Two new species of South Asian Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from the Gingee Hills, Tamil Nadu, India

We describe two new small­bodied, sympatric species of south Asian Cnemaspis belonging to the mysoriensis + adii clade from the Gingee Hills in Tamil Nadu, peninsular India. The two new species can be easily distinguished from the other eight described members of the mysoriensis + adii clade by their dorsal pholidosis, the configuration of femoral and precloacal pores in males, a number of meristic characters and subtle differences in colouration, beside 6.7–20.8 % uncorrected pairwise ND2 sequence diver­ gence. The two species represent different ecomorphs, one a stouter, microhabitat generalist and the other a more slender, elongate rock specialist. The discovery of two new species from granite boulder habitats and Tropical Dry Evergreen Forests is indicative of the importance of these areas for biodiversity. It is likely that similar rocky habitats across southern peninsular India will harbour many more undescribed species.


Introduction
South Asian Cnemaspis Strauch, 1887 (SAC) is an an cient, diverse radiation of gekkonid lizards with a West ern Ghats origin, including >130 species distributed in peninsular India, northeast India, Sri Lanka, and in and around the Andaman Sea (Agarwal et al. 2020c(Agarwal et al. , 2021a;;Pal et al. 2021;Uetz et al. 2023).Mitochondrial phyloge nies of the group reveal the presence of 13 broad clades: two in Sri Lanka (kandiana which includes the Andaman Sea subclade, and podihuna), assamensis in lowland northeast India, and ten in peninsular India (Agarwal et al. 2020c(Agarwal et al. , 2021a;;Pal et al. 2021).The center of diversi ty in peninsular India is the cool Western Ghats region, where eight clades are represented (Agarwal et al. 2020c;Pal et al. 2021).Warmer, more arid habitats outside the Western Ghats include three clades of SAC: bangara, comprising four species distributed in and around the southern edge of the Mysore Plateau; mysoriensis + adii, including eight species distributed across the Mysore Plateau and hills off the southeastern edge of the Mysore Plateau (Fig. 1); and gracilis which includes 10 species outside the Western Ghats and three distributed in the Palghat Gap and the eastern slopes of the Western Ghats (Srinvasulu et al. 2015;Khandekar 2019;Khandekar et al. 2019Khandekar et al. , 2022;;Agarwal et al. 2020cAgarwal et al. , 2022;;Sayyed et al. 2023).The species Cnemaspis umashaankeri Narayanan & Aravind, 2022 from the Biligiri Rangan Hills, Karnata ka is morphologically similar to members of the gracilis clade, with sexual dimorphism, but its relationships with other SAC clades are not well resolved (Narayanan and Aravind 2022).
The mysoriensis + adii clade began diversifying in the late Oligocene to early Miocene, indicating a long history outside the Western Ghats (Agarwal et al. 2020c).All the species in the mysoriensis + adii clade are associated with granite boulders, and the Mysore Plateau or other upland areas, a combination of which has allowed them to persist in the warmest habitats occupied by any SAC (Agarwal et al. 2020c).As part of a larger project on the lizards of Tamil Nadu, we sampled Pakkamalai, Gingee Hills, Viluppuram District, during which time we collected two sympatric species of smallbodied Cnemaspis.One lin eage is morphologically allied to C. adii and the other to C. otai.We describe these two lineages as new species based on morphology, with the mitochondrial ND2 gene confirming their distinctiveness.

Taxon sampling
Specimens were handcollected and euthanized using iso flurane and liver tissues or tail tips of 1-3 individuals and one or two entire juveniles of each new species were collected in molecular grade ethanol and subsequently stored at -20° C for genetic analysis.Specimens were fixed in 8% formaldehyde for ~12 hours, washed in wa ter and transferred to 70% ethanol for longterm storage.Specimens are deposited in the Museum and Research Collection Facility at National Centre for Biological Sci ences, Bengaluru (NRCAA).

Molecular data and analyses
We extracted whole genomic DNA from a total of eight tissue samples of the two new species (Table 1) using the Qiagen DNeasy Blood & Tissue Kit.We generated se quences for the mitochondrial ND2 gene using the prim ers L4437 + H5540/ H5934 (Macey et al. 1997) with PCR and sequencing carried out by Barcode Biosciences, Ban galore.Bidirectional sequences were assembled in MEGA 5.2 (Tamura et al. 2011) and combined with 40 published sequences for the mysoriensis + adii clade and represen tatives of other South Asian Cnemaspis (after Khandekar et al. 2022; Table 1).Sequence alignment was carried out using default settings in ClustalW (Thompson et al. 1994) and uncorrected pairwise sequence divergence with the pairwise deletion option calculated in MEGA.A parti tioned Maximum Likelihood analysis was carried out on the IQTREE webserver (Nguyen et al. 2015;Trifinopou los et al. 2016), with partitioning scheme and substitution models selected in ModelFinder (Kalyaanamoorthy et al. 2017) that partitioned the data by codon position with the TVM+F+I+G4, TIM3+F+G4, and TIM+F+I+G4 models for positions 1-3).Analysis parameters were at default settings including edgelinked partitions and bootstrap support (BS) evaluated with 1000 ultrafast bootstraps (Hoang et al. 2018).We ran a partitioned Bayesian analy sis in MrBayes 3.2 (Ronquist and Huelsenbeck 2003) us ing the best-fit models and partitions from ModelFinder (TIM and TVM were replaced with GTR).Analyses were run for 1,000,000 generations, sampling every 100 gen erations, with two parallel runs and four chains each (one cold and three hot).Convergence was determined based on the standard deviation of split frequencies (<<0.01) and ESS (>200), and a consensus tree built using the sumt command with the first 25% of trees discarded as burn-in.
We did not conduct any explicit species delimitation analyses but use the 3.7 % ND2 pdistance cutoff sug gested by Agarwal et al. (2017) as well as the lowest di vergence between described species of the mysoriensis + adii clade as indicative of specieslevel divergence.
Morphological analyses were conducted in R 4.1.3(R Core Team 2018) and used using the following men sural variables: AGL, CL, ES, HL, HW, SVL.We used Thorpe's (1975) size correction equation that standard izes variables by SVL as implemented in the R package GroupStruct (available at https://github.com/chankinonn/GroupStruct) (Chan and Grismer 2021Grismer , 2022)).We pooled sexes and used a global mean SVL for the two new spe cies.A principal components analysis (PCA) was then conducted using the ez_pca function to visualize the sep aration of the two species in multivariate morphospace using the sizecorrected data (excluding SVL).

Phylogenetic relationships
The final ND2 alignment was 1047 base pairs (bp; in cluding a nine bp insertion in C. tigris, with newly added sequences ranging in length from 447-1047 bp and 1-3 complete ND2 sequences per species).We recovered the same broad relationships within SAC as previous authors, with successive basal splits separating the wynadensis clade, and then the beddomei clade from the remaining clades of the SAC (Fig. 2

Morphological analyses
Two factors with eigen values >1 were retained that cu mulatively explained 89.2 % of the variation in the data set.Factor 1 explains 69.3 % variance and loads strongly (factors not listed are < 0.40) for CL (0.52), ES (0.51), and HL (0.47).Factor 2 explains 19.8 % variance and loads strongly for AGL (-0.76) and HW (-0.55).The two diver gent lineages are completely separated across these two PCA factors, with the lineage allied to C. adii showing up as having longer limbs, a longer and narrower head, longer snout and shorter trunk (Fig. 2).We describe the two genetically and morphologically divergent lineages as new species below.3-5 Chresonymy.Cnemaspis otai Ganesh et al. (2018)    pralabial I; two rows of scales separate the orbit from the supralabials (Fig. 4A, C).Mental enlarged, subtriangular, marginally wider (1.6 mm) than long (1.2 mm); two pairs of postmentals, inner pair slightly larger than outer pair, roughly rectangular, in median contact with each other below mental; inner pair bordered by mental, infralabi al I, outer postmental and three enlarged chin shields on either side; outer postmentals roughly square, bordered by inner postmentals, infralabial I and II, and three en larged chin shields on either side; three enlarged gular scales prevent contact of left and right outer postmentals; chin shields bordering postmentals flat, smooth, smaller than outermost postmentals, rest flattened, smooth, even smaller (Fig. 4B).Infralabials bordered below by a row or two of slightly enlarged scales, decreasing in size posteri orly.Seven supralabials up to angle of jaw on either side, and six at midorbital position on each side; supralabial I largest, decreasing in size posteriorly; seven infralabials up to angle of jaw, five at midorbital position on left and six on right side; infralabial I largest, infralabials decreas ing in size posteriorly (Fig. 4C).Body relatively slender (BW/AGL 0.52), trunk less than half of SVL (AGL/SVL 0.42) without ventrolateral folds; three spine-like scales on left flank and two on right flank.Dorsal pholidosis heterogeneous; weakly keeled, granular scales intermixed with a few scattered enlarged keeled tubercles on vertebral and paravertebral region and about three irregularly arranged rows of large, weak ly keeled, tubercles on each side of flank (Fig. 5A-C).Scales on occiput and nape slightly smaller than those on paravertebral rows and weakly keeled; scales on flank slightly larger than those on dorsum, weakly keeled, coni cal or spinelike.Ventral scales much larger than granular scales on dorsum, those on belly smooth, subcircular su bimbricate, equal from chest to vent except for those on precloacal region which slightly larger; midbody scale rows across belly 25; 112 scales from mental to anteri or border of cloaca (Fig. 5B).Scales on throat slightly smaller than those on belly, imbricate; gular region with much smaller, flattened scales with those on chin border ing postmentals, enlarged, juxtaposed and flattened (Fig. 4B).Two femoral pores on each thigh separated by 11 poreless scales on left and nine on right (count incom plete due to injury) from two continuous precloacal pores (Fig. 4D).

Cnemaspis pakkamalaiensis sp. nov.
Scales on dorsal aspect of manus heterogenous, up per arm scales much larger than dorsal granules, strong ly keeled, imbricate; those near forelimb insertion much smaller than scales on upper arm; dorsal aspect of lower arm and elbow with scales much smaller than those on upper arm, weakly keeled, flat, roughly rounded; dorsal aspect of hand predominantly bearing large, flattened, weakly keeled, imbricate scales.Ventral aspect of upper arm with smooth, roughly rounded, subimbricate scales; scales on lower arm and wrist large, smooth, imbricate; scales on palm and sole smooth, flat and subcircular.Scales on anteriodorsal aspect of thigh much larger than enlarged scales on body dorsum, strongly keeled, and imbricate except those near hindlimb insertion which are granular, much smaller than dorsal granules and conical; scales on posteriodorsal aspect smaller, weekly keeled, granular.Scales on dorsal aspect of knee and shank small er than those on dorsum of thigh, subimbricate, weak ly keeled; dorsal aspect of foot predominantly bearing small, flattened, strongly keeled, imbricate scales; scales on ventral aspect of thigh and shank larger than those on midbody ventrals, smooth, subimbricate on thigh and imbricate on shank (Fig. 3A, B).
Tail entire and original except for extreme tip which is regenerated, subcylindrical, relatively slender, flattened beneath, slightly longer than snoutvent length (TL/SVL 1.18) (Fig. 3C-E).Dorsal scales at tail base granular, sim ilar in size and shape to those on midbody dorsals, gradu ally becoming larger, flatter, subimbricate posteriorly, in termixed with much enlarged, strongly keeled, distinctly pointed, conical tubercles forming whorls; six tubercles on first nine whorls.Scales on ventral aspect of original tail much larger than those on dorsal aspect, subimbri cate, smooth, with a series of three enlarged subcaudal scales of which the median series is slightly larger than adjunct two rows, roughly pentagonal; those on tail base much smaller, imbricate and smooth, a single enlarged, smooth postcloacal spur on each side (Fig. 3D).
Colouration in life (Fig. 6A).Dorsal ground colour of head, body, limbs and tail strawbrown; head mottled with fine dark speckles and larger dark blotches.Fine dark brown preorbital streak runs from snout to orbit, two fine dark brown postorbital streaks extend till neck; labials light grey/ cream with lighter and darker bars.A straw-coloured mid-dorsal streak that is formed by five fused elongate chainlinks runs from occiput to tail base; a dark ocellus fringed by a few orange scales anterior to forelimb insertions forms the posterior boundary of the first chain-link, flanked on either side by a wishbone shaped marking opening on neck and near forelimb in sertions; followed by four pairs of dark brown blotches, one pair at the posterior boundary of each chain link with a spot on either side.Flank with smaller dark spots and lightyellow markings.Tail with nine incomplete bands and no black tip.Dorsum of forelimbs and femur with few light and dark blotches, tibia with brown bands, two strong dark streaks on the posterior of femur, dig its with alternating dark and light bands.Ventral surfaces dull white, limbs finely speckled with brown especially  prominent under forelimbs, throat with a pair of broken up light grey longitudinal stripes on each side, no dark markings on belly, underside of tail lined by a fine dark border.Pupil black, iris silver with an orange streak to ward the anteroposterior of the pupil.
Variation and additional information from paratype series.Mensural, meristic and additional character state data for the paratype series is given in Tables 3-5 respec tively.There are four males and a single female ranging in size from 26.9-28.9mm (Fig. 7A, B).All paratypes re semble the holotype except as follows: inner postmental separated from each other below mental by an enlarged chin scale in NRCAA1285; outer postmental separat ed from each other by two enlarged chin scales in NRC AA1281, outer postmental bordered by four chin scales on left and three on right side in NRCAA1283.Three paratypes-NRCAA1285, NRCAA1281, and NRC AA1282 with original and complete tail, marginally or slightly longer than body (TL/SVL 1.04, 1.23, and 1.21 respectively); NRCAA1283 without tail; and NRC AA1284 with half of the tail missing (Fig. 7A, B).All paratypes agree with the holotype in overall colouration except dorsal tail colouration of female paratype (NRC AA1285) is overall duller than rest of the male types (Fig. 6B).
Etymology.The specific epithet is an adjective formed from the Latin "caverna' for cave and "cola" meaning in habitant or dweller, as the species is only known to occur in caves and crevices below large granite boulders.

Suggested Common Name. Cavedwelling dwarf gecko.
Diagnosis.A smallsized Cnemaspis, snout to vent length less than 34 mm (n = 5).Dorsal pholidosis heterogeneous; weakly keeled, granular scales in vertebral and paraverte bral region, intermixed with about two or three regularly arranged rows of large, weakly keeled tubercles on each side of flank, tubercles in lowest row largest and spinelike; 4-6 rows of dorsal tubercles; ventral scales smooth, subcircular, subimbricate, 28-32 scales across belly, 116-125 longitudinal scales from mental to cloaca; subdigital scansors smooth, entire, unnotched; 10-12 total lamellae under digit I of manus and pes, 15-19 lamellae under dig it IV of manus and 18-21 lamellae under digit IV of pes; males (n = 3) with one or two femoral pores on each thigh separated on either side by 8-10 poreless scales from a continuous series of three precloacal pores; tail with en larged, strongly keeled, distinctly pointed, conical tuber cles forming whorls; a median row of subcaudals smooth, distinctly enlarged.Dorsal colouration greybrown with a single medial dark spot on nape followed by four light blotches from forelimb insertions to tail base, tail with 12-14 alternating light and dark bars.tween left and right supraciliaries at midorbit (Fig. 10C).Earopening deep, oval, small (EL/HL 0.04); eye to ear distance greater than diameter of eye (EE/ED 1.62; Fig. 10C).Rostral twice wider (1.3 mm) than long (0.6 mm), incompletely divided dorsally by a strongly developed rostral groove for more than half of its length; a sin gle enlarged supranasal on each side, marginally larger than postnasals, separated from each other by a single

Comparison with members of C. mysoriensis
Tubercles linearly arranged (1) or more random (0) Spinelike tubercles on flank present (1) or absent (0) Lateral caudal furrows present (1) or absent (0) Median subcaudal scale row enlarged (1) or slightly enlarged (0) Enlarged femoral scales present (1) or absent (0) enlarged internasal and a smaller scale on snout; rostral in contact with nostril, supralabial I, supranasal and in ternasal; nostrils oval, each surrounded by postnasals, supranasal, rostral and supralabial I; two rows of scales separate the orbit from the supralabials (Fig. 10C).Men tal enlarged, subtriangular, marginally wider (1.5 mm) than long (1.2 mm); two pairs of postmentals, inner pair slightly larger than outer pair, roughly square, separated from each other below mental by an enlarged chin scale; inner pair bordered by mental, infralabial I and II, outer postmental and three enlarged chin shields on either side; outer postmentals roughly square, bordered by inner post mentals, infralabial II, and four enlarged chin shields on either side; three enlarged gular scales prevent contact of left and right outer postmentals; chin shields bordering postmentals flat, smooth, smaller than outermost post mentals, rest flattened, smooth, even smaller (Fig. 10B).
Infralabials bordered below by a row or two of slightly enlarged scales, decreasing in size posteriorly.Seven su pralabials up to angle of jaw on left and eight on right, six at midorbital position; supralabial I largest, decreasing in size posteriorly; seven infralabials up to angle of jaw, six at midorbital position on left and five on right; infralabial I largest, infralabials decreasing in size posteriorly (Fig. 10C).Body relatively slender (BW/AGL 0.52), trunk less than half of SVL (AGL/SVL 0.41) without ventrolater al folds; three spine-like scales on either side of flank.Dorsal pholidosis heterogeneous; weakly keeled, granu lar scales on the vertebral and paravertebral region, in termixed with about three irregularly arranged rows of large, weakly keeled, tubercles on each side of flank (Fig. 11A-C).Scales on occiput and nape much smaller and weakly keeled than those on paravertebral rows; scales on flank slightly larger than those on dorsum, weakly keeled, conical or spinelike.Ventral scales much larger than granular scales on dorsum, those on belly smooth, subcircular subimbricate, equal from chest to vent except for those on precloacal region which slightly larger; mid body scale rows across belly 29; 123 scales from mental to anterior border of cloaca (Fig. 11B).Scales on throat slightly smaller than those on belly, imbricate; gular re gion with much smaller, flattened scales with those on chin bordering postmentals, enlarged, juxtaposed and flattened (Fig. 10B).A single femoral pore on left thigh and two on right separated by eight poreless scales on left and 10 on right from a continuous series of three preclo acal pores (Fig. 10D).
Scales on dorsal aspect of manus heterogenous, upper arm with scales much larger than dorsal granules, weak ly keeled, imbricate; those near forelimb insertion much smaller than scales on upper arm; dorsal aspect of lower arm and elbow with scales much smaller than those on upper arm, weakly keeled, flat, roughly rounded; dorsal aspect of hand predominantly bearing large, flattened, smooth to weakly keeled, imbricate scales.Ventral aspect of upper arm with smooth, roughly rounded, subimbricate scales; scales on lower arm and wrist large, smooth, im bricate; scales on palm and sole smooth, flat and subcir cular.Scales on anterodorsal aspect of thigh much larger than those on dorsal granules, weakly keeled, imbricate except those near hindlimb insertion which are granular, much smaller than dorsal granules, conical; scales on posterodorsal aspect smaller, smooth to weekly keeled, granular.Scales on dorsal aspect of knee and shank fair ly smaller than those on dorsum of thigh, subimbricate, weakly keeled; dorsal aspect of foot predominantly bear ing small, flattened, smooth to weakly keeled, imbricate scales; scales on ventral aspect of thigh and shank more or less equal to those on midbody ventrals, smooth, sub circular, subimbricate on thigh and imbricate on shank (Fig. 9A, B).
Tail entire except for extreme tip which is missing, original, subcylindrical, relatively slender, flattened be neath, marginally shorter than snoutvent length (TL/ SVL 0.96) (Fig. 9C-E).Dorsal scales at tail base granu lar, similar in size and shape to those on midbody dorsals, gradually becoming larger, flatter, subimbricate poste riorly, intermixed with much enlarged, strongly keeled, distinctly pointed, conical tubercles forming whorls; six tubercles on first eight whorls.Scales on ventral aspect of original tail much larger than those on dorsal aspect, sub imbricate, smooth, with a series of three enlarged subcau dal scales of which the median series is distinctly larger than adjunct two rows, covering almost entire portion of the tail; those on tail base much smaller, imbricate and smooth, a single enlarged, smooth postcloacal spur on each side (Fig. 9D).
Colouration in life (Fig. 12A).Dorsal ground colour of head, body, limbs and tail greybrown; head mottled, with a few dark blotches; anterior of brille yellow.Fine dark brown preorbital streak runs from nasal to orbit, two fine dark brown postorbital streaks extend till neck; labi als yellow with darker bars.Dorsal markings consist of a dark ocellus outlined by brown just anterior to forelimb insertions followed by four offwhite blotches from fore limb insertions to tail base; rest of dorsum strongly mot tled with light grey blotches and fine black spots.Flank with smaller dark spots and light grey markings.Tail more grey than body, with 14 alternating light and dark bands, tail tip black.Dorsum of limbs with scattered light grey, brown and black markings; two strong dark streaks on the posterior of femur, digits with alternating dark and light bands.Ventral surfaces dull white, fine speckling under forelimbs, throat with a grey longitudinal stripe on each side with some dark markings below eye and angle of jaw, no dark markings on belly, underside of tail lined by a fine dark border.Pupil black, iris silver with an or ange streak toward the posterior of the pupil.
Variation and additional information from paratype series.Mensural, meristic and additional character states evaluation data for the paratype series is given in Tables 3-5 respectively.There are two males and two females ranging in size from 28.2-34.0mm (Fig. 13A, B).All   postmental bordered by four gular scales on left and three on right side in NRC-AA-1289; and by five gular scales on left and three on right side in NRCAA1290.Two paratypes-NRCAA1287 and NRCAA1290 with original and complete tail, marginally and slightly longer than body (TL/SVL 1.10 and 1.21 respectively); NRC AA1289 with half original half regenerated tail; NRC AA1288 with half of the tail missing (Fig. 13A, B).All paratypes agree with the holotype in overall colouration except for unvouchered hatchling and a juvenile which are overall duller than adults and have orange tail tips.(Fig. 12B-D).
Distribution and natural history.Cnemaspis cavernicola sp.nov. is known only from its type locality (Pak kamalai Reserve Forest, Gingee Hills in Viluppuram dis trict, Tamil Nadu), at elevations of ca.400-480 m asl.(Fig. 1).Cnemaspis cavernicola sp.nov.was encoun tered during a single day of fieldwork each on two differ ent fieldtrips in Pakkamalai.Individuals were observed in small numbers (>10 in 2 hrs) and only in the higher reaches of Pakkamalai in shaded and relatively cooler areas among very large granite boulders.A few individu als were seen moving around in the morning (0930-1130 hrs) on granite rocks ≥ 2 m above the ground and NRC-AA1287 and a juvenile specimen were seen inactive in a small rock crevice located inside a granite cave in the evening (Fig. 8A, B).Sympatric lizards recorded by us at the type locality include Cnemaspis pakkamalaiensis sp.nov.Calodactylodes aureus, Hemidactylus frenatus, H. pakkamalaiensis, H. whitakeri, Eutropis carinata, and Psammophilus dorsalis.

Discussion
The high rate of discovery of South Asian Cnemaspis in peninsular India, especially from regions outside the Western Ghats, shows no signs of abating.The discov ery of these two species takes the number of peninsu lar Indian Cnemaspis known from outside the Western Ghats to 24, all but one of which have been described since the turn of the century (Jerdon 1853;Das and Bau er 2000;Khandekar 2019;Khandekar et al. 2019, 2020, 2022, Agarwal et al. 2020a, 2020b, 2020c;2021b, 2022;Narayanan et al. 2022Narayanan et al. , 2023b)).Tamil Nadu is turning out to be particularly diverse, with 13 endemic species of Cnemaspis outside the Western Ghats (Das and Bauer 2000;Khandekar 2019;Khandekar et al. 2019, Agarwal et al. 2020c, 2021b, 2022;Sayyed et al. 2023).
The discovery of two more lineages of SAC from pen insular India outside the Western Ghats, including the relict Cnemaspis cavernicola sp.nov.that has no close extant relatives, is indicative of the longterm persistence of Cnemaspis in refugia provided by boulder habitats (Srinivasulu et al. 2015;Agarwal et al. 2020c).This em phasizes the fact that at least for lizards, the true biogeo graphic unit is peninsular India (and Sri Lanka), though past climate change has wiped out many cool adapted species from regions outside the Western Ghats, except in refugia linked with boulder habitats, high elevations and or forest (Agarwal et al. 2019(Agarwal et al. , 2020c)).Granite boulders in particular have been shown to allow the persistence of Cnemaspis outside the Western Ghats, and it is likely that there is high biodiversity across other taxa linked to these refugia (Agarwal et al. 2020c).Rocky, granite habitats are not recognised by the general public or forest department as important habitats with endemic biodiversity and are under immense anthropogenic pressure from quarrying.
Tropical Dry Evergreen Forests (TDEFs) are an im portant habitat type along the southeast coast of India (Champion and Seth 1968;MeherHomji 2007).Tropi cal dry evergreen habitats have been lost across most of their original extent and are restricted to small patches around temples (sacred groves) and a few larger reserve forest areas (Parthasarathy et al. 2008).Previous studies have focused on the floristic diversity of TDEFs, but re cent discoveries show the presence of endemic species of lizards including Cnemaspis avasabinae, Cyrtodactylus irulaorum Agarwal, Thackeray &Khandekar, 2023 andCy. relictus Agarwal, Thackeray &Khandekar, 2023, and Hemidactylus pakkamalaiensis (Agarwal et al. 2020a(Agarwal et al. , 2023;;Narayanan et al. 2023b).Pakkamalai and the asso ciated Gingee Hills are unique for their extremely rocky formations and the presence of TDEFs, with a unique flo ristic assemblage including a number of Western Ghats species (Balachandran and Rajendiran 2018).These hills also have numerous endemic species including two spe cies of trees (Ramachandran et al. 2015;Balachandran and Rajendiran 2016) and the recently described Hemidactylus pakkamalaiensis (Narayanan et al. 2023b), in addition to the two new species described herein.
Though both new species are sympatric at the type lo cality, Cnemaspis pakkamalaiensis sp.nov. is morpholog ically typical of mysoriensis clade species -with a rel atively short and broad head, short snout and limbs, and longer trunk; and is distributed between 200-400 m asl.and found < 2 m from ground level on rocks and on the ground.On the other hand, Cnemaspis cavernicola sp.nov.has the typical morphology of a rock specialist, with a longer and narrower head, elongate snout and limbs and short trunk (Grismer et al. 2015); and has only been re corded from 400-480 m asl.and found >2 m from ground level on rocks.A detailed morphological comparison is needed, but members of the bangara clade, C. adii and C. cavernicola sp.nov.are all rock specialists and share a similar morphology, with a relatively elongate head and limbs, and short trunk -adaptations seen in other cave and karst adapted geckos (e.g., Grismer et al. 2015).Cnemaspis species from divergent clades are sympatric at many localities in the Central and Southern Western Ghats; while in regions of peninsular India outside the Western Ghats, rockdwelling (C.adii, bangara clade spe cies, C. cavernicola sp.nov.) and generalist ecomorphs (mysoriensis clade) are sympatric wherever the former occur, including multiple localities on the Mysore Plateau and Pakkamalai (e.g.Agarwal et al. 2017Agarwal et al. , 2020cAgarwal et al. , 2021bAgarwal et al. , 2022)).Much sampling remains to be carried out to first understand patterns of diversity and distribution within this hyperdiverse genus, and to then try and unravel ecol ogy and behaviour of individual species and communities.

Figure 1 .
Figure 1.Elevational map of peninsular India showing our sampling of the mysoriensis + adii clade which includes typical or to potypical for all species.

Figure 3 .
Figure 3. Cnemaspis pakkamalaiensis sp.nov.(holotype, NRCAA1280): A dorsal view of body, B ventral view of body, C dorsal view of tail, D ventral view of tail, E lateral view of tail.Scale bars 10 mm; photos by Akshay Khandekar.

Figure 4 .
Figure 4. Cnemaspis pakkamalaiensis sp.nov.(holotype, NRCAA1280): A dorsal view of head, B ventral view of head, C lateral view of head on right, D view of cloacal region showing femoral and precloacal pores, E ventral view of left manus, F ventral view of left pes.Scale bars 5 mm; photos by Akshay Khandekar.

Figure 8 .
Figure 8. Habitat of Cnemaspis pakkamalaiensis sp.nov., and C. cavernicola sp.nov. at the type locali ty: A general view showing granite boulders surrounded by tropical dry evergreen forests, B granite boulder cave from where individuals of the new species were collected.Photos by Akshay Khandekar

Figure 9 .
Figure 9. Cnemaspis cavernicola sp.nov.(holotype, NRCAA1286): A dorsal view of body, B ventral view of body, C dorsal view of tail, D ventral view of tail, E lateral view of tail.Scale bars 10 mm; photos by Ishan Agarwal (A), and Akshay Khandekar (B).

Figure 10 .
Figure 10.Cnemaspis cavernicola sp.nov.(holotype, NRCAA1286): A dorsal view of head, B ventral view of head, C right side lateral view of head, D view of cloacal region showing femoral and precloacal pores, E ventral view of left manus, F ventral view of left pes.Scale bars 5 mm; photos by Akshay Khandekar.

Table 1 .
List of Cnemaspis sequences used in this study (sequences generated in this study marked by *).Museum abbreviations are as follows: BNHS, Bombay Natural History Society; AK/AK-R, Akshay Khandekar field series; VG, Varad Giri field series; CES, Centre for Ecological Sciences, Bangalore; NRC-AA, National Centre for Biological Sciences; AA, Rohan Pethiyagoda field series; SB, Sayantan Biswas field series; USNM, United States National Museum; ZSI-R, Zoological Survey of India, Kolkata.
C. umashaankeri and C. adii.The two new lineag es from Pakkamalai are nested within a poorly supported mysoriensis + adii clade.Cnemaspis adii forms the sister taxon to a wellsupported clade (BS = 100, PP = 1) within which the lineage allied to C. adii from Pakkamalai forms the sister taxon to the C. mysoriensis clade.The new spe cies allied to C. otai is sister to a clade which includes C.
Maximum likelihood phylogeny of South Asian Cnemaspis based on the mitochondrial ND2 gene.Posterior probability (≥0.95) and bootstrap support (≥70) shown at nodes.Inset, plot of the first two principal components for mensural data of the two new species.
flank, tubercles in lowest row largest, spine-like; six rows of dorsal tubercles; ventral scales smooth, subcircular, subimbricate, 25-27 scales across belly, 100-112 longi tudinal scales from mental to cloaca; subdigital scansors smooth, entire, unnotched; 8-11 total lamellae under digit I of manus and pes, 14-16 lamellae under digit IV of ma nus and 17-21 lamellae under digit IV of pes; males (n = 5) with two femoral pores on each thigh separated on either side by 8-11 poreless scales from two continuous precloacal pores; tail with enlarged, strongly keeled, dis tinctly pointed, conical tubercles forming whorls; a me dian row of subcaudals smooth, slightly enlarged.Dorsal colouration straw brown with a broad, light middorsal streak formed by five or six fused elongate chain-links from occiput to tail base, single medial dark spot on nape, dark paired spots on either side of middorsal streak, four pairs between forelimb insertions and tail base, tail with nine alternating light and dark markings.Comparison with members of C. mysoriensis + adii clade.Cnemaspis pakkamalaiensis sp.nov.canbedistinguishedfromalleightmembers of the mysoriensis + adii clade on the basis of the following differing or nonover lapping characters: males with two femoral pores on each thigh separated on either side by 8-11 poreless scales from two continuous precloacal pores (versus femoral pores absent, continuous series of 2-5 precloacal pores in C. avasabinae; three femoral pores on each thigh separat ed on either side by nine or 10 poreless scales from con tinuous series of four precloacal pores in C. otai; a single femoral pore on each thigh, separated on either side by 10 poreless scales from continuous series of three precloacal pores in C. rishivalleyensis; three femoral pores on each thigh, separated by five or six poreless scales from two continuous precloacal pores in C. yercaudensis); six rows of dorsal tubercles at midbody (versus dorsal pholido sis homogeneous in C. adii, dorsal tubercles irregularly arranged at midbody in C. avasabinae, 7-10 rows of dorsal tubercles at midbody in C. stellapulvis); 25-27 ventral scales across belly at midbody (versus 17-20 ventral scales across belly at midbody in C. avasabinae, 18 ventral scales across belly at midbody in C. otai, 20 or 21 ventral scales across belly at midbody in C. mysoriensis, 20-22 ventral scales across belly at midbody in stellapulvis, 18-20 ventral scales across belly at mid body in C. yercaudensis); spinelike tubercles present on flank (versus spine-like tubercles absent on flank in C. adii, and C. avasabinae); a single distinct black dorsal ocellus on middorsal streak just anterior to forelimb in sertions (versus distinct black dorsal ocellus just anteri or to forelimb insertions absent in C. mysoriensis and C. yercaudensis); a light mid-dorsal streak formed by five or six fused, elongate chainlinks that run from occiput to tail base (versus middorsal streak absent in C. adii, a continuous light middorsal streak runs from occiput onto tail base in C. mysoriensis, C. stellapulvis, C. tigris, and C. yercaudensis).Cnemaspis pakkamalaiensis sp.nov. is diagnosed against the second new species as part of its description below.Description of the holotype.Adult male in good state of preservation except for tail tip slightly bent towards the right, partially everted hemipenis on the left, tail mar gi nally detached just posterior to tail base on the left (Fig.3A-E)).SVL 29.0 mm, head short (HL/SVL 0.26), wide (HW/HL 0.70), not strongly depressed (HD/HL 0.42), distinct from neck.Loreal region slightly inflated, canthus rostralis not prominent.Snout marginally less than half the head length (ES/HL 0.48), more than twice eye diameter (ES/ED 2.31); scales on snout and canthus ros tralis large, subcircular, smooth to weakly keeled; much larger than those on forehead and interorbital region; oc cipital and temporal region with much smaller, weakly keeled granules (Fig.4A).Eye small (ED/HL 0.20); with round pupil; orbit with extrabrillar fringe scales that are largest anteriorly; supraciliaries not elongate; six interor bital scale rows across narrowest point of frontal; 25 or 26 scale rows between left and right supraciliaries at midor bit (Fig.4A, C).Earopening deep, oval, small (EL/HL 0.05); eye to ear distance greater than diameter of eye (EE/ED 1.60; Fig.4C).Rostral twice as wide (1.3 mm) than long (0.6 mm), incompletely divided dorsally by a strongly developed rostral groove for more than half its length; a single enlarged supranasal on each side, margin ally larger than postnasals, separated from each other by a single enlarged internasal; rostral in contact with nos tril, supralabial I, supranasal and internasal; nostrils oval, each surrounded by postnasals, supranasal, rostral and su

Table 2 .
Pairwise uncorrected ND2 sequence divergence within the mysoriensis + adii clade, numbers in bold along diagonal rep resent intraspecific genetic diversity (-indicates only a single sequence available).

Table 3 .
Mensural (mm)data for the type series of the new species.Abbreviations are listed in Materials and Methods; * = tail incomplete.
region absent, about two or three regularly arranged rows of large, weakly keeled tubercles on each side of flank (versus dorsal pholidosis homogeneous in C. adii, gran ular scales in the vertebral and paravertebral region with a few scattered enlarged keeled tubercles in C. mysoriensis, C. otai, C. pakkamalaiensis sp.nov., C. stellapulvis, C. tigris, and C. yercaudensis); a middorsal streak absent (versus a continuous light middorsal streak runs from occiput onto tail base in C. mysoriensis, C. sion in sternal region for tissue collection (Fig.9A-E).SVL 32.2 mm, head short (HL/SVL 0.25), wide (HW/ HL 0.61), not strongly depressed (HD/HL 0.33), distinct from neck.Loreal region slightly inflated, canthus ros tralis not prominent.Snout marginally less than half the head length (ES/HL 0.45), more than twice eye diameter (ES/ED 2.37); scales on snout and canthus rostralis large, subcircular, smooth; much larger than those on forehead

Table 4 .
Meristic data for the type series of the new species.Abbreviations are listed in Materials and Methods except for: L&R = Left & Right; irr = irregularly arranged; abs.= absent; * = damaged.

Table 5 .
Additional morphological characters for the type series of the new species.abs.= absent; / = data unavailable.