A new species of day gecko (Reptilia, Gekkonidae, Cnemaspis Strauch, 1887) from Sri Lanka with an updated ND2 gene phylogeny of Sri Lankan and Indian species

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, heterogenous 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 4th toe. The species described herein is categorised as Critically Endangered (CR) under the IUCN Red List Criteria. The major threats for the new species 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.

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 renaissance (Deraniyagala 1953;Bauer et al. 2007;Batuwita et al. 2019;de Silva et al. 2019;Karunarathna et al. 2019b;Amarasinghe and Karunarathna 2020). Recent molecular 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 subclades 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;. During recent field excursions to Badulla District 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.

Field sampling and specimens
We conducted field surveys in 165 different locations distributed 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 is deposited in the National Museum of Sri Lanka (NMSL), Colombo. Specimens were caught by hand and were pho-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 storage. Tail tips were collected as tissue samples before fixation 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 National Museum of Sri Lanka (NMSL), The Natural History Museum, London (BMNH) and specimens collected by Anslem de Silva (bearing the field codes ADS, Aaron Bauer (bearing the field codes AMB) and Suranjan Karunarathna (bearing the field codes SSK), which have been deposited in the NMSL (Appendix 1). Specimens that formerly belonged to the Wildlife Heritage Trust (WHT) collection which bears WHT numbers are currently deposited at the NMSL, catalogued under their original numbers.
Additional information on the morphology and natural history of Sri Lankan Cnemaspis species was extracted from the relevant literature (Bauer et al. 2007;Manamendra-Arachchi et al. 2007;Wickramasinghe and Munindradasa 2007;Vidanapathirana et al. 2014;Wickramasinghe 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 molecular 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 collected 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 available 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 isolation kit (Valencia, CA, USA) following the manufacture's protocols. The quality of the isolated DNA was determined through gel electrophoresis in ethidium bromide stained 1% Agarose gel. The concentration of the isolated DNA samples was quantified using a Nabi Nano-spectrophotometer (MicroDigital Company Ltd, Korea). We PCR amplified a 1040 bp fragment of the ND2 gene using already-published primers L4437a, AAGCTTTCG-GGCCCATACC and H5934, AGRGTGCCAATGTCT-TTGTGRTT (Macey et al. 1997). The PCR was carried out in 25 μl reactions with a primer concentration of 0.4 μM for each primer employing 35 cycles with an annealing temperature of 50 °C (Macey et al. 1997) following standard PCR protocols with Promega PCR master mix (Promega Corporation, Madison, Wisconsin, USA). The success of the PCR amplification and size of the amplified fragment 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 Genetech 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 sequences for Cnemaspis species of the South Asian radiation (Appendix 1). We did not include the Southeast Asian Cnemaspis as they are known to be a separate unrelated 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 ) 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 species from India and one species from Southeast Asia. Calodactylodes 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) methods. Partitioning schemes and best-fit substitution models for each partition were assessed using the Bayesian Information 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 implemented in RAxML 7.2.6. (Stamatakis et al. 2008) with 200 independent ML searches using the rapid hill-climb-ing algorithm. Branch support was estimated using 1000 bootstrap pseudoreplicates. Partitioned Bayesian analysis was performed in MrBayes 3.2.6 (Ronquist and Huelsenbeck 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 frequencies (< 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 average site cut-off of 95% (Kumar et al. 2018).
Species delimitation analysis using Poisson Tree Process (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 generations, 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 dissecting 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 nostril 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; interorbital 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 posterior 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-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 elbow; lower-arm length (LAL), the distance from the elbow 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 axilla 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 microscopes on both the left (L) and the right ( 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 addition, 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 characteristics, such as appearance of the caudal scales (except in specimens with regenerated tails). Colouration was determined 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 natural history of the focal species were observed through opportunistic and non-systematic means. The ambient temperature and the substrate temperature were measured using a standard thermometer and a N19 Q1370 infrared thermometer (Dick Smith Electronics, Shanghai, 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 species 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).
Dorsal surface of the trunk with smooth scales intermixed with keeled heterogeneous granules, 132 paravertebral granules; 148 smooth, mid-ventral scales; 95 midbody scales; 6/5 weakly-developed tubercles on the flanks; ventrolateral scales small, irregular; granules on snout 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 interorbital region circular and keeled; short tubercles present both on the sides of the neck and around the ear; ear opening vertically oval, slanting from anterodorsal to posteroventral, 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 posterior 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 supranasals. 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, posteriorly in contact with 3 enlarged postmentals (smaller 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 st infralabial; ventral scales smaller than chin scales. Smooth, rounded, juxtaposed scales on the chin and the gular region; pectoral and abdominal scales smooth, subimbricate towards precloacal region, abdominal scales slightly larger than dorsals; 17 belly scales across ventre; smooth scales around vent and base of tail, subimbricate; 3 precloacal 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, subrhomboidal, arranged in a single median series (Fig. 2).
Colour in life. Dorsum of head, body and limbs generally 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; dorsum 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 infralabials 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 (lokugei) in the masculine genitive singular, honouring Mr. Ajith Nethkelum Lokuge, a pioneer ecologist, analogue forestry specialist and a senior member of Young Zoologist's Association of Sri Lanka, for his significant 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 Province), which are situated in the central highlands of Sri  (Fig. 4). Tropical sub-montane and montane forests 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 habitats and tea plantations. It is very likely that the species occurs in the intervening regions between these two locations as there are similar habitats scattered between the two locations. However, this needs to be verified through a thorough field survey. These locations lie between 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 is 26.1-28.9 ºC. Both areas are rich in granite rock boulders with 40 identified caves. Cnemaspis lokugei sp. nov. appears to be a common species in the two locations as we recorded more than 50 indi-viduals from both locations during a two-day survey. This species was observed in granite caves and relatively old buildings on vertical surfaces, about 2 m from ground within the forested area (Fig. 5). The granitic cave microhabitat of C. lokugei sp. nov. was poorly illuminated (light intensity: 392-476 Lux), relatively moist (relative humidity: 76-92%), well shaded (canopy cover: 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, Hemidactylus frenatus, H. parvimaculatus and Hemiphyllodactylus 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 characteristic of Cnemaspis species and as there were no other Cnemaspis species observed in these habitats, it was presumed that the eggs most likely belong to C. lokugei sp. nov. Conservation status. Application of the IUCN Red List Criteria indicates that C. lokugei sp. nov. is Critically Endangered (CR) due to its having an area of occupancy (AOO) < 10 km 2 (3.84 km 2 in total assuming a 100 m radius around the seven georeferenced locations), severely fragmented habitat and a projected decline in the area, extent and the quality of habitat [Applicable criteria B2ab (iii)].

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 microhabitat specialists with narrow niches limited to moist, cool, Figure 4. Currently known distribution of Cnemaspis lokugei sp. nov. (holotype-red star, paratype-red circle) and its closely-related species (C. butewai -white circl, and C. pulchra -yellow circle) in Sri Lanka. canopy-shaded rock outcrops, granite caves, trees, abandoned buildings, buildings associated with caves, wattle and daub houses and semi-naturalised rock walls, where their cryptic morphology and body colouration camouflage them in the environment (Smith 1935;Karunarathna et al. 2019c). Further, Cnemaspis species prefer narrow (~ 3-4 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 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 sister to a clade comprising Cnemaspis sp. 3, Cnemaspis sp. 4, C. pulchra and C. butawаi. The taxonomic status of Cnemaspis sp. 3 and Cnemaspis sp. 4 needs to be further investigated. Two additional species of Cnemaspis,  C. rammalensis and C. rajakarunai, which were placed in phylogenetic analyses for the first time, were recovered 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. podihuna clade (Fig. 1) indicating speciation in the isolated mountains (Vidanapathirana et al. 2014;Wickramasinghe 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 intermediate bioclimatic zone (see Fig. 4). Our studies illustrate that Cnemaspis are distributed throughout all bioclimatic zones of the Island; however, the majority, i.e. 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 m a. s. l. (Fig. 4) Amarasinghe and Karunarathna 2020). In addition to this, our on-going studies indicate that there are at least another 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-Arachchi et al. (2007), Gymnodactylus malabarica Jerdon, 1853 (= Cnemaspis malabaricus) described from the forests of Malabar [Kerala State] is a valid species restricted to India. Although Kluge (2001) listed Cnemaspis malabarica (Jerdon, 1853) in the synonymy of C. kandiana, according 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 (Manamendra-Arachchi et al. 2007). Therefore, we consider these two species to be distinct. Similarly, C. lokugei 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 Sri Lanka and that C. malabarica is a species restricted to India, we believe the name C. malabarica is not applicable to C. lokugei sp. nov. Due to the presence of smooth ventrals in C. lokugei sp. nov., (vs. keeled ventrals in C. tropidogaster) 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 described 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 morphologically and genetically distinct and allopatric, we believe that Gymnodactylus humei (= Cnemaspis humei) 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 distinctly different from a suit of morphological characters (see Table 4) from C. lokugei sp. nov. and is also genetically 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 2 (i.e. AOO < 10 km 2 , EOO < 100 km 2 ) 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 localities 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 cultivation and invasive species. Most Cnemaspis species, like Cnemaspis lokugei sp. nov. described here are restricted to forests in mountains (Fig. 5a). Therefore, the conservation of such forests and other mountainous habitats are imperative to ensure the future survival of these species.