Cnemaspis flaviventralis, a new species of gecko (Squamata: Gekkonidae) from the Western Ghats of Maharashtra, India

 

 

 

 

 

doi: http://dx.doi.org/10.11609/jott.2599.8.14.9619-9629 | ZooBank: urn:lsid:zoobank.org:pub:B5F65768-6FF2-4670-95B5-8C2E80B8D97E

Editor: Olivier S.G. Pauwels, Royal Belgian Institute of Natural Sciences, Brussels, Belgium. Date of publication: 26 December 2016 (online & print)

Manuscript details: Ms # 2599 | Received 03 March 2016 | Final received 09 December 2016 | Finally accepted 12 December 2016

Citation: Sayyed, A., R.A. Pyron & N. Dahanukar (2016). Cnemaspis flaviventralis, a new species of gecko (Squamata: Gekkonidae) from the Western Ghats of Maharashtra, India. Journal of Threatened Taxa 8(14): 9619–9629; http://dx.doi.org/10.11609/jott.2599.8.14.9619-9629

Copyright: © Sayyed et al. 2016. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use of this article in any medium, reproduction and distribution by providing adequate credit to the authors and the source of publication.

Funding: RAP is funded by U.S. NSF grants DBI-0905765 and DEB-1441719. ND is funded by DST-INSPIRE Research Grant [IFA12-LSBM-21].

Competing interests: The authors declare no competing interests. Funding sources had no role in study design, data collection, results interpretation and manuscript writing.

Author Details: Amit Sayyed is head of research department at Wildlife Protection and Research Society. He works on faunal diversity, ecology, taxonomy, distribution and evolution of Reptiles and Amphibians. Robert Alexander Pyron is an Assistant Professor of Biology at The George Washington University. He works on theoretical and applied methods in statistical phylogenetics, using reptiles and amphibians as model groups. Neelesh Dahanukar works in ecology and evolution with an emphasis on mathematical and statistical analysis. He is also interested in taxonomy, distribution patterns and molecular phylogeny of ichthyo- and herpetofauna.

Author Contribution: AS conducted field work, took photographs, studied museum specimens, diagnosed the species and performed taxonomic studies; ND performed phylogenetic analysis; AS, RAP and ND wrote the paper.

Acknowledgements: AS is thankful to Hemant Ghate, Varad Giri, Anil Mahabal, TSN Murthy and Anilkumar Khaire for inspiration and to Zeeshan Mirza, Amod Zhambre, Harshal Bhosle and Vivek Sharma for motivation. Fieldwork would not have been possible without the help of Kaka Bhise and member of WLPRS: Abhijit Nale, Jitendra Patole, Rohit Katke, Kapil Taple, Rahul Thombre, Vinay Chavan, Datta Chavan, Mangesh Karve, Devendra Bhosle, Santosh Chavan, Aman Adsul, Nikhil Mutha, Mudassar Narwade and Chaitanya Shukla. We are thankful to Patrick Campbell, Natural History Museum, London; Deepak Apte, director, and Rahul Khot, museum curator, Bombay Natural History Society, Mumbai; and Kaushik Deuti and Varada Raju, Zoological Survey of India, HQ, Kolkata, and P.S. Bhatnagar, the officer-in-charge, Shrikant Jadhav and Sameer Pati, Zoological Survey of India, Western Regional Center, Pune, for their help in museum related work.

 

 

 

 

 

Abstract: Cnemaspis flaviventralis, a new species of day gecko, is described from the forests of Amboli, Sindhudurg District, Maharashtra State, northern Western Ghats, India. The new species was previously confused with the sympatric species Cnemaspis girii, C. indraneildasii, C. kolhapurensis and C. goaensis. It is distinguished from C. giri by having spine-like tubercles on flanks, granular dorsal scales intermixed with large, depressed, slightly keeled scales (vs. lack of spine-like tubercles on flanks, granular dorsal scales, intermixed with large smooth scales); from C. indraneildasii by having dorsal scales heterogeneous (vs. homogenous), lacking a series of enlarged median sub-caudal scales, and 28–29 (vs. 20) ventral scales across mid-body; from C. kolhapurensis by having heterogeneous (vs. homogenous) dorsal scalation, lacking spine-like tubercles on flanks and lacking pre-cloacal pores (vs. 24–28 pre-cloacal-femoral pores); and from C. goaensis by lacking pre-cloacal pores and lacking a series of enlarged median sub-caudal scales. We further provide partial mitochondrial 16S rRNA gene sequences for the new species and for the sympatric species C. girii, C. kolhapurensis and C. goaensis, and show that the new species is genetically distinct.

Keywords: Amboli, Maharashtra, molecular phylogeny, new species, taxonomy.

 

INTRODUCTION

The species-rich lizard genus Cnemaspis Strauch, 1887 in the family Gekkonidae includes at least 120 Old World species (Uetz & Hošek 2015) distributed from Africa to South-east Asia. As presently understood, the widespread genus Cnemaspis appears to be polyphyletic (Gamble et al. 2012; Pyron et al. 2013; Grismer et al. 2014). Further, cryptic and underestimated diversity is also very high in Cnemaspis (e.g., Manamendra-Arachchi et al. 2007; Grismer et al. 2014; Amarasinghe et al. 2015). The Western Ghats-Sri Lanka biodiversity hotspot is becoming a hotspot for Cnemaspis with several new species described in recent years (e.g., Manamendra-Arachchi et al. 2007; Giri et al. 2009; Cyriac & Umesh 2014; Mirza et al. 2014; Vidanapathirana et al. 2014; Srinivasulu et al. 2015). Given the extreme morphological similarity among species within the group, their small size and the generally overlooked nature of the South Asian herpetofauna, discovering new species is not surprising.

Based on a new collection from the northern Western Ghats of Maharashtra, India, we describe here a new species of Cnemaspis to make a name available and facilitate their conservation. We also provide a molecular analysis based on 16S rRNA genomic sequences that may facilitate future studies in the region.

MATERIALS AND METHODS

 

Specimen collection

Specimens of the type series were collected from Amboli (15.9600N & 73.9990E, elevation 735m), Sindhudurg District, Maharashtra, India. Four males and four female specimens were collected during night. Two specimens of Cnemaspis girii Mirza, Pal, Bhosale & Sanap, 2014 were collected from Kaas plateau (17.7240N & 73.8190E), the type locality of the species; two specimens of C. kolhapurensis Giri, Bauer & Gaikwad, 2009 were collected from Amboli (15.9710N & 73.9790E); and two specimens of C. goaensis Sharma, 1976, were collected from Goa (15.2900N & 74.0060E). They were anaesthetized, fixed in formalin, and preserved in 70% ethyl alcohol. Geographic coordinates and atmospheric temperature readings were taken at the collection localities with the aid of a Kestrel 4500 weather meter.

 

Museum details

Specimens from the current study are deposited in the collections of the Bombay Natural History Society, Mumbai (BNHS) and the Zoological Survey of India, Western Regional Center (ZSI-WRC), Pune, Maharashtra. Comparative materials were examined at the Zoological Survey of India, Kolkata (ZSI-K).

 

Morphometric and meristic data collection

Following measurements were taken using a Yamayo digimatic caliper to the nearest 0.1mm following Giri et al. (2009) and Mirza et al. (2014), and include snout vent length (SVL, from tip of snout to vent), trunk length (TRL, distance from axilla to groin measured from posterior edge of forelimb insertion to anterior edge of hind limb insertion), trunk width (TrW, maximum width of body), tail length (TL, from vent to tip of tail), tail width (TaW, 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 depth of head, from occiput to underside of jaws), forearm length (FL, from base of palm to elbow), tibia length (TBL, knee to tarsal), eye to nares distance (E-N, distance between anteriormost point of eye and nostril), eye to snout distance (E-S, distance between anteriormost point of eye and tip of snout), eye to ear distance (E-E, distance from anterior edge of ear opening to posterior corner of eye), Ear length (EL, greatest diameter of ear opening), internarial width (IN, distance between nares), orbital diameter (OD, greatest diameter of orbit), inter orbital distance (IO; greatest distance between left and right supraciliary scale rows). Following meristic data were taken for all specimens under a Leica stereo microscope: supralabial scales (SupraL), infralabial scales (InfraL), femoral pores (FPores), lamellae under digits of manus (MLam) and pes (PLam) for both left (L) and right (R) sides (lamellae counts taken from the scale just behind claw to first interphalangeal joint excluding large scansors), longitudinal scale rows across the belly (in between the lowest rows of lateral granular scales (MVS).

 

Genetic analysis

Muscle tissue harvested from two fresh specimens of the new species (ZSI-WRC 1042, 1043) collected from same locality of holotype, two specimens of C. girii (BNHS 2445, 2446), two specimens of C. goaensis (ZSI R/1044 and 1045) and two specimens of C. kolhapurensis (BNHS 2447, 2448). Tissue samples were digested at 550C using STE buffer (50 mM Tris-HCl, 20 mM EDTA and 50μl of 10% SDS) with 10 μL of 20 mg/ml Proteinase K. RNase treatment was given for two hours at 370C. Final extraction process was done using phenol-chloroform method. Polymerase Chain Reaction was performed for amplification of mitochondrial 16S rRNA gene using the primer pair 16SF (5’-CCC GCC TGT TTA CCA AAA ACA-3’) and 16SR (5’-GGT CTG AAC TCA GAT CAC GTA-3’). PCR reaction was performed in a 25μl reaction volume containing 5μl of template DNA (~200ng), 2.5μl of 10X reaction buffer (100 mM Tris pH 9.0, 500 mM KCl, 15 mM MgCl2, 0.1% Gelatin), 2μl of 25 mM MgCl2, 1μl of 10 mM dNTPs, 1μl of each primer, 1μl Taq polymerase and 16.5μl nuclease free water. The thermal profile was 10 minutes at 950C, and 35 cycles of one minute at 940C, one minute at 520C and two minutes at 720C, followed by extension of 10 minutes at 720C. Amplified DNA fragments were purified using the ‘Promega Wizard Gel and PCR clean up’ system. The purified PCR products were sequenced using ABI prism 3730 sequencer (Applied Biosystems, USA) and Big dye terminator sequencing kit (ABI Prism, USA). Sequences were analyzed by BLAST tool (Altschul et al. 1990) for similar sequences in the NCBI (www.ncbi.gov) database. These sequences have been deposited in GenBank under the accession numbers KX269819-KX269826.

Additional sequences for phylogenetic analysis were downloaded from NCBI database. Eublepharis macularius (Blyth, 1854), a member of family Eublepharidae, was used as an outgroup. Gene sequences were aligned using MUSCLE (Edgar 2004). Best fit nucleotide substitution model was selected from 56 models available in PhyML (Guindon et al. 2010) using TOPALi v2 (Milne et al. 2008) based minimum Bayesian Information Criterion (BIC) value (Schwarz 1978; Nei & Kumar 2000). Best nucleotide substitution matrix was used to perform maximum likelihood analysis using PhyML (Guindon et al. 2010). Reliability of the phylogenetic tree was estimated using bootstrap values run for 1000 iterations. Phylogenetic tree was edited in FigTree v1.4.2 (Rambaut 2009). Raw (p) distances between pairs of sequences were calculated in MEGA 6 (Tamura et al. 2013).

RESULTS

Cnemaspis flaviventralis sp. nov.

(Images 1–3)

urn:lsid:zoobank.org:act:2C8F9BA1-BB95-4728-8C11-3F5CA09F4978

 

 

Holotype: BNHS 2442, 21.x.2012, adult male, India: Maharashtra: Sindhudurg District, Amboli (15.9600N & 73.9990E; 735m), coll. Amit Sayyed and Jitendra Patole.

Paratypes: BNHS 2443 (male), BNHS 2444 (female), ZSI-WRC R/1039 (male) and ZSI-WRC R/1040 (female), 9.xi.2013, same locality as holotype, collected by Amit Sayyed, Abhiji Nale, Jitendra Patole, Mangesh Karve, and Kapil Taple; 23.x. 2015, ZSI-WRC R/1042 (male), ZSI-WRC R/1041 (female) and ZSI-WRC R/1043 (female), same locality as holotype, coll. Amit Sayyed , Abhiji Nale, Jitendra Patole, Mangesh Karve, and Kapil Taple.

Diagnosis: A medium-sized Cnemaspis, maximum known SVL less than 37.0mm (30.4–36.4, n = 8), granular dorsal scales on trunk heterogeneous, intermixed with large depressed slightly keeled scales, large keeled conical tubercles present on flanks, postcloacal spurs in both sexes, ventral scales larger than dorsal scales, imbricate, smooth, without a series of enlarged median sub-caudal scales, 28–29 scales across the belly, 8–9 lamellae under first digit and 10–11 lamellae under fourth digit of manus, 8–9 lamellae under first digit and 10–12 lamellae under fourth digit of pes, males with three femoral pores on each side and no pre-cloacal pores.

Description of holotype BNHS 2442 (Image 1): Adult male, in good condition with original tail. Medium sized, SVL 31.8 mm. Head moderately short (HL/SVL = 0.20), wide (HW/HL = 0.98), fairly burly (HD/HL = 0.63), distinct from neck. Loreal region not inflated, canthus rostralis not prominent. Snout slightly long; much longer than eye diameter (OD/ES = 0.29); scales on snout, canthus rostralis and forehead region granular; interorbital and occipital region with smaller, granular scales (Image 2). Eye relatively small (OD/HL = 0.23); pupil rounded; supraciliary scales granular. Ear opening semicircular, oblique, minute (EL/HL = 0.03); eye to ear distance much greater than diameter of eyes (E-E/OD = 2.73). Rostral wider (1.2mm) than deep (0.9mm), swollen, partially divided mid-dorsally by well developed rostral groove; internasals rounded; nostrils not in contact with first supralabial; rostral in contact with first supralabial, two rows of scales separate eye from supralabial row. Mental sub-triangular, slightly wider (1.6mm) than long (1.4 mm); three pairs of postmentals, primary postmentals enlarged, each surrounded laterally by first infralabial; a single enlarged gular scale separates postmentals (Image 2b). Infralabials bordered by a row of slightly enlarged scales, decreasing in size posteriorly. Enlarged supralabials to angle of jaw 8 (right) – 9 (left) (Image 2a), to mid-orbit position 6 (on both sides); infralabials 8 (on both sides). Body relatively slender, not elongate (TRL/SVL = 0.47) without a ventrolateral fold. Granular dorsal scales on trunk heterogeneous, intermixed with large, depressed, slightly keeled scales; large keeled conical tubercles on flanks (Image 3a). Ventral scales larger than dorsal scales, smooth; 28–29 mid-body scale rows across belly (Image 1b). Three femoral pores on each side (Image 3c, 4b). Forelimbs and hind limbs relatively short, slender; forearm and tibia short (FL/SVL = 0.16; TBL /SVL = 0.19), interdigital webbing absent. Sub-digital lamellae under fourth digit of right manus, 8,9,11,11,9 (Image 3d), fourth digit of right pes, 8,9,11,11,11 (Image 3e); relative length of digits (measurements in mm): IV (3.93) > III (3.61) > V (3.14) > II (2.77) > I (2.07) (right manus); IV (3.99) > III (3.83) > V (3.76) > II (3.38) > I (1.73) (right pes) (Image 3d,e). Tail sub-cylindrical, tail longer than SVL (TL/SVL = 1.45); tail base swollen when viewed ventrally (Image 3b); post cloacal spurs present (Image 1a,b, 3b,c, 4b); dorsal scales of tail granular, with carinated enlarged tubercles, ventral scales imbricate, smooth, without a series of enlarged median sub-caudal scales (Image 3b).

Coloration: In life (Image 4a), dorsum ground color reddish-brown; dorsal and lateral tubercles, including flank tubercles, bright yellow. Transverse brown line present in interorbital area. Irregular dark brown patches scattered on head dorsally. Lateral view of head with a faded brown line from posterior border of eye to ear. Curved black marking surrounded by orange scales present on nape. Black arrowhead shape patch present on posterior side of neck. Supraciliaries brown. Pupil black with orange surrounding. Supralabials in a yellow background with faded black spots. Throat white. Ventral view of arm dust yellow; belly including thighs bright yellow (Image 4b). Mid-dorsal area of body reddish-brown, with three faded ‘W’ shaped brownish-black markings on thoracic vertebral region, between fore- and hind limbs; horseshoe shaped brownish-black markings present on lumbar vertebral region (Image 4a). Black and yellow colored spots on each digit of fore limbs and hind limbs. Tail dorsally brown, with faded yellow transverse marks; ventral tail greyish. In preservative (Image 1a, b), dorsum ground color brown, dorsal tubercles dusky white; ventral sides white with scattered black spots.

 

 

Variation

Variation in mensural and meristic data as in Table 1. Some paratypes are duller than the holotype in overall appearance.

 

 

Etymology

The species epithet is a combination name derived from the Latin words ‘flavus’ meaning yellow and ‘ventralis’ meaning ventral or pertaining to the belly, referring to the distinct yellow colouration of the ventral part of the body of the new species.

Suggested common name: Yellow-bellied Day Gecko.

 

 

Distribution

Cnemaspis flaviventralis sp. nov. is known only from its type locality (Image 5).

Natural history

Cnemaspis flaviventralis sp. nov. is a semi-arboreal species. Four specimens were collected from a tree trunk about two metres above ground level, another two found in leaf litter, and two specimens were collected from rock surfaces. All specimens were collected in October 2012 and in November 2013 between 2200 and 0200 hr (atmospheric temperature was 22–27 0C). Individuals were found to be more active between 220C and 250C at dusk. The specimens were observed in a dense forest (Image 6) in Amboli. Field observations suggest that September to November is the breeding season for the species and egg laying occurs in October and November. After mating females deposit two eggs under rocks, on boulders, or in tree holes. Co-occurring herpetofauna include: Bungarus caeruleus (Elapidae); Trimeresurus malabaricus (Viperidae); Lycodon travancoricus and Macropisthodon plumbicolor (Colubridae); Uropeltis sp. (Uropeltidae); Hemidactylus prashadi, H. cf. brookii, Geckoella albofasciatus and Cnemaspis kolhapurensis (Gekkonidae); Raorchestes ghatei, Pseudophilautus amboli and Rhacophorus malabaricus (Rhacophoridae); Indirana chiravasi (Ranixalidae); and Xanthophryne tigerina (Bufonidae).

 

 

Molecular analysis

Model selection suggested general time reversible with gamma distribution (GTR+G, lnL = -2672.85, BIC = 5548.48, G = 0.283, df = 32, n = 565) as the best fit model for nucleotide substitution. Maximum likelihood analysis (Image 7) suggested that Cnemaspis flaviventralis sp. nov. forms a monophyletic group with other species of Cnemaspis found in sympatry, with C. girii as its sister taxon. Uncorrected raw genetic distance between the two specimens of C. flaviventralis was 0.6%, whereas the distances between C. flaviventralis and other northern Western Ghats congeners was 7.5–8.1% with C. girii, 9.1–10.1% with C. goaensis and 15.4–15.7% with C. kolhapurensis.

Comparisons

Cnemaspis flaviventralis sp. nov. can be distinguished from its congeners from the northern Western Ghats on the basis of the following characters: presence of spine-like tubercles on flanks, granular dorsal scales intermixed with large, depressed, slightly keeled scales (vs. lack of spine-like tubercles on flanks, granular dorsal scales, intermixed with large smooth scales in C. girii); dorsal scales heterogeneous, 28 or 29 ventral scales across mid-body (vs. dorsal scales homogenous, 20 ventral scales across mid-body in C. indraneildasii Bauer, 2002); dorsal scales heterogeneous, lacking spine-like tubercles on flanks and lacking pre-cloacal pores (vs. dorsal scales homogenous, spine-like tubercles absent on flanks, a continuous series of pre-cloacal femoral pores in C. kolhapurensis); and pre-cloacal pores and series of enlarged median sub-caudal scales absent (vs. present in C. goaensis). From its other congeners from India, Cnemaspis flaviventralis can be distinguished based on characters such as: presence of spine-like tubercles on flanks (vs. absent in C. heteropholis Bauer, 2002, C. anaikattiensis Mukherjee, Bhupathy & Nixon, 2005, C. adii Srinivasulu, Kumar & Srinivasulu, 2015); males with femoral pores (vs. pre-anal and femoral pores absent in C. boiei (Gray, 1842), C. assamensis Das & Sengupta, 2000); dorsal scales heterogeneous (vs. dorsal scales homogenous in C. indica (Gray, 1831), C. boiei, C. mysoriensis (Jerdon, 1853), C. littoralis (Jerdon, 1854), C. jerdonii (Theobald, 1868), C. wynadensis (Beddome, 1870), C. sisparensis (Theobald, 1876), C. nilagirica Manamendra-Arachchi, Batuwita & Pethiyagoda, 2007); dorsal scales of tail granular, small keeled scales with carinated enlarged tubercles, three femoral pores on each side (vs. dorsal scales of tail granular and smooth, four or five femoral pores on each side in C. kottiyoorensis Cyriac & Umesh, 2014); pre-cloacal pores absent (vs. present in C. gracilis (Beddome, 1870), C. wicksi (Stoliczka, 1873), C. tropidogaster (Boulenger, 1885), C. andersonii (Annandale, 1905), C. otai Das & Bauer, 2000, C. yercaudensis Das & Bauer, 2000, C. australis Manamendra-Arachchi, Batuwita & Pethiyagoda, 2007, C. monticola Manamendra-Arachchi, Batuwita & Pethiyagoda, 2007, C. nilagirica); femoral pores present while pre-cloacal pores absent (vs. femoral pores absent while pre-cloacal pores present in C. ornata (Beddome, 1870), C. beddomei (Theobald, 1876), and C. nairi Inger, Marx & Koshy, 1984); series of enlarged median sub-caudal scales absent (vs. smooth enlarged median row of sub-caudals in C. indica, C. boiei, C. jerdonii, C. gracilis, C. wynadensis, C. ornata, C. sisparensis, C. nairi, C. heteropholis, C. nilagirica).

 

 

 

DISCUSSION

It is already established that the Cnemaspis species in the Western Ghats and Sri Lanka are distinct (Manamendra-Arachchi et al. 2007). Nevertheless, it is essential to note that Giri & Vithoba (2001) reported Cnemaspis kandiana (Kelaart, 1852), a species restricted to Sri Lanka (Manamendra-Arachchi et al. 2007), from Amboli, the type locality of the new species. Manamendra-Arachchi et al. (2007) have already suggested that records of C. kandiana outside Sri Lanka are based on misidentifications. Further, Mirza et al. (2014) suggested that reports of C. kandiana from southern Maharashtra should be attributed either to C. indraneildasii or are likely to belong to an undescribed species. Cnemaspis flaviventralis sp. nov. cannot be confused with either C. kandiana or C. indraneildasii. Cnemaspis flaviventralis differs from C. kandiana in having 28–29 ventral scales across mid-body (vs. 19); conical (vs. spine-like) tubercles present on flanks; pre-cloacal pores absent (vs. present); and sub-digital lamellae on digit IV of pes 10–12 (vs. 18–20). Cnemaspis flaviventralis differs from C. indraneildasii based on characters provided in the comparison section above.

As presently understood, 28 species of Cnemaspis are known from India (Uetz & Hošek 2015). The description of a new species has elevated the number of Indian species to 29. Within the genus Cnemaspis, more work remains to be done at both the species and the generic level to resolve their taxonomy. In our genetic analysis (Image 7), Cnemaspis species from northern Western Ghats formed a monophyletic clade that was well separated from Southeast Asian C. limi Das & Grismer, 2003, with a species of Hemidactylus in between. Although the current placement could be an artifact of limited genetic data for 16S rRNA, previous molecular studies (Pyron et al. 2013) with other genetic markers have also shown similar trends. More extensive molecular work and comprehensive taxonomic revisions are therefore essential to understand and to resolve the taxonomy of this group. Moreover, the species diversity within the Western Ghats/Sri Lanka biodiversity hotspot is obviously high.

Given that several species of Western Ghats Cnemaspis are globally threatened (Srinivasulu et al. 2014), understanding both the unknown diversity and assessment of distribution of previously recognized species is essential to design and implement future conservation.

 

Comparative material and data sources

Cnemaspis girii: Holotype, BNHS 2299 (male), India: Maharashtra: Satara District, Kaas Plateau (17.7240N & 73.8190E, elevation 1,203m), coll. A. Sayyed. Paratypes, BNHS 2081 (male) and BNHS 2078 (female), collection data same as holotype; India: Maharashtra: Satara District, Kaas Plateau (17.7240N & 73.8190E), coll. A. Sayyed.

Cnemaspis goaensis: Holotype, ZSI-K 22110 (male), India: Goa: “ca. 3 km S. of Forest Rest House, Canacona (Poinguinim), Goa”. Paratypes, ZSI-K 22213–22216 (4 ex.), same data as holotype; ZSI R/1044 (male) and ZSI R/1045 (female), India: Goa (15.2900N & 74.0060E), coll. A. Sayyed.

Cnemaspis gracilis: BNHS 1182 (male), Goa.

Cnemaspis indica: BNHS 1252-10 (male) and BNHS 1252-1 (female), India: Tamil Nadu: Parson Valley, Nilgiris.

Cnemaspis kolhapurensis: Holotype, BNHS 1855 (male), India: Maharashtra: Kolhapur district, Dajipur; BNHS 2447–2448 (both males), India: Maharashtra: Sindhudurg District, Amboli (15.9710N & 73.9790E), coll. A. Sayyed.

Cnemaspis littoralis: BNHS 1150 (male), India: Nilambur, Malabar.

Cnemaspis mysoriensis: Neotype, BNHS 1830 (male), India: Karnataka: Agara Village near Bangalore (designated by Giri et al. 2009).

Cnemaspis wynadensis: BNHS 1042 (male) and BNHS 1043 (male), Kerala: Mannarghat, Palghat.

Data for C. adii from Srinivasulu et al. (2015); for C. australis, C. monticola, C. nilagirica, C. beddomei, C. boiei, C. ornata, C. andersonii, C. jerdonii, C. wicksi and C. sisparensis from Manamendra-Arachchi et al. (2007); for C. indraneildasii from Bauer (2002); for C. heteropholis from Bauer (2002) and Ganesh et al. (2011); for C. otai and C. yercaudensis from Das & Bauer (2000); for C. anaikattiensis from Mukherjee et al. (2005); for C. kottiyoorensis from Cyriac & Umesh (2014); for C. assamensis from Das & Sengupta (2000); and for C. nairi and C. tropidogaster from Inger et al. (1984).

 

 

 

References

Altschul, S.F., W. Gish, W. Miller, E.W. Myers & D.J. Lipman (1990). Basic local alignment search tool. Journal of Molecular Biology 215: 403–410; http://dx.doi.org/10.1016/S0022-2836(05)80360-2

Amarasinghe, A.A.T., M.B. Harvey, A. Riyanto & E.N. Smith (2015). A new species of Cnemaspis (Reptilia: Gekkonidae) from Sumatra, Indonesia. Herpetologica 71: 160–167; http://dx.doi.org/10.1655/HERPETOLOGICA-D-14-00034

Bauer, A.M. (2002). Two new species of Cnemaspis (Reptilia: Squamata: Gekkonidae) from Gund, Uttara Kannada, India. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 99: 155–167.

Cyriac, V.P. & P.K. Umesh (2014). Description of a new ground-dwelling Cnemaspis Strauch, 1887 (Squamata: Gekkonidae), from Kerala, allied to C. wynadensis (Beddome, 1870). Russian Journal of Herpetology 21(3): 187–194.

Das, I. & A.M. Bauer (2000). Two new species of Cnemaspis (Sauria: Gekkonidae) from Tamil Nadu, southern India. Russian Journal of Herpetology 7(1): 17–28.

Das, I. & S. Sengupta (2000). A new species of Cnemaspis (Sauria: Gekkonidae) from Assam, northeastern India. Journal of South Asian Natural History 5(1): 17–23.

Edgar, R.C. (2004). MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32(5): 1792–1797; http://dx.doi.org/10.1093/nar/gkh340

Gamble, T., E. Greenbaum, T.R. Jackman, A.P. Russell & A.M. Bauer (2012). Repeated origin and loss of adhesive toepads in geckos. PLoS ONE 7(6): e39429; http://dx.doi.org/10.1371/journal.pone.0039429

Ganesh, S.R., R. Sreekar, S.P. Pal, G. Ramchandra, C. Srinivasulu & B. Srinivasulu (2011). Discovery and first description of male Cnemaspis heteropholis Bauer, 2002 (Reptilia: Gekkonidae) from Agumbe, central Western Ghats, India. Journal of Threatened Taxa 3(8): 2023–2027; http://dx.doi.org/10.11609/JoTT.o2614.2023-7

Giri, V.B., I. Agarwal & A.M. Bauer (2009). Designation of a neotype for Cnemaspis mysoriensis (Jerdon 1853) (Sauria: Gekkonidae), with a redescription and notes on its distribution and habitat. Russian Journal of Herpetology 16(4): 256–264.

Giri, V.B., A.M. Bauer & S.K. Gaikwad (2009). A new ground-dwelling species of Cnemaspis Strauch (Squamata: Gekkonidae) from the northern Western Ghats, Maharashtra, India. Zootaxa 2164: 49–60.

Giri, V. & H. Vithoba (2001) Occurrence of Cnemaspis kandiana, family Gekkonidae, in Amboli, Maharashtra. Journal of the Bombay Natural History Society 98: 454–455.

Grismer, L.L., P.J. Wood, S. Anuar, A. Riyanto, N. Ahmad, M.A. Muin, M. Sumontha, J.L. Grismer, C.K. Onn, E.S. Quah & O.S.G. Pauwels (2014). Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia. Zootaxa 3880(1): 001–147; http://dx.doi.org/10.11646/zootaxa.3880.1.1

Guindon, S., J.F. Dufayard, V. Lefort, M. Anisimova, W. Hordijk & O. Gascuel (2010). New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology 59: 307–321; http://dx.doi.org/10.1093/sysbio/syq010

Inger, R.F., H. Marx & M. Koshy (1984). An undescribed species of gekkonid lizard (Cnemaspis) from India with comments on the status of C. tropidogaster. Herpetologica 40(2): 149–154.

Manamendra-Arachchi, K., S. Batuwita & R. Pethiyagoda (2007). A taxonomic revision of the Sri Lankan day-geckos (Reptilia: Gekkonidae: Cnemaspis), with description of new species from Sri Lanka and southern India. Zeylanica 7(1): 9–122.

Milne, I., D. Lindner, M. Bayer, D. Husmeier, G. McGuire, D.F. Marshall & F. Wright (2008). TOPALi v2: a rich graphical interface for evolutionary analyses of multiple alignments on HPC clusters and multi-core desktops. Bioinformatics 25: 126–127; http://dx.doi.org/10.1093/bioinformatics/btn575

Mirza, Z.A., S. Pal, H.S. Bhosale & R.V. Sanap (2014). A new species of gecko of the genus Cnemaspis Strauch, 1887 from the Western Ghats, India. Zootaxa 3815: 494–506; http://dx.doi.org/10.11646/zootaxa.3815.4.2

Mukherjee, D., S. Bhupathy & A.M.A. Nixon (2005). A new species of day gecko (Squamata, Gekkonidae, Cnemaspis) from the Anaikatti Hills, Western Ghats, Tamil Nadu, India. Current Science 89(8): 1326–1328.

Nei, M. & S. Kumar (2000). Molecular Evolution and Phylogenetics. Oxford University Press, New York, 333pp.

Pyron, R.A., F.T. Burbrink & J.J. Wiens (2013). A phylogeny and updated classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13: 93; http://dx.doi.org/10.1186/1471-2148-13-93

Rambaut, A. (2009). FigTree, ver. 1.4.2. [Online]. Available: http:/tree.bio.ed.ac.uk/software/figtree/. Accessed on 28 February 2015.

Schwarz, G. (1978). Estimating the dimension of a model. Annals of Statistics 6: 461–464.

Srinivasulu, C., B. Srinivasulu & S. Molur (Compilers) (2014). The Status and Distribution of Reptiles in the Western Ghats, India. Conservation Assessment and Management Plan (CAMP), Wildlife Information Liaison Development Society, Coimbatore, Tamil Nadu, 148pp.

Srinivasulu, C., G.C. Kumar & B. Srinivasulu (2015). A new species of Cnemaspis (Sauria: Gekkonidae) from Northern Karnataka, India. Zootaxa 3947(1): 85–98; http://dx.doi.org/10.11646/zootaxa.3947.1.5

Strauch, A.A. (1887). Bemerkungen über die Geckoniden-Sammlung im zoologischen Museum der kaiserlichen Akademie der Wissenschaften zu St. Petersburg. Mémoires de l’Académie impériale des Sciences de St. Pétersbourg 35(2): 1–72.

Tamura, K., G. Stecher, D. Peterson, A. Filipski & S. Kumar (2013). MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution 30: 2725–2729; http://dx.doi.org/10.1093/molbev/mst197

Uetz, P. & J. Hošek (eds.) (2015). The Reptile Database, http://www.reptile-database.org, accessed 1 March 2016

Vidanapathirana, D.R., M.D. Rajeev, N. Wickramasinghe, S.S. Fernando & L.J. Wickramasinghe (2014). Cnemaspis rammalensis sp. nov., Sri Lanka’s largest day-gecko (Sauria: Gekkonidae: Cnemaspis) from Rammalakanda Man and Biosphere Reserve in southern Sri Lanka. Zootaxa 3755: 273–286; http://dx.doi.org/10.11646/zootaxa.3755.3.5