Research Article |
Corresponding author: Ishan Agarwal ( ishan.agarwal@gmail.com ) Academic editor: Anthony Herrel
© 2024 Akshay Khandekar, Tejas Thackeray, 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, Agarwal I (2024) Two new species of the Cnemaspis galaxia complex (Squamata, Gekkonidae) from the eastern slopes of the southern Western Ghats. ZooKeys 1196: 209-242. https://doi.org/10.3897/zookeys.1196.117947
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Two new species allied to Cnemaspis galaxia are described from the eastern slopes of the south Western Ghats, Tamil Nadu, India. Both new species are members of the ornata subclade within the beddomei clade. The two new species can be easily distinguished from all other members of the beddomei clade and each other by a combination of nonoverlapping morphological characters such as small body size, distinct colouration of both sexes, the number of dorsal tubercles around the body, the number or arrangement of paravertebral tubercles, the number of midventral scales across the belly and longitudinal ventral scales from mental to cloaca, besides uncorrected pairwise ND2 and 16S sequence divergence of ≥ 7.4% and ≥ 2.7%. The two new species are distributed from low elevation, deciduous forests of Srivilliputhur, and add to the five previously known endemic vertebrates from Srivilliputhur-Megamalai Tiger Reserve.
Asia, biodiversity hotspot, dwarf geckos, integrative taxonomy, phylogeny, species complex
The beddomei clade is the only clade of South Asian Cnemaspis restricted to the southern Western Ghats (
The most diverse of the three subclades of the beddomei clade is the ornata subclade which includes nine valid species distributed from low elevations on the eastern slopes to high elevations (~ 200–1000+ m a.s.l.) in the Western Ghats south of Srivilliputhur (Fig.
Maximum likelihood phylogeny of the beddomei clade (ND2 + 16S concatenated, 1610 base pairs) with photographs of the new species and C. galaxia (not to scale); numbers at nodes represent bootstrap support/ posterior probability > 70/0.99 (not shown close to terminal nodes). Inset, elevation map of the southern Western Ghats showing type and sampled localities for the ornata subclade.
As part of a project on the lizards of Tamil Nadu, we surveyed the southern Western Ghats from 2018–2022, specifically targeting known species of Cnemaspis as well as potential habitats that had not been previously sampled. We were able to collect most described species of the ornata subclade as well as multiple unnamed divergent lineages, two of which were subsequently described as C. rashidi and C. sundara (
Surveys were conducted in the early morning until a few hours after dark, specimens were observed on rocks, tree trunks, and collected by hand, followed by euthanasia using isoflurane after taking colour photos in life. Liver or tail tissues of at least two individuals of each new species/per locality were collected in molecular grade ethanol and subsequently stored at –20 °C for genetic analysis. Specimens were fixed in 8% formalin for ~ 12–24 h, washed and kept in tap water for ~ 24 h, and transferred to 70% ethanol for long-term storage. Collection permit was issued by the Tamil Nadu Forest Department (see acknowledgements), and collection protocols cleared by an inhouse ethics committee. Specimens are deposited in the Museum and Research Collection Facility at National Centre for Biological Sciences, Bengaluru (NRC-AA).
We generated new sequences for 25 individuals representing five known species and three divergent lineages of the ornata subclade from ~ 18 localities (Fig.
Sequences used in this study. Museum abbreviations are as follows:
Species | Voucher | Locality | ND2 | 16S | Subclade |
---|---|---|---|---|---|
Cnemaspis aaronbaueri | AS 214 | KL: Kollam District, Thenmala | OR714926 | OR708521 | ornata |
C. azhagu | NRC-AA1171 | TN: Tirunelveli District, Thirukurungudi range | – | PP382789 | ornata |
C. azhagu | NRC-AA1172 | TN: Tirunelveli District, Thirukurungudi range | ON494554 | PP382790 | ornata |
C. galaxia | AK-R 1323 | TN: Virudhunagar District, Shenbaga Thopu | PP387688 | PP382791 | ornata |
C. galaxia | AK-R 1324 | TN: Virudhunagar District, Shenbaga Thopu | PP387689 | PP382792 | ornata |
C. galaxia | AK-R 1422 | TN: Virudhunagar District, Vyankateshpuram RF | PP387690 | PP382793 | ornata |
C. galaxia | AK-R 1428 | TN: Virudhunagar District, Shenbaga Thopu | – | PP382794 | ornata |
C. galaxia | AK-R 1470 | TN: Virudhunagar District, Atti Kovil falls | PP387691 | PP382795 | ornata |
C. galaxia | AK-R 1471 | TN: Virudhunagar District, Atti Kovil falls | PP387692 | PP382796 | ornata |
C. galaxia | AK-R 1473 | TN: Virudhunagar District, Atti Kovil falls | PP387693 | PP382797 | ornata |
C. galaxia | CESL 511 | TN: Virudhunagar District, Shenbaga Thopu | MZ701818 | MZ291589 | ornata |
C. nairi | AK-R 894 | TN: Tenkasi District, Courtallam | PP387700 | PP382804 | ornata |
C. nairi | AK-R 895 | TN: Tenkasi District, Courtallam | PP387701 | PP382805 | ornata |
C. nairi | CESL 712 | KL: Kollam District, Shendurney | – | MZ291607 | ornata |
C. nairi | CESL 715 | KL: Kollam District, Shendurney | – | MZ291608 | ornata |
C. nigriventris | AK-R 1265 | TN: Tenkasi District, Courtallam | PP387702 | PP382806 | ornata |
C. nigriventris | AK-R 1275 | TN: Tenkasi District, Courtallam | PP387703 | PP382807 | ornata |
C. nigriventris | CESL 264 | KL: Kollam District, Achankovil RF | MZ291609 | MZ701808 | ornata |
C. nigriventris | CESL 265 | KL: Kollam District, Achankovil RF | MZ291610 | – | ornata |
C. ornata | CESL 276 | TN: Tirunelveli District, Devarmalai Hills | MZ701809 | MZ291613 | ornata |
C. rashidi | AK-R 1439 | TN: Virudhunagar District, Srivilliputhur-Megamalai Tiger Reserve, higher elevations of Shenbaga Thopu | PP387704 | PP382808 | ornata |
C. rashidi | AK-R 1440 | TN: Virudhunagar District, Srivilliputhur-Megamalai Tiger Reserve, higher elevations of Shenbaga Thopu | PP387705 | PP382809 | ornata |
C. rashidi | AK-R 1692 | TN: Theni District, Srivilliputhur-Megamalai Tiger Reserve, Vellimalai | PP387706 | PP382810 | ornata |
C. rashidi | AK-R 1693 | TN: Theni District, Srivilliputhur-Megamalai Tiger Reserve, Vellimalai | PP387707 | PP382811 | ornata |
C. rashidi | AS 226 | TN: Virudhunagar District, Kottamalai estate | OR714921 | – | ornata |
C. rashidi | AS 228 | TN: Virudhunagar District, Kottamalai estate | OR714922 | – | ornata |
C. regalis | CESL 487/ 488 | TN: Tirunelveli District, Kalakad Mundanthurai Tiger Reserve | MZ701816/ MZ701817 | MZ291615 | ornata |
C. sathuragiriensis sp. nov. | AK-R 1510 | TN: Virudhunagar District, Sathuragiri Hills | PP387694 | PP382798 | ornata |
C. sathuragiriensis sp. nov. | AK-R 1511 | TN: Virudhunagar District, Sathuragiri Hills | PP387695 | PP382799 | ornata |
C. sundara | AK-R 1254 | TN: Tenkasi District, Mohan’s resort | PP387708 | – | ornata |
C. sundara | AK-R 1266 | TN: Tenkasi District, Mohan’s resort | PP387709 | – | ornata |
C. sundara |
|
TN: Tenkasi District, Mekkarai | OR714924 | – | ornata |
C. sundara |
|
TN: Tenkasi District, Mekkarai | OR714925 | – | ornata |
C. vangoghi sp. nov. | AK-R 1351 | TN: Virudhunagar District, Srivilliputhur-Megamalai Tiger reserve, Ayyanar Kovil, | PP387696 | PP382800 | ornata |
C. vangoghi sp. nov. | AK-R 1352 | TN: Virudhunagar District, Srivilliputhur-Megamalai Tiger reserve, Ayyanar Kovil, | PP387697 | PP382801 | ornata |
C. vangoghi sp. nov. | AK-R 1373 | TN: Virudhunagar District, Srivilliputhur-Megamalai Tiger reserve, Settur RF | PP387698 | PP382802 | ornata |
C. vangoghi sp. nov. | AK-R 1374 | TN: Virudhunagar District, Srivilliputhur-Megamalai Tiger reserve, Settur RF | PP387699 | PP382803 | ornata |
C. anamudiensis | CESL 232 | KL: Idukki District | MZ701805 | MZ291574 | anamudiensis |
C. nimbus | CESL 252 | KL: Idukki District, Mathikettan Shola NP | MZ701807 | MZ291612 | anamudiensis |
C. wallaceii | CESL 377 | TN: Coimbatore District, Anaimalai, Andiparai Shola | MZ701813 | MZ291619 | anamudiensis |
C. beddomei | CESL 379 | TN: Tirunelveli District, Kalakkad-Mundanthurai Tiger Reserve | MZ701814 | MZ291581 | beddomei |
C. cf. maculicollis | CESL 709 | KL: Kollam District, Shendurney WLS | MZ701825 | MZ291582 | beddomei |
C. rubraoculus | CESL 114 | KL: Idukki District, Periyar Tiger Reserve | ON494559 | MZ291616 | beddomei |
C. smaug | CESL 251 | KL: Idukki District, Mathikettan Shola NP | MZ701806 | MZ291618 | beddomei |
C. chengodumalaensis | CESL 624 | KL: Kozhikode District, Chengodumala | MZ701822 | MZ291584 | wynadensis |
C. kolhapurensis | CESL 868 | MH: Sindhudurg District, Amboli | MZ701829 | MZ291599 | wynadensis |
C. wynadensis | CESL 630 | KL: Wayanad District, Wayanad | MZ701823 | MZ291620 | wynadensis |
We restricted morphological comparisons to the beddomei clade (see Results). Morphological data were collected from 12 specimens of the two new species and from 42 specimens of the beddomei clade including type material of C. azhagu, C. nimbus, C. smaug, and C. wallaceii; type as well as topotypic and/ or additional materials for C. galaxia, C. nigriventris, C. regalis, and C. rubraoculus; and additional materials for C. beddomei, C. nairi, C. rashidi, and C. sundara (all listed in Appendix
We recovered the three subclades of the beddomei clade, anamudiensis, beddomei, and ornata, each of which received high support (Fig.
Uncorrected % sequence divergence within the C. ornata subclade of the beddomei clade. Above diagonal, 16S; below diagonal, ND2; along diagonal in bold, maximum intraspecific ND2 divergence; – indicates no data.
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | ||
---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | C. sathuragiriensis sp. nov. | – | 2.7 | – | 7.3 | 3.4 | 12.8 | 12.0 | 12.6 | 14.2 | 8.1 | – |
2 | C. vangoghi sp. nov. | 10.7 | 2.3 | – | 8.4 | 3.1 | 12.9 | 12.4 | 12.7 | 14.2 | 9.0 | – |
3 | C. aaronbaueri | 19.4 | 17.6 | – | – | – | – | – | – | – | – | – |
4 | C. azhagu | 20.4 | 17.5 | 12.3 | – | 8.8 | 15.0 | 14.6 | 14.5 | 15.6 | 2.8 | – |
5 | C. galaxia | 10.1 | 7.4 | 19.6 | 18.2 | 1.7 | 12.7 | 12.1 | 12.5 | 14.7 | 9.0 | – |
6 | C. nairi | 31.1 | 30.3 | 28.7 | 29.8 | 30.9 | 0.1 | 2.2 | 7.2 | 10.0 | 15.3 | – |
7 | C. nigriventris | 31.2 | 29.8 | 28.8 | 29.9 | 30.1 | 6.3 | 0.6 | 7.7 | 9.7 | 15.0 | – |
8 | C. ornata | 29.4 | 28.4 | 28.1 | 28.8 | 29.0 | 21.6 | 19.0 | – | 4.8 | 15.2 | – |
9 | C. rashidi | 31.6 | 30.4 | 30.3 | 30.2 | 29.8 | 20.3 | 17.7 | 10.8 | 3.0 | 16.5 | – |
10 | C. regalis | 21.3 | 20.0 | 13.1 | 11.1 | 20.6 | 27.8 | 28.2 | 26.6 | 28.0 | – | – |
11 | C. sundara | 17.5 | 15.6 | 16.9 | 18.0 | 15.6 | 28.5 | 28.7 | 28.4 | 29.5 | 19.1 | 3.0 |
Holotype. NRC-AA-8342 (AK-R 1356), adult male, from near Neer Katha Ayyanar Kovil (9.5108°N, 77.4529°E; ca 250 m a.s.l.), Srivilliputhur-Meghamalai Tiger Reserve, Virudhunagar district, Tamil Nadu state, India; collected by Akshay Khandekar, Ishan Agarwal, Swapnil Pawar and team on 16 April 2022. Paratypes. NRC-AA-8343 (AK-R 1351), NRC-AA-8344 (AK-R 1352), adult males, same data as holotype; NRC-AA-8345 (AK-R 1358), adult female, from near Ayyanar Kovil waterfalls (9.5200°N, 77.4478°E; ca 400 m a.s.l.), same data as holotype; NRC-AA-8346 (AK-R 1373), NRC-AA-8347 (AK-R 1374), NRC-AA-8348 (AK-R 1380), adult males, from Settur Reserve Forest (9.4036°N, 77.3721°E; ca 350 m a.s.l.), same data as holotype except collected on 17 April 2022.
A small-sized Cnemaspis, snout to vent length ≤ 34 mm (n = 7). Dorsal pholidosis heterogeneous; smooth to weakly keeled granular scales intermixed with fairly regularly arranged rows of enlarged, weakly keeled, conical tubercles; 10 rows of dorsal tubercles at midbody, 7–14 tubercles in paravertebral rows; ventral scales subequal from chest to vent, smooth, subcircular and subimbricate with rounded end; 29–31 midventral scales across belly, 125–140 longitudinal ventral scales from mental to cloaca; subdigital scansors smooth, unnotched, some divided and others entire, a distinct enlarged metacarpal scale below digit I; 11–14 lamellae under digit I of manus and 11–13 under digit I of pes, 19–22 lamellae under digit IV of manus and 18–25 lamellae under digit IV of pes; males with continuous series of six or seven precloacal pores (n = 6); scales on non-regenerated tail dorsum heterogeneous; small, smooth, subcircular, flattened, subimbricate scales intermixed on anterior one third portion with enlarged, weakly keeled, and weakly conical tubercles forming seven whorls; six tubercles on first three whorl, four tubercles on fourth to seventh whorls, only a pair of paravertebral tubercles each on eighth to 11th whorls; rest of the tail lacking enlarged tubercles; median row of subcaudals smooth, roughly rectangular, distinctly enlarged, with condition of two enlarged scales alternating with a divided scale. Males with ochre anterior 1/2 of body, single central black dorsal ocellus on neck, a white ocellus on ventrolateral side of neck and one on throat posterior to jaw, venter off-white with dark throat, tail unbanded, females and juveniles brown, juveniles with indistinct mid-dorsal streak.
Cnemaspis vangoghi sp. nov. can be easily distinguished from all 16 members of the beddomei clade as well as from C. boiei by a combination of the following differing or non-overlapping characters: A small-sized Cnemaspis, snout to vent length ≤ 34 mm (vs medium-sized Cnemaspis, snout to vent length 40–49 mm in C. nairi, C. nimbus, C. ornata, C. rashidi, C. rubraoculus, and C. wallaceii; large-sized Cnemaspis, snout to vent length > 50 mm in C. anamudiensis, C. beddomei, C. maculicollis, and C. smaug; snout to vent length ≤ 38 mm in C. azhagu, C. boiei, and C. nigriventris); ten rows of dorsal tubercles at midbody (vs only a few enlarged scattered tubercles at midbody dorsum in C. anamudiensis, two or three rows of dorsal tubercles at midbody in C. azhagu, eight in C. galaxia, 16–18 in C. nairi, 13 or 14 in C. nigriventris, 12–14 in C. nimbus and C. ornata, 7–9 in C. regalis, 19–22 in C. smaug, six in C. sundara, 14 or 15 in C. wallaceii); 125–140 longitudinal ventral scales from mental to cloaca (vs 151–171 longitudinal ventral scales from mental to cloaca in C. azhagu, 154–161 in C. beddomei, 153–159 in C. galaxia, 143–147 in C. nairi, 154–159 in C. nigriventris, 157–165 in C. ornata, 170–172 in C. rashidi, 148–154 in C. regalis, 142–150 in C. smaug, 156–160 in C. sundara, 154–156 in C. wallaceii); 7–14 tubercles in paravertebral rows (vs paravertebral tubercles either absent or irregular in C. anamudiensis, C. azhagu, and C. sundara, 18 or 19 tubercles in paravertebral rows in C. aaronbaueri and C. beddomei, 16 or 17 in C. nimbus, 21–23 in C. ornata, 27–30 in C. smaug, 18–20 in C. wallaceii); 29–31 midventral scales across belly (vs 34–44 midventral scales across belly in C. azhagu, 32 or 33 in C. nairi, 38–40 in C. nigriventris, 26 or 27 in C. nimbus, 40–44 in C. regalis, 33–37 in C. rubraoculus, 35 or 36 in C. sundara); a distinct white ocellus on ventrolateral sides of neck present in males (vs white ocellus on ventrolateral sides of neck absent in C. aaronbaueri, C. anamudiensis, C. azhagu, C. beddomei, C. maculicollis, C. nimbus, C. nimbus, C. regalis, C. rubraoculus, C. smaug, C. wallaceii); tail unbanded (tail distinctly banded in C. nairi, C. nigriventris, C. ornata, C. rashidi, C. smaug, C. sundara). Cnemaspis vangoghi sp. nov. is diagnosed against the other new species as part of its description below.
Adult male in good state of preservation except tail marginally bent towards left and tip is missing, hemipenis partially everted on right and fully on left side, and a 3.1 mm long incision in sternal region for tissue collection (Fig.
Body relatively slender (BW/AGL 0.37), trunk < 1/2 of SVL (AGL/SVL 0.42) without spine-like tubercles on flank (Fig.
Scales on palm and soles granular, smooth, rounded, and flattened, a distinct enlarged metacarpal scale on palm below digit I; scales on dorsal aspects of limbs heterogeneous in shape and size; scales on upper arm and thigh much larger than granular scales on body dorsum, elongate, subimbricate with pointed ends; scales on lower arm and shank granular, similar in size to granular scales on body dorsum, smooth, rounded, gradually becoming larger, flattened and subimbricate anterolaterally and posteriorly, largest on anterolateral aspect of the hands and feet; scales on ventral aspect of upper arm smooth, granular, much smaller than granular scales on body dorsum, scales on ventral aspect of lower arm with much larger scales than those on upper arm, smooth, subcircular and flattened scales; ventral aspect of thigh and shank with enlarged, smooth, flattened, subimbricate scales, much larger than midventrals (Fig.
Tail original, subcylindrical, slender, not entire, tail tip is detached and missing, TL = 27.2 mm (Fig.
(Fig.
(Figs
Mensural (mm) data for the new species. Abbreviations are listed in Materials and methods, * = tail incomplete.
Museum number | Cnemaspis vangoghi sp. nov. | Cnemaspis sathuragiriensis sp. nov. | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
Holotype | Paratypes | Holotype | Paratypes | |||||||||
NRC-AA-8342 | NRC-AA-8343 | NRC-AA-8344 | NRC-AA-8345 | NRC-AA-8346 | NRC-AA-8347 | NRC-AA-8348 | NRC-AA-8349 | NRC-AA-8350 | NRC-AA-8351 | NRC-AA-8352 | NRC-AA-8353 | |
Sex | Male | Male | Male | Female | Male | Male | Subadult male | Male | Male | Male | Male | Subadult female |
SVL | 32.1 | 33.6 | 33.6 | 32.9 | 31.3 | 32.8 | 28.6 | 32.8 | 32.9 | 31.2 | 33.0 | 26.7 |
TL | 27.2* | 16.9* | 29.8* | 33.7* | 22.2* | 26.9* | 35.4 | 37.6 | 38.6 | 32.1* | 3.1* | 1.9* |
TW | 3.2 | 3.1 | 3.2 | 3.4 | 3.3 | 3.0 | 2.7 | 3.0 | 2.7 | 3.6 | 3.2 | 2.3 |
FL | 4.7 | 4.6 | 5.0 | 5.2 | 4.9 | 4.7 | 4.6 | 4.8 | 4.9 | 4.9 | 5.1 | 4.0 |
CL | 6.2 | 5.9 | 6.3 | 6.2 | 6.3 | 6.2 | 5.3 | 6.5 | 6.2 | 6.0 | 6.7 | 4.9 |
AGL | 13.7 | 13.0 | 12.8 | 13.2 | 12.6 | 13.6 | 11.5 | 12.9 | 14.2 | 12.4 | 12.7 | 11.1 |
BH | 3.5 | 3.7 | 3.4 | 4.2 | 3.6 | 3.4 | 3.7 | 3.6 | 3.5 | 3.3 | 4.2 | 3.9 |
BW | 5.2 | 7.0 | 5.9 | 6.3 | 6.2 | 5.7 | 5.6 | 6.5 | 5.6 | 6.1 | 6.9 | 5.1 |
HL | 8.2 | 8.5 | 8.8 | 9.1 | 8.4 | 8.8 | 7.0 | 9.0 | 8.1 | 8.5 | 8.9 | 6.6 |
HW | 5.6 | 6.1 | 5.9 | 5.9 | 5.8 | 5.7 | 5.4 | 6.7 | 5.9 | 6.1 | 5.9 | 4.7 |
HD | 3.3 | 3.5 | 3.7 | 4.6 | 3.78 | 3.7 | 3.3 | 4.7 | 3.4 | 3.7 | 3.8 | 2.7 |
ED | 1.6 | 1.6 | 1.9 | 1.8 | 1.6 | 1.5 | 1.4 | 1.8 | 1.7 | 1.8 | 1.9 | 1.5 |
EE | 2.6 | 2.9 | 2.7 | 2.8 | 2.8 | 2.9 | 2.3 | 2.9 | 2.6 | 2.7 | 2.7 | 2.3 |
ES | 4.0 | 3.9 | 4.3 | 4.3 | 3.9 | 4.2 | 3.3 | 4.0 | 4.0 | 4.0 | 4.3 | 3.1 |
EN | 3.1 | 3.1 | 3.4 | 3.2 | 3.1 | 3.3 | 2.6 | 3.2 | 3.0 | 3.2 | 3.5 | 2.5 |
IN | 1.1 | 1.1 | 1.1 | 1.2 | 1.1 | 1.0 | 0.8 | 1.1 | 1.0 | 1.1 | 1.2 | 1.0 |
IO | 2.1 | 2.3 | 2.5 | 2.6 | 2.5 | 2.1 | 2.1 | 2.4 | 2.4 | 1.8 | 2.3 | 1.9 |
EL | 0.5 | 0.6 | 0.5 | 0.6 | 0.8 | 0.5 | 0.5 | 0.5 | 0.7 | 0.6 | 0.7 | 0.4 |
Meristic data for the new species. Abbreviations are listed in Materials and methods, * = lamellae damaged, L&R = left & right, A = absent.
Museum number | Cnemaspis vangoghi sp. nov. | Cnemaspis sathuragiriensis sp. nov. | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
Holotype | Paratypes | Holotype | Paratypes | |||||||||
NRC-AA-8342 | NRC-AA-8343 | NRC-AA-8344 | NRC-AA-8345 | NRC-AA-8346 | NRC-AA-8347 | NRC-AA-8348 | NRC-AA-8349 | NRC-AA-8350 | NRC-AA-8351 | NRC-AA-8352 | NRC-AA-8353 | |
SL L&R | 9&9 | 9&8 | 9&9 | 8&8 | 7&7 | 9&10 | 8&7 | 8&8 | 8&8 | 8&8 | 8&8 | 8&8 |
IL L&R | 8&7 | 6&7 | 7&7 | 8&7 | 6&7 | 7&7 | 8&8 | 7&7 | 7&7 | 7&7 | 7&7 | 7&7 |
SL M L&R | 5&5 | 5&5 | 6&5 | 5&6 | 5&5 | 5&6 | 5&5 | 5&5 | 5&5 | 5&5 | 5&5 | 5&5 |
IL M L&R | 4&5 | 4&4 | 5&5 | 4&5 | 5&4 | 4&5 | 4&4 | 5&5 | 4&4 | 4&4 | 4&4 | 5&4 |
PVT L&R | 12&14 | 7&7 | 7&8 | 7&9 | 12&12 | 11&13 | 7&7 | A | A | irr | A | irr |
DTR | 10 | 10 | 10 | 10 | 10 | 10 | 10 | 6 | 7 | 8 | 6 | 7 |
MVSR | 31 | 31 | 30 | 30 | 29 | 29 | 31 | 30 | 28 | 30 | 28 | 29 |
VS | 138 | 125 | 137 | 126 | 134 | 140 | 134 | 132 | 131 | 137 | 130 | 131 |
DLAMF1 L&R | 11&11 | 10&10 | 11&11 | 9&10 | 10&11 | 12&12 | 12&12 | 10&10 | 10&10 | 10&10 | 10&10 | 11&12 |
BLAMF1 L&R | 2&1 | 2&2 | 2&2 | 2&2 | 2&2 | 2&2 | 2&1 | 1&1 | 1&1 | 3&3 | 2&2 | 1&1 |
DLAMF4 L&R | 15&15 | 13&13 | 14&15 | 13&14 | 14&14 | 16&10* | 15&15 | 14&14 | 15&14 | 15&15 | 13&13 | 15&15 |
BLAMF4 L&R | 6&6 | 6&6 | 6&6 | 6&6 | 6&6 | 6&6 | 7&6 | 6&6 | 3&4 | 6&6 | 6&6 | 6&5 |
DLAMT1 L&R | 10&10 | 9&9 | 10&10 | 9&9 | 10&10 | 11&11 | 11&10 | 10&10 | 10&10 | 9&9 | 9&9 | 10&10 |
BLAMT1 L&R | 2&2 | 2&2 | 2&2 | 2&2 | 2&2 | 2&2 | 2&2 | 1&1 | 2&2 | 2&2 | 2&2 | 2&2 |
DLAMT4 L&R | 16&16 | 15&15 | 15&15 | 14&15 | 14&14 | 17&16 | 16&17 | 15&16 | 16&16 | 16&16 | 14&14 | 16&16 |
BLAMT4 L&R | 8&7 | 7&6 | 6&6 | 4&6 | 6&7 | 8&8 | 7&7 | 8&8 | 7&7 | 8&8 | 9&9 | 7&7 |
DLAMT5 L&R | 16&16 | 15&14 | 15&15 | 15&14 | 15&15 | 17&16 | 16&17 | 15&16 | 15&16 | 15&16 | 13&14 | 16&16 |
BLAMT5 L&R | 3&4 | 2&2 | 2&2 | 2&2 | 3&3 | 2&2 | 2&2 | 2&2 | 2&2 | 2&2 | 2&3 | 2&2 |
TLAMF1 L&R | 13&12 | 12&12 | 13&13 | 11&12 | 12&13 | 14&14 | 14&13 | 11&11 | 11&11 | 13&13 | 12&12 | 12&13 |
TLAMF4 L&R | 21&21 | 19&19 | 20&21 | 19&20 | 20&20 | 22&16* | 22&21 | 20&20 | 18&18 | 21&21 | 19&19 | 21&20 |
TLAMT1 L&R | 12&12 | 11&11 | 12&12 | 11&11 | 12&12 | 13&13 | 13&12 | 11&11 | 12&12 | 11&11 | 11&11 | 12&12 |
TLAMT4 L&R | 24&23 | 22&21 | 21&21 | 18&21 | 20&21 | 25&24 | 23&24 | 23&24 | 23&23 | 24&24 | 23&23 | 23&23 |
TLAMT5 L&R | 19&20 | 17&16 | 17&17 | 17&16 | 18&18 | 19&18 | 18&19 | 17&18 | 17&18 | 17&18 | 15&17 | 18&18 |
PP | 6 | 7 | 7 | A | 7 | 7 | 7 | 7 | 8 | 7 | 8 | A |
SB PP | A | A | A | A | A | A | A | A | A | A | A | A |
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 | 1&1 |
Additional morphological characters of the new species. A = absent, / = data unavailable.
Museum number | Cnemaspis vangoghi sp. nov. | Cnemaspis sathuragiriensis sp. nov. | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
Holotype | Paratypes | Holotype | Paratypes | |||||||||
NRC-AA-8342 | NRC-AA-8343 | NRC-AA-8344 | NRC-AA-8345 | NRC-AA-8346 | NRC-AA-8347 | NRC-AA-8348 | NRC-AA-8349 | NRC-AA-8350 | NRC-AA-8351 | NRC-AA-8352 | NRC-AA-8353 | |
Anterior extra-brillar fringe scales enlarged (0) or not enlarged (1) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Occipital ocellus present (0) or absent (1) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Dorsal pholidosis homogeneous (0) or heterogeneous (1) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Dorsal tubercles weakly keeled (0) or smooth (1) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Tubercles linearly arranged (0) or more random (1) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 1 | 1 |
Spine-like tubercles on flank present (0) or absent (1) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Gular scales keeled (0) or smooth (1) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Pectoral scales keeled (0) or smooth (1) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Ventral scales keeled (0) or smooth (1) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Precloacal pores continuous (0) or separated (1) | 0 | 0 | 0 | A | 0 | 0 | 0 | 0 | 0 | 0 | 0 | A |
Precloacal pores elongate (0) or round (1) | 0 | 0 | 0 | A | 0 | 0 | 0 | 0 | 0 | 0 | 0 | A |
Enlarged femoral scales present (0) or absent (1) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Subtibial scales keeled (0) or smooth (1) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Lateral caudal furrows present (0) or absent (1) | 1 | / | 1 | / | 1 | 1 | 1 | 1 | 1 | 1 | / | / |
Caudal tubercles encircle tail (0) or not (1) | 1 | / | 1 | / | 1 | 1 | 1 | 1 | 1 | 1 | / | / |
Subcaudals keeled (0) or smooth (1) | 1 | / | 1 | / | 1 | 1 | 1 | 1 | 1 | 1 | / | / |
Median subcaudal scale row not enlarged (0) or enlarged (1) | 1 | / | 1 | / | 1 | 1 | 1 | 1 | 1 | 1 | / | / |
The new species is strongly sexually dimorphic and also shows ontogenetic colour variation (Fig.
The specific epithet is a patronym for Dutch painter Vincent Van Gogh (1853–1890). The colouration of the new species is reminiscent of one of Van Gogh’s most iconic paintings, The Starry Night. Suggested common name is Van Gogh’s starry dwarf gecko.
Cnemaspis vangoghi sp. nov. is known only from two closely spaced localities (Ayyanar Kovil and Settur Reserve Forest, both in Meghamalai-Srivilliputhur Tiger Reserve, Tamil Nadu) within < 15 km straight line distance (Fig.
Holotype. NRC-AA-8349 (AK-R 1510), adult male, from near Sathuragiri entry point (9.7093°N, 77.6307°E; ca 250 m a.s.l.), Sathuragiri, Virudhunagar district, Tamil Nadu state, India; collected by Akshay Khandekar, Ishan Agarwal, Swapnil Pawar and team on 26 April 2022. Paratypes. NRC-AA-8350 (AK-R 1511), NRC-AA-8351 (AK-R 1512), adult males, same data as holotype; NRC-AA-8352 (AK-R 1513), adult male, NRC-AA-8353 (AK-R 1515), subadult female, from near Vazhukkuparai Saptur Reserve Forest (9.7174°N, 77.6244°E; ca 400 m a.s.l.), Sathuragiri; same data as holotype.
A small-sized Cnemaspis, snout to vent length ≤ 33 mm (n = 5). Dorsal pholidosis heterogeneous; smooth to weakly keeled granular scales intermixed with irregularly arranged rows of enlarged, weakly keeled, conical tubercles; 6–8 rows of dorsal tubercles at midbody, paravertebral tubercles either absent or irregular; ventral scales subequal from chest to vent, smooth, subcircular and subimbricate with rounded end; 28–30 midventral scales across belly, 130–137 longitudinal ventral scales from mental to cloaca; subdigital scansors smooth, unnotched, some divided and others entire, a distinct enlarged metacarpal scale below digit I; 11–13 lamellae under digit I of manus and 11 or 12 under digit I of pes, 18–21 lamellae under digit IV of manus and 23 or 24 lamellae under digit IV of pes; males with continuous series of seven or eight precloacal pores (n = 4); scales on non-regenerated tail dorsum heterogeneous; small, smooth, subcircular, flattened, subimbricate scales intermixed on anterior one third portion with enlarged, weakly keeled, and weakly conical tubercles forming eight whorls; six tubercles on first whorl, four tubercles on second to fourth whorls, only a pair of paravertebral tubercles each on fifth to eighth whorls; rest of the tail lacking enlarged tubercles; median row of subcaudals smooth, roughly subcircular, distinctly enlarged than rest, with condition of two enlarged scales alternating with a divided scale. Males with ochre dorsum, single central black dorsal ocellus on neck, a white ocellus on ventrolateral side of neck and one on throat posterior to jaw, venter off-white with dark throat, tail unbanded, females and juveniles brown with a prominent mid-dorsal streak.
Cnemaspis sathuragiriensis sp. nov. can be easily distinguished from all 16 members of the beddomei clade as well as from C. boiei by a combination of the following differing or non-overlapping characters: A small-sized Cnemaspis, snout to vent length ≤ 33 mm (vs medium-sized Cnemaspis, snout to vent length 40–49 mm in C. nairi, C. nimbus, C. ornata, C. rashidi, C. rubraoculus and C. wallaceii; large-sized Cnemaspis, snout to vent length > 50 mm in C. anamudiensis, C. beddomei, C. maculicollis, and C. smaug; snout to vent length ≤ 38 mm in C. azhagu, C. boiei, and C. nigriventris); 6–8 rows of dorsal tubercles at midbody (vs only a few enlarged scattered tubercles at midbody dorsum in C. anamudiensis, two or three rows of dorsal tubercles at midbody in C. azhagu, 16–18 in C. nairi, 13 or 14 in C. nigriventris, 12–14 in C. nimbus and C. ornata, 19–22 in C. smaug, 10 in C. vangoghi sp. nov., 14 or 15 in C. wallaceii); 130–137 longitudinal ventral scales from mental to cloaca (vs 151–171 longitudinal ventral scales from mental to cloaca in C. azhagu, 154–161 in C. beddomei, 153–159 in C. galaxia, 143–147 in C. nairi, 154–159 in C. nigriventris, 157–165 in C. ornata, 170–172 in C. rashidi, 148–154 in C. regalis, 142–150 in C. smaug, 156–160 in C. sundara, 154–156 in C. wallaceii); paravertebral tubercles either absent or irregular (vs 18 or 19 tubercles in paravertebral rows in C. aaronbaueri and C. beddomei, 16 or 17 in C. nimbus, 21–23 in C. ornata, 27–30 in C. smaug, 7–14 in C. vangoghi sp. nov., 18–20 in C. wallaceii); 28–30 midventral scales across belly (vs 34–44 midventral scales across belly in C. azhagu, 32 or 33 in C. nairi, 38–40 in C. nigriventris, 26 or 27 in C. nimbus, 40–44 in C. regalis, 33–37 in C. rubraoculus, 35 or 36 in C. sundara); a distinct white ocellus on ventrolateral sides of neck present in males (vs white ocellus on ventrolateral sides of neck absent in C. aaronbaueri, C. anamudiensis, C. azhagu, C. beddomei, C. maculicollis, C. nimbus, C. regalis, C. rubraoculus, C. smaug, C. wallaceii); tail unbanded (tail distinctly banded in C. nairi, C. nigriventris, C. ornata, C. rashidi, C. smaug, C. sundara).
Adult male in good state of preservation except tail marginally bent towards right, hemipenis fully everted on right, and a 4.1 mm long incision in sternal region for tissue collection (Fig.
Body relatively slender (BW/AGL 0.50), trunk < 1/2 of SVL (AGL/SVL 0.39) without spine-like tubercles on flank (Fig.
Scales on palm and soles granular, smooth, rounded, and flattened, a distinct enlarged metacarpal scale on palm below digit I; scales on dorsal aspects of limbs heterogeneous in shape and size; scales on upper arm and thigh much larger than granular scales on body dorsum, smooth and a few feebly keeled, slightly elongate, subimbricate with weakly pointed ends; scales on lower arm and shank granular, similar in size to granular scales on body dorsum, smooth, rounded, gradually becoming larger, flattened and subimbricate anterolaterally and posteriorly, largest on anterolateral aspect of the hands and feet; scales on ventral aspect of upper arm smooth, granular, much smaller than granular scales on body dorsum, scales on ventral aspect of lower arm with much larger scales than those on upper arm, smooth, subcircular and flattened scales; ventral aspect of thigh and shank with enlarged, smooth, flattened, subimbricate scales, much larger than midventrals (Fig.
Tail mostly original with regenerated tip, entire, subcylindrical, slender, marginally longer than body (TL/SVL = 1.14) (Fig.
(Fig.
(Figs
The new species is strongly sexually dichromatic and shows ontogenetic colour variation (Fig.
The specific epithet is a toponym for the type locality of the new species, Sathuragiri mountain in Srivilliputhur-Megamalai Tiger Reserve (SMTR), Virudhunagar District, Tamil Nadu. Suggested Common name is Sathuragiri dwarf gecko.
Cnemaspis sathuragiriensis sp. nov. is known only from its type locality (Sathuragiri hills in Virudhunagar district, Tamil Nadu) between elevations of 250–400 m a.s.l. on eastern slopes of the Western Ghats (Fig.
1 | White ocellus on ventrolateral sides of neck in males | 2 |
– | No white ocellus on ventrolateral sides of neck in males | 7 |
2 | Original tail unbanded | 3 |
– | Original tail banded | 5 |
3 | ≤ 140 longitudinal scales from mental to cloaca | 4 |
– | 153–159 longitudinal scales from mental to cloaca | C. galaxia |
4 | Entire dorsum ochre in adult males | C. sathuragiriensis sp. nov. |
– | Anterior 1/2 of dorsum ochre in adult males | C. vangoghi sp. nov. |
5 | 150–165 longitudinal scales from mental to cloaca | 6 |
– | 143–147 longitudinal scales from mental to cloaca | C. nairi |
– | 170–172 longitudinal scales from mental to cloaca | C. rashidi |
6 | Paravertebral tubercles absent | C. sundara |
– | 15 or 16 paravertebral tubercles | C. nigriventris |
– | 21–23 paravertebral tubercles | C. ornata |
7 | Paravertebral tubercles irregular or absent | C. azhagu |
– | 18 or 19 paravertebral tubercles | C. aaronbaueri |
– | 15 or 16 paravertebral tubercles | C. regalis |
The ornata subclade now has 11 known valid species (including the two new species described in this paper) in a small geographic area spanning < 1° longitude and 1.5° latitude. At the southern extreme of the Western Ghats, the region is incredibly heterogeneous, with altitudinal variation from close to sea level to > 1,500 m a.s.l. and strong east-west gradients in total annual precipitation and seasonality. Habitats range from thorny scrub forest on the lower eastern slopes of the mountains to evergreen forest at higher elevations and on the western slopes. This subclade is distributed across the Shencottah Gap (SG), a relatively low elevation pass through the Western Ghats. All 11 members of the clade are strongly sexually dichromatic, and sexual selection may at least in part be a driver of the high diversity in this clade, as has been speculated for members of the C. gracilis clade (
Though sampling of the ornata subclade likely remains incomplete as this vast mountainous landscape has a number of higher elevations we could not access in our rapid surveys, there are some geographic patterns that emerge based on available data. The only two high elevation species are the sister pair C. ornata + C. rashidi that are distributed north of the SG, together forming the sister taxon to the low elevation species C. nairi + C. nigriventris that are distributed around the SG. These two sister pairs are deeply divergent from one another, indicative of potentially more undiscovered species in the intervening areas. The subclade containing C. aaronbaueri, C. azhagu and C. regalis is distributed entirely south of the SG and mainly on the eastern slopes, while the final subclade includes the low to mid elevation C. sundara which is distributed close to the SG, and the galaxia complex with three low elevation species in Srivilliputhur. Large sampling gaps exists between the distribution of C. sundara and C. galaxia as well as between C. nairi and C. regalis.
This last section is a note on violations of Principle 2 of the Code of Ethics prescribed by The Code (appendix A; Anonymous 1999) which states
“A zoologist should not publish a new name if he or she has reason to believe that another person has already recognized the same taxon and intends to establish a name for it (or that the taxon is to be named in a posthumous work). A zoologist in such a position should communicate with the other person (or their representatives) and only feel free to establish a new name if that person has failed to do so in a reasonable period (not less than a year).”
One of the authors of Cnemaspis rashidi accompanied us in the field in 2022 when we collected the then unnamed and distinctively coloured species, and multiple co-authors including the first author were aware that we were working in Tamil Nadu on Cnemaspis among other lizards (
We thank the Tamil Nadu Forest Department for permits to carry out this study (permit no. 53/2018), Teja Bhargava (former DFO, Jawadhu) for his help and co-ordination throughout fieldwork, and the then Field Director SMTR Deepak Bilgi. Fieldwork assistance was provided by Swapnil Pawar, Vaibhav Patil, Satpal Gangalmale, Vivek Waghe, and Satheesh Kumar. We are thankful to Tarun Karmakar (NCBS field station and museum facility, Bengaluru for help with specimen registration, Uma Ramakrishnan for lab support at NCBS, Navendu Page for help with forest classification, R. Chaitanya for edits in the discussion, and Azhar Hotel Devaki for sustaining us on a high protein diet through our time at Srivilliputhur.
The authors have declared that no competing interests exist.
The research project was cleared by an in-house animal ethics committee and was carried out with permits from the Tamil Nadu Forest Department.
We thank the Tamil Nadu Forest Department for permits to carry out this study (permit no. 53/2018).
Conceptualization: AK, TT, IA. Data curation: AK, IA. Formal analysis: AK, IA. Funding acquisition: TT. Investigation: IA, AK. Resources: TT. Software: IA. Writing - original draft: AK, IA. Writing - review and editing: AK, TT, IA.
Akshay Khandekar https://orcid.org/0000-0002-7956-089X
Tejas Thackeray https://orcid.org/0000-0002-9981-8763
Ishan Agarwal https://orcid.org/0000-0001-9734-5379
All of the data that support the findings of this study are available in the main text.
Material examined.
Institutional and field series abbreviations are as follows: National Centre for Biological Sciences, Bengaluru (NCBS-AU/ NCBS-BH/NRC-AA); Bombay Natural History Society, Mumbai (
Cnemaspis azhagu: holotype, NRC-AA-1170 (adult male); paratypes, NRC-AA-1171, NRC-AA-1172, NRC-AA-1174,
Cnemaspis beddomei: AK-R 578, AK-R 579, AK-R 581–583 (adult males), AK-R 580 (adult female), from Kakachi, Kalakkad-Mundanthurai Tiger Reserve, Tirunelveli District, India.
Cnemaspis galaxia: holotype,
Cnemaspis nairi: AK-R 894, AK-R 895 (adults), from Thenmala, Courtallam, Tenkasi District, Tamil Nadu, India.
Cnemaspis nigriventris: holotype,
Cnemaspis nimbus: holotype,
Cnemaspis rashidi: AK-R 1439 and AK-R 1440 (adults), from high elevation of Shenbaga Thopu; AK-R 1692 and AK-R 1693, from Vellimalai, Srivilliputhur-Megamalai Tiger Reserve, Virudhunagar and Theni Districts, Tamil Nadu, India.
Cnemaspis regalis: holotype,
Cnemaspis rubraoculus: holotype,
Cnemaspis smaug: holotype,
Cnemaspis sundara: AK-R 1254 and AK-R 1266 (adults), from Mohan’s resort, Tenkasi District, Tamil Nadu, India.
Cnemaspis wallaceii: holotype,