Three more novel species of South Asian Cnemaspis Strauch, 1887 (Squamata, Gekkonidae) from Kalakad Mundanthurai Tiger Reserve, Tamil Nadu, India

We describe three distinct, small­bodied, scansorial species of south Asian Cnemaspis from Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu, India—Cnemaspis azhagu sp. nov. from Thirukurungudi forest range, Cnemaspis mundanthuraiensis sp. nov. from Mundanthurai forest range and Cnemaspis kalakadensis sp. nov. from Kalakad forest range. Phylogenetic analyses using a partial sequence of the mitochondrial ND2 gene and general morphology places each of the three new species in the beddomei, gracilis and littoralis clades, respectively. The three new species are diagnosed from all other described members of their respective clades by a suite of differing morphological characters including snout vent length, number of dorsal tubercle rows at mid-body, number of paravertebral tubercles, presence or absence of spine-like scales on flanks, number of ventral scales across belly at mid­body, number of ventral scales from mental to anterior border of cloaca, number of lamellae under digit IV of pes, number of femoral and/or precloacal pores and poreless scales separating these series, as well as subtle colouration differences. We also provide some novel characters of tail tuberculation of the three new species described herein. With the discovery of these three new species, eight species of geckos including five Cnemaspis are now known to be endemic to KMTR.


Introduction
The South Asian clade within the polyphyletic genus Cnemaspis Strauch, 1887 diversified from a Western Ghats origin during the PaleoceneEocene and today in cludes over 100 species disjunctly distributed in peninsu lar India, Sri Lanka, parts of Southeast Asia, with a single species in northeast India (Iskandar et al. 2017;Lee et al. 2019;Agarwal et al. , 2021aAmarasinghe et al. 2021;Pal et al. 2021;Uetz et al. 2022). A recent series of contributions on the taxonomy and biogeography of South Asian Cnemaspis have resulted in the placement of almost all described species from India and Sri Lanka into phylogenies, redescriptions of several poorly known species, and descriptions of a large number of new species (Sayyed et al. 2018(Sayyed et al. , 2021Cyriac et al. 2018Cyriac et al. , 2020 Ten well supported, broad clades are now recognised within South Asian Cnemaspis in peninsular India based on ND2 and 16S mitochondrial phylogenies (Sayyed et al. 2018;Khandekar et al. 2019a;Agarwal et al. 2020aAgarwal et al. , 2021bCyriac et al. 2020;Pal et al. 2021). Members of the bangara and mysoriensis clades are restricted to rela tively arid, rocky landscapes on the Mysore Plateau; the girii clade to the northern Western Ghats, the wynadensis and goaensis clades to the central and northern Western Ghats, beddomei to the southern Western Ghats, and the indica, littoralis, and monticola clades are distributed in the southern and central Western Ghats (Pal et al. 2021;Khandekar et al. 2022). The gracilis clade is among the most widely distributed, found in the central and southern Western Ghats, the southern Eastern Ghats and the My sore Plateau.
During a recent survey in Kalakad Mundanthurai Tiger Reserve, Tamil Nadu, India as part for an ongoing proj ect on the systematics and taxonomy of peninsular Indian geckos, we collected multiple unidentified small-bodied Cnemaspis specimens from a few forest ranges in the Tiger reserve. Based on preliminary morphological ex amination, the specimens collected from Thirukurungudi, Mundanthurai, and Kalakad forest ranges represent three species that fall within the beddomei, gracilis and littoralis clades, respectively, as defined by Pal et al. (2021). Detailed taxonomic examination revealed that the three species differ from other members of their respective clades in a number of nonoverlapping morphological characters and represent three undescribed species. Mo lecular analysis based on ND2 mitochondrial gene frag ments of these samples confirmed their placements in respective clades as well as their distinctiveness. In this paper, we describe these three new species based on their distinctive morphology.

Taxon sampling
Surveys were conducted in both day and night time, spec imens were spotted on rocks, tree trunks and collected by hand and euthanized using isoflurane after taking colour photos in life. Liver/ tail tissues of at least three individu als of each new species were collected in molecular grade ethanol and subsequently stored at -20°C for genetic analysis. Specimens were fixed in 8% formalin for ~12 hours, washed and kept in tap water for ~12 hours, and transferred to 70% ethanol for long-term storage. Spec imens are deposited in the museum and research collec tion facility at the National Centre for Biological Scienc es, Bengaluru (NRC) and at the Bombay Natural History Society, Mumbai (BNHS).

Molecular data
We extracted genomic DNA from liver/ tail biopsies us ing the Qiagen DNeasy Blood & Tissue extraction kit for samples of the three new species as well as a few topo typical Cnemaspis littoralis (Jerdon, 1853) (Table 1). We targeted the protein coding mitochondrial ND2 gene us ing the primers L4437 & H5934 (Macey et al. 1997), with L4437 used for sequencing. PCR and sequencing were outsourced to Medauxin, Bangalore. These sequences were combined with published sequences including rep resentatives of all clades of South Asian Cnemaspis and all known species of the beddomei, gracilis and littoralis clades (after Agarwal et al. 2020;Pal et al. 2021; Table  1). ND2 sequences were not available for C. aaronbaueri Sayyed, Grismer, Campbell andDileepkumar, 2019 andC. nairi Inger, Marx andKoshy, 1984 of the beddomei clade, and C. flavigularis Pal, Mirza, Dsouza and Shanker, 2021 and C. palakkadensis Sayyed, Cyriac and Dileepkumar, 2020 of the littoralis clade. Additionally, we used a chimeric sequence for C. regalis Pal, Mirza, Dsouza and Shanker, 2021 as the two published sequenc es (that are of individuals from the same locality) have only 70 nucleotides of overlap. Sequences were aligned using default settings in ClustalW (Thompson et al. 1994) as implemented in MEGA 5.2 (Tamura et al. 2011). Un corrected pairwise pdistance with the partial deletion option was calculated in MEGA and the best fit models of sequence evolution were picked using the Bayesian Information Criteria in Partitionfinder 2 (Lanfear et al. 2012

Phylogenetic relationships
The ND2 sequences ranged in length from 490-1038 nu cleotides and we translated these to amino acids to ensure that there were no erroneous internal stop codons. We re covered the same broad ten clades as Pal et al. (2021), within which we recognise the littoralis clade to be ex clusive of the Sri Lankan species of the podihuna clade and C. assamensis (Fig. 2) Etymology. The specific epithet, azhagu (a-lha-gu, also sometimes transliterated as alaku), is the Tamil word for beauty (அழகு) and is used as a noun in apposition for this beautiful new species.

Diagnosis.
A smallsized Cnemaspis, snout to vent length less than 38 mm (n=9). Dorsal pholidosis hetero geneous; smooth granular scales intermixed with a fair ly regularly arranged row of enlarged, weakly keeled, conical tubercles on either side of flank; granular scales gradually increasing in size towards each flank, largest on mid-flank; spine-like scales absent on the flank; two (rarely three, n=1/9) rows of dorsal tubercles at mid body, enlarged tubercles in paravertebral region absent (rarely a few present, n=2/9); ventral scales subcircular, smooth, subimbricate, and subequal from chest to vent, 34-44 scales across belly at midbody, 151-171 longi tudinal scales from mental to cloaca; subdigital scansors smooth, some divided and others entire; 13-16 lamellae under digit I of manus and 12-14 lamellae under digit I of pes, 20-25 lamellae under digit IV of manus and 24-27 lamellae under digit IV of pes; males (n=7/9) with a continuous series of 6-8 precloacal pores (n= 5/7), rare ly divided medially by a single poreless scale (n=2/7), femoral pores absent; tail with enlarged, weakly keeled, pointed, and weakly conical tubercles forming four whorls only on anterior third; followed by a row of three paravertebral tubercles on either side; rest of tail without enlarged tubercles; a median row of subcaudals smooth, regularly arranged with condition of two slightly larg er scales alternating with a large divided scale. Males with ochre head and grey body, females more uniform brown; single central black ocellus on occiput flanked by lighter markings, collar formed by thick black spots/ streak flanked posteriorly by white band just anterior to forelimb insertions; tail without strong markings; iris red with light orange ring surrounding pupil.
Body relatively slender (BW/AGL 0.44), trunk less than half of SVL (AGL/SVL 0.40) without ventrolateral folds and spine-like scales on flank (Fig. 5B, C). Dorsal pholidosis heterogeneous; smooth granular scales inter mixed with a fairly regularly arranged row of enlarged, weakly keeled, conical tubercles on either side of flank; granular scales gradually increasing in size towards each flank, largest on mid-flank; granular scales on occiput and nape slightly smaller than paravertebral granules (Fig. 5A). Ventral scales much larger than granular scales on dorsum, subcircular, smooth, subimbricate, subequal from chest to vent; scales on precloacal region and on last two or three rows on femur distinctly enlarged; mid body scale rows across belly 40; 159 scales from mental to anterior border of cloaca (Fig. 3B). Scales on base of neck similar to those on belly, marginally smaller; gular region with much smaller, smooth, granular scales, those bordering postmentals enlarged, smooth, subcircular, and flattened (Fig. 4B). A continuous series of six precloacal pores, femoral pores absent (Fig. 4D). Scales on palm and soles granular, smooth, rounded, and flattened; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small granules, slightly larger than body dor sum and flattened, smooth (few weakly keeled), subim bricate scales which are twice the size of granules on the body dorsum, largest on anterolateral aspect of the hands and feet; posterolateral aspect of limbs with small granu lar scales; ventral aspect of forelimbs with small granular scales, slightly larger on lower arm than upper arm; ven tral aspect of hindlimb with enlarged, smooth, flattened,   3A, B). Forelimbs and hindlimbs slightly long, slender (LAL/ SVL 0.15); (CL/SVL 0.18); digits long, with a strong, re curved claw, distinctly inflected, distal portions laterally compressed conspicuously. Digits with both paired and unpaired lamellae, separated into a basal and narrower distal series by single enlarged lamella at inflection; 1-7 most basal paired on basal series and 1-5 paired lamel lae above the inflection; basal lamellae series: (2-6-6-6-4 right manus, 27784 right pes), (26664 left manus, Tail original, entire, subcylindrical, slender, slightly longer than snoutvent length (TL/SVL 1.30; Fig. 3C-E). Dorsal pholidosis on tail heterogeneous; small, smooth, subcircular, flattened, subimbricate scales intermixed on anterior one third portion with enlarged, weakly keeled, and weakly conical tubercles forming four whorls; eight tubercles on first whorl, six tubercles on second, five tu bercles on third and four tubercles on fourth; followed by a row of three paravertebral tubercles on either side; rest of the tail lacking enlarged tubercles (Fig. 3C, E). Scales on tail venter much larger than those on dorsal aspect, smooth, roughly subcircular, flattened, sub-imbricate; median series slightly larger than rest, regularly arranged with condition of two slightly larger scales alternating with a large divided scale (Fig. 3D). Scales on tail base slightly smaller, smooth, imbricate; a single enlarged, smooth, subcircular, and weakly conical postcloacal spur on each side (Fig. 3D, E). Fig. 6A). Dorsal ground colour of body, limbs and tail grey; entire head and region anteri or to forelimb insertions ochre. Indistinct light preorbital streak runs from nostril to orbit; three light postorbital streaks, uppermost terminating in parietal region, middle at occiput and lowermost continuing until ear opening. A single large central black ocellus on occiput enclosed within a Ushaped light marking and in between upper most and middle postorbital streaks; a single smaller black ocellus on the right side anterior to forelimb inser tions; an incomplete collar at anterior edge of forelimb insertions consisting of a pair of dark blotches flanked posteriorly by a narrower white band on centre of back and small black spot flanked by white band at anterior base of forelimb insertion. Five white spots on vertebral region between forelimbs and tail base, scattered white spots of similar or smaller size on dorsum and femur, larger irregular black blotches scattered on dorsolateral aspect of back (two on left side and one on right). Origi nal tail with indistinct lighter bands, digits with numerous light grey bands. Ventral surfaces of body, limbs and tail light grey with some scattered darker scales especially in centre of belly, throat and underside of neck ochre with scattered darker scales. Pupil black, iris reddish with a light orange ring lining pupil.

Colouration in life (
Variation and additional information from type series. Mensural, meristic and additional character state data for the type series is given in Tables (Fig. 7).  (1)  Distribution and natural history. Cnemaspis azhagu sp. nov. is currently known only from around its type locality (Thirukurungudi forest range ca. 200-400 m asl.), Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu (Fig. 1). Individuals of the new spe cies were observed active during the daytime (morning to afternoon, 0930-1300 hrs) on rocks (< 2 m high from the base) inside dry deciduous forest patches ( Fig. 8A-C). A large number of individuals (n = >30) were observed at all three locations indicating high abundance. The species moved inside rock crevices rapidly when approached. The holotype and two paratypes were collected during the day from rock crevices, while the rest of the type se ries were collected at nighttime (1930-2100 hrs) from the other two closely spaced localities. All the paratypes collected at night were observed inactive, located openly on the rocks and did not try to escape when approached. Suggested Common Name. Mundanthurai dwarf gecko.

Diagnosis.
A smallsized Cnemaspis, snout to vent length less than 33 mm (n=9). Dorsal pholidosis hetero geneous; weakly keeled, weakly conical, granular scales intermixed with irregularly arranged rows of enlarged, strongly keeled, conical tubercles which are gradually in creasing in size towards flank; last two rows of enlarged tubercles on flank largest, spine-like; 6-8 rows of dorsal tubercles at midbody, tubercles in paravertebral region ir regular; ventral scales subcircular, smooth, subequal from chest to vent, 30-34 scales across belly at midbody, 115-128 longitudinal scales from mental to cloaca; subdigital scansors smooth, unpaired, unnotched; 10 or 11 lamellae under digit I of manus and 9-12 lamellae under digit I of pes, 14-18 lamellae under digit IV of manus and 18-22 lamellae under digit IV of pes; males (n=7/9) with 3-5 femoral pores on each thigh separated by 9-11 poreless scales from two precloacal pores, precloacal pores sep arated medially by 2-4 poreless scales (n=4/7), preclo acal pores sometimes absent entirely and femoral pores on each thigh separated by 23-25 poreless scales (n=3/7); tail with enlarged, strongly keeled, pointed, and spinelike tubercles forming whorls; six tubercles on first six whorl, five tubercles on seventh, and four on eighth whorl; rest of the tail with tubercles present only on the paravertebral rows; median row of subcaudals smooth, roughly pentag onal, and distinctly enlarged. Dorsum pale brown with numerous light blotches and red patches (more prominent in males); single small central black ocellus on occiput and larger one anterior to forelimb insertions; original tail with 8-10 alternating black and light grey bands; ventral surfaces off-white, dark streaks on posterior margin of throat and sternal region (males with underside of hind limbs and margin of belly suffused with yellow).
Description of the holotype. Adult male in good state of preservation except tail tip marginally bent towards right (Fig. 9A, B); SVL 31.2 mm, head short (HL/SVL 0.24), wide (HW/HL 0.68), not strongly depressed (HD/HL 0.42), distinct from neck. Loreal region marginally in flated, canthus rostralis indistinct. Snout marginally larg   er than half the head length (ES/HL 0.52), slightly larg er than 2.5 times eye diameter (ES/ED 2.66); scales on snout and canthus rostralis subcircular, subequal, weakly keeled, twice the size of those on forehead and interorbit al region; scales on forehead similar to those on snout and canthus rostralis except much smaller and weakly coni cal; scales on interorbital region, still smaller, granular; scale occipital, and temporal region with small, smooth granular scales intermixed with slightly enlarged, weakly keeled, conical tubercles (Fig. 10A). Eye small (ED/HL 0.19); with round pupil; supraciliaries short, larger anteri orly; seven interorbital scale rows across narrowest point of frontal bone; 30 or 31 scale rows between left and right supraciliaries at midorbit (Fig. 10A). Earopening deep, oval, small (EL/HL 0.06); eye to ear distance greater than diameter of eye (EE/ED 1.53) (Fig. 10C). Rostral twice as wide (1.4 mm) as high (0.7 mm), incompletely divided dorsally by a strongly developed rostral groove and by an internasal scale for more than half of its height; a sin gle enlarged supranasal on each side, slightly larger than upper postnasal, separated from each other by a small er, elongated internasal on the snout; rostral in contact with supralabial I, nostril, supranasal, internasal, and a lower postnasal on either side; nostrils oval, surrounded by two postnasals, supranasal, and rostral on either side; two postnasals on either side, both almost half the size than supranasal; a single row of scales separate orbit from supralabials (Fig. 10C). Mental enlarged, subtriangular, slightly wider (1.8 mm) than high (1.3 mm); two pairs of postmentals, inner pair roughly rectangular, much shorter (0.6 mm) than mental, separated from each other below mental by a single enlarged median chin shield; inner pair bordered by mental, infralabial I, outer postmental and two enlarged chin shields on either side; outer postmen tals roughly square, marginally smaller (0.5 mm) than in ner pair, bordered by inner postmentals, infralabial I and II, and three enlarged chin shields on either side; two en larged gular scales between left and right outer postmen tals; all chin scales bordering postmentals flat, subcircu lar, smooth, smaller than outermost postmentals; scales on rest of the throat granular, smooth, decreasing in size posteriorly (Fig. 10B). Infralabials bordered below by a row or two of slightly enlarged, much elongated scales, decreasing in size posteriorly and laterally. Six supralabi als up to angle of jaw on left and seven on right side, and six at midorbital position on each side; supralabial I largest, gradually decreasing in size posteriorly; seven infralabials up to angle of jaw, six at midorbital position on either side; infralabial I largest, gradually decreasing in size posteriorly (Fig. 10C). Body relatively slender (BW/AGL 0.43), trunk less than half of SVL (AGL/SVL 0.41) without ventrolater al folds (Fig. 11B, C). Dorsal pholidosis heterogeneous; weakly keeled, weakly conical granular scales intermixed with irregularly arranged rows of enlarged, strongly keeled, conical tubercles which are gradually increasing in size towards flank; last two rows of enlarged tubercles on flank largest, spine-like; approximately six longitudinal rows at midbody; only a few scattered enlarged tubercles in paravertebral region, not forming rows; granular scales on nape slightly smaller than paravertebral granules, still smaller and smooth on occiput (Fig. 11A). Ventral scales much larger than granular scales on dorsum, subcircular, smooth, subequal from chest to vent; scales on precloacal region slightly enlarged; midbody scale rows across bel ly 30; 115 scales from mental to anterior border of cloaca (Fig. 9B). Scales on throat marginally smaller than those on belly, gular region with still smaller, smooth scales, those bordering postmentals enlarged, smooth, subcircu lar and flattened (Fig. 10B). Three femoral pores on left thigh and four on right, separated by 10 poreless scales on left and nine on right side from two precloacal pores, pre cloacal pores separated medially by two poreless scales (Fig. 10D). Scales on palm and soles small, smooth, su bimbricate; scales on dorsal aspects of limbs heteroge neous is size, those on upper arm and thigh large, strongly keeled and imbricates except those near limb insertions that are much smaller, smooth and granular, posterolat eral aspect of thigh with small, smooth to weakly keeled granular scales; dorsal aspect of lower arm and shank with scales smaller than those on upper arm and thigh, strongly keeled, imbricate; ventral aspect of upper arm with small, granular scales, lower arm with slightly larg er, smooth, subimbricate scales; ventral aspect of thigh with scales similar to midbody ventrals, scales on shank slightly larger than those on thigh, smooth, flattened and imbricate (Fig. 9A, B).
Forelimbs and hindlimbs slightly long, slender (LAL/ SVL 0.15); (CL/SVL 0.18); digits long, with a strong, re curved claw, distinctly inflected, distal portions laterally compressed conspicuously. Digits with unpaired lamel lae, separated into a basal and narrower distal series by single enlarged lamella at inflection; basal lamellae se Tail original, entire, subcylindrical, slender, slight ly longer than snoutvent length (TL/SVL 1.39; Fig.   9C-E). Dorsal pholidosis on tail heterogeneous; small, weakly keeled, flattened, and subimbricate scales (be coming larger, elongated and imbricates posteriorly) intermixed with enlarged, strongly keeled, pointed, and spine-like tubercles forming whorls; six tubercles on first six whorl, five tubercles on seventh, and four on eight whorl; rest of the tail (tail tip) with tubercles only on the paravertebral rows (Fig. 9C, E). Scales on tail venter much larger than those on tail dorsal, smooth, flattened, and subimbricate; with a series of three enlarged sub caudal scales of which the median series almost twice the size of adjunct two rows, roughly pentagonal (Fig.  9D). Scales on tail base much smaller, imbricate, and smooth; a single enlarged, conical postcloacal spur on each side (Fig. 9D, E). (Fig. 12A). Dorsal ground colour of head, body, limbs and tail pale brown. Head with indis tinct reddish markings and light grey blotches; anterior edge of brille yellow; upper labials dull white to yellow with dark blotches. Indistinct dark brown preorbital streak; three dark brown postorbital streaks, upper merg ing with light grey blotches on occiput, middle continu ing until ear opening and lower to throat. Single central small black ocellus on occiput followed by a larger white marking and subsequently a larger dark central spot just anterior to forelimb insertions. About four light blotch es on vertebral region between forelimbs and tail base, reddish patches and light spots scattered across dorsum; dorsum of forelimbs and hindlimbs with light blotches and bands, dark blotches speckled with yellow scales, digits with alternating dark and light yellow bands; dorsum of original tail with eight black and nine light  Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu (Fig. 1). Like most other members of the gracilis clade, the new species seems to be diurnal, rupicolous, and locally abundant. At all three collection sites, many individuals (n = >25) were observed active during the daytime (0900-1230 hrs) on rocks below 2 m height in moist deciduous forest patches (Fig. 14A-C (1) or not enlarged (  Etymology. The specific epithet is a toponym for the Ka lakad forest range of KalakadMundanthurai Tiger Re serve in Tirunelveli district of Tamil Nadu, the type and currently only known locality for this species.

Colouration in life
Suggested Common Name. Kalakad dwarf gecko.

Diagnosis.
A smallsized Cnemaspis, snout to vent length less than 33 mm (n=7). Dorsal pholidosis hetero geneous; smooth, subcircular, weakly conical granular scales intermixed with irregularly arranged rows of en larged, smooth, laterally compressed, spinelike scales on flanks; four or five rows of dorsal tubercles at midbody, enlarged scales or tubercles absent in paravertebral region; ventral scales subcircular, smooth, subimbricate, subequal from chest to vent; 28-34 scales across belly at midbody, 122-134 longitudinal scales from mental to cloaca; subdigital scansors smooth, unpaired, unnotched; basal scansors distinctly enlarged, plate like; 11 or 12 lamellae under digit I of manus and pes, 16-19 lamellae under digit IV of manus and 19-22 lamellae under digit IV of pes; males (n=3/7) with a series of 12-14 femo ral pores on each thigh separated medially by 10 or 11 poreless scales; tail dorsum with enlarged, smooth, flat tened scales only in the paravertebral rows, not forming whorls; 16-18 scales in paravertebral rows on either side, only 2-4 enlarged scales in the lateral row on either side; median row of subcaudals smooth, roughly subcircular, and distinctly enlarged. Dorsum greybrown with indis tinct dark bands; single longitudinally placed hourglass shaped black marking anterior to forelimb insertions; tail with ~12 alternating dark and pale bands; underside of body, limbs and tail in males pale yellow with precloacal and femoral region off-white, gular region bright yellow; ventral surfaces in females off-white to cream.
Comparison with members of C. littoralis clade. Cnemaspis kalakadensis sp. nov. is a member of the littoralis clade and can be easily distinguished from all four members of the clade by combination of following dif fering or non-overlapping characters: four or five rows of dorsal tubercles at mid-body restricted to flanks, laterally compressed, spinelike (versus few scattered tubercles on the flank, reduced, subconical in C. flavigularis; a few scattered tubercles on flanks, spine-like tubercles only in the lowermost row on flanks in C. littoralis; conical or spine-like tubercles absent on flanks in C. palakkaden-sis; a few scattered spinose tubercles on the flanks in C. palanica Pal, Mirza, Dsouza and Shanker, 2021); 28-34 scales across belly at midbody (versus 21-23 scales across belly at midbody in C. flavigularis, and 16-18 in C. palanica); 122-134 longitudinal scales from mental to cloaca (versus 104-108 longitudinal scales from mental to cloaca in C. flavigularis, 143-157 in C. littoralis, and 103-106 in C. palanica); 19-22 lamellae under digit IV of pes (versus 16-18 lamellae under digit IV of pes in C. flavigularis, 16-18 in C. littoralis, 14-17 in C. palakkadensis, and 17 or 18 in C. palanica); males with a series 12-14 femoral pores on each thigh separated medially by 10 or 11 poreless scales (versus males with a series of 15-18 femoral pores separated by 14-16 poreless scales in C. littoralis, and 15 or 16 femoral pores separated by at least 15 poreless scales in C. palakkadensis).
Description of the holotype. Adult male in good state of preservation except head marginally bent towards right and fully everted hemipenis on left side (Fig. 15A, B) and digit V of left manus incomplete (indicated by *). SVL 29.6 mm, head short (HL/SVL 0.25), slightly wid er (HW/HL 0.61), not strongly depressed (HD/HL 0.38), distinct from neck. Loreal region marginally inflated, canthus rostralis indistinct. Snout marginally shorter than half the head length (ES/HL 0.46), 2.5 times larger than eye diameter (ES/ED 2.50); scales on snout and canthus rostralis smooth, subcircular, subequal, hemispherical and protrudent, slightly larger than those on forehead and almost twice the size than those on interorbital region and occiput; scales on forehead similar to those on snout and canthus rostralis except slightly smaller and weakly pro trudent; scales on interorbital region, occipital and tem poral region smooth, still smaller, granular (Fig. 16A). Eye small (ED/HL 0.18); with round pupil; supraciliaries short, larger anteriorly; seven interorbital scale rows across narrowest point of frontal bone; 28 or 29 scale rows between left and right supraciliaries at midorbit (Fig. 15A). Earopening deep, oval, small (EL/HL 0.05); eye to ear distance greater than diameter of eye (EE/ED 1.78) (Fig. 16C). Rostral marginally more than twice as wide (1.3 mm) as high (0.6 mm), incompletely divided dorsally by a strongly developed rostral groove for more than half of its height; a single enlarged supranasal on each side, almost twice the size of upper postnasal, sep arated from each other by a smaller, elongated internasal on the snout; rostral in contact with supralabial I, nos tril, supranasal, internasal and a lower postnasal on either side; nostrils subcircular, surrounded by two postnasals, supranasal, and rostral on either side; two postnasals on either side, similar in size to each other, both almost half the size than supranasal; one or two rows of scales sepa rate orbit from supralabials (Fig. 16C). Mental enlarged, subtriangular, slightly wider (1.7 mm) than high (1.2 mm); two pairs of postmentals, inner pair roughly rectan gular, much shorter (0.5 mm) than mental, separated from each other below mental by a single enlarged median chin shield; inner pair bordered by mental, infralabial I, outer postmental and three enlarged chin shields on left side and bordered by mental, infralabial I & II, outer post   mental and three enlarged chin shields on right side; outer postmentals roughly square, much smaller (0.3 mm) than inner pair, bordered by inner postmentals, infralabial I, and two enlarged chin shields on left side and bordered by inner postmentals, infralabial I & II, and two enlarged chin shields on right side; five enlarged, gular scales be tween left and right outer postmentals; all chin scales bordering postmentals hemispherical and protrudent, subcircular, smooth, smaller or equal in size to outermost postmentals; scales on rest of the gular with smooth, sub circular, hemispherical, and protrudent scales decreasing in size posteriorly (Fig. 16B). Infralabials bordered below by a row or two of slightly enlarged, somewhat circular scales, decreasing in size posteriorly and laterally. Eleven supralabials up to angle of jaw on left and 10 on right side and eight at midorbital position on each side; supralabial I largest, gradually decreasing in size posteriorly; 10 infral abials up to angle of jaw on left side and nine or right, and seven at midorbital position on left and six on right side; infralabial I largest, gradually decreasing in size posteri orly (Fig. 16C). Body relatively slender (BW/AGL 0.40), trunk less than half of SVL (AGL/SVL 0.42) without ventrolater al folds (Fig. 17B, C). Dorsal pholidosis heterogeneous; smooth, subcircular, weakly conical granular scales in termixed with irregularly arranged rows of enlarged, smooth, laterally compressed, spine-like scales on flank; approximately five longitudinal rows spine-like scales at midbody; enlarged scales absent on paravertebral region; granular scales on nape similar in size to those on paravertebral region, slightly smaller on occiput (Fig.  17A). Ventral scales much larger than granular scales on dorsum, subcircular, smooth, subimbricate, subequal from chest to vent; midbody scale rows across belly 30; 122 scales from mental to anterior border of cloaca (Fig.  15B). Scales on throat marginally smaller than those on belly, gular region with still smaller, smooth and pro trudent scales, those on chin bordering postmentals, en larged, smooth, subcircular, and protrudent (Fig. 16B). A series of fourteen femoral pores each thigh, separated from each other 11 poreless scales (Fig. 16D). Scales on palm and soles small, smooth, subimbricate; scales on dorsal aspects of limbs heterogeneous is size, those on upper arm and thigh larger, smooth, subcircular and su bimbricate except those near limb insertions are much smaller, smooth and granular; posterolateral aspect of thigh with small, and smooth granular scales; dorsal aspect of lower arm and shank with scales smaller than those on upper arm and thigh, smooth and subimbricate; ventral aspect of upper arm with small, granular scales, lower arm with slightly larger, smooth, subimbricate scales; ventral aspect of thigh with scales similar to mid body ventrals, scales on shank marginally larger than those on thigh, smooth, flattened and subimbricate (Fig.  15A, B).
Digits with unpaired lamellae (except 1-3 paired lamellae at the base), separated into a basal and narrower distal series by single enlarged lamella at inflection, basal lamellae much enlarged, plate like; basal lamellae series: Tail original, entire, subcylindrical, slender, margin ally longer than snoutvent length (TL/SVL 1.12; Fig.  15C-E). Dorsal pholidosis on tail heterogeneous; small, smooth, flattened, subcircular and subimbricate scales (becoming slightly larger and elongated posteriorly) in termixed with enlarged, smooth, flattened scales only in the paravertebral rows, not forming whorls; 16 scales in paravertebral rows on either side, only two enlarged scales in the lateral row on either side (Fig. 15C, E). Scales on tail venter much larger than those on tail dor sum, smooth, flattened, and sub-imbricate; with a series of three enlarged subcaudal scales of which the median series almost twice the size of adjunct two rows, roughly subcircular (Fig. 15D). Scales on tail base small, smooth and subimbricate; a single enlarged, smooth, weakly con ical postcloacal spur on each side (Fig. 15D, E). (Fig. 18A). Dorsal ground colour of head, body, limbs and tail pale grey-brown; finely speckled with darker markings. Anterior edge of brille and snout tip faintly suffused with yellow; labials dull white to yellow, finely speckled with dark markings, dark markings prominent on outer margins. Indistinct dark brown preorbital streak runs from nostril to orbit, postorbital streaks indistinct. A single longitudinal ly placed, hourglassshaped black marking anterior to forelimb insertions, flanked by a poorly defined lighter blotch anteriorly and posteriorly. About four poorly de fined dark bands between axilla and tail base of which the first is most prominent, narrowest in vertebral region and flaring out on flanks, alternating with lighter, elon gate middorsal blotches; dorsum of forelimbs, hind limbs and digits mottled with small dark blotches and bands; dorsum of original tail with about 12 alternating dark and pale bands. Underside of body, limbs and tail pale yellow with a few scattered grey scales, only preclo acal and femoral region off-white; gular region almost entirely covered by a bright yellow patch mottled with a few black scales and very few larger grey blotches. Pupil black, iris greyish.

Colouration in life
Variation and additional information from type series. Mensural, meristic and additional character state data for the type series is given in Tables 9, 10 & 11 respectively. There are two adult males and four adult females ranging in size from 27.4-32.8 mm (Fig. 19). All paratypes resemble the holotype except as follows: supranasals marginally in contact with each other anteri    (0)  Distribution and Natural history. Cnemaspis kalaka densis sp. nov. is currently known only from in and around its type locality (Sengaltheri in Kalakadu reserve forest, ca. 900-1060 m asl.) in Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu (Fig. 1). Individuals of the new species were seen active during the daytime (1030-1600 hrs) on tree trunks (2-5 m above the ground) inside wet evergreen forest patches ( Fig.  20A-C). Cnemaspis kalakadensis sp. nov. appears to be a tree specialist as they were only observed on tree trunks, some individuals were seen effortlessly climbing above approximately 4 m on trees when disturbed. They were observed to be fairly common (n = >15) at all three col lection sites indicating good abundance. The new species were recorded in good numbers only at places with high canopy cover and were seen only during the daytime.

Discussion
South Asian Cnemaspis continue to be named at an un precedented rate from peninsular India, the description of these three new species taking the number of species known from peninsular India to 63, and the Western Ghats alone to 47 (Pal et al. 2021;Khandekar et al. 2020;Uetz et al. 2022). A staggering 71 % of known species (45/63) have been described in the last decade, with almost ev ery area surveyed yielding at least one new species, again highlighting the fact that we are far from uncovering the true diversity of this exceptionally diverse and ancient clade. Each of the three new species is the southernmost representative of its respective clade, with Cnemaspis azhagu sp. nov. being the most southern representative of the genus in mainland India. All three new species are from the Kalakad Mundan thurai Tiger Reserve (KMTR), in southern Tamil Nadu; which forms part of the Agasthyamalai Biosphere Re serve along with the contiguous Neyyar, Peppara and Shendurney Wildlife Sanctuaries in Kerala. Spread across 895 km 2 , KMTR spans an elevational range from <100 m to >1800 m and includes both westfacing and eastfac ing slopes, receiving rainfall during the summer (south west) and winter (northeast) monsoons (see Ganesh et al. 1996). These features result in a high diversity of habi tats, from open scrub to deciduous, tropical dry evergreen and moist evergreen forests. A host of lizard species are endemic to KMTR, including the geckos Cnemaspis beddomei, C. regalis, Dravidogecko douglasadamsi Chait anya, Giri, Deepak, DattaRoy, Murthy & Karanth, 2019, Hemidactylus acanthopholis Mirza and Sanap, 2014and Hemiphyllodactylus peninsularis Agarwal, Bauer, Pal, Srikanthan and Khandekar, 2020(Chaitanya et al. 2019Pal et al. 2021) in addition to the three new species described in this paper.
Four clades of South Asian Cnemaspis are distribut ed south of the Shencottah Gap -the beddomei clade with six known species, and the gracilis, littoralis and monticola clades with a single species each. Cnemaspis azhagu sp. nov. is the 14 th member to be described from the beddomei clade, 11 of which have been described in the last four years, and all of which are diurnal, sexually dimorphic and distributed south of the Palghat Gap in the Western Ghats with narrow distributional ranges (except C. regalis which is known from two localities about 25 km apart from each other) (Cyriac et al. 2018;Sayyed et al. 2019;Pal et al. 2021). Cnemaspis azhagu sp. nov. is most similar to C. aaronbaueri, C. galaxia, C. nigriventris, C. ornata and C. regalis, in superficial morphology and colour pattern; with all these species known from low to mid elevations (< 400 m asl. except C. ornata which is known from 900-1000 m asl.) in dry deciduous forests along the eastern slopes of the southern Western Ghats. The distributional range of Cnemaspis azhagu sp. nov. is 36 km south from its geographically closest congener C. regalis (holotype locality) and just 15 km south from the additional paratype locality in straightline distance. A vast area of suitable habitat along the eastern slopes of the southern Western Ghats remains to be explored and it is almost certain that additional fieldwork in these areas will reveal additional undescribed species within the beddomei clade across both low and high elevational forests.
Cnemaspis mundanthuraiensis sp. nov. is a member of the gracilis clade and is only the third species from this clade known from the Western Ghats after C. gracilis and C. jackieii, the other four species distributed in gran ite boulder habitats in southern India outside the Western Ghats (Khandekar 2019;Khandekar et al. 2019a;Agarw al et al. 2020a;Pal et al.2021). The distributional range of Cnemaspis mundanthuraiensis sp. nov. is approximately 55 km south from its closest described member (C. jackieii) of the clade, which also spans the Shencottah Gap -though these are both low elevation species and much more sampling is needed to understand the role of the gap in speciation. Members of this clade are all diurnal, large ly rupicolous and sexually dimorphic. We take this oppor tunity to correct an error in a previous publication on the gracilis clade; in the original description of Cnemaspis shevaroyensis, Khandekar et al. (2019a) accidentally list ed the same museum number for a female paratype as the holotype (NCBSBH 674) in the species description but provided the correct number in table 5 (NCBSBH 676).
Cnemaspis kalakadensis sp. nov. is the fifth member of the littoralis clade and second to be described from Tamil Nadu state. Three out of four Indian species from this clade were described in the last few years Pal et al. 2021). Cnemaspis kalakadensis sp. nov. extends the known distribution of the clade by 160 km south from its closest described member (C. flavigularis). Indian members of this clade appear to be tree spe cialists and are unique among peninsular Indian conge ners in having much enlarged, platelike basal lamellae. Cnemaspis kalakadensis sp. nov. is superficially similar in morphology to one of the only two Indian Cnemaspis species that lack precise type locality data -C. jerdonii in having similar dorsal pholidosis, presence of spinelike scales on flanks, absence of enlarged lateral tubercles on tail and possessing only femoral pores in males. Howev er, it can be easily distinguished from the latter by having higher number of femoral pores on each thigh (12-14) in males versus eight femoral pores on each thigh in males; maximum SVL 33 mm versus maximum SVL 38.1 mm; scales on lateral sides of tail smooth versus scales on lat eral side of the tail carinate. Based on superficial simi lar morphology, it is likely that the C. jerdonii could be a member of the littoralis clade and most probably de scribed from the Western Ghats of either Kerala or Tamil Nadu.