Research Article |
Corresponding author: Ishan Agarwal ( ishan.agarwal@gmail.com ) Academic editor: Uwe Fritz
© 2023 Akshay Khandekar, Tejas Thackeray, Ayuthavel Kalaimani, Ishan Agarwal.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Khandekar A, Thackeray T, Kalaimani A, Agarwal I (2023) Two new species of South Asian Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from the Gingee Hills, Tamil Nadu, India. Vertebrate Zoology 73: 887-913. https://doi.org/10.3897/vz.73.e110512
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Abstract
We describe two new small-bodied, 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 divergence. 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.
Endemic species, granite boulders, integrative taxonomy, micro-endemism, southern India
South Asian Cnemaspis Strauch, 1887 (SAC) is an ancient, diverse radiation of gekkonid lizards with a Western Ghats origin, including >130 species distributed in peninsular India, northeast India, Sri Lanka, and in and around the Andaman Sea (
Cnemaspis mysoriensis (Jerdon, 1853) was the only species known from peninsular India outside the Western Ghats till the turn of the century, when
The mysoriensis + adii clade began diversifying in the late Oligocene to early Miocene, indicating a long history outside the Western Ghats (
Specimens were hand-collected and euthanized using isoflurane 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 water and transferred to 70% ethanol for long-term storage. Specimens are deposited in the Museum and Research Collection Facility at National Centre for Biological Sciences, Bengaluru (NRC-AA).
We extracted whole genomic DNA from a total of eight tissue samples of the two new species (Table
List of Cnemaspis sequences used in this study (sequences generated in this study marked by *). Museum abbreviations are as follows:
Species | Voucher | Locality | GenBank Accession number |
---|---|---|---|
C. adii |
|
India, Karnataka, Ballari District, Hampi | MT188142 |
C. agarwali | AK 107 | India, Tamil Nadu, Salem District, Sankari |
MK792466
|
C. amboliensis | VG 394 | India, Maharashtra, Sindhudurg District, Amboli | MK792481 |
C. cf. assamensis |
|
India, Meghalaya, Ri Bhoi District, Saiden | MW3674 |
C. avasabinae | CES G416 | India, Andhra Pradesh, Nellore District, Penchalakona | MT773207 |
C. azhagu | NRC-AA-1172 (AKR 671) | India, Tamil Nadu, Tirunelveli District, Thirukurungudi forest range | ON494554 |
C. bangara |
|
India, Karnataka, Kolar District, Paparajanahalli | MT188143 |
C. cavernicola sp. nov. | AK-R 134 | India, Tamil Nadu, Viluppuram District, Pakkamalai RF | OR415867* |
C. cavernicola sp. nov. |
NRC-AA-1286 (AK-R 2339) |
India, Tamil Nadu, Viluppuram District, Pakkamalai RF | OR415868* |
C. cavernicola sp. nov. |
NRC-AA-1287 (AK-R 135) |
India, Tamil Nadu, Viluppuram District, Pakkamalai RF | OR415869* |
C. cavernicola sp. nov. |
NRC-AA-1289 (AK-R 2340) |
India, Tamil Nadu, Viluppuram District, Pakkamalai RF | OR415870* |
C. cavernicola sp. nov. | AK-R 2343 | India, Tamil Nadu, Viluppuram District, Pakkamalai RF | OR415871* |
C. flaviventralis | VG 354 | India, Maharashtra, Sindhudurg District, Amboli | MK792495 |
C. goaensis | VG 385 | India, Karnataka, Uttara Kannada District, Gund | MK792475 |
C. gracilis | CES G385 | India, Kerala, Palakkad District, near Chittur River | MK792465 |
C. graniticola |
|
India, Andhra Pradesh, Chittoor District, Horsley hills | MT188145 |
C. indica | CES L291 | India, Tamil Nadu, Nilgiris | MZ701810 |
C. kallima | AA 82 | Sri Lanka, Matale District, Rattota, Gammaduwa | KY037970 |
C. kolhapurensis | unvouchered | India, Maharashtra, Kolhapur District, Dajipur | MK792501 |
C. koynaensis | CES G349 | India, Maharashtra, Satara District, Humbarli | MK792490 |
C. littoralis | SB 151 | India, Kerala, Thrissur District, Athirappilly Falls | KY038013 |
C. magnifica | unvouchered | India, Karnataka, Hassan District, Sakleshpur | MK792503 |
C. mysoriensis | unvouchered | India, Karnataka, Bangalore Urban District, IISc Campus | MK792474 |
C. mysoriensis | AK 570 | India, Karnataka, Bangalore Urban District, NCBS Campus | MT773208 |
C. mysoriensis | AK 676 | India, Karnataka, Mysore District, Hunsur | MT773209 |
C. mysoriensis | AK 851 | India, Karnataka, Kolar District, Kolar | MT773210 |
C. mysoriensis | AK 852 | India, Karnataka, Kolar District, Kolar | MT773211 |
C. mysoriensis | AK 979 | India, Karnataka, Bangalore Rural District, near Thathaguni | MT773212 |
C. otai | AK 668 | India, Tamil Nadu, Vellore District, Vellore Fort | MT188146 |
C. otai | AK 823 | India, Tamil Nadu, Vellore District, Jawadhu Hills | MT773213 |
C. otai | AK 935 | India, Tamil Nadu, Vellore District, Jawadhu Hills | MT773214 |
C. pakkamalaiensis sp. nov. | AK-R 121 | India, Tamil Nadu, Viluppuram District, Pakkamalai RF | OR415864* |
C. pakkamalaiensis sp. nov. |
NRC-AA-1284 (AK-R 2344) |
India, Tamil Nadu, Viluppuram District, Pakkamalai RF | OR415865* |
C. pakkamalaiensis sp. nov. |
NRC-AA-1285 (AK-R 122) |
India, Tamil Nadu, Viluppuram District, Pakkamalai RF | OR415866* |
C. punctata | AA 80 | Sri Lanka, Matale District, Rattota, Gammaduwa | KY038007 |
C. rishivalleyensis | AK 659 | India, Andhra Pradesh, Chittoor District, Rishi Valley School | MT773218 |
C. rishivalleyensis | AK 660 | India, Andhra Pradesh, Chittoor District, Rishi Valley School | MT773219 |
C. rubraoculus | CES L114 | India, Kerala, Idukki District, Upper Manalar | ON494559 |
C. cf. schalleri | SB 048 | India, Karnataka, Kodagu District, Kumarahalli | KY037995 |
C. stellapulvis | AK 846 | India, Karnataka, Mandya District, Yadiyur | MT773215 |
C. stellapulvis | AK 847 | India, Karnataka, Mandya District, Yadiyur | MT773216 |
C. thayawthadangyi | USNM 595052 | Myanmar, Tanintharyi Region, Myeik Archipelago, southeast side of Linn Lune Kyun | MN104950 |
C. tigris | NRC-AA-1160 (AK 884) | India, Karnataka, Chickballapur District, Kaiwara | OK424590 |
C. tigris | NRC-AA-1159 (AK 885) | India, Karnataka, Chickballapur District, Kaiwara | OK424591 |
C. umashaankeri |
|
India, Karnataka, Chamarajnagar District, BR Hills | OP358455 |
C. umashaankeri |
|
India, Karnataka, Chamarajnagar District, BR Hills | OP358454 |
C. vijayae |
|
India, Karnataka, Kodagu District, Honey Valley Estate | OK424592 |
C. yercaudensis | CES G133 | India, Tamil Nadu, Namakkal District, Kollimalai massif | MK792473 |
C. yercaudensis | AK 767 | India, Tamil Nadu, Salem District, Yercaud massif | MT773217 |
We did not conduct any explicit species delimitation analyses but use the 3.7 % ND2 p-distance cutoff suggested by
Morphological data were collected from a total of 11 specimens of the two new species. We restricted comparisons to the mysoriensis + adii clade as molecular data placed the two lineages within this clade and no other Cnemaspis clades occur close to the Gingee Hills. Comparative morphological data of all other members of the mysoriensis + adii clade were obtained from the specimens listed in Appendix
Morphological analyses were conducted in R 4.1.3 (R Core Team 2018) and used using the following mensural variables: AGL, CL, ES, HL, HW, SVL. We used Thorpe’s (1975) size correction equation that standardizes variables by SVL as implemented in the R package GroupStruct (available at https://github.com/chankinonn/GroupStruct) (Chan and Grismer 2021, 2022). We pooled sexes and used a global mean SVL for the two new species. A principal components analysis (PCA) was then conducted using the ez_pca function to visualize the separation of the two species in multivariate morphospace using the size-corrected data (excluding SVL).
The final ND2 alignment was 1047 base pairs (bp; including 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.
Pairwise uncorrected ND2 sequence divergence within the mysoriensis + adii clade, numbers in bold along diagonal represent intraspecific genetic diversity (– indicates only a single sequence available).
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | |
1 | C. cavernicola sp. nov. | 0.0 | |||||||||
2 | C. pakkamalaiensis sp. nov. | 16.0 | 0.0 | ||||||||
3 | C. adii | 18.9 | 17.2 | – | |||||||
4 | C. avasabinae | 16.0 | 8.7 | 18.5 | – | ||||||
5 | C. mysoriensis | 20.8 | 19.4 | 22.6 | 20.2 | 1.0 | |||||
6 | C. otai | 17.3 | 6.7 | 18.5 | 8.3 | 20.0 | 1.0 | ||||
7 | C. rishivalleyensis | 17.6 | 9.0 | 19.2 | 5.6 | 21.6 | 8.3 | 0.0 | |||
8 | C. stellapulvis | 15.6 | 15.3 | 18.9 | 15.9 | 21.4 | 15.2 | 15.8 | 0.0 | ||
9 | C. tigris | 16.9 | 16.2 | 20.4 | 15.6 | 23.4 | 16.5 | 17.2 | 12.1 | 0.0 | |
10 | C. yercaudensis | 18.5 | 18.3 | 20.2 | 17.9 | 22.6 | 18.3 | 18.4 | 13.4 | 14.5 | 2.0 |
Two factors with eigen values >1 were retained that cumulatively explained 89.2 % of the variation in the dataset. 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 divergent 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.
Cnemaspis otai
NRC-AA-1280 (AK-R 124), adult male (SVL = 29.0 mm), from Pakkamalai Reserve Forest (12.17224°N, 79.31907°E; elevation ca. 400 m asl.), Gingee Hills, Viluppuram district, Tamil Nadu state, India, collected by Akshay Khandekar, Swapnil Pawar and team, on 3rd April 2021.
NRC-AA-1281 (AK-R 123) and NRC-AA-1282 (AK-R 126), adult males, same collection data as holotype; NRC-AA-1283 (AK-R 131) and NRC-AA-1284 (AK-R 2344), adult males, and NRC-AA-1285 (AK-R 122), adult female, from another side of Pakkamalai Reserve Forest (12.16936°N, 79.30614°E; elevation ca. 400 m asl.), Gingee Hills, Viluppuram district, Tamil Nadu state, India, collected by Akshay Khandekar, Ishan Agarwal, Swapnil Pawar and team, on 19th September 2022.
AK-R 121, juvenile specimen, from Pakkamalai Reserve Forest (12.17343°N, 79.31772°E; elevation ca. 400 m asl.), same collection data as holotype.
The specific epithet is a toponym for Pakkamalai, Gingee Hills in Viluppuram district of Tamil Nadu state, the type and only known locality for this species.
Pakkamalai dwarf gecko.
A small-sized Cnemaspis, snout to vent length less than 29 mm (n = 6). 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, weakly keeled, tubercles on each side of 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 longitudinal 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 manus 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, distinctly pointed, conical tubercles forming whorls; a median row of subcaudals smooth, slightly enlarged. Dorsal colouration straw brown with a broad, light mid-dorsal 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 mid-dorsal streak, four pairs between forelimb insertions and tail base, tail with nine alternating light and dark markings.
Cnemaspis pakkamalaiensis sp. nov. can be distinguished from all eight members of the mysoriensis + adii clade on the basis of the following differing or non-overlapping 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 separated on either side by nine or 10 poreless scales from continuous 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 mid-body (versus dorsal pholidosis homogeneous in C. adii, dorsal tubercles irregularly arranged at mid-body in C. avasabinae, 7–10 rows of dorsal tubercles at mid-body in C. stellapulvis); 25–27 ventral scales across belly at mid-body (versus 17–20 ventral scales across belly at mid-body in C. avasabinae, 18 ventral scales across belly at mid-body in C. otai, 20 or 21 ventral scales across belly at mid-body in C. mysoriensis, 20–22 ventral scales across belly at mid-body in stellapulvis, 18–20 ventral scales across belly at mid-body in C. yercaudensis); spine-like tubercles present on flank (versus spine-like tubercles absent on flank in C. adii, and C. avasabinae); a single distinct black dorsal ocellus on mid-dorsal streak just anterior to forelimb insertions (versus distinct black dorsal ocellus just anterior to forelimb insertions absent in C. mysoriensis and C. yercaudensis); a light mid-dorsal streak formed by five or six fused, elongate chain-links that run from occiput to tail base (versus mid-dorsal streak absent in C. adii, a continuous light mid-dorsal 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.
Adult male in good state of preservation except for tail tip slightly bent towards the right, partially everted hemipenis on the left, tail marginally detached just posterior to tail base on the left (Fig.
Cnemaspis pakkamalaiensis sp. nov. (holotype, NRC-AA-1280): 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.
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, weakly keeled, tubercles on each side of flank (Fig.
Scales on dorsal aspect of manus heterogenous, upper arm scales much larger than dorsal granules, strongly 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 smaller than those on dorsum of thigh, subimbricate, weakly 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 mid-body ventrals, smooth, subimbricate on thigh and imbricate on shank (Fig.
Fore-limbs and hind-limbs long, slender (LAL/SVL 0.14); (CL/SVL 0.17); digits long, with a strong, recurved claw, distinctly inflected, distal portions laterally compressed conspicuously. Series of unpaired lamellae on basal portion of digits, separated from unpaired, narrower distal lamellae by a single large scale at the inflection; basal lamellae series: 1-3-3-4-4 (right manus), 3-4-4-7-4 (right pes), 1-3-3-4-4 (left manus; Fig.
Tail entire and original except for extreme tip which is regenerated, subcylindrical, relatively slender, flattened beneath, slightly longer than snout-vent length (TL/SVL 1.18) (Fig.
Dorsal ground colour of head, body, limbs and tail straw-brown; 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 chain-links 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 insertions; 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 light-yellow 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, digits 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 toward the antero-posterior of the pupil.
Mensural, meristic and additional character state data for the paratype series is given in Tables
Mensural (mm) data for the type series of the new species. Abbreviations are listed in Materials and Methods; * = tail incomplete.
Species | Cnemaspis pakkamalaiensis sp. nov. | Cnemaspis cavernicola sp. nov. | |||||||||
Type | Holotype | Paratypes | Holotype | Paratypes | |||||||
Specimen number | NRC-AA-1280 | NRC-AA-1281 | NRC-AA-1282 | NRC-AA-1283 | NRC-AA-1284 | NRC-AA-1285 | NRC-AA-1286 | NRC-AA-1287 | NRC-AA-1288 | NRC-AA-1289 | NRC-AA-1290 |
Sex | Male | Male | Male | Male | Male | Female | Male | Male | Male | Female | Female |
SVL | 29.0 | 27.9 | 27.5 | 26.9 | 28.9 | 28.1 | 32.2 | 32.1 | 28.2 | 34.0 | 30.6 |
TL | 34.3 | 34.5 | 33.4 | 1.9* | 18.1* | 29.5 | 31.2 | 35.5 | 11.0* | 23.7 | 37.1 |
TW | 3.1 | 3.0 | 2.6 | 2.9 | 2.9 | 2.9 | 2.8 | 3.1 | 2.4 | 2.9 | 2.4 |
LAL | 4.1 | 4.3 | 4.2 | 3.8 | 4.1 | 4.5 | 4.8 | 5.1 | 4.1 | 4.7 | 4.2 |
CL | 5.2 | 5.3 | 5.3 | 4.5 | 5.1 | 5.4 | 5.6 | 6.1 | 5.4 | 5.8 | 5.7 |
AGL | 12.3 | 11.5 | 11.7 | 11.8 | 13.3 | 12.1 | 14.2 | 13.3 | 12.7 | 14.6 | 14.6 |
BH | 3.5 | 2.9 | 3.2 | 2.7 | 3.6 | 2.3 | 2.6 | 2.6 | 2.8 | 2.8 | 2.9 |
BW | 6.5 | 5.0 | 5.1 | 6.0 | 5.7 | 5.2 | 5.9 | 5.7 | 5.1 | 6.3 | 5.5 |
HL | 7.7 | 7.7 | 7.6 | 6.9 | 7.8 | 7.5 | 8.3 | 8.0 | 7.1 | 8.4 | 7.6 |
HW | 5.4 | 4.6 | 4.5 | 4.8 | 5.1 | 4.6 | 5.1 | 4.7 | 4.4 | 5.0 | 4.5 |
HD | 3.3 | 2.6 | 2.7 | 2.9 | 3.1 | 2.6 | 2.8 | 3.1 | 2.5 | 2.8 | 2.8 |
ED | 1.6 | 1.5 | 1.6 | 1.5 | 1.5 | 1.5 | 1.6 | 1.6 | 1.5 | 1.7 | 1.4 |
EE | 2.6 | 2.5 | 2.4 | 2.1 | 2.4 | 2.6 | 2.6 | 2.3 | 2.2 | 2.4 | 2.3 |
ES | 3.7 | 3.6 | 3.5 | 2.6 | 3.7 | 3.6 | 3.8 | 4.0 | 3.2 | 3.9 | 3.6 |
EN | 3.0 | 2.8 | 2.7 | 2.7 | 3.0 | 2.8 | 2.9 | 3.1 | 2.6 | 3.3 | 2.8 |
IN | 1.0 | 0.8 | 0.9 | 0.9 | 0.9 | 0.9 | 0.9 | 1.0 | 0.8 | 0.8 | 0.8 |
IO | 1.6 | 1.6 | 1.7 | 1.5 | 2.2 | 1.7 | 2.5 | 1.9 | 1.8 | 2.3 | 2.0 |
EL | 0.4 | 0.4 | 0.7 | 0.3 | 0.5 | 0.6 | 0.4 | 0.6 | 0.4 | 0.6 | 0.3 |
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.
Species | Cnemaspis pakkamalaiensis sp. nov. | Cnemaspis cavernicola sp. nov. | |||||||||
Types | Holotype | Paratypes | Holotype | Paratypes | |||||||
Specimen number | NRC-AA-1280 | NRC-AA-1281 | NRC-AA-1282 | NRC-AA-1283 | NRC-AA-1284 | NRC-AA-1285 | NRC-AA-1286 | NRC-AA-1287 | NRC-AA-1288 | NRC-AA-1289 | NRC-AA-1290 |
Sex | Male | Male | Male | Male | Male | Female | Male | Male | Male | Female | Female |
SL L&R | 7&7 | 7&7 | 8&7 | 7&7 | 7&7 | 8&7 | 7&8 | 8&8 | 8&8 | 8&7 | 8&8 |
IL L&R | 7&7 | 7&7 | 7&7 | 6&6 | 6&7 | 7&7 | 7&7 | 8&8 | 7&7 | 7&7 | 7&8 |
SL M L&R | 6&6 | 6&6 | 7&6 | 7&6 | 7&7 | 6&6 | 6&6 | 7&7 | 7&6 | 7&6 | 6&6 |
IL M L&R | 5&6 | 5&6 | 6&5 | 5&5 | 5&6 | 5&5 | 6&5 | 6&6 | 5&5 | 6&6 | 5&5 |
PVT | irr | irr | irr | irr | irr | irr | abs. | abs. | abs. | abs. | abs. |
DTR | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 4 | 6 | 6 | 4 |
MVSR | 25 | 25 | 27 | 27 | 25 | 27 | 29 | 32 | 29 | 29 | 28 |
VS | 112 | 111 | 105 | 107 | 100 | 105 | 123 | 125 | 116 | 118 | 125 |
DLam1F L&R | 8&8 | 8&8 | 8&8 | 8&7 | 7&7 | 9&9 | 9&9 | 10&10 | 9&10 | 8&9 | 10&10 |
BLam1F L&R | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&2 | 2&2 | 2&2 | 2&2 | 2&2 | 2&2 |
DLam4F L&R | 11&11 | 12&12 | 11&12 | 11&12 | 10&10 | 11&11 | 13&12 | 12&12 | 12&12 | 11&12 | 12*&13 |
BLam4F L&R | 4&4 | 4&4 | 4&4 | 4&4 | 4&4 | 4&4 | 5&5 | 4&4 | 5&4 | 4&4 | 4&6 |
DLam1T L&R | 8&8 | 8&9 | 8&8 | 8&7 | 7&7 | 8&8 | 9&9 | 9&9 | 9&9 | 8&8 | 9&10 |
BLam1T L&R | 3&3 | 2&2 | 1&1 | 2&2 | 2&1 | 2&2 | 2&2 | 2&2 | 2&2 | 2&2 | 2&2 |
DLam4T L&R | 11&12 | 12&12 | 13&13 | 13&13 | 11&11 | 12&12 | 14&14 | 13&13 | 13&12 | 12&* | 14&14 |
BLAM4T L&R | 6&7 | 8&8 | 8&8 | 7&7 | 7&6 | 7&7 | 6&6 | 7&7 | 7&6 | 8&5* | 7&7 |
DLAM5T L&R | 12&12 | 12&11 | 12&13 | 11&12 | 11&11 | 12&12 | 13&13 | 12&13 | 13&13 | 12&* | 13&13 |
BLam5T L&R | 4&4 | 5&5 | 6&6 | 6&6 | 4&3 | 6&6 | 5&6 | 6&6 | 3&4 | 3&3 | 4&5 |
PP L&R | 2 | 2 | 2 | 2 | 2 | abs. | 4 | 3 | 3 | abs. | abs. |
FP L&R | 2&2 | 2&2 | 2&2 | 2&2 | 2&2 | abs. | 1&2 | 2&2 | 2&1 | abs. | abs. |
SB PP&FP L&R | 11&9* | 9&9 | 9&8 | 11&11 | 9&10 | abs. | 8&10 | 10&9 | 9&10 | abs. | abs. |
PCT L&R | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 |
Additional morphological characters for the type series of the new species. abs. = absent; / = data unavailable.
Species | Cnemaspis pakkamalaiensis sp. nov. | Cnemaspis cavernicola sp. nov. | |||||||||
Types | Holotype | Paratypes | Holotype | Paratypes | |||||||
Specimen number | NRC-AA-1280 | NRC-AA-1281 | NRC-AA-1282 | NRC-AA-1283 | NRC-AA-1284 | NRC-AA-1285 | NRC-AA-1286 | NRC-AA-1287 | NRC-AA-1288 | NRC-AA-1289 | NRC-AA-1290 |
Sex | Male | Male | Male | Male | Male | Female | Male | Male | Male | Female | Female |
Anterior extra-brillar fringe scales enlarged (1) or not enlarged (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Ventral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Gular scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Pectoral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Precloacal pores continuous (1) or separated (0) | 1 | 1 | 1 | 1 | 1 | abs. | 1 | 1 | 1 | abs. | abs. |
Precloacal pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | 1 | abs. | 1 | 1 | 1 | abs. | abs. |
Femoral pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | 1 | abs. | 1 | 1 | 1 | abs. | abs. |
Dorsal pholidosis homogeneous (1) or heterogeneous (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Dorsal tubercles keeled (1) or not keeled (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Tubercles linearly arranged (1) or more random (0) | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 1 |
Spine-like tubercles on flank present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Lateral caudal furrows present (1) or absent (0) | 0 | 0 | 0 | / | 0 | 0 | 0 | 0 | 0 | / | 0 |
Subcaudals keeled (1) or smooth (0) | 0 | 0 | 0 | / | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Single median row of keeled subcaudals (1) or smooth (0) | 0 | 0 | 0 | / | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Caudal tubercles encircle tail (1) or not (0) | 1 | 1 | 1 | / | 1 | 1 | 1 | 1 | 1 | / | 1 |
Median subcaudal scale row enlarged (1) or slightly enlarged (0) | 0 | 0 | 0 | / | 0 | 0 | 1 | 1 | 1 | 1 | 1 |
Enlarged femoral scales present (1) or absent (0) | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 0 |
Subtibial scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Cnemaspis pakkamalaiensis sp. nov. is known only from its type locality (Pakkamalai Reserve Forest, Gingee Hills in Viluppuram district, Tamil Nadu), at elevations of ca. 200–400 m asl., though all specimens were collected at ~ 400 m asl. (Fig.
Habitat of Cnemaspis pakkamalaiensis sp. nov., and C. cavernicola sp. nov. at the type locality: 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
NRC-AA-1286 (AK-R 2339), adult male (SVL 32.2 mm), from Pakkamalai Reserve Forest (12.17224°N, 79.31907°E; elevation ca. 400 m asl.), Gingee Hills, Viluppuram district, Tamil Nadu state, India, collected by Akshay Khandekar, Ishan Agarwal, Swapnil Pawar and team, on 19th September 2022.
NRC-AA-1287 (AK-R 135) and NRC-AA-1288 (AK-R 2341), adult males, same collection data as holotype except for NRC-AA-1287 which was collected by Akshay Khandekar, Swapnil Pawar and team on 3rd April 2021; NRC-AA-1289 (AK-R 2340) and NRC-AA-1290 (AK-R 2342), adult females, from near Sri Vanadurgai Amman Temple, Pakkamalai Reserve Forest (12.16991°N, 79.30865°E; elevation ca. 480 m asl.), Gingee Hills, Viluppuram district, Tamil Nadu state, India, same collection data as holotype.
AK-R 134, AK-R 2343, juveniles, from Pakkamalai Reserve Forest (12.16825°N, 79.31971°E; elevation ca. 400 m asl.), same collection data as holotype.
The specific epithet is an adjective formed from the Latin “caverna’ for cave and “cola” meaning inhabitant or dweller, as the species is only known to occur in caves and crevices below large granite boulders.
Cave-dwelling dwarf gecko.
A small-sized Cnemaspis, snout to vent length less than 34 mm (n = 5). Dorsal pholidosis heterogeneous; weakly keeled, granular scales in vertebral and paravertebral 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 spine-like; 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 digit 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 enlarged, strongly keeled, distinctly pointed, conical tubercles forming whorls; a median row of subcaudals smooth, distinctly enlarged. Dorsal colouration grey-brown 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.
Cnemaspis cavernicola sp. nov. can be distinguished from all eight members of the mysoriensis + adii clade on the basis of the following differing or non-overlapping characters: 28–32 scales across belly at mid-body (versus 22–26 ventral scales across belly at mid-body in C. adii, 17–20 ventral scales across belly at mid-body in C. avasabinae, 20 or 21 ventral scales across belly at mid-body in C. mysoriensis, 18 ventral scales across belly at mid-body in C. otai, 25–27 ventral scales across belly at mid-body in C. pakkamalaiensis sp. nov., 23–25 ventral scales across belly at mid-body in C. rishivalleyensis, 20–22 ventral scales across belly at mid-body in C. stellapulvis, 23–25 scales across belly in C. tigris, and 18–20 ventral scales across belly at mid-body in C. yercaudensis); 116–125 longitudinal scales from mental to cloaca (versus 102–114 longitudinal scales from mental to cloaca in C. avasabinae, 93–98 longitudinal scales from mental to cloaca in C. otai, 100–112 longitudinal scales from mental to cloaca in C. pakkamalaiensis sp. nov., 102–112 longitudinal scales from mental to cloaca in C. rishivalleyensis, 90–107 longitudinal scales from mental to cloaca in C. stellapulvis, 91–107 longitudinal scales from mental to cloaca in C. tigris, and 90–105 longitudinal scales from mental to cloaca in C. yercaudensis); a median row of subcaudals distinctly enlarged (versus a median row of subcaudals slightly enlarged in C. avasabinae, C. mysoriensis, C. otai, C. pakkamalaiensis sp. nov., C. rishivalleyensis, C. stellapulvis, C. tigris, and C. yercaudensis); enlarged tubercles in vertebral and paravertebral 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, granular 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 mid-dorsal streak absent (versus a continuous light mid-dorsal streak runs from occiput onto tail base in C. mysoriensis, C. stellapulvis, C. tigris, and C. yercaudensis; a light mid-dorsal streak formed by seven fused, elongate chain-links that runs from occiput to tail base in C. avasabinae, C. otai, C. pakkamalaiensis sp. nov., and C. rishivalleyensis).
Adult male in good state of preservation except extreme tail tip missing, partially everted hemipenis on left side, and a 4.0 mm long incision in sternal region for tissue collection (Fig.
Cnemaspis cavernicola sp. nov. (holotype, NRC-AA-1286): 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.
Body relatively slender (BW/AGL 0.52), trunk less than half of SVL (AGL/SVL 0.41) without ventrolateral folds; three spine-like scales on either side of flank. Dorsal pholidosis heterogeneous; weakly keeled, granular scales on the vertebral and paravertebral region, intermixed with about three irregularly arranged rows of large, weakly keeled, tubercles on each side of flank (Fig.
Scales on dorsal aspect of manus heterogenous, upper arm with scales much larger than dorsal granules, weakly 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, imbricate; scales on palm and sole smooth, flat and subcircular. 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 fairly smaller than those on dorsum of thigh, subimbricate, weakly keeled; dorsal aspect of foot predominantly bearing small, flattened, smooth to weakly keeled, imbricate scales; scales on ventral aspect of thigh and shank more or less equal to those on mid-body ventrals, smooth, subcircular, subimbricate on thigh and imbricate on shank (Fig.
Fore-limbs and hind-limbs moderately long, slender (LAL/SVL 0.14); (CL/SVL 0.17); digits long, with a strong, recurved claw, distinctly inflected, distal portions laterally compressed conspicuously. Series of unpaired lamellae on basal portion of digits, separated from unpaired, narrower distal lamellae by a single large scale at the inflection; basal lamellae series: 2-3-3-5-4 (right manus), 2-4-6-6-6 (right pes), 2-3-4-5-4 (left manus; Fig.
Tail entire except for extreme tip which is missing, original, subcylindrical, relatively slender, flattened beneath, marginally shorter than snout-vent length (TL/SVL 0.96) (Fig.
Dorsal ground colour of head, body, limbs and tail grey-brown; 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; labials yellow with darker bars. Dorsal markings consist of a dark ocellus outlined by brown just anterior to forelimb insertions followed by four off-white blotches from forelimb insertions to tail base; rest of dorsum strongly mottled 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 orange streak toward the posterior of the pupil.
Mensural, meristic and additional character states evaluation data for the paratype series is given in Tables
Cnemaspis cavernicola sp. nov. is known only from its type locality (Pakkamalai Reserve Forest, Gingee Hills in Viluppuram district, Tamil Nadu), at elevations of ca. 400–480 m asl. (Fig.
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 discovery of these two species takes the number of peninsular Indian Cnemaspis known from outside the Western Ghats to 24, all but one of which have been described since the turn of the century
The discovery of two more lineages of SAC from peninsular India outside the Western Ghats, including the relict Cnemaspis cavernicola sp. nov. that has no close extant relatives, is indicative of the long-term persistence of Cnemaspis in refugia provided by boulder habitats (
Tropical Dry Evergreen Forests (TDEFs) are an important habitat type along the southeast coast of India (
Though both new species are sympatric at the type locality, Cnemaspis pakkamalaiensis sp. nov. is morphologically typical of mysoriensis clade species — with a relatively 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 (
We are thankful to the Tamil Nadu Forest Department for permits (permit no. 53/2018), and Teja Bhargava (TN FD) for all his help with coordination. Fieldwork wouldn’t have been possible without the help of Satpal Gangalmale, Swapnil Pawar, Vaibhav Patil and Vivek Waghe. Satpal and Vivek helped with morphological data collection. We thank Uma Ramakrishnan for lab support and Yeshwant HM, Vivek Ramachandran and Tarun Karmakar (NCBS field station and museum facility, Bengaluru) for specimen registrations. Suranjan Karunarathna and two anonymous reviewers provided valuable inputs on the manuscript.
Specimens examined.
Museum abbreviations are as follows: Bombay Natural History Society, Mumbai (
Cnemaspis adii:
Cnemaspis avasabinae: holotype, NCBS-BH754 (adult male); paratypes, NCBS-BH755, NCBS-BH756, (adult males) from Penchalakona, Nellore District, Andhra Pradesh, India.
Cnemaspis mysoriensis: AK569, AK570, AK 571 from National Centre for Biological Sciences (NCBS) campus, Bengaluru, Bengaluru District; AK 676 from Hunsur, Mysore District; AK 851, AK 852 from Kolar, Kolar District;
Cnemaspis otai:
Cnemaspis rishivalleyensis: holotype, NCBS-BH723 (adult male); paratypes NCBS-BH724, NCBS-BH725, NCBS-BH726, NCBS-BH727 (adult females), from Cave Rock Hill, near Rishi Valley School, Chittoor District, Andhra Pradesh, India.
Cnemaspis stellapulvis: holotype, NCBS-BH757 (adult male); paratypes, NCBS-BH759, NCBS-BH664 (adult males), NCBS-BH758, NCBSBH760, NCBS-BH761, NCBS-BH762 (adult females), from near Haddina Kallu Anjaneya Temple, at the base of granite rocky hillock near Yadiyur, Mandya District, Karnataka, India.
Cnemaspis tigris: holotype, NRC-AA-1159 (adult male); paratypes, NRC-AA-1160,
Cnemaspis yercaudensis: NCBS-BH678 (adult male), and NCBS-BH677 (adult female), AK 767, from near Grange resort, Yercaud town, in the Shevaroys, Salem District; NCBS-BH679,