventralis, C. goaensis, C. jerdonii, C. littoralis, C. monticola and C. nilagirica); scales on dorsal aspect of trunk heterogeneous (versus scales on dorsal aspect of trunk homogeneous in C. adii, C. assamensis, C. australis, C. boiei, C. indica, C. jerdonii, C. kolhapurensis, C. littoralis, C. nilagirica and C. sisparensis); tail with median row of sub-caudal scales smooth and enlarged (versus median row of sub-caudal scales smooth and not enlarged in C. ajijae, C. flaviventralis, C. girii, C. limayei; C. monticola, C. australis with keeled subcaudals); absence of keeled scales on the venter or gular regions (versus keeled scales on the venter or gular region in C. beddomei and C. goaensis); males with 5–9 femoral pores on each thigh, separated on either side by 1–6 poreless scales from five or six precloacal pores; precloacal pores separated medially by one or two poreless scales (versus precloacal pores absent, femoral pores present in C. ajijae, C. anaikattiensis, C. flaviventralis, C. girii, C. indica, C. jerdonii, C. kottiyoorensis, C. limayei, C. littoralis, C. mahabali, C. sisparensis, C. heteropholis, C. wynadensis; only precloacal pores present in C. anamudiensis, C. beddomei, C. maculicollis, C. nairi, C. ornata; both femoral and precloacal pores absent in C. boiei, C. assamensis; three femoral and four precloacal pores present in C. otai; two femoral and two precloacal pores on each thigh in C. adii; four or five femoral and three precloacal pores in C. australis; 2–4 femoral and three precloacal pores in C. goaensis; 3–5 femoral pores on each thigh, separated on either side by 7–10 poreless scales from two precloacal pores; precloacal pores separated medially by two or three poreless scales in C. gracilis; 4– 6 femoral pores on each thigh, separated on either side by eight poreless scales from four precloacal pores; precloacal pores separated medially by a single poreless scale in C. agarwali; two femoral and two precloacal pores in C. mysoriensis; three femoral and two precloacal pores in C. yercaudensis; a continuous series of 26–28 precloacal- femoral pores in C. kolhapurensis). C. thackerayi sp. nov. closely resembles C. gracilis and C. agarwali. However, it can be distinguished from both species by the presence of 5–9 femoral pores on each thigh and five or six precloacal pores separated by one or two poreless scales (versus 3–5 femoral pores on each thigh and two precloacal pores separated medially by two or three poreless scales in C. gracilis; 4–6 femoral pores on each thigh and four precloacal pores separated medially by a single poreless scale in C. agarwali); 1–6 poreless scales between femoral and precloacal pores (versus eight poreless scales between femoral and precloacal pores in C. agarwali); maximum SVL 41 mm (versus maximum SVL 34 mm in C. gracilis; 33 mm in C. agarwali); presence of a single dorsal ocellus between forelimb insertions (versus presence of two single dorsal ocellus on occiput and between forelimb insertions in C. gracilis; presence of two single dorsal ocellus on occiput and between forelimb insertions, two pairs on either side just anterior and posterior to forelimb insertions in C. agarwali). Additionally, C. thackerayi sp. nov. is 12.9% divergent in ND2 sequence from C. gracilis, 16.6 % divergent from C. cf. gracilis and 13.3% divergent from C. agarwali (Table 2). Diagnosis against Cnemaspis shevaroyensis sp. nov. is provided after its description. Description of the holotype. Adult male in good state of preservation except tail slightly bent to the right and a 4.1 mm long incision in the sternal region for tissue collection as artefacts of preservation (Fig. 3 A, B). SVL 38.1 mm, head short (HL/SVL ratio 0.25), wide (HW/HL ratio 0.68), not strongly depressed (HD/HL ratio 0.40), distinct from neck. Loreal region slightly inflated, canthus rostralis not prominent. Snout just less than half head length (ES/HL ratio 0.48), 2.5 X eye diameter (ED/ES ratio 0.39); scales on snout and canthus rostralis large, weakly keeled, juxtaposed; larger than those on forehead and interorbital region; occipital and temporal region with much smaller granules, intermixed with larger, roughly rounded, tubercles (Fig. 4 A). Eye small (ED/HL ratio 0.19); with round pupil; orbit with extra-brillar fringe scales that are largest anteriorly; supraciliaries not elongate. Ear-opening deep, vertical, small (EL/HL ratio 0.08); eye to ear distance greater than diameter of eye (EE/ED ratio 1.57) (Fig. 4 C). Rostral much wider (1.8 mm) than long (0.8 mm), incompletely divided dorsally by a strongly developed rostral groove for more than half of its length; single enlarged supranasal on each side, slightly larger than postnasals, separated from each other by smaller single internasal and two smaller scales on the snout; rostral in contact with supralabial I, nasal, supranasal and internasal; nostrils oval, each surrounded by postnasal, supranasal, rostral and supralabial I; a single row of scales separate the orbit from the supralabials (Fig. 4 C).

BNHS 2528

BNHS 2527 ……continued on the next page

BNHS 2526 BNHS NCBS- BH673 Abbreviations are listed Materials in and Methods. = tail/ digit * NCBS- BH672 sp. nov. sp. NCBS- BH671 Cnemaspis thackerayi Cnemaspis BH670 BH670

Measurements (mm) and datameristic for the type series of incomplete or missing, /= data not available. available. not data /= or missing, incomplete TABLE 4. No.Specimen Sex SVL TL TW LAL CL AGL BH BW HL HW M M F F F M F HD ED EE NCBS- ES L/R 3.6 EN 37.8 L/R 38.2 38.1 36.5L/R 36.2 37.4 40.8 3.9 IN 4 4.3 L/R 4 4 3.5 IO 44.1 M 34.6* 12.2* 49.5 16.2 6 EL 53.4 50.2 49.1 6 6.1 16.1 6.4 M 5.9 5.8 6.1 SL 18.4 7.3 7.2 7 7.4 15.8 15 16 16 IL 6.9 7.1 7.4 3.6 SL 4.9 3.6 3.4 3.1 4.3 7.2 IL 7.5 7.1 5.7 6.4 6.7 8.1 9.4 9.8 PVT 9.8 8.3 8.3 9.4 10.1 6.5 6.9 RBS L/R 6.9 6.5 6.2 6.5 6.7 RVS 3.9 4 4.2 4.4 3.7 3.3 3.8 VS 7/7 7/6 2 1.9 1.9 1.8 2 2.1 6/7 8/7 6/6 1.1 6/6 8/7 1 0.9 LamF1 2.1 0.9 1.1 1.1 6/6 1.1 3 3 7/6 3.1 7/7 6/6 7/6 5/5 6/5 2.9 2.99/8 2.9 3.2 4.5 8/8 4.9 4.8 4.4 4.3 4.4 4.9 3.4 6/7 8/7 7/7 8/8 9/10 3.9 3.8 3.3 3.3 3.4 3.88/8 7/7 7/8 7/7 8/8 6/7 7/7 2.4 2.7 2.7 2.4 2.6 2.5 2.6 0.6 0.7 0.8 0.7 0.6 0.6 0.7 11/11 11/11 11/11 12/12 12/13 11/11 12/12 12 14 12 12 14 13 14 13 14 14 11 13 11 25 25 25 22 24 105 106 122108 109 108 113 Holotype Paratypes Holotype

BNHS 2528

BNHS 2527

BNHS 2526 BNHS NCBS- BH673 NCBS- BH672 NCBS- BH671 BH670 BH670 (Continued) (Continued) Holotype Paratypes Holotype Dorsal pholidosis homogeneous (1) or heterogeneous (0) 0 0 0 0 0 0 0 (0) 0 0 tubercles heterogeneous TABLE 4. L/R No.Specimen L/R L/R LamF4 postcloacal L/R or (0) L/R LamT1L/R 0 0 LamT4 0 0 0 0 0 PP&FP LamT5 1 L/R 1 1 1 1 1 1 PP of (1) absent SBPP SB 17/16 16/17 FP 17/19 17/18 19/19 19/19 19/18 No. homogeneous or 21/21 21/22 scales Anterior 21/22 enlarged (1) extra-brillar fringe not or (0) 21/22 22/22 21/22 24/23 18/19 19/19 (0) smooth or (1) keeled Ventral 19/19 scales 19/20 20/19 19/20 20/19 11/11 0 3/6 1/4 11/11 (0) 0 smooth or (1) keeled scales 11/10 Gular 12/12 11/10 11/11 11/11 0 0 4/5 keeled Pectoral scales (1) smooth (0) or pholidosis (1) separated (0) or (1) continuous Precloacal pores 1 NCBS- (1)Precloacal or pores elongate round (0) 0 3/3 4/2 0 0 pores elongate (1)femoral round (0) or 0 3/2 Dorsal present 0 (0) 6/5 Dorsal keeled ortubercles keeled (1) not 9/6 1 0 0 0 0 linearly random arranged (1) (0)Tubercles more or 1 6/5 2 0 0 Spine-like present on (0) or(1) tubercles flanks absent 0 (0) flanks lower on (0) absent or (1) present Tubercles scales 0 0 present caudal orLateral (0) (1) furrows absent 1 Caudal inor(1) tubercles not furrow lateral (0) 1 (0) 0 smooth or (1) keeled Subcaudals 0 1 1 keeled of Single row subcaudals median (1) (0) or smooth 0 subtibial 0 0 0 1 Caudal encircle tail or tubercles (1) not (0) 1 (1)row Enlarged ornot subcaudal (0) median scale 0 0 presentscales Enlarged (1) femoral absent or (0) - 1 1 1 Shield-like 0 0 0 1 0 1 1 Subtibial keeled (1) smooth (0) or scales 1 - 0 - 1 1 1 - 0 0 0 0 1 0 1 1 1 1 0 - 0 - 1 1 1 - 0 0 0 0 1 0 0 1 1 1 0 1 - 0 0 1 0 1 1 0 - 0 0 0 0 1 1 1 0 1 0 1 - 1 0 0 1 1 1 0 0 0 1 0 1 1 0 1 - 1 0 1 - 0 1 0 1 0 1 1 0 1 1 0 1 0 1 1 0 1 1 0 1 0 1 1

TWO NEW CNEMASPIS FROM TAMIL NADU, INDIA Zootaxa 4609 (1) © 2019 Magnolia Press · 81 FIGURE 3. Cnemaspis thackerayi sp. nov. (holotype, NCBS-BH670), dorsal view (A), ventral view (B), dorsal view of tail (C), and ventral view of tail (D). Scale bars 10 mm.

82 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. FIGURE 4. Cnemaspis thackerayi sp. nov. (holotype, NCBS-BH670), dorsal view of head (A), ventral view of head (B), lateral view of head (C), view of cloacal region showing precloacal and femoral pores (D), ventral view of right manus (E), and ventral view of right pes (F). Scale bars 5 mm.

TWO NEW CNEMASPIS FROM TAMIL NADU, INDIA Zootaxa 4609 (1) © 2019 Magnolia Press · 83 FIGURE 5. Type series of Cnemaspis thackerayi sp. nov. (from left to right, NCBS-BH671, NCBS-BH672, NCBS-BH673, BNHS 2526, BNHS 2527, BNHS 2528). Scale bar 10 mm.

Mental enlarged, subtriangular, wider (2.1 mm) than long (1.5 mm); two pairs of postmentals, inner pair large, roughly rectangular, bordered by mental, infralabial I, outer postmentals and three enlarged chin shields; outer postmentals slightly smaller than inner postmentals, bordered by infralabials I, II, inner postmentals and three enlarged chin shields; three enlarged gular scales prevent contact of left and right outer postmentals; chin shields bordering postmentals flat, smooth, smaller than outermost postmentals, rest granular, much smaller, smooth. Infralabials bordered below by a row of slightly enlarged scales, decreasing in size posteriorly (Fig. 4 B). Supralabials to angle of jaw and at midorbit eight (L) — seven (R); supralabial I largest, supralabials decreasing in size posteriorly; infralabials to angle of jaw seven (L) — eight (R), and seven on either side at midorbit; infralabial I largest, infralabials decreasing in size posteriorly (Fig. 4 C). Extra-brillar fringe scales seven or eight on each side, interorbital scale rows across narrowest point of frontal seven or eight; 24 or 25 scale rows between left and right supraciliaries at midorbit (Fig 4 A). Body relatively slender, trunk less than half of SVL (AGL/SVL ratio 0.41) without ventrolateral folds or spine- like scales on flanks. Dorsal scales on trunk heterogeneous, weakly keeled; granular scales intermixed with much larger, strongly keeled, conical tubercles; tubercles in approximately 14 longitudinal rows at mid-body; 14 tubercles in paravertebral row from above forelimb insertion to the hind limb insertion (Fig. 7 A). Scales on nape slightly smaller than those on paravertebral rows, smaller still on occiput (Fig. 4 A). Ventral scales slightly larger than dorsals, those on belly smooth, subimbricate, slightly rounded, subequal from chest to vent; midbody scale rows across belly 25; 122 scales from mental to anterior border of cloaca (Fig. 7 B). Scales on throat and pectoral region slightly smaller than those on belly, flat and imbricate; gular region with much smaller, flattened granules with those on chin bordering postmentals, enlarged, juxtaposed and flattened (Fig. 4 B ). Six femoral pores on right thigh and nine on left; six precloacal pores separated medially by two poreless scales (Fig. 4 D). Scales on palm and sole smooth, flat and roughly circular; scales on dorsal aspect of manus and pes heterogenous, upper arm with scales slightly larger than dorsal granules, weakly keeled, subimbricate; those near forelimb insertion much smaller, granular; dorsal aspect of forearm with scales smaller than those on upper arm, weakly keeled, flat, roughly rounded; scales on elbow weakly keeled, subimbricate and similar in size to those on upper arm; dorsal aspect of hand predominantly bearing small, flattened weakly keeled, imbricate scales. Ventral aspect of upper arm with smooth, roughly rounded, weakly conical granules; scales on forearm and wrist with larger, smooth, weakly imbricate scales. Scales on dorsal aspect of thigh larger than those on dorsal granules, weakly keeled, imbricate except those near hindlimb insertion which are much smaller, roughly rounded, conical granules. Scales on dorsal aspect of knee and shank slightly smaller than those on dorsum of thigh, subimbricate, weakly keeled; dorsal aspect of foot predominantly bearing small, flattened, weakly keeled, imbricate scales;

84 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. Scales on ventral aspect of thigh similar to those on midbody ventrals; those on ventral surface of shank slightly smaller than those on thigh, smooth, imbricate (Fig. 3 B). Fore and hind limbs moderately long, slender (LAL/SVL ratio 0.15); (CL/SVL ratio 0.18); digits long, with a strong, recurved claw, distinctly inflected, distal portions laterally compressed conspicuously. Series of unpaired lamellae on basal portion of digits, separated from narrower distal lamellae by a single large scale at the inflection; proximal lamellae series: 1–3–3–4–4 (right manus; Fig. 4 E), 1–4–6–9–5 (right pes; Fig. 4 F), 1–4–3–5–4 (left manus), 1–5–5–9–5 (left pes); distal lamellae series: 11–12– 14–14–11 (right manus; Fig. 4 E), 10–13–15–14–14 (right pes; Fig. 4 F), 11–13–15–15–13 (left manus), 10–12– 15–15–15 (left pes). Relative length of digits (measurements in mm in parentheses): IV (3.55) > III (3.06) > V (3.04) > II (2.86) > I (2.19) (right manus); IV (5.11) > V (4.79) > III (4.60) > II (3.75) > I (2.36) (right pes). Tail entire except the tip, which is regenerated, cylindrical, relatively slender, flattened beneath, tail slightly longer than snout-vent length (TL/SVL ratio 1.15) (Fig. 3 C, D). Dorsal scales at tail base granular, similar in size and shape to those on midbody dorsum, gradually becoming larger, flatter, pointed, subimbricate posteriorly, intermixed with slightly enlarged, strongly keeled, conical tubercles forming whorls; six or seven tubercles on first 2–5 whorls. Scales on ventral aspect of original tail much larger than those on dorsal, imbricate, smooth, with a series of three enlarged subcaudal scales of which the median series is almost twice the size of adjunct two rows, roughly hexagonal; those on tail base much smaller, imbricate and smooth, a single enlarged postcloacal spur on each side (Fig. 3 D).

FIGURE 6. Photos in life of (A) Cnemaspis yercaudensis, male NCBS-BH678, (B) Cnemaspis yercaudensis, female BNHS 2534, (C) Cnemaspis thackerayi sp. nov., holotype male NCBS-BH670, (D) Cnemaspis thackerayi sp. nov., female, uncollected individual, (E) Cnemaspis shevaroyensis sp. nov., holotype male NCBS-BH674, (F) Cnemaspis shevaroyensis sp. nov., paratype female BNHS 2529; all from Yercaud on the Shevaroy massif, Salem district, Tamil Nadu, India.

TWO NEW CNEMASPIS FROM TAMIL NADU, INDIA Zootaxa 4609 (1) © 2019 Magnolia Press · 85 FIGURE 7. Details of dorsal pholidosis at midbody of (A) Cnemaspis thackerayi sp. nov., holotype, NCBS-BH670, (C) Cnemaspis shevaroyensis sp. nov., holotype, NCBS-BH674 and details of ventral scales at midbody (B) Cnemaspis thackerayi sp. nov., holotype, NCBS-BH670, and (D) Cnemaspis shevaroyensis sp. nov., holotype, NCBS-BH674. Scale bars 5 mm.

Colouration in life. (Fig. 6 C) Dorsum of head and body light brown, limbs yellowish brown; tail base similar to body dorsum. Tail with eight alternating black and whitish-grey bands with a black regenerated tail tip. Head with numerous grey blotches and light bands on labials. Two orangish-brown postorbital streaks, the upper one merging with its counterpart from the other orbit to form a thick W-shaped zig-zag band on nape, the lower streak terminating anterior to forelimb insertion. A single large black ocellus with a margin of orangish scales at the anterior of forelimb insertion. Dorsum with six light grey vertebral blotches forming an indistinct vertebral stripe extending from neck to tail base, orangish-brown blotches interspersed with smaller yellowish-grey spots on rest of dorsum and flank. Dorsum of limbs with lighter spots, more prominent on hindlimbs, digits with alternating dark and light bands. Ventral surfaces off-white (including neck, belly, limbs and tail), with black speckles on the gular region. Variation and additional information from type series. Mensural data for the type series is given in Table 4. There are three male and four female specimens ranging in SVL from 36.2 mm to 40.8 mm. All paratypes resemble the holotype except as follows: the number of lamellae on digit I of the manus ranges from 11–13 and on digit IV

86 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. from 16–19; on digit I of the pes ranges from 10–12, on digit IV from 21–24 and on digit V from 18–20; 6–10 supralabials to angle of jaw; 11–13 tubercles in paravertebral rows; number of ventrals in longitudinal and transverse series vary from 105–113 and 22–24 in NCBS-BH673 and BNHS 2526 respectively. Five or six femoral pores on either side, 2–4 precloacal pores separated by one or two poreless scales; 3–6 poreless scales between precloacal and femoral pores. Complete tail slightly longer than body (TL/SVL ratio 1.29–1.32). The two male paratypes match the holotype in overall colouration, while the four female paratypes are duller. Distribution and Natural history. Cnemaspis thackerayi sp. nov. is known only from the type locality (near Grange resort, Yercaud town, in the Shevaroys), at an elevation of ca. 1390 m asl. (Fig. 1). The species was found in abandoned and decrepit houses along the roadside. The species seems to be crepuscular or nocturnal as we did not encounter individuals at the site during daytime (0900–1400h), and observed juveniles and sub-adults emerging from refugees at around sunset (1800h). After complete darkness (2000h), we encountered many active adults on the walls both inside and outside the abandoned houses. They seemed to avoid light as we did not encounter this species in inhabited buildings near the abandoned house which had electric lighting. Individuals were found in high density with as many as 10–15 individuals in a single small room (approximately 2.5 x 3 m). This species may be specialised to high elevations as captured individuals became lethargic and sluggish and one individual died when moved to lower elevations (~700 to 1100 m asl.). Sympatric lizards at the type locality include Hemidactylus frenatus, Hemidactylus parvimaculatus, Hemiphyllodactylus aurantiacus, Cnemaspis yercaudensis, Eutropis carinata, Eutropis macularia, and Lygosoma albopunctata. Note. Cnemaspis gracilis was described by Beddome (1870) from ”Palghat hills”. Manamendra-Arachchi et al. (2007) designated a lectotype and provided a description of the type series and we assigned a specimen from near Chittur, Kerala in the Palghat Gap and other genetically identical individuals from Valparai Town (presumably introduced there from Palghat, about 50 km straight-line distance away and 1000 m lower in altitude) to C. gracilis based on these specimens. An expanded redescription of C. gracilis sensu stricto, and the descriptions of more species from the complex is forthcoming (Khandekar, Pal & Agarwal, in prep.).

Cnemaspis shevaroyensis sp. nov. Figs. 6 E&F, 7 C&D, 8, 9, 10; Table 5.

Holotype. NCBS-BH674, adult male, from near Kottachedu Kari Raman Temple, Yercaud, in the Shevaroys (11.819° N 78.270° E; ca. 811 m asl.), Salem district, Tamil Nadu state, India, collected by A. Khandekar, I. Agarwal and N. Gaitonde on 13 September 2018. Paratypes. BNHS 2530, BNHS 2531, adult males, NCBS-BH675, NCBS-BH674, BNHS 2529, adult females, same data as holotype. Etymology. The specific epithet shevaroyensis is a toponym for the Shevaroy massif in Salem district of Tamil Nadu, the type and only known locality for this species. Suggested Common Name. Shevaroy dwarf gecko Diagnosis and comparison with Indian congeners: A medium-sized Cnemaspis, snout to vent length less than 35 mm. Dorsal pholidosis heterogeneous, weakly keeled, granular scales intermixed with large, strongly keeled, conical scales, 10–14 rows of dorsal scales, 13–17 scales in paravertebral rows; spine-like scales absent on flanks. Ventral scales smooth, subimbricate, 21–24 scales across belly, 111–118 longitudinal scales from mental to cloaca. Subdigital scansors smooth, entire, unnotched; lamellae under IV digit of pes 16–20. Males with four femoral pores on each thigh, separated on either side by 7–9 poreless scales from 2–4 precloacal pores; precloacal pores separated medially by two or three poreless scales. Tail with enlarged, strongly keeled, conical tubercles forming whorls; a median row of sub-caudals smooth, enlarged. Dorsum with 6–8 light grey vertebral blotches between neck to tail base; two single dorsal ocelli on occiput and between forelimb insertions, two pairs on either side just anterior and posterior to forelimb insertions; throat off-white, strongly suffused yellow on lateral margins with three black longitudinal streaks running parallel to each other; original tail in males with alternating black and whitish-grey bands, regenerated tail orange. Cnemaspis shevaroyensis sp. nov. can be distinguished from all other Indian congeners on the basis of the following differing or non-overlapping characters: spine-like tubercles absent on flanks (versus spine-like tubercles present on flanks in C. amboliensis, C. assamensis, C. flaviventralis, C. goaensis, C. jerdonii, C. littoralis, C. monticola, C. mysoriensis and C. nilagirica); scales on dorsal aspect of trunk heterogeneous (versus scales on dorsal

TWO NEW CNEMASPIS FROM TAMIL NADU, INDIA Zootaxa 4609 (1) © 2019 Magnolia Press · 87 aspect of trunk homogeneous in C. adii, C. assamensis, C. australis, C. boiei, C. indica, C. jerdonii, C. kolhapurensis, C. littoralis, C. nilagirica and C. sisparensis); tail with median row of sub-caudal scales smooth and enlarged (versus median row of sub-caudal scales smooth and not enlarged in C. ajijae, C. flaviventralis, C. girii, C. limayei; and C. monticola; C. australis with keeled sub-caudals), absence of keeled scales on the venter or gular regions (versus keeled scales on the venter or gular region in C. beddomei and C. goaensis), males with four femoral pores on each thigh, separated on either side by 7–9 poreless scales from 2–4 precloacal pores; precloacal pores separated medially by two or three poreless scales (versus precloacal pores absent, femoral pores present in C. ajijae, C. anaikattiensis, C. flaviventralis, C. girii, C. indica, C. jerdonii, C. kottiyoorensis, C. limayei, C. littoralis, C. mahabali, C. sisparensis, C. heteropholis, C. wynadensis; only precloacal pores present in C. anamudiensis, C. beddomei, C. maculicollis, C. nairi, C. ornata; both femoral and precloacal pores absent in C. boiei, C. assamensis; three femoral and four precloacal pores present in C. otai; two femoral and two precloacal pores on each thigh in C. adii; four or five femoral and three precloacal pores in C. australis; 2–4 femoral and three precloacal pores in C. goaensis; 4–6 femoral pores on each thigh, separated on either side by eight poreless scales from four precloacal pores; precloacal pores separated medially by a single poreless scale in C. agarwali; 5–9 femoral pores on each thigh, separated on either side by 1–6 poreless scales from five or six precloacal pores; precloacal pores separated medially by one or two poreless scales in C. thackerayi sp. nov.; two femoral and two precloacal pores in C. mysoriensis; three femoral and two precloacal pores in C. yercaudensis; a continuous series of 26–28 precloacal-femoral pores in C. kolhapurensis). Cnemaspis shevaroyensis sp. nov. closely resembles C. gracilis, C. agarwali, and C. thackerayi sp. nov. However, it can be distinguished from all three by presence of four femoral pores on each thigh and 2–4 precloacal pores separated medially by two or three poreless scales (versus 4–6 femoral pores on each thigh and four precloacal pores separated medially by a single poreless scale in C. agarwali; 5–9 femoral pores on each thigh and five or six precloacal pores separated medially by one or two poreless scales in C. thackerayi sp. nov.); lamellae under digit IV of pes 16–20 (versus lamellae under digit IV of pes 22 in C. gracilis; 21–24 in C. thackerayi sp. nov.); maximum snout to vent length 34 mm (versus maximum SVL 41 mm in C. thackerayi sp. nov.); by having 10–14 rows of dorsal scales (versus 9–11 rows of dorsal scales in C. agarwali); presence of two single dorsal ocelli on occiput and between forelimb insertions, two pairs on either side just anterior and posterior to forelimb insertions (versus presence of two single dorsal ocelli on occiput and between forelimb insertions in C. gracilis; presence of single dorsal ocellus between forelimb insertions in C. thackerayi sp. nov.). Additionally, Cnemaspis shevaroyensis sp. nov. is 10.3 % divergent from C. gracilis; 13.3 % divergent from C. cf. gracilis 7.9 % divergent from C. agarwali and 13.6 % divergent from C. thackerayi sp. nov. (Table 2). Description of the holotype. Adult male in good state of preservation except body fixed in slightly sigmoid manner and a 3.3 mm long incision in the sternal region for tissue collection as artefacts of preservation (Fig. 8 A, B). SVL 30.4 mm, head short (HL/SVL ratio 0.25), wide (HW/HL ratio 0.68), not strongly depressed (HD/HL ratio 0.41), distinct from neck. Loreal region slightly inflated, canthus rostralis not prominent. Snout just less than half head length (ES/HL ratio 0.49), 2.5 X eye diameter (ED/ES ratio 0.41); scales on snout and canthus rostralis large, weakly keeled, juxtaposed; larger than those on forehead and interorbital region; occipital and temporal region with much smaller granules, intermixed with larger, roughly rounded, tubercles (Fig. 9 A). Eye small (ED/HL ratio 0.20); with round pupil; orbit with extra-brillar fringe scales that are largest anteriorly; supraciliaries not elongate. Ear- opening deep, vertical, small (EL/HL ratio 0.05); eye to ear distance greater than diameter of eye (EE/ED ratio 1.62) (Fig. 9 C). Rostral much wider (1.5 mm) than long (0.8 mm), incompletely divided dorsally by a strongly developed rostral groove for more than half of its length; single enlarged supranasal on each side, roughly similar in size to postnasals, separated from each other by smaller single internasal and a single smaller scale on the snout; rostral in contact with supralabial I, nasal, supranasal and internasal; nostrils oval, each surrounded by postnasal, supranasal, rostral and supralabial I; a single row of scales separate the orbit from the supralabials (Fig. 9 C). Mental enlarged, subtriangular, slightly wider (1.6 mm) than long (1.2 mm); two pairs of postmentals, inner pair large, roughly rectangular, strongly in contact with each other, bordered by mental, infralabial I, outer postmentals and two enlarged chin shields; outer postmentals slightly smaller than inner postmentals, bordered by infralabials I, II, inner postmentals and four enlarged chin shields; two enlarged gular scales prevent contact of left and right outer postmentals; chin shields bordering postmentals flat, smooth, smaller than outermost postmentals, rest granular, much smaller, smooth. Infralabials bordered below by a row of slightly enlarged scales, decreasing in size posteriorly (Fig. 9 B). Supralabials to angle of jaw seven (L) — eight (R) and six on either side at midorbit; supralabial I

BNHS 2531

BNHS 2530 ……continued on the next page

BNHS 2529 BNHS NCBS- BH676 BH675 BH675 = tail/ Materials or anddigit incomplete Abbreviations are listed Methods. in * sp. nov. 7.9 8.6 7.1 8.5 8.2 7.5 7.9 8.2 8.6 7.1 8.5 1.5 1.6 1.6 1.8 1.5 1.6 2.9 3.1 2.9 3.5 2.8 3.2 6/5 5/5 6/5 5/5 5/5 4/4 6/5 5/5 5/5 6/5 5/5 40 19* 35.1 42.6 4.2* 14.1* 4.2* 42.6 35.1 40 19* 3.3 3.4 3.4 2.6 3.8 3.3 2.4 2.6 2.7 2.6 2.4 2.8 0.4 0.4 0.6 0.4 0.3 0.6 6/6 7/7 7/5 6/7 7/7 7/7 6/6 6/6 6/6 6/6 5/5 5/5 6/6 5/5 6/6 6/6 13 13 12 10 14 11 M F F F M M M M F 113 117 118 114 111 118 Holotype Paratypes 3.1 3 3 5.2 2.8 3.1 5.8 5.8 5.3 6.1 5.6 5.6 5.8 5.6 5.8 5.3 6.1 6.3 8 5.2 8 6.2 6.1 6.2 8 6.3 8 5.2 3.8 3.9 3.8 4.4 3.9 4.2 24 22 21 24 24 23 23 24 24 24 21 22 30.4 33.2 28.6 34.1 31.3 29.7 30.4 31.3 33.2 28.6 34.1 15 17 13 16 16 16 5.4 5.7 4.9 5.8 5.4 5.7 5.7 5.4 5.4 5.8 4.9 5.7 0.9 1 0.9 1 0.9 0.9 0.9 0.9 1 1 0.9 0.9 2.5 2.5 2.4 2.9 2.1 2.5 12.7 15 11.4 14.9 12.6 11.6 11.6 12.6 14.9 15 12.7 11.4 3.3 3.8 3 3.8 3.3 3.3 3.3 3.3 3 3.8 3.8 3.3 7/8 7/7 7/8 7/7 7/7 6/6 6/6 7/87/7 7/7 7/8 7/7 4.9 4.9 5 4.7 4.7 4.9 5.2 Cnemaspis shevaroyensis Measurements (mm) and data (mm) the series for Measurements of type meristic TABLE 5. missing, /= data not available. No.Specimen Sex SVL TL TW LAL CL AGL BH BW HL HW HD ED EE ES EN IN IO NCBS-BH674 EL NCBS- L/R SL L/RIL L/R M SL M IL L/R PVT RBS L/R RVS VS LamF1 9/9 10/10 10/9 8/9 10/10 10/10

BNHS 2531 BNHS

BNHS 2530

BNHS 2529 NCBS- BH676 BH675 BH675 Holotype Paratypes 2 0 0 0 2 3 2 0 1/1 0 0 0 1/1 2/2 2/2 1/1 0 1/1 4/4 0 0 0 4/4 4/4 4/4 0 0 0 4/4 (Continued) (Continued) . 5 SB PP&FP L/R L/R PP&FP SBPP SB FP L/R No. tubercles postcloacal of scales Anteriorenlarged extra-brillar(1) fringe not or enlarged (0) (0) smooth or (1) keeled Ventral scales scales keeled (1)Gular (0) smooth or 7/7 keeled Pectoral scales (1) smooth (0) or 8/9 0 separated (0) or (1) continuous Precloacal pores 9/8 (1)Precloacal or pores elongate round (0) pores elongate (1)femoral round (0) or 1 Dorsal homogeneous heterogeneous (0) or pholidosis (1) (0)Dorsal keeled ortubercles keeled (1) not linearly random arranged (1) (0)Tubercles more or Spine-like present on (0) or(1) tubercles flanks absent 1 (0) flanks lower on (0) absent or (1) present Tubercles present caudal orLateral (0) (1) furrows absent Caudal inor(1) tubercles not furrow lateral (0) Subcaudals(0) (1) keeled smooth or 0 1 0 1 keeled of Single row subcaudals median (1) (0) or smooth 0 0 0 Caudal encircle tail or tubercles (1) not (0) (1)row Enlarged ornot subcaudal (0) median scale 1 presentscales Enlarged (1) femoral absent or (0) 0 1 1 (0) absent or (1) present subtibial scales Shield-like 1 - 0 1 1 0 0 1 Subtibial keeled (1) smooth (0) or scales 0 - 1 0 1 - 1 0 1 - 0 0 0 1 1 0 1 1 - 0 1 1 - 1 1 - 0 0 0 0 0 1 0 1 1 1 1 0 - 0 1 0 1 1 1 0 0 0 - 0 0 1 0 1 1 1 1 1 0 0 1 1 0 1 1 0 0 1 1 1 0 0 1 1 0 1 1 1 1 0 1 1 1 1 1 0 1 1 1 1 0 1 0 1 0 1 0 1 1 0 1 0 0 1 1 0 1 1 0 1 1 TABLE L/R No.Specimen L/R LamF4 L/R LamT1L/R LamT4 LamT5 PP L/R 16/15 16/16 15/15 16/16 15/15 16/16 17/17 19/19 16/17 16/15 20/18 20/19 20/18 14*/18 17/16 18/17 18/17 16/16 NCBS-BH674 8/9 NCBS- 8/8 9/9 10/10 10/10 10/10

90 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. FIGURE 8. Cnemaspis shevaroyensis sp. nov. (holotype, NCBS-BH674), dorsal view (A), ventral view (B), dorsal view of tail (C), and ventral view of tail (D). Scale bars 10 mm.

TWO NEW CNEMASPIS FROM TAMIL NADU, INDIA Zootaxa 4609 (1) © 2019 Magnolia Press · 91 FIGURE 9. Cnemaspis shevaroyensis sp. nov. (holotype, NCBS-BH674), dorsal view of head (A), ventral view of head (B), lateral view of head (C), view of cloacal region showing precloacal and femoral pores (D), ventral view of right manus (E), and ventral view of right pes (F). Scale bars 5 mm.

92 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. FIGURE 10. Type series of Cnemaspis shevaroyensis sp. nov. (from left to right, NCBS-BH675, NCBS-BH676, BNHS 2529, BNHS 2530 and BNHS 2531). Scale bar 10 mm. largest, decreasing in size posteriorly; seven infralabials on either side up to angle of jaw and six(L) — five(R) at midorbit position; infralabial I largest, decreasing in size posteriorly (Fig. 9 C). Extra-brillar fringe scales 16 or 17 on each side, interorbital scale rows across narrowest point of frontal six or seven; 27 or 28 scale rows between left and right supraciliaries at midorbit (Fig 9 A). Body relatively slender, trunk less than half of SVL (AGL/SVL ratio 0.41) without ventrolateral folds or spine- like tubercles on flanks. Dorsal scales on trunk heterogeneous, weakly keeled, granular scales intermixed with much larger, strongly keeled, conical tubercles; tubercles in approximately 14 longitudinal rows at mid-body; 13 tubercles in paravertebral row from above forelimb insertion to the hind limb insertion (Fig. 7 C). Scales on nape slightly smaller than those on paravertebral rows, smaller still on occiput (Fig. 9 A). Ventral scales slightly larger than dorsals, those on belly smooth, subimbricate, slightly rounded, subequal from chest to vent; midbody scale rows across belly 24; 117 scales from mental to anterior border of cloaca (Fig. 7 D). Scales on throat and pectoral region slightly smaller than those on belly, flat and imbricate; gular region with much smaller, flattened granules with those on chin bordering postmentals, enlarged, juxtaposed and flattened (Fig. 9 B). Four femoral pores on each thigh and two precloacal pores separated medially by two poreless scales (Fig. 9 D). Scales on palm and sole smooth, flat and roughly circular; scales on dorsal aspect of manus and pes heterogenous, upper arm with scales slightly larger than dorsal granules, weakly keeled, subimbricate; those near forelimb insertion much smaller, granular; dorsal aspect of forearm with scales smaller than those on upper arm, weakly keeled, flat, roughly rounded; scales on elbow weakly keeled, subimbricate and similar in size to those on upper arm; dorsal aspect of hand predominantly bearing small, flattened weakly keeled, imbricate scales. Ventral aspect of upper arm with smooth, roughly rounded, weakly conical granules; scales on forearm and wrist with larger, smooth, weakly imbricate scales. Scales on dorsal aspect of thigh larger than those dorsal granules, weakly keeled, imbricate except those near hindlimb insertion which are much smaller, roughly rounded, conical granules. Scales on dorsal aspect of knee and shank slightly smaller than those on dorsum of thigh, subimbricate, weakly keeled; dorsal aspect of foot predominantly bearing small, flattened, weakly keeled, imbricate scales. Scales on ventral aspect of thigh similar to those on midbody ventrals; those on ventral surface of shank roughly larger than those on thigh, smooth, imbricate (Fig. 8 B). Fore and hind limbs moderately long, slender (LAL/SVL ratio 0.16); CL/SVL ratio 0.19; digits long, with a strong, recurved claw, distinctly inflected, distal portions laterally conspicuously compressed. Series of unpaired lamellae on basal portion of digits, separated from narrower distal lamellae by a single large scale at the inflection; proximal lamellae series: 1–3–3–3–3 (right manus; Fig. 9 E), 1–3– 5–5–5 (right pes; Fig. 9 F), 1–3–3–4–4 (left manus), 1–4–4–4–5 (left pes); distal lamellae series: 8–9–11–12–10 (right manus; Fig. 9 E), 8–10–12–12–11 (right pes; Fig. 9 F), 8–9–12–11–10 (left manus), 7–10–12–12–12 (left

TWO NEW CNEMASPIS FROM TAMIL NADU, INDIA Zootaxa 4609 (1) © 2019 Magnolia Press · 93 pes). Relative length of digits (measurements in mm in parentheses): IV (2.79) > III (2.65) > V (2.55) > II (2.29) > I (1.80) (right manus); IV (3.98) > V (3.69) > III (3.33) > II (2.53) > I (1.71) (right pes). Tail entire, cylindrical, relatively slender, flattened beneath, longer than snout-vent length (TL/SVL ratio 1.31) (Fig. 8 C, D). Dorsal scales at tail base granular, similar in size and shape to those on midbody dorsum, gradually becoming larger, flatter, pointed, subimbricate posteriorly, intermixed with slightly enlarged, strongly keeled, conical tubercles forming whorls; 6–8 tubercles on first 2–5 whorls. Scales on ventral aspect of original tail much larger than those on dorsal, imbricate, smooth, with a series of three enlarged subcaudal scales of which the median series is almost twice the size of adjunct two rows, roughly hexagonal; those on tail base much smaller, imbricate and smooth, a single enlarged postcloacal spur on each side (Fig. 8 D). Colouration in life. (Fig. 6 E) Dorsum of head and body brown, limbs suffused with yellow; tail base similar to body dorsum, rest of tail with eight alternating black and light grey/white bands. Head with numerous greyish blotches, brille yellow. Radiating lines from orbit forming an indistinct preorbital streak on snout; yellow and dark grey and light bands on labials; two postorbital streaks, the lower black and fine and the upper thicker and red with some black terminating anterior to forelimb insertion in a small black streak. Small dark spot on occiput flanked by larger light grey blotches; spot and blotches with orange-red markings. A single large central black ocellus at anterior of forelimb insertion, two pairs of smaller spots on each side just anterior and posterior to forelimb insertions that are larger than occipital spot; all ocelli outlined by some orange-red scales. Dorsum with seven light grey vertebral blotches between limb insertions and one on tail base; smaller light grey spots on rest of dorsum and flanks; three indistinct, small orange blotches on flanks forming a discontinuous line as a continuation of dorsal ocelli, first anterior to midbody, one anterior to hindlimb insertions and the last at tail base. Dorsum of limbs with lighter spots, more prominent on hindlimbs, digits with alternating dark and light bands. Ventral surfaces off-white, suffused with yellow under limbs and tail base, less so on belly and most strongly on lateral margins of gular region; a dark broken longitudinal streak outlines yellow of throat, followed by two more prominent streaks within yellow; no dark markings on underside of tail. Variation and additional information from type series. Mensural data for the type series is given in Table 5. There are two male and three female specimens ranging in SVL from 28.6–34.1 mm. All paratypes resemble the holotype except as follows: the number of lamellae on digit I of the manus ranges from 8–10 and on digit IV from 15 or 16; on digit IV of the pes ranges from 17–20 and on digit V from 15–18; 5–7 supralabials to angle of jaw; 5– 7 infralabials; 15–17 tubercles in paravertebral rows; 10–13 dorsal tubercle rows; ventral scale counts in longitudinal and transverse series vary from 111–118 and 21–23, respectively. Males with one or two precloacal pores on either side separated by two or three poreless scales; seven or eight poreless scales between precloacal and femoral pores. Complete tail slightly longer than body (TL/SVL ratio 1.22–1.24). Two male paratypes match the holotype in overall colouration; the three female paratypes are duller in appearance without oranges blotches on dorsum, tail light yellow with light brown bands and lacking ashy-grey colouration. Distribution and Natural history. Cnemaspis shevaroyensis sp. nov. is so far known only from the type locality (Kottachedu Kari Raman Temple, Yercaud, in the Shevaroys), at an elevation of ca. 900 m asl. (Fig. 1). The species was encountered during a few hours of fieldwork in perennial stream during the day (1230h) on rocky boulders situated in a teak plantation. The forest type was dry deciduous with partial canopy cover (Fig. 11 C). The active male holotype (NCBS-BH674; Fig. 6 E) and paratypes were collected from a dry culvert inside a large cement pipe. The culvert was also a communal oviposition site and contained a few hatched and unhatched eggs along with the gravid female paratype (BNHS 2529; Fig. 6 F). The species seems to prefer shaded areas as all collected specimens were from dark and cool spots during the day. Sympatric lizards at the type locality include Hemidactylus frenatus, Hemidactylus parvimaculatus, Eutropis carinata, Eutropis macularia, Lygosoma albopunctata, and Calotes versicolor.

Discussion

We performed a rapid survey along an elevation gradient (800–1500 m asl.) of the Shevaroy massif and found four species of Cnemaspis: C. yercaudensis and C. thackerayi sp. nov. at high elevations (>1300 m asl.), and two other species at allopatric mid elevation sites, C. cf. gracilis on the southwestern slope and C. shevaroyensis sp. nov. on the southeastern slope of the hill, all within 10 km straight line distance from each other. The seemingly

94 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. FIGURE 11. Habitat photos of (A) Cnemaspis yercaudensis, (B) Cnemaspis thackerayi sp. nov., and (C) Cnemaspis shevaroyenesis sp. nov., from the Shevaroy massif, Salem district, Tamil Nadu, India.

sp. nov. shevaroyensis C. sp. nov. sp.

thackerayi C. yercaudensis . C from Yercaud. Colour characters binary were scored as present (1) or absent (0) and Cnemaspis Diagnostic colour pattern characters separating three species of depending upon choicethe between two colours as one (1) or other the (0). TABLE 6. Species dimorphic Dorsal (1)or colour pattern not sexually (0) (1) Ventralpatterndimorphic not or (0) sexually (0) not or (1) yellow Head (1) scales orAnterioryellow extra-brillar fringe not (0) (0) not or (1) body and head on blotches Orangish (1) on reticulum Dense orange neck occiput and sideof ornot (0) not (0) or occiput (1) on ocelli dark Dorsal insertion Dorsal forelimb dark ocelli between not or (1) (0) not (0) or insertion(1) to forelimb ocelli anterior dark Lateral insertion dark ocelli Lateral posterior not (0) or (1) to forelimb on (1)Thin neck of ornot orange reticulum (0) side (1) or Dorsal (0) brown yellow ground colour orabsent(0) (1) present stripe vertebral Light 1 present Squarish, markings (1) vertebral dorsal absent or (0) roundSmall, (1) yellow spots on flanks or (0) white (1) region Gular oryellow not (0) 0 1 0 gular orLineate present (1) absent markings (0) (1) or Throat not region yellow (0) 0 (1) Pectoral or(0) region not yellow 0 (0) not or (1) yellow Abdomen 0 1 0 (1) Forelimbs oryellow not (0) (1)Hindlimbs yellow ornot (0) 0 1 blotches/bands (1)Orange not or on (0) limbs 0 0 1 of (1)Ventralforelimbs ornot (0) yellow surfaces 0 (0) not or (1) yellow limbs hind of surfaces Ventral 0 1 Original tailor banded not (1) (0) 1 0 orange tail Regenerated in (1) (0) males brown or 0 0 1 1 1 0 1 1 0 1 1 1 1 1 1 1 0 1 1 1 0 1 0 0 0 0 1 0 1 0 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1

96 · Zootaxa 4609 (1) © 2019 Magnolia Press KHANDEKAR ET AL. specialised elevational range of these Cnemaspis species is indicative of conservatism in thermal tolerance, while the occurrence of non-sister Cnemaspis species along the elevation gradient of the Shevaroy hills indicates the region has received multiple dispersals of distinct Cnemaspis lineages. Interestingly, C. yercaudensis is distributed along the high elevations of Shevaroys and Kollimalai, whereas the two new species and C. cf. gracilis include two mid elevation lineages and a high elevation point endemic, which are replaced by different species in Kollimalai (Khandekar & Agarwal unpubl. data). This counterintuitive pattern, with one high elevation species distributed across the isolated high elevations of the Shevaroys and Kollimalai is suggestive of a recent cool and or wet phase that may have allowed C. yercaudensis to disperse between the hill ranges. Apart from having specific elevation ranges, the different species have distinct microhabitat preferences. C. yercaudensis was found to be semi- terrestrial, C. thackerayi sp. nov. occupied human-modified habitats (on walls < 5 m), whereas C. shevaroyensis sp. nov. and C. cf. gracilis are strictly rock-dwelling. The eastern part of southern India has several hill ranges east of the Western Ghats that rise to elevations over 1000 m asl., including the Shevaroys, Kollimalai, Pachamalai, Chitteri hills, Biligirangan, Malemahadeshwar, and Mettur hills. These ranges have often been considered part of the Eastern Ghats, but are now emerging as being more closely related to the Western Ghats (e.g. Raheem et al. 2014; Ramachandran et al. 2016; Agarwal et al. 2019). The Shevaroy hills rise up to >1600 m asl. with high elevation plateaus with tropical semi evergreen forests and deciduous forests along the slopes which transition into dry scrub at lower elevation (Kadavul & Parthasarathy 2004). Thus, the hills harbour distinct habitats and host a wide variety of taxa, many of which are endemic to the region (Nayaka et al. 2013; Ganesh et al. 2014; Ganesh & Arumugam 2016; Agarwal et al. 2019). The tall hills abruptly rising from the low-lying plains serve as habitat islands aiding isolation, and have possibly given rise to the high diversity observed in some gecko lineages such as Cnemaspis, Cyrtodactylus, Hemiphyllodactylus and Hemidactylus (Agarwal 2016; Agarwal et al. 2017; Khandekar 2019). The unique sky island-like habitat at higher elevations of the hills separated by low lying dry deciduous scrub forests has led to narrow distribution and habitat specialization in species, giving rise to many point endemics in the region. Therefore, dedicated and extensive sampling along with genetic investigations are needed to unravel the true diversity of the diverse gekkonid lineages in this understudied region. The deep genetic divergence within South Asian Cnemaspis (>30 %) is comparable to that seen within other highly diverse gekkonid genera including Southeast Asian Cnemaspis (Grismer et al. 2014), Cyrtodactylus (Wood et al. 2012; Agarwal et al. 2014), and Hemidactylus (Bauer et al. 2010). This suggests a deep history of the group in this landscape, and that species diversity may be much higher than currently thought. In addition to flagging many divergent lineages, we found that topotypical C. indraneildasii is genetically identical to C. goaensis (as did Sayyed et al. 2018)—and a preliminary examination of morphology suggests that the two are in fact conspecific, with the older valid name C. goaensis.

Acknowledgements

We thank Varad Giri, Swapnil Pawar, Tejas Thackeray, Aniruddha Datta-Roy, Krishna R. Chaitanya, and Anuradha Batabyal for helping with fieldwork at Yercaud, Tamil Nadu, and Tejas Thackeray, Hemant Ogale, Sujit Borkar, Santosh Saste, Shankar Desai, Arjun Kamdar, Swapnil Pawar, Vijay Sandge, Bhaskar Mahadik, Motiram, Rajendra Kadam, Robin Bansode, Sanjay Patil, Santosh Chalke, Shivaji Zunjar, Vaibabh Patil and Ninad Gosavi for assistance in fieldwork in Maharashtra. AK is grateful to Tejas Thackeray and Santosh Saste for their constant support and encouragement. V. Deepak helped with extractions. Partial funding came from the Ministry of Environment and Forests, the Department of Atomic Energy (2012/21/06/BRNS to Uma Ramakrishnan), and the Department of Science and Technology (DST grant SR/SO/AS-57/2009 to Praveen Karanth), Govt. of India. We thank Krushnamegh Kunte for letting us examine specimens using his lab microscopes and Uma Ramakrishnan for lab support. Krushnamegh Kunte and Rahul Khot helped register specimens at NCBS and BNHS museums respectively. We are thankful to the Forest Departments of Maharashtra and Tamil Nadu for permits and logistic support.

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Cnemaspis agarwali: holotype NCBS-AU486 (adult male), paratypes NCBS-AU487, BNHS 2337, NCBS-AU488, NCBS- AU490, and BNHS 2338, (adult males), NCBS-AU485, BNHS 2336, and BNHS 2339, (adult females), from near Kidayur Road, Sankari, Salem District, Tamil Nadu state, India. Cnemaspis indraneildasii: AK 177, AK 178, AK 179, AK 180, AK 182, AK 183, AK 184, from Gund, Uttara Kanada District, Karnataka; AK 185, from Ganeshgudi, Uttara Kanada District, Karnataka. AK 195, AK 196, AK 197, AK 198, AK 199, VG 403, VG 404, VG 405, VG 406, from Sangli, Sangli district, Maharashtra, India. Cnemaspis amboliensis: AK 181, AK 190, AK 191, AK 192, AK 194 from Parpoli, Sindhudurg District, Maharashtra, India. Cnemaspis flaviventralis: AK 186, AK 187, AK 188, AK 189, AK 193, from Amboli, Sindhudurg District, Maharashtra, India. Cnemaspis ajijae: AK 321, AK 322, AK 323, AK 324, AK 325, AK 326, AK 327, AK 328, AK 329, AK 330, AK 331, AK 332, AK 357, AK 429, AK 430, AK 431, AK 432, AK 433, AK 434, AK 435, AK 436, AK 437, AK 438 from Mahabaleshwar, Satara District, Maharashtra, India. Cnemaspis goaensis: AK 358, AK 359, AK 360, AK 361, from Cancona, South Goa district, Goa, India. Cnemaspis limayei: AK 362, AK 363, AK 364, AK 365, AK 366, AK 367, AK 368, AK 369, AK 372, AK 373, AK 374, AK 375, AK 376, AK 377, AK 378, AK 379, AK 380, AK 381 from Dajipur, Kolhapur District, Maharashtra, India. Cnemaspis kolhapurensis: AK 370, AK 371 from Dajipur, Kolhapur district, Maharashtra, India. Cnemaspis girii: AK 439, AK 440, AK 441, AK 442 from Kaas Plateau, Satara District, Maharashtra, India. Cnemaspis gracilis: CESG385 from near Chittur River, Palakkad District, Kerala, India. AK 133, AK 134, AK 135, AK 136, AK 137, AK 138, AK 139, AK 140, AK 141, AK 142, AK 143, AK 144, from Valparai town, Coimbatore District, Tamil Nadu, India. Cnemaspis mysoriensis: AK569, AK570, AK 571 from National Centre for Biological Sciences (NCBS) campus, Bengaluru, Bengaluru District, Karnataka, India.