Zoosyst. Evol. 97 (1) 2021, 191-209 | DOI 10.3897/zse.97.60099 Gane Te BERLIN A new species of day gecko (Reptilia, Gekkonidae, Cnemaspis Strauch, 1887) from Sri Lanka with an updated VD2 gene phylogeny of Sri Lankan and Indian species Suranjan Karunarathna!, Anslem De Silva*, Dinesh Gabadage*, Madhava Botejue?*, Majintha Madawala®, Kanishka D.B. Ukuwela® Nature Explorations and Education Team, No: B-1 / G-6, De Soysapura Flats, Moratuwa 10400, Sri Lanka Amphibia and Reptile Research Organization of Sri Lanka, 15/1, Dolosbage Road, Gampola, Sri Lanka Biodiversity Conservation Society, 150/6, Stanley Thilakaratne Mawatha, Nugegoda 10250, Sri Lanka Central Environmental Authority, 104, Denzil Kobbekaduwa Mawatha, Battaramulla 10120, Sri Lanka Victorian Herpetological Society, P.O. box 4208, Ringwood, VIC 3134, Australia Department of Biological Sciences, Faculty of Applied Sciences, Rajarata University of Sri Lanka, Mihintale 50300, Sri Lanka Nn oO FP W DY http://zoobank. org/401B344F-6CA0-48C5-BC22-7E28B345F3CD Corresponding authors: Suranjan Karunarathna (suranjan.karu@gmail.com); Kanishka D.B. Ukuwela (kanishkauku@gmail.com) Academic editor: Rafe Brown @ Received 23 April 2020 # Accepted 24 February 2021 Published 25 March 2021 Abstract A new day gecko of the genus Cnemaspis Strauch, 1887 is described from the intermediate bioclimatic zone (Haputale Forest and Idalgashinna Forest in Badulla District) of Sri Lanka. The new species belongs to the Cnemaspis kandiana clade and was recorded from granite caves and abandoned buildings within forested areas. The region in which these habitats are located, receives relatively high annual rainfall (2500-3500 mm) and has fairly cool, moist and well-shaded conditions. The new species is medium in size (30.2—32.9 mm SVL) and can be differentiated from all other Sri Lankan Cnemaspis by the presence of small subcaudals, heteroge- nous dorsal scales, smooth pectoral and ventral scales, 7 or 8 supralabials and infralabials, 143—159 ventral scales, 15—17 belly scales, 95-103 mid-body scales, 122—132 paravertebrals, 3 pre-anal pores, 4 or 5 femoral pores and 17 or 18 lamellae on 4" toe. The species described herein is categorised as Critically Endangered (CR) under the IUCN Red List Criteria. The major threats for the new spe- cies are habitat loss due to expansion of commercial-scale agriculture and illicit forest encroachments. Therefore, we recommend relevant authorities to take immediate conservation action to ensure the protection of these forest areas in Haputale and Idalgashinna along with the buffer zone in the near future. Key Words Conservation, genetic distance, granite caves, mtDNA, montane rainforests, species delimitation, taxonomy Introduction The tropical island of Sri Lanka has a rich and diverse as- semblage of reptiles that comprises a total of 238 species, of which 155 (65%) are endemic and 107 are threatened with extinction (MoE-SL 2012; Batuwita 2016; Batuwita and Edirisinghe 2017; de Silva and Ukuwela 2020; Wick- ramasinghe et al. 2017, 2019, 2020; Karunarathna et al. 2019a, 2019b, 2020; Batuwita et al. 2020). Amongst the diverse reptile community of the Island, the diversity of geckos (Family Gekkonidae) are remarkable; 59 species have been recognised so far which accounts for 25% of the overall reptilian richness (Karunarathna et al. 2019b; Amarasinghe and Karunarathna 2020). Forty-nine (~ 83%) of them are endemic to the Island (Batuwita and Udugampola 2017; de Silva and Ukuwela 2020; Batuwi- ta et al. 2019; de Silva et al. 2019; Karunarathna et al. 2019a, 2019b, 2019c; Karunarathna and Ukuwela 2019; Copyright Suranjan Karunarathna et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Nie Pd Amarasinghe and Karunarathna 2020) and 31(~53%) are threatened with extinction (MoE-SL 2012). Though the Sri Lankan gecko fauna consists of seven genera: Cal- odactylodes, Cnemaspis, Cyrtodactylus, Gehyra, Hemi- dactylus, Hemiphyllodactylus and Lepidodactylus, none is endemic to the country (Somaweera and Somaweera 2009; de Silva and Ukuwela 2020). With 37 nominal spe- cies in Sri Lanka, Cnemaspis is considered as the most speciose reptile genus in the country, with 100% species endemism (Karunarathna et al. 2019b; Amarasinghe and Karunarathna 2020). Cnemaspis are diurnal geckos distributed in Africa and Asia comprising three distinct paraphyletic groups in Africa, Indian subcontinent and Sri Lanka and Southeast Asia (Gamble et al. 2015). During the past two decades, the number of species recognised in the genus Cnemaspis in Sri Lanka has grown rapidly with more than a nine-fold increase (from 4 to 37 species) as a result of the recent taxonomic renais- sance (Deraniyagala 1953; Bauer et al. 2007; Batuwita et al. 2019; de Silva et al. 2019; Karunarathna et al. 2019b; Amarasinghe and Karunarathna 2020). Recent molecu- lar phylogenetic analyses have indicated two distinct Sri Lankan clades of Cnemaspis, namely: C. kandiana and C. podihuna (Agarwal et al. 2017) and eight subclades (Karunarathna et al. 2019c) in the two clades; four sub- clades in the C. podihuna clade and four sub-clades in the C. kandiana clade. The use of molecular phylogenetics, detailed elucidation of morphological characters, as well as their polarity, greater access to remote locations and enhanced knowledge on geology and geography of the region have contributed to the taxonomic advances of Cnemaspis in Sri Lanka (Batuwita et al. 2019; de Silva et al. 2019). During recent field excursions to Badulla Dis- trict of Sri Lanka, a Cnemaspis species which had been previously confused with C. kandiana (Kelaart 1852) was discovered from Haputhale and Idalgashinna. Here, we describe this as a new species using a combination of morphological and molecular data. Methods Field sampling and specimens We conducted field surveys in 165 different locations dis- tributed across several bioclimatic regions (e.g. dry zone, intermediate zone and wet zone) in Sri Lanka as a part of an on-going island-wide survey of lizards under permit number WL/3/2/42/18 (a & b), issued by the Department of Wildlife Conservation and permit number R&E/RES/ NFSRCM/2019-04, issued by the Forest Department of Sri Lanka. At each location, we surveyed and documented gecko species found with special attention on the focal genus Cnemaspis. On average, per location, we spent 12 man-hours per survey. Museum acronyms follow Uetz et al. (2019). The type material discussed in this paper 1s de- posited in the National Museum of Sri Lanka (NMSL), Colombo. Specimens were caught by hand and were pho- zse.pensoft.net Karunarathna, S. et al.: A new species of Cnemaspis from Sri Lanka tographed in life. They were euthanised using halothane and fixed in 10% formaldehyde for two days, washed in water and transferred to 70% ethanol for long-term stor- age. Tail tips were collected as tissue samples before fix- ation and were stored in 95% ethanol. For comparison, we examined 458 Cnemaspis specimens (catalogued and uncatalogued) representing all recognised Sri Lankan species, including all type specimens housed at the Na- tional Museum of Sri Lanka (NMSL), The Natural His- tory Museum, London (BMNH) and specimens collected by Anslem de Silva (bearing the field codes ADS, Aaron Bauer (bearing the field codes AMB) and Suranjan Karu- narathna (bearing the field codes SSK), which have been deposited in the NMSL (Appendix 1). Specimens that for- merly belonged to the Wildlife Heritage Trust (WHT) col- lection which bears WHT numbers are currently deposit- ed at the NMSL, catalogued under their original numbers. Additional information on the morphology and nat- ural history of Sri Lankan Cnemaspis species was ex- tracted from the relevant literature (Bauer et al. 2007; Manamendra-Arachchi et al. 2007; Wickramasinghe and Munindradasa 2007; Vidanapathirana et al. 2014; Wick- ramasinghe et al. 2016; Agarwal et al. 2017; Batuwita and Udugampala 2017; Batuwita et al. 2019; de Silva et al. 2019; Karunarathna et al. 2019a; Karunarathna et al. 2019b; Karunarathna et al. 2019c; Karunarathna and Ukuwela 2019; Amarasinghe and Karunarathna 2020). Assignment of unidentified specimens to the new species was based on their morphometric, meristic and molecu- lar characters, colour patterns and the level of geographic isolation. The new species described in the present paper has been included in previous phylogenies of the genus as Cnemaspis sp. 5 (NMSL AA87 and AA87B collect- ed from Haputhale, Sri Lanka) in Agarwal et al. (2017) and Cnemaspis sp. 4 in Karunarathna et al. (2019c). In this paper, we initially refer to this species as Cnemaspis sp. 5 following Agarwal et al. (2017). The tissue voucher (bearing the Field number SK5) was sampled from one of the paratypes collected from Idalgashinna, Sri Lanka. DNA-based species delimitation To determine the genetic distinction of the new species to already-known species of Cnemaspis, we examined the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene. ND2 gene is commonly used as a barcode marker for geckos and the majority of DNA sequences availa- ble on GenBank for Sri Lankan and Indian Cnemaspis species are of this gene. Additionally, we included two Cnemaspis (C. rammalensis [n = 2] and C. rajakarunai [n = 3]) species that have not been included in previous phylogenies. Genetic distinction was determined through examining the haplotype clusters through phylogenetic analysis (Wiens and Penkrot 2002), uncorrected pairwise genetic distances and species delimitation analyses. Whole genomic DNA was isolated from the tissue samples using a Qiagen DNeasy blood and tissue DNA Zoosyst. Evol. 97 (1) 2021, 191-209 isolation kit (Valencia, CA, USA) following the manufac- ture’s protocols. The quality of the isolated DNA was de- termined through gel electrophoresis in ethidium bromide stained 1% Agarose gel. The concentration of the isolated DNA samples was quantified using a Nabi Nano-spec- trophotometer (MicroDigital Company Ltd, Korea). We PCR amplified a 1040 bp fragment of the ND2 gene us- ing already-published primers L4437a, AAGCTTTCG- GGCCCATACC and H5934, AGRGTGCCAATGTCT- TTGTGRITT (Macey et al. 1997). The PCR was carried out in 25 wl reactions with a primer concentration of 0.4 uM for each primer employing 35 cycles with an annealing temperature of 50 °C (Macey et al. 1997) following stand- ard PCR protocols with Promega PCR master mix (Prome- ga Corporation, Madison, Wisconsin, USA). The success of the PCR amplification and size of the amplified frag- ment was checked through gel electrophoresis in ethidium bromide stained 1% Agarose gel using a Promega 100 bp ladder (Promega Corporation, Madison, Wisconsin, USA). The PCR products of the successfully amplified samples were purified and sequenced in both directions at the Ge- netech Sri Lanka Pvt. Ltd., Colombo, Sri Lanka. Consensus sequences from forward and reverse reads were assembled in Geneious v.5.6 software (Drummond et al. 2009). We downloaded all the available ND2 se- quences for Cnemaspis species of the South Asian ra- diation (Appendix 1). We did not include the Southeast Asian Cnemaspis as they are known to be a separate unre- lated lineage from the South Asian Cnemaspis. However, C. modiglianii, C. tanintharyi and C. thayawthadangyi (Agarwal et al. 2017; Lee et al. 2019) are known to be nested within the South Asian Cnemaspis radiation and are closely related to each other (Lee et al. 2019) and thus C. modiglianii has been included in the phylogenetic analyses. The total dataset included 104 taxa comprising 27 of the 37 Cnemaspis species known from Sri Lanka, four putative species from Sri Lanka, 17 Cnemaspis spe- cies from India and one species from Southeast Asia. Cal- odactylodes illingworthorum was used as the outgroup since it has been shown to be the sister lineage of the South Asian Cnemaspis radiation (Agarwal et al. 2017). DNA sequences were aligned using Geneious alignment (Drummond et al. 2009) in Geneious v.5.6 software using default settings and refined manually. The sequences were translated to amino acid sequences using the vertebrate mitochondrial genetic code to check for premature stop codons that might indicate amplification of pseudogenes and to determine the correct reading frame. The mitochondrial ND2 gene tree was reconstructed using Bayesian and Maximum Likelihood (ML) meth- ods. Partitioning schemes and best-fit substitution models for each partition were assessed using the Bayesian Infor- mation Criterion (BIC) implemented in Partitionfinder 2 (Lanfear et al. 2017). BIC indicated three partitions based on the three codon positions with GTR+I+G substitution model for each partition. Partitioned ML analysis was 1m- plemented in RAXML 7.2.6. (Stamatakis et al. 2008) with 200 independent ML searches using the rapid hill-climb- 193 ing algorithm. Branch support was estimated using 1000 bootstrap pseudoreplicates. Partitioned Bayesian analysis was performed in MrBayes 3.2.6 (Ronquist and Huelsen- beck 2003) with unlinked model parameters using default priors for 80 million generations with two independent runs and four chains (one hot and three cold chains) sampling every 10000 generations. Convergence of the independent runs was assessed by examining split fre- quencies (< 0.01) of clades across runs, effective sample sizes (ESS values) and likelihood plots in Tracer v.1.4.1 (Rambaut et al. 2018). An all-compatible consensus tree was built after first 25% of sampled trees were discarded as burn-in. Uncorrected pairwise distances (p-distances) between species were calculated in MEGA X with an av- erage site cut-off of 95% (Kumar et al. 2018). Species delimitation analysis using Poisson Tree Pro- cess (PTP) (Zhang et al. 2013) was conducted using the rooted Bayesian tree as input tree (ML and Bayesian). The calculations were performed on the PTP web server (http://species.h-its.org/ptp/), with 200,000 MCMC gen- erations, thinning set to 100 and burn-in set at 25% and performing a Bayesian search. The probability of each node to represent a species node was calculated in both Bayesian and Maximum Likelihood methods. Morphometric characters Forty morphometric measurements were taken using a Mitutoyo digital Vernier calliper (to the nearest 0.1 mm) and detailed observations of scales and other structures were made through Leica Wild M3Z and Leica EZ4 dis- secting microscopes. The following symmetrical meristic characters were taken on the left side of the body: eye diameter (ED), horizontal diameter of eye ball; orbital diameter (OD), the greatest diameter of orbit; eye to nos- tril length (EN), the distance between anteriormost point of the orbit and the posterior border of the nostril; snout length (ES), the distance between anteriormost point of the orbit and the tip of snout; snout to nostril length (SN), the distance between tip of snout and the anteriormost point of the nostril; nostril width (NW), the maximum horizontal width of the nostrils; eye to ear distance (EE), the distance between the posterior border of eye and the anteriormost point of ear opening; snout to axilla distance (SA), the distance between axilla and tip of snout; ear length (EL), the maximum length of the ear opening; in- terorbital width (IO), the shortest distance between the left and right supraciliary scale rows; inter-ear distance (IE) the distance across the head between the two ear openings; head length (HL), the distance between pos- terior edge of mandible and the tip of the snout; head width (HW), the maximum width of the head in-between the ears and the orbits; head depth (HD), the maximum height of the head at the level of the eye; jaw length (JL), the distance between the tip of snout and the corner of the mouth; internarial distance (IN), the smallest distance between the inner margins of nostrils; snout to ear dis- zse.pensoft.net 194 tance (SED), the distance between the tip of snout and anteriormost point of the ear; upper-arm length (UAL), the distance between the axilla and the angle of the el- bow; lower-arm length (LAL), the distance from the el- bow to the wrist with palm flexed; palm length (PAL), the distance between the wrist (carpus) and the tip of longest finger excluding the claw; length of digits I-V of manus (DLM), the distance between the juncture of the basal phalanx with the adjacent digit and the tip of the digit, excluding the claw; snout-vent length (SVL), the distance between tip of snout and the anterior margin of vent; trunk length (TRL), the distance between the axil- la and the groin; trunk width (TW), the maximum width of body; trunk depth (TD), the maximum depth of body; femur length (FEL), the distance between the groin and the knee; tibia length (TBL), the distance from the knee to the heel with ankle dorsiflexed; heel length (HEL), the distance between ankle (tarsus) and the tip of longest toe (excluding the claw) with both foot and tibia flexed; length of pedal digits I-V (DLP), the distance between the juncture of the basal phalanx with the adjacent digit and the digit tip, excluding the claw; tail length (TAL), the distance between the anterior margin of the vent and the tail tip; tail base depth (TBD), the maximum height of the tail base; tail base width (TBW), the widest point of the tail base. Meristic characters Thirty discrete characters were observed and recorded using Leica Wild M3Z and Leica EZ4 dissecting micro- scopes on both the left (L) and the right (R) side of the body (reported in the form L/R): number of supralabials (SUP) and infralabials (INF) between the first labial scale and the corner of the mouth; number of interorbital scales (INOS) between the left and right supraciliary scale rows; number of postmentals (PM) bounded by chin scales, 1* infralabial on the left and right and the mental; number of chin scales (CHS) touching medial edge of infralabials and mental between juncture of 1* and 2" infralabials on the left and right; number of supranasal (SUN) scales be- tween nares; presence of the postnasal (PON) scales pos- terior to the naris; presence of the internasal (INT) scale between supranasals; number of supraciliary scales (SUS) above the eye; number of scales between the eye and tym- panum (BET) from posterior-most point of the orbit to anterior-most point of the tympanum; number of canthal scales (CAS), number of scales from posterior-most point of naris to anteriormost point of the orbit; total lamellae on manus I-V (TLM) counted from first proximal enlarged scansor, greater than twice width of the largest palm scale, to distalmost lamella at tip of digits; number of dorsal para- vertebral granules (PG) between pelvic and pectoral limb insertion points along a straight line immediately left of the vertebral column; number of mid-body scales (MBS) from the centre of mid-dorsal row diagonally towards the ventral scales; number of mid-ventral scales (MVS) from zse.pensoft.net Karunarathna, S. et al.: A new species of Cnemaspis from Sri Lanka the first scale posterior to the mental to last scale anterior to the vent; number of belly scales (BLS) across the ventre between the lowest rows of granular dorsal scales; total lamellae on pes I-V (TLP), counted from first proximal enlarged scansor greater than twice the width of the largest heel scale, to distalmost lamella at tip of digits; number of precloacal pores (PCP) anterior to the cloaca; number of femoral pores (FP) present on the femur; numbers of non-pored proximal femoral scales (PFS) counted from proximal ends of femoral pore rows to precloacal pores; numbers of non-pored distal femoral scales (DFS) counted from distal ends of femoral pore rows to knee. In addi- tion, we also evaluated the texture [keeled (KD) or smooth (SM)] of the ventral scales, the texture [heterogeneous (HET) or homogeneous (HOM)] of the dorsal scales, the number of spinous scales on the flanks (FLSP) and char- acteristics, such as appearance of the caudal scales (except in specimens with regenerated tails). Colouration was de- termined from digital images of living specimens and also from direct observations in the field. Distribution and natural history During the surveys, behavioural and other aspects of nat- ural history of the focal species were observed through opportunistic and non-systematic means. The ambient temperature and the substrate temperature were mea- sured using a standard thermometer and a N19 Q1370 infrared thermometer (Dick Smith Electronics, Shang- hai, China), respectively. The relative humidity and light intensity were measured with a QM 1594 multifunction environment meter (Digitek Instruments Co., Ltd., Hong Kong, China). To record elevation and georeference spe- cies locations, an eTrex 10 GPS (Garmin) was used. Sex was determined by the presence of hemipenial bulges, precloacal and femoral pores in males (M) or absence of the above in females (F). The conservation status of the species was evaluated using IUCN Red List Categories and Criteria version 14 (IUCN 2019). Results DNA-based species delimitation Both ML (Supplementary Fig. 1) and Bayesian analyses recovered highly similar topologies and branch lengths and hence only the Bayesian tree is shown (Fig. 1). Cnemaspis sp. 5 was recovered in the C. kandiana clade (Agarwal et al. 2017) and was sister to a clade compris- ing Cnemaspis sp. 3, Cnemaspis sp. 4, C. pulchra and C. butewai (BPP = 0.98, BS = 70). All three sequenc- es of the new species were monophyletic (BPP = 1.0, BS = 100) (Fig. 1). The average uncorrected pair-wise genetic distance between Cnemaspis sp. 5 and other taxa in the C. po- dihuna clade was 26.3% (range 23.9-28.3%), while it Zoosyst. Evol. 97 (1) 2021, 191-209 195 ‘oy U0 + vu ma] — U Cnemaspis sp. 11 AMB7507 SSK22.2 SSK22.0 SSk22.1 - AA81 C. podihuna SSK17.0 clade SSK25.0 WHT268 SSK027 SSK029 SSKO006 SSkK004 SSK005 KY038006 JX041328 58A 70A WHT7334 Cnemaspis alwisi ; Cnemaspis kohukumburai Cnemaspis punctata | Cnemaspis sp. 9 KY038009 KY038010 SSK3.0 WHT591 SSK3.1 SSK3.2 MK792498 MK792499 MK792496 MK792497 MK792492 MK792493 a Cnemaspis giril MK792489 MK792490 MK792490 MK792486 MK792487 MK792488 MK792475 MK792478 MK792472 3 | MK792473 Cnemaspis thackerayi MK792465 MK792470 MK792468 MK792469 MK792466 MK792467 7 Cnemaspis cf. gracilis Cnemaspis sp. 7 Chemashs sp. 6 KY03/977 é KY037978 . j nemaspis sp. C. kandiana clade Cnemaspis samanalensis KY037980 KY037981 KY037979 SSK24.1 KY037986 KY037988 KY037987 SSK24.0 . Cnemaspis kallima. ; Cnemaspis retigalensis KY037973 = SSK18.0 KY037971 KY037972 Cnemaspis ingerorum Cnemaspis silvula SSK16.0 SSK16.1 : : : Cnemaspis gotaimbarai KY037975 a SSK15.1 KY037974 SSK15.0 SSK15.2 SSK2.0 SSK2.1 Cnemaspis latha SSK4.0 SSK4.1 SSK4.2 SK5 B AA87 AA87B Cnemaspis sp. 3 SSK9.1 SSK9.0 SSK9.20 Cnemaspis sp. 4 SSK1.2 SSK1.0 SSK1.1 0.09 Figure 1. Bayesian all compatible ND2 gene tree of South Asian Cnemaspis lineage. Dark circles depict nodes with Bayesian poste- rior probability > 0.95 and Bootstrap support > 70. The outgroup Calodactylodes illingworthorum is not shown. Scale bar indicates the number of substitutions per site. Colours of the branches indicate the geographical origin of the taxa where green, blue and brown depict Sri Lankan, Indian and Southeast Asian taxa, respectively. Results of molecular species delimitation analyses (bPTP and PTP) are shown in grey bars on the right. zse.pensoft.net 196 Karunarathna, S. et al.: A new species of Cnemaspis from Sri Lanka Table 1. Uncorrected pairwise genetic distances in the VD2 gene between the Cnemaspis sp.5 (Cnemaspis sp. 4 in Karunarathna et al. 2019c) and the members of the Cnemaspis kandiana clade, the clade to which Cnemaspis sp. 5 species belongs. 2 3 4 5 6 7 8 1. C. butewai - 2. C. kumarasinghei 9.5 3. C. gotaimbarai Ts 2 ae 855 - 4. C. ingerorum S08 Fil ww BO : 5. C. kallima 74 66 88 54 - 6. C. kandiana 82 7.0 89 62 46 : 7. C. kohukumburai 744 8:0 “9:99 6:3 -b.l” 528 : 8. C. latha Sib Bai “Sikes Tee ST e245 9. C. modiglianii 1006" OS 1 FeeBsl= pind Fao Oe. Sel 10. C. nandimithrai 86 65 76 59 55 61 #62 6.7 11. C. pava Bok” WT 223. -6:3¢ £5754 59. © 633. = 6.1 12. C. pulchra Bree = SO) 13. C. retigalensis CAB ened. 14. C.samanalensis 9.8 8.7 15. C. silvula 0.4.8 SO. - 9:0" 153d, "6.02 6th? -b7-. AS 16. Cnemaspissp.1 82 7.0 94 64 45 63 6.1 6.7 IA} CHEMASDISSD:. 35 5i8.- “QS 18. Cnemaspis sp.4 5.1 9.1 19. Cnemaspis sp.5 8.3 8.6 20. C. upendrai Si 2 eh et Oe eee tle EOS sb OSG 9 LO S00 2 1S alder Lhe er 71S 19 8.4 7.6 C8 ie oP 7 FS (65) 67 - 1.5 of 735 64 : 1025" Bio fesel e238) sotly Gism corn SZ - FOS 16 3 GG Pile aBiS> Gide G6: 3.6 - 9.0" 6:6 8:25-6:9 -7.5.°°9:27 483° 6.6 e7.1- 74 = Tid tl On Oud fT SOB De oF 2d” “Tie Che dS Bold numbers indicate uncorrected pairwise genetic distance between Cnemaspis sp. 5 and other members of the C. kandiana clade. was 20.0% (range 22.1—34.3%) between Cnemaspis sp. 5 and Indian Cnemaspis species exclusive of the mem- bers of the C. kandiana clade. The average uncorrected pair-wise genetic distance between Cnemaspis sp. 5 and other taxa in the C. kandiana clade was 7.8% (range 6.6— 10.4%) (Table 1) which was greater than the mean (5.6%, range 3.7—8.5%) uncorrected pair-wise genetic distanc- es between sister species pairs in the C. kandiana clade. Cnemaspis nandimithrai shares the lowest uncorrected pair-wise genetic distance (6.6%) with Cnemaspis sp. 5. Species delimitation analyses using PTP implementing the Maximum Likelihood (PTP) approach indicated the presence of two species within Cnemaspis sp. 5, but with low support (ML support: 0.29). However, Bayesian ap- proach (bPTP) indicated the presence of a single species within Cnemaspis sp. 5 (Bayesian support: 0.71). Systematics Cnemaspis lokugei sp. nov. http://zoobank.org/91469423-CEBA-4BB2-BAF0-386C9572A588 Figs 2-3; Tables 2-3 Lokuge’s day gecko (English) Lokugege diva-seri hoona (Sinhala) Cnemaspis sp. 5 Agarwal et al. 2017 Cnemaspis sp. 4 Karunarathna et al. 2019c Holotype. NMSL.2021.01.01, adult male, 32.9 mm SVL, collected from a granite cave bordering a stream, Haputale, Badulla District, Uva Province, Sri Lanka (6.7753°N; 80.9667°E, WGS1984; elevation 1510 m; around 10:00 hrs) on 18 December 2019 by Suranjan Karunarathna. zse.pensoft.net Paratypes. NMSL.2021.01.02, adult female, 30.8 mm SVL and NMSL.2021.01.03, adult female, 30.2 mm SVL, collected from a granite cave bordering a stream, Idalgashinna, Badulla District, Uva Province, Sri Lanka (6.7791°N; 80.8967°E, WGS1984; elevation 1565 m; around 09:00 hrs) on 19 December 2019 by Suranjan Karunarathna. Diagnosis. Cnemaspis lokugei sp. nov., can be readily distinguished from its Sri Lankan congeners by a com- bination of the following morphological and meristic characteristics: maximum SVL 32.9 mm; dorsum scala- tion heterogeneous, mixed with smooth and keeled large granular scales; 1/1 supranasals, | internasal, 1/1 postna- sal; 3 enlarged postmentals; postmentals bounded by 5 enlarged chin scales; chin, gular, pectoral and abdominal scales smooth, subimbricate; 15—17 belly scales across mid-body; 5 or 6 feebly-developed tubercles on posterior flank; 122-132 paravertebral granules linearly arranged; 3 precloacal pores, 4 or 5 femoral pores in males, sepa- rated by 8 or 9 proximal femoral scales lacking pores, 7 or 8 distal femoral scales lacking pores; 143-159 ven- tral scales; 95-103 mid-body scales; smooth subcaudals, median row comprising an irregular series of diamond shaped, small scales; 7 or 8 supralabials; 7 or 8 infralabi- als; 15 or 16 total lamellae on fourth digit of manus and 17 or 18 total lamellae on fourth digit of pes. Description of holotype. An adult male, 32.9 mm SVL and 36.9 mm TAL (regenerated). Body slender, relatively short (TRL/SVL ratio 39.9%). Head rela- tively small (AL/SVL ratio 27.7% and HL/TRL ratio 69.5%), narrow (HW/SVL ratio 14.2% and HW/HL ra- tio 51.2%), depressed (HD/SVL ratio 10.1% and HD/ HL ratio 36.4%) and distinct from neck. Snout relatively long (ES/HW ratio 75.5% and ES/HL ratio 38.6%), less than twice the eye diameter (ED/ES ratio 53.4%), more Zoosyst. Evol. 97 (1) 2021, 191-209 Loy: Eh ae SEM 1 ee POE. Figure 2. Cnemaspis lokugei sp. nov. male holotype (NMSL.2021.01.01), (a) dorsal head, (b) lateral head, (c) ventral head, (d) heter- ogeneous scales on dorsal surface of trunk, (e) lateral surface of trunk, (f) smooth ventral scales, (g) cloacal characters with precloacal and femoral pores (h) subdigital lamellae on manus, (i) subdigital lamellae on pes; female paratype (NMSL.2021.01.03), (j) dorsal side of tail, (kK) lateral side of tail, (1) subrhomboid-shaped small subcaudals. Scale bar: 1 mm (Photos: Suranjan Karunarathna). than half length of jaw (ES/JL ratio 64.8%), snout slight- (OD/EE ratio 125.8%) and equal to length of digit IV ly concave in lateral view; eye relatively small (ED/HL — of manus (OD/DLM IV ratio 100%); supraocular ridges ratio 20.6%), larger than the ear (EL/ED ratio 43.6%), | moderately developed; ear opening small (EL/HL ratio pupil round; orbit length greater than eye to ear distance 9.0%), deep, taller than wide, larger than nostrils; two zse.pensoft.net 198 Karunarathna, S. et al.: A new species of Cnemaspis from Sri Lanka at ane 28 et 2 hee, 2 . en ae - é ? Figure 3. Cnemaspis lokugei sp. nov. male holotype (NMSL.2021.01.01) in life in-situ (a) dorsal view of the full body with typical colour pattern, (b) ventral aspect showing gular and ventral colouration, (b) lateral aspect with spines on flank and labial colouration from Haputale (Photos: Suranjan Karunarathna). rows of scales separate orbit from supralabials; inter- Dorsal surface of the trunk with smooth scales inter- orbital distance is a little shorter than snout length IO/ mixed with keeled heterogeneous granules, 132 paraver- ES ratio 97.7%), shorter than head length (IO/HL ratio — tebral granules; 148 smooth, mid-ventral scales; 95 mid- 37.8%); eye to nostril distance greater than the eye to _ body scales; 6/5 weakly-developed tubercles on the flanks; ear distance (EN/EE ratio 102.8%). ventrolateral scales small, irregular; granules on snout zse.pensoft.net Zoosyst. Evol. 97 (1) 2021, 191-209 oval, keeled and raised, larger than those on interorbital and occipital regions; canthus rostralis nearly absent, 13/13 smooth oval scales from eye to nostril; scales of the inter- orbital region circular and keeled; short tubercles present both on the sides of the neck and around the ear; ear open- ing vertically oval, slanting from anterodorsal to postero- ventral, 21/20 scales between anterior margin of the ear opening and the posterior margin of the eye. Supralabials 7/7, infralabials 8/7, becoming smaller towards the posteri- or end of the mouth. Rostral scale wider than long, partially divided (80%) by a median groove and in contact with first supralabial. Nostrils separated by 1/1 enlarged supranasals with 1 internasal; few enlarged scales behind the suprana- sals. Nostrils oval, dorsolaterally orientated, not in contact with first supralabials; 1/1 postnasals, smooth, larger than nostrils, partially in contact with first supralabial (Fig. 2). Mental, sub-rhomboid in shape, as wide as long, pos- teriorly in contact with 3 enlarged postmentals (small- er than mental and lager than chin scales); postmentals in contact and bordered posteriorly by 5 unkeeled chin scales (larger than nostrils), in contact with the 1* infral- abial; ventral scales smaller than chin scales. Smooth, rounded, juxtaposed scales on the chin and the gular re- gion; pectoral and abdominal scales smooth, subimbri- cate towards precloacal region, abdominal scales slightly larger than dorsals; 17 belly scales across ventre; smooth scales around vent and base of tail, subimbricate; 3 pre- cloacal pores; 4/5 femoral pores; 8/9 proximal femoral scales lacking pores on each side; 7/8 enlarged distal femoral scales. Regenerated tail little longer than the snout-vent length (TAL/SVL ratio 112.2%); hemipenial bulge moderately swollen (TBW 2.8 mm), heterogeneous scales on the dorsal aspect of the tail directed backwards, spine-like tubercles present at the base of tail: tail with 3 or 4 enlarged flattened obtuse scales forming whorls; a large, blunt post-cloacal spur on each side, dorsoventrally flattened and narrow; subcaudals smooth and small, sub- rhomboidal, arranged in a single median series (Fig. 2). Forelimbs very short, slender (LAL/SVL ratio 11.6% and UAL/SVL ratio 13.1%); hind limbs long, tibia little longer than the femur (TBL/SVL ratio 16.7% and FEL/ SVL ratio 15.8%). Scales on anterior, upper, posterior and ventral surfaces of upper arm with keeled granules and less imbricate scales, scales of the anterior surface twice as large as those of the other surfaces; anterior, upper, pos- terior and ventral surfaces of lower arm with keeled and less imbricate scales, scales of the upper surface twice as large as those of the other surfaces. Scales on anterior, upper, posterior surfaces of femur keeled, ventral surface with smooth, subimbricate scales, scales on the ventral surface twice the size of those of other aspects; anterior, upper, posterior surfaces of tibia keeled, ventral surface with smooth, imbricate scales, scales on the upper sur- face twice the size of those of other aspects. Dorsal and ventral surfaces of manus and pes with keeled granules; dorsal surfaces of digits with granular scales (Fig. 2H, I). Digits elongate and slender with inflected distal phalan- ges, all bearing slightly recurved claws (Fig. 2). Subdigi- 199 tal lamellae entire (except divided at first interphalangeal joint), unnotched; total lamellae on manus (left/right): digit I (10/11), digit I (13/14), digit III (16/16), digit IV (16/15), digit V (15/15); total lamellae on pes (left/right): digit I (11/10), digit If (15/15), digit III (17/18), digit IV (18/18), digit V (17/17); interdigital webbing absent; rel- ative length of left manual digits: I (1.4 mm), II (1.8 mm), V (2.3 mm), II (2.5 mm), IV (3.1 mm); relative length of left pedal digits: I (1.5 mm), II (2.9 mm), III (3.4 mm), V (3.5 mm), IV (3.9 mm). Variation in the type series (Tables 2 and 3). The SVL of adult specimens in the type series of Cnemaspis lokugei sp. nov. (n = 3) ranges from 30.2 to 32.9 mm; in- terorbital scales 24—26; supraciliaries above the eye 14— 16; supralabials 7 or 8, infralabials 7 or 8; scales from eye to tympanum 19-21; canthal scales 12 or 13; tubercles on posterior flank 5 or 6; ventral scales 143-159 (Tables 2-3); mid-body scales 95-103; paravertebral granules 122-132; belly scales across ventre 15—17; femoral pores in males 4 or 5; proximal femoral scales lacking pores in males 8 or 9; distal femoral scales lacking pores in males 7 or 8; total lamellae on digit of the manus: digit I (10 or 11), digit H (13 or 14), digit II (15 or 16), digit 'V (15 or 16), digit V (14 or 15); total lamellae on digit of the pes: digit I (10 or 11), digit II (14 or 15), digit III (17 or 18), digit IV (17 or 18), digit V (16 or 17). Colour in life. Dorsum of head, body and limbs gen- erally reddish-brown; yellow spot with black outer edge on neck dorsally; broken faded, yellow vertebral stripe running from occiput to tail (Fig. 3); seven irregular blackish-brown, chevron shaped paravertebral markings present. Tail dark brown dorsally, with 10 faded brown irregular cross-bands; pupil circular and black with the surrounding iris yellow; two very faint postorbital stripes on each side; supralabials and infralabials yellowish with tiny black spots; chin and gular scales yellow, with dark spots; pectoral, abdominal, cloacal and subcaudal scales are cream and intermixed with irregular stippling; dor- sum of limbs with faded black markings; manus and pes with alternating black and cream-white cross bands. Colour in preservation. Dorsally grey brown with seven distinct dark, irregular blotches; pale spot with dark outer edge on neck dorsally; supralabials and infralabi- als dirty white; two dark postorbital stripes on each side; chin and gular scales grey; ventral surface uniformly dirty white colour with some scales on thigh, tail base and arms with dark brown margins. Etymology. The specific epithet is an eponym Latinised (/okugei) in the masculine genitive singular, honouring Mr. Ajith Nethkelum Lokuge, a pioneer ecolo- gist, analogue forestry specialist and a senior member of Young Zoologist’s Association of Sri Lanka, for his sig- nificant contribution towards environmental conservation and research in Sri Lanka. Distribution and natural history. The specimens of the type series were collected from the two locations, Haputale and Idalgashinna (Badulla District, Uva Prov- ince), which are situated in the central highlands of Sri zse.pensoft.net 200 Karunarathna, S. et al.: A new species of Cnemaspis from Sri Lanka Table 2. Morphometric measurements (mm) of holotype from Haputale and two paratypes from Idalgashinna of Cnemaspis lokugei co 99 sp. nov. from Badulla District, Sri Lanka (“—” = not applicable). Measurement NMSL.2021.01.01 Holotype (M) NMSL.2021.01.02 Paratype (F) Snout-vent length 32:9 30.8 Trunk length ce al le Trunk width 5.2 5.4 Trunk depth 3.6 3.4 Head length 9.1 8.9 Head width 4.7 4.6 Head depth x 2.8 Snout to axilla distance 15:9 14.3 Jaw length 5.4 Sto) Tail length 0:9: 35,1 Tail base width 2.8 2.4 Tail base depth 2 papal Eye diameter 1.9 1.7 Orbital diameter Sak 2.9 Eye to nostril length 2.6 2.4 Snout length 345 forall Snout to nostril length is: 12 Nostril width 0.2 O12 Eye to ear distance 2:5 2.4 Ear length 0.8 0.7 Interorbital width 3.4 Sud Inter-ear distance 3.6 5.5 Internarial distance Le" ilps: Snout to ear distance 8.3 8.2 Upper-arm length 4.3 4.3 Lower-arm length on a Palm length iia 29 Digits length of manus (i) 1.4 1.5 Digits length of manus (ii) bes: 1.8 Digits length of manus (iil) 25 2.4 Digits length of manus (iv) Sul 259 Digits length of manus (v) 23 ie Femur length Be oral Tibia length othe: 5.4 Heel length 4.1 oe Digits length of pes (i) 1e5 le3 Digits length of pes (ii) 2,9 caw Digits length of pes (iii) 3.4 5 Digits length of pes (iv) SS, 3.8 Digits length of pes (v) Foie) 3.4 Lanka (central intermediate bioclimatic zone [annual rainfall between 2000-2500 mm (Burt and Weerasinghe 2014)]) (Fig. 4). Tropical sub-montane and montane for- ests make up the dominant vegetation type (Gunatileke and Gunatileke 1990) of this area. The forest acreage in both areas is approximately 1200 ha and is relatively isolated from other forests due to anthropogenic habi- tats and tea plantations. It is very likely that the species occurs in the intervening regions between these two lo- cations as there are similar habitats scattered between the two locations. However, this needs to be verified through a thorough field survey. These locations lie be- tween an elevation of 1400 and 1700 m a.s.l. (Fig. 4). The mean annual rainfall is received mainly during the southwest monsoon (May—September), while the mean annual temperature 1s 26.1—28.9 °C. Both areas are rich in granite rock boulders with 40 identified caves. Cne- maspis lokugei sp. nov. appears to be a common species in the two locations as we recorded more than 50 indi- zse.pensoft.net NMSL.2021.01.03 Range Mean+SD Paratype (F) 30%? 30.2-32.9 31.341.4 12.4 12.3-13.1 12.6+0.4 523 5.2-5.4 5.34001 3.4 3.4-3.6 syo0-1 S9 8.9-9.1 9,020.1 4.4 4.4-4.7 4.6+0.1 2.8 2.8-3.3 SAzg3 14.4 14.3-15.9 14.9+0.9 BS 5.3-5.4 5.4+0.1 34.4 34.4-36.9 Glove mad (Po. 2.4 2.4-2.8 2.6+0.2 Pe. 2.2-2.5 2st, 2 Ls 1.7-1.9 17+0.1 2.9 2.9-3.1 3.0+0.1 2.5 2.4-2.6 2,5£071 Sel 3.1-3.5 3:340:2 1,2 1.2-1.3 1220.1 O22 - 0.2+0.0 ea 2.3-2.5 2.4+0.1 0.7 0.7-0.8 0.8+0.1 3.4 3.3-3.4 3.4+0.1 Pid 3.5-3.6 3.5+0.0 1.4 1.3-1.5 1.4+0.1 8.2 8.2-8.3 82401 4.2 4.2-4.3 4.3+0.1 3:6 3.6-3.8 S.A EE I Sul 2.9-3.2 SALEQ}1 1.5 1.4-1.5 1.5+0.1 1,9 1.8-1.9 1.8+0.1 223 2.3-2.5 2.4+0.1 3.1 2.9-3.1 320#0,4 2.1 2.1-2.3 2.2+0.1 era 5.1-5.2 523031 5:3 5.3-5.5 5.4+0.1 ee) 3.9-4.1 4.0+0.1 1.4 1.3-1.5 1.4+0.1 2.6 2.6-2.9 2.7+0.1 3:2 3.2-3.5 3.4+0.1 3.8 3.8-3.9 3.8+0.0 S03 3.3-3.5 3.4+0.1 viduals from both locations during a two-day survey. This species was observed in granite caves and rela- tively old buildings on vertical surfaces, about 2 m from ground within the forested area (Fig. 5). The granitic cave microhabitat of C. /okugei sp. nov. was poorly illu- minated (light intensity: 392-476 Lux), relatively moist (relative humidity: 76-92%), well shaded (canopy cov- er: 62-78%) and relatively cool (ambient temperature: 29.5—31.2 °C and substrate temperature: 27.4—28.7 °C). The new species is sympatric with several other gecko species: Cyrtodactylus sp., Gehyra mutilata, Hemidac- tylus frenatus, H. parvimaculatus and Hemiphyllodac- tylus typus. Pure white and almost spherical shaped (mean diameter 4.9 + 0.02 mm [n = 34]) eggs with a slightly flattened side attached to a rocky substrate were observed in cave habitats where Cnemaspis lokugei sp. nov. was observed. Since these eggs were charac- teristic of Cnemaspis species and as there were no other Cnemaspis species observed in these habitats, it was Zoosyst. Evol. 97 (1) 2021, 191-209 201 Table 3. Meristic data of holotype from Haputale and two paratypes from Idalgashinna of Cnemaspis lokugei sp. nov. from Badulla District, Sri Lanka (“—” = not applicable). Character NMSL.2021.01.01 NMSL.2021.01.02 NMSL.2021.01.03 Range Holotype (M) Paratype (F) Paratype (F) Supralabials (L, R) Vibea! 7,8 7,8 7-8 Infralabial (L, R) 8,7 Vv Teg: 7-8 Lateral spines (L, R) 6,5 5,5 56 5-6 Interorbital scales 26 24 25 24-26 Postmentals 3 3 3 - Chin scales 5 5 5 - Supranasal (L, R) 1,1 alzal 1,1 - Postnasal (L, R) ile, Je alee - Internasal 1 1 1 - Supraciliary (L, R) 15, 14 15,15 14,16 14-16 Eye to tympanum scales (L, R) 21°20 20, 20 Ls 2 19-21 Canthal scales (L, R) 13,13 Perl? V2, le. 12-13 Total lamellae on manus (i) (L, R) 10, 11 10, 10 10, 10 10-11 Total lamellae on manus (ii) (L, R) 13, 14 14, 14 13, 14 13-14 Total lamellae on manus (ili) (L, R) 16, 16 15, 16 15,15 15-16 Total lamellae on manus (iv) (L, R) 16,15 L565." 15 Los 1S 15-16 Total lamellae on manus (v) (L, R) L515 14, 14 15,15 14-15 Paravertebral granules 132 128 122 122-132 Mid-body scales 95 98 103 95-103 Mid-ventral scales 148 159 143 143-159 Belly scales Ly 17 1.5 15-17 Total lamellae on pes (i) (L, R) 1110 Ta ey 10, 10 10-11 Total lamellae on pes (ii) (L, R) 5a 14,15 15,15 14-15 Total lamellae on pes (iil) (L, R) 17,18 18, 18 T¥,.18 17-18 Total lamellae on pes (iv) (L, R) 18, 18 17,17 IE Wi 17-18 Total lamellae on pes (v) (L, R) LAY? bey 16,17 16-17 Precloacal pores 6) absent absent - Femoral pores (L, R) 45 absent absent 4-5 Proximal femoral scales (L, R) 8,9 absent absent 8-9 Distal femoral scales (L, R) 7,8 absent absent 7-8 presumed that the eggs most likely belong to C. lokugei Sp. nov. Conservation status. Application of the IUCN Red List Criteria indicates that C. /okugei sp. nov. is Critically Endangered (CR) due to its having an area of occupancy (AOO) < 10 km?(3.84 km? in total assuming a 100 m radius around the seven georeferenced locations), severely frag- mented habitat and a projected decline in the area, extent and the quality of habitat [Applicable criteria B2ab (111)]. Comparisons with other Sri Lankan species. Mor- phologically, Cnemaspis lokugei sp. nov. most closely resembles C. butewai, C. ingerorum, C. kivulegedarai, C. kallima, C. kandiana, C. kotagamai, C. menikay and C. retigalensis because of the presence of a dorsum with heterogeneous scales and smooth belly scales (see the species comparison and Table 4 for more details). Amongst species of the C. kandiana clade sensu Agar- wal et al. (2017), Cnemaspis lokugei sp. nov. differs by having heterogeneous (vs. homogeneous) dorsal scales from C. amith, C. dissanayakai, C. gotaimbarai, C. kaw- miniae, C. kumarasinghei, C. latha and C. nandimithrai. It can also be distinguished from C. butewai, C. kandi- ana, C. menikay, C. pava, C. pulchra, C. retigalensis, C. samanalensis, C. silvula, C. tropidogaster and C. upend- rai by having smooth (vs. keeled) gular scales. The new species differs from C. ingerorum and C. kivulegedarai by having more ventral scales (143-159 vs. 88-95 and 109-114) and by having more mid-body scales (95—103 vs. 62-69 and 69-76); from C. kallima by having more mid-body scales (95—103 vs. 67—74) and by having more paravertebral granules (122-132 vs. 99-107); from C. kotagamai by having more mid-body scales (95-103 vs. 79-84) and by fewer scales across belly (15-17 vs. 21-22). Amongst species of the C. podihuna clade sensu Agar- wal et al. (2017), Cnemaspis lokugei sp. nov. differs by the absence of clearly enlarged, hexagonal or subhexag- onal subcaudal scales from the following species with homogeneous dorsal scales: C. alwisi, C. anslemi, C. gemunu, C. godagedarai, C. hitihamii, C. kandambyi, C. kohukumburai, C. molligodai, C. nilgala, C. phillip- si, C. podihuna, C. punctata, C. rajakarunai, C. ram- malensis and C. scalpensis. The new species also differs from C. alwisi, C. anslemi, C. gemunu, C. godagedarai, C. hitihamii, C. kohukumburai, C. nilgala, C. phillipsi, C. punctata, C. rajakarunai, C. rammalensis and C. scal- pensis by having precloacal pores (vs. absence). Discussion Our present morphological and molecular analyses and previous studies (Agarwal et al. 2017; Karunarathna et al. 2019c) strongly indicate the presence of a novel species of Cnemaspis in Sri Lanka, adding yet another species to the growing list of Cnemaspis in Sri Lanka and increasing the total number of species to 38. These Cnemaspis species are adapted for a scansorial and crepuscular mode of life, with most being rupicolous, while a few are arboreal or ground-dwelling (Das 2005; Karunarathna et al. 2019b). Sri Lankan representatives of the genus are microhabi- tat specialists with narrow niches limited to moist, cool, zse.pensoft.net 202 Karunarathna, S. et al.: A new species of Cnemaspis from Sri Lanka Contour 600 m Climate zone boundary Dry zone Intermediate zone Wet zone N 0 25 50 Kilometers WGS 1984 UTM Zone 44N Figure 4. Currently known distribution of Cnemaspis lokugei sp. nov. (holotype— red star, paratype— red circle) and its closely-re- lated species (C. butewai — white circl, and C. pulchra — yellow circle) in Sri Lanka. canopy-shaded rock outcrops, granite caves, trees, aban- doned buildings, buildings associated with caves, wattle and daub houses and semi-naturalised rock walls, where their cryptic morphology and body colouration camou- flage them in the environment (Smith 1935; Karunarath- na et al. 2019c). Further, Cnemaspis species prefer nar- row (~ 34 mm), long (~ 100-400 mm) and deep (~ 20-180 mm) crevices as refugia and oviposition sites (Karunarathna et al. 2019b). Likewise, the new species is also exclusively recorded from vertical surfaces about 1 to 2 m from ground in poorly illuminated, relatively moist, well shaded and relatively-cool granite caves or old buildings within forested areas (see Fig. 5B). When threatened, they readily escape to narrow crevices. These observations indicate the requirement of cool and damp zse.pensoft.net environments for the survival of these geckos signifying the narrow ecological niches they occupy. This could be one of the key drivers of speciation in these geckos where narrow ecological niches most likely have been an isolating mechanism. However, most importantly, this may also highlight the fact that these species are at a very high risk of extinction, if such habitats are destroyed. Phylogenetic analyses of the ND2 gene placed the novel species in the C. kandiana clade (Agarwal et al. 2017) as expected given its strong morphological resemblance to other members of the clade. The new species was sis- ter to a clade comprising Cnemaspis sp. 3, Cnemaspis sp. 4, C. pulchra and C. butawai. The taxonomic status of Cnemaspis sp. 3 and Cnemaspis sp. 4 needs to be fur- ther investigated. Two additional species of Cnemaspis, Zoosyst. Evol. 97 (1) 2021, 191-209 203 A Figure 5. General habitat of Cnemaspis lokugei sp. nov. in the Uva Province of Sri Lanka (a) wet forest highland with short trees, viewed from Idalgashinna, (b) a granite cave habitat in Idalgashinna (c) communal egg clutches in Haputale. (Photos: Suranjan Karunarathna). Table 4. Comparison of morphological and morphometric characters of C. /okugei sp. nov. with the other congenors of the C. kan- diana clade in Sri Lanka. > o a < A 3 te 7) @ @ ee § fg 28 8 2 £ 2 & 5 i Species Se — $8 Ss 8 2 & = un 3 = ry G s 88 & £5 8 see se. © “se 5 Sage 8 se Ff FF ae ae = YH Pu oH 2 = = ng = 7 © Fa < © ° r= Yo o ° < in s £ a UG S = o € ee = a 7) =o o iz oa & Se Sa C. pava 32.4 HET KD KD Sml 7-8 6-7 139-145 22-25 64-75 83-98 9-11 24 45 16-17 18-19 C. pulchra 34.2 HET KD KD Sml 7-8 7-8 120-135 24-27 67-73 94-103 57 34 46 15-17 17-20 C. samanalensis 37.5 HET KD KD Smil 810 89 128-144 19-20 61-67 64-72 5-6 3-4 3-5 16-17 18-20 C. silvula 28.6 HET KD KD Sml 7-8 7-8 132-139 19-21 73-81 102-113 10-15 34 45 15-16 18-19 C. tropidogaster 31.7 HET KD KD Sml 7-8 7-8 132-146 21-25 92-98 99-106 5-7 34 45 16-17 18-19 C. upendrai 35.2 HET KD KD Sml 7-8 7-8 112-128 16-25 69-74 97-102 13-15 2-3 45 17-18 17-21 C. ingerorum 26.9 HET SM SM Sml 7-8 7-8 8895 17-21 62-69 93-101 7-8 2-3 45 13-16 17-18 C. kivulegedarai 31.2 HET SM SM Sml 7-8 6-7 109-114 17-19 69-76 131-133 45 23 45 13-15 14-16 C. kallima 35.1 HET SM SM Sml 7-8 7-8 131-138 19-23 67-74 99-107 12-15 34 45 16-18 18-20 C. kotagamai 29.8 HET SM SM Sml 7-8 7-8 131-137 21-22 79-84 114-119 6-7 il 4-5 13-15 17-18 C. lokugeisp. nov. 32.9 HET SM SM Sml 7-8 7-8 143-159 15-17 95-103 122-132 5-6 3 4-5 15-16 17-18 C. butewai 31.8 HET SM SM SmI 7-8 7-8 125-128 23-25 92-98 134-138 56 35 45 15-17 17-18 C. kandiana 34.6 HET SM SM Sml 89 7-8 119-138 19-20 68-75 86-99 5-7 24 34 12-14 18-20 C. menikay 28.0 HET SM SM Sml 7-9 7-8 124-138 20-26 71-79 83-98 13-15 1-2 34 14-15 15-17 C. retigalensis 30.8 HET SM SM Sml 7-8 7-8 121-128 16-20 69-77 82-86 45 1 3-4 14-15 16-20 C. amith 33.0 HOM SM SM Sml 7-8 7-8 123-131 19-21 67-74 79-84 4-5 S 3 16-17 18-19 C. dissanayakai 29.4 HOM SM SM Sml 7-8 7-8 118-120 17-19 94-98 105-107 6-7 2-3 45 21-22 21-22 C. gotaimbarai 33.7 HOM SM SM Sml 7-8 89 129-138 23-25 72-79 117-121 56 24 3-4 16-17 19-20 C. kawminiae 35.6 HOM SM SM Sml 7-8 7-8 107-114 17-21 76-78 86-92 7-8 23 34 1415 15-16 C. kumarasinghei 31.6 HOM SM SM Sml 7-8 7-8 120-134 17-21 87-94 61-68 7-9 23 3-5 15-16 16-18 C. latha 30.4 HOM SM SM Sml 7-8 7-8 109-115 13-15 69-73 72-79 5-7 23 45 15-17 17-18 C. nandimithrai 31.7 HOM SM SM Sml 5-6 6-7 108-112 25-27 87-89 95-99 34 2-4 2-4 12-13 19-20 Abbreviations: HET — Heterogenous, HOM — Homogenous, KD — Keeled, SM — Smooth, Sml — Small. Characters that can be used to diagnose C. /okugei sp. nov. from other Cnemaspis species in Sri Lanka are shown in bold text. zse.pensoft.net 204 C. rammalensis and C. rajakarunai, which were placed in phylogenetic analyses for the first time, were recov- ered in the C. podihuna clade (Agarwal et al. 2017). This is again expected because of their strong morphological similarity to other members of the clade characterised by the presence of enlarged hexagonal/subhexagonal subcaudal scales. The two species were recovered to be sister taxa forming a unique lineage in the C. podi- huna clade (Fig. 1) indicating speciation in the isolated mountains (Vidanapathirana et al. 2014; Wickramasing- he et al. 2016) in the wet zone of Sri Lanka. These findings further reinforce the importance of isolated mountains for the speciation of Sri Lankan day geckos. Cnemaspis lokugei sp. nov. was discovered from the in- termediate bioclimatic zone (see Fig. 4). Our studies il- lustrate that Cnemaspis are distributed throughout all bio- climatic zones of the Island; however, the majority, Le. 23 species (~ 60%) are recorded from the wet bioclimatic zone which thus coincides with the notion that the Island’s wet bioclimatic zone is home to high species richness and endemism (MoE-SL 2012). Further, the discovery of this new species from Haputale and Idalgashinna (1400— 1700 m a.s.l.) suggests that the occurrence of Cnemaspis genus in high elevations is also considerable making this the fifth species to be described from elevations above 1000 ma. s. |. (Fig. 4) in Sri Lanka. Since 2015, we have been conducting an island-wide survey on Cnemaspis and sampled over 165 locations using visual encounter sur- veys. Our on-going studies, based on morphological and molecular analyses, have thus far discovered ~16 new species of which 14 species have been described (Bote- jue et. al. 2019; de Silva et al. 2019; Karunarathna et. al. 2019a, 2019b, 2019c; Karunarathna and Ukuwela 2019; Amarasinghe and Karunarathna 2020). In addition to this, our on-going studies indicate that there are at least anoth- er 10 new species, potentially increasing the Cnemaspis count to more than 50 species in Sri Lanka, resulting in the highest density of Cnemaspis species per land area. More field surveys in mountainous areas and detailed studies may yield promising results in the understanding of taxonomy and biogeography of this genus. We are certain that the species that we have described here is novel and has not been previously described due to the following reasons. According to Manamendra-Arach- chi et al. (2007), Gymnodactylus malabarica Jerdon, 1853 (= Cnemaspis malabaricus) described from the forests of Malabar [Kerala State] is a valid species restricted to In- dia. Although Kluge (2001) listed Cnemaspis malabarica (Jerdon, 1853) in the synonymy of C. kandiana, accord- ing to Jerdon (1853), C. malabarica (type locality Kerala State, southern India) has homogeneous dorsal scalation. However, C. kandiana has heterogenous dorsal scalation and has a very restricted range in Sri Lanka (Manamen- dra-Arachchi et al. 2007). Therefore, we consider these two species to be distinct. Similarly, C. /okugei sp. nov., has heterogeneous dorsal scalation, while C. malabarica has homogeneous dorsal scalation. Additionally, given that C. lokugei sp. nov., is restricted to a narrow range within zse.pensoft.net Karunarathna, S. et al.: A new species of Cnemaspis from Sri Lanka Sri Lanka and that C. malabarica is a species restricted to India, we believe the name C. malabarica 1s not applicable to C. lokugei sp. nov. Due to the presence of smooth ven- trals in C. Jokugei sp. nov., (vs. keeled ventrals in C. tropi- dogaster) and many other differences (see comparison for details), the name C. tropidogaster is also not applicable to the new species described here. Gymnodactylus humei is a Species without enlarged hexagonal scales on the tail (thus a member of the C. kandiana clade), which was de- scribed from Kandy by Theobald (Theobald 1876). This species has been synonymised with C. kandiana now and C. kandiana is restricted to the Kandyan Region. Due to the fact that C. kandiana and C. lokugei sp. nov., are mor- phologically and genetically distinct and allopatric, we be- lieve that Gymnodactylus humei (= Cnemaspis humezi) is also unavailable for Cnemaspis lokugei sp. nov. The only Cnemaspis species already known from the Region is C. latha, which was described from Bandarawela, which is about 10 km from Haputhale. However, this species is dis- tinctly different from a suit of morphological characters (see Table 4) from C. /okugei sp. nov. and is also genet- ically distinct (see Figure 1). We therefore conclude that none of the available names or species in synonymy with C. kandiana is closely related, geographically proximate or relevant to the new species described here. Most of the Sri Lankan Cnemaspis are point-endemics with distribution ranges limited to < 10 km? (i.e. AOO < 10 km?, EOO < 100 km?) and the new species described here corresponds with this general pattern, which has led to categorising most species as critically endangered. This restricted distribution could be a consequence of the narrow ecological niche leading to the limitation of favourable microhabitats. The known localities of the new species, Haputale and Idalgashinna are mountainous forested areas with granite caves. Although these locali- ties are somewhat isolated from human habitations, they are susceptible to some degree of human-induced habitat degradation, including clearing and timber felling, forest fragmentation, granite mining, tea and vegetable cultiva- tion and invasive species. Most Cnemaspis species, like Cnemaspis lokugei sp. nov. described here are restricted to forests in mountains (Fig. 5a). Therefore, the conser- vation of such forests and other mountainous habitats are imperative to ensure the future survival of these species. Acknowledgements We thank to the Director General, the Research Director, the research committee and the field staff of the Depart- ment of Wildlife Conservation of Sri Lanka for grant- ing research permits (WL/3/2/42/18a, b) and assisting us during the field surveys. The Additional Conservator and field staff of the Department of Forest Conservation are acknowledged for issuing research permits (R&E/ RES/NFSRCM/2019-04) and the support provided during the field surveys. Further, we are grateful to Nan- da Wickramasinghe, Sanuja Kasthuriarachchi, Lankani Zoosyst. Evol. 97 (1) 2021, 191-209 Somaratne, Chandrika Munasinghe, Tharushi Gamage, Rasika Dasanayake, Thushari Dasanayake, Ravindra Wickramanayake and Pannilage Gunasiri at NMSL for assisting us during examining collections under their care. Thilina Surasinghe, Thasun Amarasinghe, Nirma- la Perera, Sulakshana Wickramarachchi, Hasantha Wi- jethunga, Angelo Hettige, Nimantha Aberathna, Mendis Wickramasinghe, Tharaka Kusuminda, Buddhika Mad- urapperuma, Sanjaya Bandara and Niranjan Karunarath- na provided valuable assistance in numerous stages of the study. Dimanthi Jayatilake and Danesha Nanayak- kara are thanked for providing laboratory facilities and assistance with laboratory work. Finally, we would like to thank Mark Scherz and three anonymous reviewers and the editor, Rafe Brown for constructive comments that immensely helped to improve the manuscript. References Agarwal I, Biswas S, Bauer AM, Greenbaum E, Jackman TR, de Silva A, Batuwita S (2017) Cryptic species, taxonomic inflation, or a bit of both? New species phenomenon in Sri Lanka as suggested by a phylogeny of dwarf geckos (Reptilia, Squamata, Gekkonidae, Cne- maspis). 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Evol. 97 (1) 2021, 191-209 Appendix 1 Comparative material examined from Sri Lanka Museum acronyms BMNH__ The Natural History Museum, London, UK NMSL National Museum of Sri Lanka, Colombo, Sri Lanka DWC Museum of the Department of Wildlife Con- servation, Giritale, Sri Lanka WHT Collection of the Wildlife Heritage Trust, Sri Lanka (Now at the NMSL) Cnemaspis alwisi. NMSL 2004.09.01 (holotype), NMSL 2004.09.02 (paratype), NMSL 2004.09.03 (paratype), WHT 5918, WHT 6518, WHT 6519, WHT 7336, WHT 7337, WHT 7338, WHT 7343, WHT 7344, WHT 7345, WHT 7346. C. anslemi. NMSL 2019.14.01 (holotype), NMSL 2019.14.02 (paratype), NMSL 2019.14.03 (paratype). C. amith. BMNH 63.3.19.1066A (holotype), BMNH 63.3.19.1066B (paratype), BMNH 63.3.19.1066C (paratype). C. butewai. NMSL 2019.07.01 (holotype), NMSL 2019.07.02 (paratype), NMSL 2019.07.03 (paratype). C. dissanayakai NMSL 2019.20.01 (holotype), NMSL 2019.20.02 (paratype), NMSL 2019.20.03 (paratype). C. gemunu: AMB 7495 (holotype), AMB 7507 (para- type??), WHT 7221, WHT 7347, WHT 7348, NMSL 2006.11.01, NMSL 2006.11.02, NMSL 2006.11.03, NMSL 2006.11.04. C. godagedarai. NMSL 2019.09.01 (holotype), NMSL 2019.16.01 (paratype), NMSL 2019.16.02 (paratype). C. gotaimbarai: NMSL 2019.04.01 (holotype), NMSL 2019.04.02 (paratype), NMSL 2019.04.03 (paratype). C. hitihamii: NMSL 2019.06.01 (holotype), NMSL 2019.06.02 (paratype), NMSL 2019.06.03 (paratype). C. ingerorum: WHT 7332 (holotype), WHT 7330 (para- type) WHT 7331 (paratype). C. kallima: WHT 7245 (holotype), WHT 7222 (para- type), WHT 7227 (paratype), WHT 7228 (paratype), WHT 7229 (paratype), WHT 7230(paratype), WHT 7239 (paratype), WHT 7249 (paratype), WHT 7251 (paratype), WHT 7252 (paratype), WHT 7253 (para- type), WHT 7254 (paratype), WHT 7255 (paratype). C. kandambyi. WHT 9466 (holotype), WHT 9467 (para- type). C. kandiana. BMNH_ 53.4.1.1 (lectotype), BMNH 80.2.2.119A (paralectotype), BMNH 80.2.2.119B (paralectotype), BMNH 80.2.2.119C (paralectotype), WHT 7212, WHT 7213, WHT 7267, WHT 7305, WHT 7307, WHT 7308, WHT 7310, WHT 7313, WHT 7319, WHT 7322. C. kawminiae NMSL 2019.18.01 (holotype), NMSL 2019.18.02 (paratype), NMSL 2019.18.03 (paratype). C. kivulegedarai: NMSL 2019.08.01 (holotype), NMSL 2019.08.02 (paratype), NMSL 2019.08.03 (paratype). 207 C. kohukumburai: NMSL 2019.05.01 (holotype), NMSL 2019.05.02 (paratype), NMSL 2019.05.03 (paratype). C. kotagamai NMSL 2019.15.01 (holotype), NMSL 2019.15.02 (paratype), NMSL 2019.15.03 (paratype). C. kumarasinghei: NMSL 2006.13.01 (holotype), NMSL 2006.13.02 (paratype). C. latha: WHT 7214 (holotype). C. manoae. NMSL 2019.10.01 (holotype), NMSL 2006.10.02 (paratype), NMSL 2006.10.03 (paratype). C. menikay: WHT 7219 (holotype), WHT 7218 (para- type), WHT 7349 (paratype). C. molligodai: NMSL 2006.14.01 (holotype), NMSL 2006.14.02 (paratype), NMSL 2006.14.03 (paratype), NMSL 2006.14.04 (paratype), NMSL 2006.14.05 (paratype). C. nandimithrai: NMSL 2019.01.01 (holotype), NMSL 2019.01.02 (paratype), NMSL 2019.01.03 (paratype). C. nilgala. NMSL_ 2018.07.01 (holotype), NMSL 2018.06.01 (paratype), NMSL 2018.06.02 (paratype), NMSL 2018.06.03 (paratype). C. pava: WHT 7286 (holotype), WHT 7281 (para- type), WHT 7282 (paratype), WHT 7283 (para- type), WHT 7285 (paratype), WHT 7288 (paratype), WHT 7289 (paratype), WHT 7290 (paratype), WHT 7291 (paratype), WHT 7292 (paratype), WHT 7293 (paratype), WHT 7294 (paratype), WHT 7295 (para- type), WHT 7296 (paratype), WHT 7297 (paratype), WHT 7298 (paratype), WHT 7299 (paratype), WHT 7300 (paratype), WHT 7301 (paratype), WHT 7302 (paratype). C. phillipsi. WHT 7248 (holotype), WHT 7236 (para- type); WHT 7237 (paratype); WHT 7238 (paratype). C. podihuna: BMNH 1946.8.1.20 (holotype), NMSL 2006.10.02, NMSL 2006.10.03, NMSL 2006.10.04. C. pulchra. WHT 7023 (holotype), WHT 1573a (para- type), WHT 7011 (paratype), WHT 7021 (paratype), WHT 7022 (paratype). C. punctata: WHT 7256 (holotype), WHT 7223 (para- type), WHT 7226 (paratype), WHT 7243 (paratype), WHT 7244 (paratype). C. rajakarunai: NMSL 2016.07.01 (holotype), DWC 2016.05.01 (paratype), DWC 2016.05.02 (paratype). C. rammalensis: NMSL 2013.25.01 (holotype), DWC 2013.05.001. C. retigalensis. NMSL 2006.12.01 (holotype), NMSL 2006.12.02 (paratype), NMSL 2006.12.03 (paratype), NMSL 2006.12.04 (paratype). C. samanalensis: NMSL 2006.15.01 (holotype), NMSL 2006.15.02 (paratype), NMSL 2006.15.03 (paratype), NMSL 2006.15.04 (paratype), NMSL 2006.15.05 (paratype). C. scalpensis: NMSL 2004.01.01 (neotype), NMSL 2004.02.01, NMSL 2004.03.01, NMSL 2004.04.01, WHT 7265, WHT 7268, WHT 7269, WHT 7274, WHT 7275, WHT 7276, WHT 7320. zse.pensoft.net 208 C. silvula: WHT 7208 (holotype), WHT 7206 (paratype), WHT 7207 (paratype), WHT 7209 (paratype), WHT 7210 (paratype), WHT 7216 (paratype), WHT 7217 (paratype), WHT 7018, WHT 7027, WHT 7202, WHT 7203, WHT 7220, WHT 7354, WHT 7333. C. tropidogater. BMNH 71.12.14.49 (lectotype), NMSL 5152, NMSL 5151, NMSL 5159, NMSL 5157, NMSL 5970, NMSL 5974. Karunarathna, S. et al.: A new species of Cnemaspis from Sri Lanka C. upendrai: WHT 7189 (holotype), WHT 7184 (para- type), WHT 7187 (paratype), WHT 7188 (paratype), WHT 7181 (paratype), WHT 7182 (paratype), WHT 7183 (paratype), WHT 7185 (paratype), WHT 7190 (paratype), WHT 7191 (paratype), WHT 7192 (para- type), WHT 7193 (paratype), WHT 7194 (paratype), WHT 7195 (paratype), WHT 7196 (paratype), WHT 7197 (paratype), WHT 7260 (paratype). Specimens, voucher numbers and GenBank accession numbers of the taxa used for the DNA-based species delimitation in this Appendix 2 study. Species Field No./ Museum Voucher No. Calodactylodes illingworthorum AMB7415 Cnemaspis agarwali AK107 AK108 Cnemaspis ajijae AK429 AK432 Cnemaspis alwisi AMB7447 Cnemaspis butewai Cnemaspis ctf. flaviventralis Cnemaspis cf. gracilis Cnemaspis cf. kumarasinghei Cnemaspis cf. mahabali Cnemaspis cf. gemunu Cnemaspis cf. podihuna Cnemaspis lokugei sp. nov. Cnemaspis gemunu Cnemaspis girii Cnemaspis goaensis Cnemaspis gotaimbarai Cnemaspis gracilis Cnemaspis hitihamii Cnemaspis ingerorum Cnemaspis kallima Cnemaspis kandiana Cnemaspis kivulegedarai Cnemaspis kohukumburai Cnemaspis kumarasinghei Cnemaspis latha Cnemaspis limayei zse.pensoft.net SSK1.0/ NMSL.2019.07.01 SSK1.1/ NMSL.2019.07.02 SSK1.2/ NMSL.2019.07.03 AK518 AK517 AK213 AA13/NMSL AK398 AK389 AMB7507/NMSL AMB7449/NMSL AA87/ NMSL AA87B/ NMSL SK5/NMSL.2021.01.02 SSK22.0/ ADS217/ NMSL SSK22.1/ ADS216/ NMSL SSK22.2/ ADS218/ NMSL AK439 VG385 VG399 SSK6.0/NMSL.2019.04.01 CES G385 AK135 SSK3.0/ NMSL.2019.06.01 SSK3.1/ NMSL.2019.06.02 WHT591/ NMSL SSK3.2/ NMSL.2019.06.03 WHT7331 AA82/ NMSL AA57/ NMSL AMB7487/ NMSL AAO1/ NMSL SSK18.0/ NMSL SSK4.0/ NMSL.2019.08.01 SSK4.1/ NMSL.2019.08.02 SSK4.2/ NMSL.2019.08.03 SSK23.0/ NMSL.2019.05.01 AMB7431/ NMSL SSK15.0/ NMSL SSK15.1/ NMSL SSK15.2/ NMSL SSK2.0/ NMSL SSK2.1/ NMSL AA13/ NMSL WHT214/ NMSL AK DAJ DV40.5 GenBank Accession No. Source JX041318 Gamble et al. 2012 MK792466 Khandekar et al. 2019 MK792467 Khandekar et al. 2019 MK792498 Khandekar et al. 2019 MK792499 Khandekar et al. 2019 KY037997 Agarwal et al. 2017 MK562351 Karunarathna et al. 2019c MK562352 Karunarathna et al. 2019c MK562353 Karunarathna et al. 2019c MK792496 Khandekar et al. 2019 MK792497 Khandekar et al. 2019 MK792464 Khandekar et al. 2019 KY037975 Agarwal et al. 2017 MK792492 Khandekar et al. 2019 MK792493 Khandekar et al. 2019 KYO38000 Agarwal et al. 2017 KYO38006 Agarwal et al. 2017 KY037993 Agarwal et al. 2017 KY037994 Agarwal et al. 2017 MW594290 This study MK562340 Karunarathna et al. 2019c MK562341 Karunarathna et al. 2019c MK562342 Karunarathna et al. 2019c MK792491 Khandekar et al. 2019 MK792475 Khandekar et al. 2019 MK792478 Khandekar et al. 2019 MK562364 Karunarathna et al. 2019c MK792465 Khandekar et al. 2019 MK792470 Khandekar et al. 2019 MK562337 Karunarathna et al. 2019c MK562338 Karunarathna et al. 2019c KY038012 Agarwal et al. 2017 MK562339 Karunarathna et al. 2019c KY037990 Agarwal et al. 2017 KY037970 Agarwal et al. 2017 KY037971 Agarwal et al. 2017 KYOS7972 Agarwal et al. 2017 KY037973 Agarwal et al. 2017 MK562347 Karunarathna et al. 2019c MK562348 Karunarathna et al. 2019c MK562349 Karunarathna et al. 2019c MK562350 Karunarathna et al. 2019c MK562336 Karunarathna et al. 2019c KY037974 Agarwal et al. 2017 MK562358 Karunarathna et al. 2019c MK562359 Karunarathna et al. 2019c MK562357 Karunarathna et al. 2019c MK562360 Karunarathna et al. 2019c MK562361 Karunarathna et al. 2019c KYOS7975 Agarwal et al. 2017 KY037976 Agarwal et al. 2017 MK792485 Khandekar et al. 2019 MK792486 Khandekar et al. 2019 Zoosyst. Evol. 97 (1) 2021, 191-209 Species Cnemaspis modiglianii Cnemaspis mysoriensis Cnemaspis nandimithrai Cnemaspis nilgala Cnemaspis pava Cnemaspis phillipsi Cnemaspis podihuna Cnemaspis pulchra Cnemaspis rajakarunai Cnemaspis rammalensis Cnemaspis retigalensis Cnemaspis samanalensis Cnemaspis scalpensis Cnemaspis shevaroyensis Cnemaspis silvula Cnemaspis sp. 1 (Sri Lanka) Cnemaspis sp. 3 (Sri Lanka) Cnemaspis sp. 4 (Sri Lanka) Cnemaspis sp. 6 (India) Cnemaspis sp. 7 (India) Cnemaspis sp. 9 (Sri Lanka) Cnemaspis sp. 11 (India) Cnemaspis sp. (India) Cnemaspis sp. (India) Cnemaspis sp. (India) Cnemaspis sp. (India) Cnemaspis thackerayi Cnemaspis upendrai Cnemaspis yercaudensis Field No./ Museum Voucher No. MVZ239314 MVZ239315 NA SSK16.0/ NMSL.2019.03.01 SSK16.1/ NMSL.2019.03.02 AMB7418/ NMSL AMB7436/ NMSL WHT7261/ NMSL AMB7494/ NMSL AA19/ NMSL SSK24.1/ ADS214/ NMSL AA81/ NMSL SSK17.0/ ADS220/ NMSL 70A/ NMSL AMB7449/ NMSL 58A/ NMSL WHT 7334/ NMSL SSK9.0/ ADS205 SSK9.1/ ADS206 SSK9.2/ ADS207 AA80/ NMSL SSKOO4 SSKO05 SSKO06 SSKO27 SSKO029 AMB7448/ NMSL AMB7505/ NMSL SSK25.0/ ADS219 WHT7268/ NMSL AK204 AK205 AA88/ NMSL AA17/ NMSL AMB7508/ NMSL AMB7529/ NMSL SBO048 JB2Z39 47A/ NMSL SB151 G349 VG407 VG408 AK470 CES G143 AA83 AA12 AMB7488 SSK24.0/ ADS213 AK280 G133 GenBank Accession No. KY037977 KY037978 MK792474 MK562362 MK562363 KY038009 KY038010 KYOS4979 KY037980 KY037981 MK562346 KY038001 MK562343 KY038002 JX041328 KY038005 KY038004 MK562354 MK562355 MK562356 KY038007 MW594285 MW594286 MW594287 MW594288 MW594289 KY037982 KY037983 MK562344 KY038008 MK792468 MK792469 KY037984 KY037989 KY037991 KY037992 KYO37995 KY037996 KY038011 KY038013 MK792490 MK792487 MK792488 MK792489 MK792471 KY037986 KY037987 KY037988 MK562345 MK792472 MK792473 209 Source Agarwal et al. 2017 Agarwal et al. 2017 Khandekar et al. 2019 Karunarathna et al. 2019c Karunarathna et al. 2019c Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Karunarathna et al. 2019c Agarwal et al. 2017 Karunarathna et al. 2019c Agarwal et al. 2017 Gamble et al. 2012 Agarwal et al. 2017 Agarwal et al. 2017 Karunarathna et al. 2019c Karunarathna et al. 2019c Karunarathna et al. 2019c Agarwal et al. 2017 This study This study This study This study This study Agarwal et al. 2017 Agarwal et al. 2017 Karunarathna et al. 2019c Agarwal et al. 2017 Khandekar et al. 2019 Khandekar et al. 2019 Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Khandekar et al. 2019 Khandekar et al. 2019 Khandekar et al. 2019 Khandekar et al. 2019 Khandekar et al. 2019 Agarwal et al. 2017 Agarwal et al. 2017 Agarwal et al. 2017 Karunarathna et al. 2019c Khandekar et al. 2019 Khandekar et al. 2019 zse.pensoft.net