Research Article |
Corresponding author: Akshay Khandekar ( akshaykhandekar555@gmail.com ) Corresponding author: Ishan Agarwal ( ishan.agarwal@gmail.com ) Academic editor: Uwe Fritz
© 2022 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 (2022) Three more novel species of South Asian Cnemaspis Strauch, 1887 (Squamata, Gekkonidae) from Kalakad Mundanthurai Tiger Reserve, Tamil Nadu, India. Vertebrate Zoology 72: 385-422. https://doi.org/10.3897/vz.72.e82343
|
We describe three distinct, small-bodied, 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 mid-body, 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.
Biodiversity hotspot, dwarf geckos, endemic species, southern India, taxonomy, Western Ghats
The South Asian clade within the polyphyletic genus Cnemaspis Strauch, 1887 diversified from a Western Ghats origin during the Paleocene-Eocene and today includes over 100 species disjunctly distributed in peninsular India, Sri Lanka, parts of Southeast Asia, with a single species in northeast India (
Ten well supported, broad clades are now recognised within South Asian Cnemaspis in peninsular India based on ND2 and 16S mitochondrial phylogenies (
During a recent survey in Kalakad Mundanthurai Tiger Reserve, Tamil Nadu, India as part for an ongoing project 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 examination, 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
Surveys were conducted in both day and night time, specimens 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 individuals 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. Specimens are deposited in the museum and research collection facility at the National Centre for Biological Sciences, Bengaluru (NRC) and at the Bombay Natural History Society, Mumbai (BNHS).
Morphological data were collected from a total of 25 specimens of three new species. We restricted morphological comparisons to the clade each new species was assigned to (see Results). Comparative morphological data of all other members of beddomei, gracilis, and littoralis clades were collected either from the type and/or topotypic specimens listed in the Appendix
We extracted genomic DNA from liver/ tail biopsies using the Qiagen DNeasy Blood & Tissue extraction kit for samples of the three new species as well as a few topotypical Cnemaspis littoralis (Jerdon, 1853) (Table
List of Cnemaspis sequences used in this study. Museum abbreviations are as follows: AA, Rohan Pethiyagoda field series; AK/ AK-R, Akshay Khandekar field series; AMB; Aaron Bauer field series; BNHS, Bombay Natural History Society, Mumbai; CESL, CESG, Centre for Ecological Sciences, Bangalore; MVZ, Museum of Vertebrate Zoology. Berkeley; NRC-AA, National Centre for Biological Sciences, Bangalore; SB, Sayantan Biswas field series; VG, Varad Giri field series; ZM, Zeeshan Mirza field series; ZSI-WRC, Zoological Survey of India, Pune. Accession numbers with * indicate 16S sequences.
Species | Voucher | Locality | GenBank Accession number |
---|---|---|---|
C. adii | BNHS 2465 | India, Karnataka, Bellary, Hampi | KX753654* |
C. agarwali | AK 107 | India, Tamil Nadu, Salem District, Sankari | MK792466 |
C. ajijae | AK 432 | India, Maharashtra, Satara District, Mahabaleshwar | MK792499 |
C. ajijae | ZSI-WRC-R/1060 | India, Maharashtra, Satara District, Mahabaleshwar | KX753649* |
C. amba | NCBS-BH691 (VG408) | India, Tamil Nadu, Kolhapur District, Amba | MK792488 |
C. amboliensis | VG 394 | India, Maharashtra, Sindhudurg District, Amboli | MK792481 |
C. amboliensis | BNHS2508 | India, Maharashtra, Sindhudurg District, Amboli | MH174356* |
C. anamudiensis | CESL 232 | India, Kerala, Idukki District, Anamudi | MZ701805 |
C. anandani | CESL 297 | India, Tamil Nadu, Nilgiris | MZ701811 |
C. cf. assamensis | BNHS 2808 | India, Meghalaya, Ri Bhoi District, Saiden | MW3674 |
C. australis | ZM003 | India, Kerala, Peppara | MZ701834 |
C. azhagu sp. nov. | NRC-AA-1172 (AK-R 671) | India, Tamil Nadu, Tirunelveli District, Thirukurungudi forest range | ON494554 |
C. balerion | CESL 415 | India, Karnataka, Chikkmagaluru District, Kemmangudi | MZ701815 |
C. bangara | BNHS 2584 (AK 292) | India, Karnataka, Kolar District, Paparajanahalli | MT188143 |
C. beddomei | CESL 379 | India, Tamil Nadu, Tirunelveli District, Kalakad Mundanthurai Tiger Reserve | MZ701814 |
C. chengodumalaensis | CESL 626 | India, Kerala, Kozhikode District, Chengodumala |
MT217043
|
C. flavigularis | CESL 247 | India, Kerala, Idukki District, Mathikettan Shola National Park | MZ291585* |
C. cf. flavigularis | ZM002 | India, Kerala, Idukki District, Mathikettan Shola National Park | MZ291569/ MZ291569* |
C. flaviventralis | VG 354 | India, Maharashtra, Sindhudurg District, Amboli | MK792495 |
C. galaxia | CESL 511 | India, Tamil Nadu, Virudhunagar District, Srivilliputhur | MZ701818 |
C. goaensis | AK 360 | India, Goa, South Goa District, Canacona | MK792479 |
C. gracilis | CES G385 | India, Kerala, Palakkad District, near Chittur River | MK792465 |
C. cf. gracilis | AK 209 | India, Tamil Nadu, Salem District, Yercaud | MK792462 |
C. indica | CESL 291 | India, Tamil Nadu, Nilgiris | MZ701810 |
C. jackieii | CESL 192 | India, Tamil Nadu, Vairavankulam RF, near Karuppanadhi dam | MZ701804 |
C. kalakadensis sp. nov. | NRC-AA-1181 (AK-R 643) | India, Tamil Nadu, Tirunelveli District, Kalakadu forest range | ON494555/ ON521706* |
C. kalakadensis sp. nov. | NRC-AA-1182 (AK-R 644) | India, Tamil Nadu, Tirunelveli District, Kalakadu forest range | ON494556/ ON521707* |
C. kolhapurensis | unvouchered | India, Maharashtra, Kolhapur District, Dajipur | MK792501 |
C. koynaensis | CES G349 | India, Maharashtra, Satara District, Humbarli | MK792490 |
C. krishnagiriensis | NRC-AA-1120 (AK 896) | India, Tamil Nadu, Krishnagiri District, Krishnagiri fort | MW580961 |
C. littoralis | AK 955 | India, Kerala, Kozhikode District, Kozhikode | ON494557 |
C. littoralis | AK 956 | India, Kerala, Kozhikode District, Kozhikode | ON494558 |
C. littoralis | AK 963 | India, Kerala, Kozhikode District, Kozhikode | ON494559 |
C. littoralis | SB 151 | India, Kerala, Thrissur District, Athirappilly Falls | KY038013 |
C. littoralis | ZM001 | India | MZ701831 |
C. littoralis | BNHS 2517 | India, Kerala, Kozhikode | MH174367* |
C. littoralis | BNHS 2518 | India, Kerala, Kozhikode | MH174368* |
C. cf. maculicollis | CESL 709 | India, Kerala, Kollam District, Shendurney Wildlife Sanctuary | MZ701825 |
C. magnifica | BNHS 2547 (AK855) | India, Karnataka, Hassan District, Sakleshpur | MT180437 |
C. modigliani | MVZ239314 | Sumatra, Kecematan Enggano, Pulau Enggano, near Malakoni |
KY037977
|
C. monticola | CESL 044 | India, Kerala, Wayanad District, Manikunjmalai | MZ701803 |
C. mundanthuraiensis sp. nov. | NRC-AA-1176 (AK-R 443) | India, Tamil Nadu, Tirunelveli District, Mundanthurai forest range | ON494557 |
C. mundanthuraiensis sp. nov. | NRC-AA-1177 (AK-R 445) | India, Tamil Nadu, Tirunelveli District, Mundanthurai forest range | ON494558 |
C. mysoriensis | unvouchered | India, Karnataka, Bangalore Urban District, IISc Campus | MK792474 |
C. nigriventris | CESL 264 | India, Kerala, Achankovil Reserve Forest | MZ701808 |
C. nimbus | CESL 252 | India, Kerala, Idukki District, Mathikettan Shola National Park | MZ701807 |
C. ornata | CESL 276 | India, Tamil Nadu, Tirunelveli District, Vairavakulam Reserve Forest | MZ701809 |
C. palanica | CESL 341 | India, Tamil Nadu, Dindigul District, Palani Hills | MZ701812/MZ291614 |
C. palakkadensis | BNHS 2790 | India, Kerala, Palakkad | MT762366* |
C. cf. palakkadensis | CESL 221 | India, Kerala, Palakkad | MZ291583* |
C. phillipsi | AA81 | Sri Lanka, Matale District, Gammaduwa, Rattota | KY038001 |
C. podihuna | 58A | Sri Lanka, Moneragala District, Kukulagoda | KY038005 |
C. regalis | CESL 487/488 | India, Tamil Nadu, Tirunelveli District, Kalakad Mundanthurai Tiger Reserve |
MZ701816/ MZ701817 |
C. rubraoculus | CESL 114 | India, Kerala, Periyar Tiger Reserve, Upper Manalar | ON494559 |
C. shevaroyensis | NCBS-BH 674 (AK 204) | India, Tamil Nadu, Salem District, Yercaud | MK792468 |
C. smaug | CESL 251 | India, Kerala, Idukki District, Mathikettan Shola National Park | MZ701806 |
C. thackerayi | CESG 143 | India, Tamil Nadu, Salem District, Yercaud | MK792471 |
C. tropidogaster | AMB 7505 | Sri Lanka, Nuwara Eliya District, Upcot tea estate | KY037983 |
C. wallaceii | CESL 377 | India, Tamil Nadu, Anamalai, Andiparai Shola | MZ701813 |
C. yercaudensis | CES G133 | India, Tamil Nadu, Namakkal District, Kollimalai massif | MK792473 |
The ND2 sequences ranged in length from 490–1038 nucleotides and we translated these to amino acids to ensure that there were no erroneous internal stop codons. We recovered the same broad ten clades as
Maximum likelihood (ML) phylogeny of South Asian Cnemaspis based on the mitochondrial ND2 gene. Bootstrap support >70 shown at nodes; beddomei, gracilis and littoralis clades marked by text. Roman numerals indicate other broad clades: I, australis; II, mysoriensis; III, goaensis; IV, girii; V, bangara; VI, indica; VII, podihuna; VIII, assamensis; IX, wynadensis. Top left inset is a 16S ML phylogeny of the littoralis clade (outgroups not shown).
Pairwise uncorrected ND2 sequence divergence within the beddomei (only four closest species included), gracilis and littoralis clades of South Asian Cnemaspis.
Species | 1 | 2 | 3 | 4 | beddomei clade | |||
1 | azhagu | |||||||
2 | galaxia | 11.1 | ||||||
3 | nigriventris | 32.2 | 28.8 | |||||
4 | ornata | 28.3 | 26.6 | 18.5 | ||||
5 | regalis | 19.6 | 20.0 | 32.1 | 29.0 | |||
Species | 1 | 2 | 3 | 4 | 5 | 6 | gracilis clade | |
1 | mundanthuraiensis | |||||||
2 | agarwali | 8.8 | ||||||
3 | gracilis | 7.6 | 8.8 | |||||
4 | cf. gracilis | 13.9 | 11.3 | 13.9 | ||||
5 | jackieii | 6.1 | 9.0 | 7.7 | 11.3 | |||
6 | shevaroyensis | 10.3 | 7.9 | 10.3 | 13.4 | 9.7 | ||
7 | thackerayi | 13.3 | 13.1 | 13.1 | 16.7 | 12.3 | 13.5 | |
Species | 1 | 2 | 3 | littoralis clade | ||||
1 | kalakadensis | |||||||
2 | cf. flavigularis | 19.5 | ||||||
3 | littoralis | 14.4 | 19.9 | |||||
4 | palanica | 19.3 | 11.7 | 20.2 |
Mensural (mm) data for the type series of Cnemaspis azhagu sp. nov.. Abbreviations are listed in Materials and Methods.
Type | Holotype | Paratypes | |||||||
Specimen no. | NRC-AA-1170 | NRC-AA-1171 | NRC-AA-1172 | NRC-AA-1173 | NRC-AA-1174 | BNHS 2818 | BNHS 2819 | BNHS 2820 | BNHS 2821 |
Sex | Male | Male | Male | Female | Male | Male | Male | Male | Female |
SVL | 37.5 | 37.3 | 34.9 | 27.4 | 37.2 | 37.0 | 35.4 | 34.3 | 38.0 |
TL | 48.8 | 42.5 | 42.0 | 35.4 | 40.7 | 49.5 | 49.4 | 38.0 | 46.7 |
TW | 3.9 | 4.1 | 3.4 | 2.5 | 3.5 | 3.7 | 4.0 | 3.5 | 3.7 |
LAL | 5.9 | 5.5 | 6.1 | 4.7 | 6.1 | 5.9 | 5.5 | 5.6 | 5.8 |
CL | 7.0 | 7.1 | 7.2 | 5.3 | 7.8 | 7.1 | 6.5 | 6.6 | 7.0 |
AGL | 15.2 | 14.3 | 12.7 | 11.2 | 16.0 | 15.8 | 14.9 | 14.5 | 16.0 |
BH | 3.7 | 4.0 | 3.9 | 2.6 | 4.6 | 5.5 | 4.7 | 4.4 | 3.8 |
BW | 6.8 | 6.9 | 6.0 | 5.0 | 7.9 | 8.2 | 7.7 | 7.1 | 7.4 |
HL | 9.6 | 9.8 | 9.4 | 7.1 | 10.2 | 9.5 | 9.0 | 8.7 | 10.7 |
HW | 6.2 | 6.2 | 6.1 | 5.0 | 6.5 | 6.5 | 6.3 | 5.9 | 6.7 |
HD | 4.1 | 4.0 | 3.9 | 3.1 | 4.3 | 4.2 | 4.2 | 4.0 | 4.2 |
ED | 1.9 | 1.9 | 1.9 | 1.5 | 2.1 | 1.9 | 1.9 | 1.8 | 2.0 |
EE | 2.8 | 3.0 | 2.7 | 2.1 | 3.3 | 2.8 | 2.9 | 2.7 | 2.8 |
ES | 4.7 | 4.5 | 4.4 | 3.4 | 4.7 | 4.4 | 4.3 | 4.4 | 4.9 |
EN | 3.6 | 3.5 | 3.4 | 2.7 | 3.7 | 3.5 | 3.4 | 3.4 | 3.9 |
IN | 1.0 | 1.2 | 1.0 | 1.0 | 1.3 | 1.1 | 1.1 | 1.1 | 1.2 |
IO | 2.3 | 2.3 | 2.3 | 1.5 | 2.6 | 2.4 | 1.3 | 2.2 | 2.8 |
EL | 0.4 | 0.5 | 0.5 | 0.4 | 0.8 | 0.6 | 0.7 | 0.5 | 0.7 |
Meristic data for the type series of Cnemaspis azhagu sp. nov.. Abbreviations are listed in Materials and Methods except for: L&R = Left & Right; irr = irregular; * = ventral scales, precloacal pores, lamellae, and tail tuberculation incomplete; abs. = absent.
Type | Holotype | Paratypes | |||||||
Specimen no. | NRC-AA-1170 | NRC-AA-1171 | NRC-AA-1172 | NRC-AA-1173 | NRC-AA-1174 | BNHS 2818 | BNHS 2819 | BNHS 2820 | BNHS 2821 |
Sex | Male | Male | Male | Female | Male | Male | Male | Male | Female |
SL (L&R) | 8&8 | 8&8 | 10&10 | 9&8 | 10&10 | 9&8 | 9&9 | 9&9 | 9&9 |
IL (L&R) | 7&7 | 8&7 | 8&9 | 8&8 | 8&8 | 8&7 | 8&8 | 8&8 | 9&8 |
SL M (L&R) | 6&6 | 7&6 | 6&6 | 6&6 | 5&6 | 7&6 | 6&6 | 6&6 | 6&6 |
IL M (L&R) | 6&5 | 6&5 | 5&5 | 6&6 | 5&5 | 6&5 | 6&5 | 5&5 | 6&5 |
PVT (L&R) | abs. | abs. | irr | abs. | irr | abs. | abs. | abs. | abs. |
DTR | 2 | 2 | 2 | 2 | 3 | 2 | 2 | 2 | 2 |
MVSR | 40 | 39 | 38 | 40 | 37 | 44 | 38 | 34 | 43 |
VS | 159 | 162 | 168 | 95* | 151 | 171 | 169 | 168 | 169 |
LamF1 (L&R) | 15&15 | 15&14 | 15&15 | 14&14 | 14&13 | 16&15 | 15&14 | 14&16 | 13&13 |
LamF4 (L&R) | 21&21* | 20&22 | 25&23 | 25&24 | 24&24 | 23&22 | 24&24 | 22&23 | 22&24 |
LamT1 (L&R) | 13&14 | 13&14 | 14&14 | 13&13 | 12&12 | 12&13 | 13&14 | 14&14 | 13&12 |
LamT4 (L&R) | 25&25 | 24&24 | 25&25 | 26&25 | 27&26 | 26&26 | 26&27 | 24&24 | 25&24 |
LamT5 (L&R) | 21&22 | 21&20 | 22&22 | 23&20 | 21&24 | 21&20 | 20&20 | 20&20 | 20&19 |
PP (L&R) | 6 | 8 | 3&3 | abs. | 3&3 | 8 | 7 | 6* | abs. |
SBPP | abs. | abs. | 1 | abs. | 1 | abs. | abs. | abs. | abs. |
PCT (L&R) | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 |
PVTT (L&R) | 8&7 | 7&8 | 10&9 | 6&7 | 1*&1* | 8&9 | 7&6 | 6&7* | 7&8 |
LTT (L&R) | 4&4 | 6&5 | 6&5 | 3&3 | 1*&1* | 3&2 | 4&4 | 4&5 | 4&5 |
VLTT (L&R) | 3&3 | 2&2 | 3&3 | 2&2 | 1*&1* | 2&2 | 3&3 | 3&3 | 3&4 |
NRC-AA-1170 (AK-R 678), adult male, from Thirukurungudi forest range (8.4069°N, 77.5485°E; ca. 200 m asl.), Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu state, India; collected by Akshay Khandekar and team on 06 May 2021.
NRC-AA-1171 (AK-R 670), NRC-AA-1172 (AK-R 671), NRC-AA-1174 (AK-R 673), BNHS 2818 (AK-R 674), BNHS 2819 (AK-R 675), BNHS 2820 (AK-R 676), adult males; BNHS 2821 (AK-R 677), adult female; NRC-AA-1173 (AK-R 672), subadult female. AK-R 670 and AK-R 671 collected from the same locality as holotype; AK-R 673, AK-R 674, and AK-R 675 collected from 8.4049°N, 77.5376°E, ca. 280 m asl.; AK-R 676, AK-R 677, and AK-R 672 collected from 8.4142°N, 77.5323°E, ca. 400 m asl.; all localities from Thirukurungudi forest range, Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu state, India; same collectors as holotype except collected on 07 May 2021.
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.
Thirukurungudi dwarf gecko.
A small-sized Cnemaspis, snout to vent length less than 38 mm (n=9). Dorsal pholidosis heterogeneous; smooth granular scales intermixed 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; 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 mid-body, 151–171 longitudinal 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), rarely 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 larger 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.
Cnemaspis azhagu sp. nov. is a member of the beddomei clade and can be easily distinguished from all 13 members of the clade by a combination of the following differing or non-overlapping characters: two (rarely three) rows of dorsal tubercles at mid-body (versus 8–10 rows of dorsal tubercles at mid-body in C. aaronbaueri; 10–12 in C. beddomei (Theobald, 1876), eight in C. galaxia; 16–18 in C. nairi; 13 or 14 in C. nigriventris Pal, Mirza, Dsouza and Shanker, 2021; 12–14 in C. nimbus Pal, Mirza, Dsouza and Shanker, 2021; and C. ornata (Beddome, 1870); 7–9 in C. regalis; 8–10 in C. rubraoculus Pal, Mirza, Dsouza and Shanker, 2021; 19–22 in C. smaug Pal, Mirza, Dsouza and Shanker, 2021; and 14–15 in C. wallaceii Pal, Mirza, Dsouza and Shanker, 2021); paravertebral tubercle rows absent (versus paravertebral tubercle rows present in C. aaronbaueri, C. beddomei, C. galaxia, C. nairi, C. nigriventris, C. nimbus, C. maculicollis, C. ornata, C. regalis, C. rubraoculus, C. smaug, and C. wallaceii); A small-sized Cnemaspis SVL 38 mm (versus medium-sized Cnemaspis SVL 40–50 mm: C. nairi; C. nimbus; C. ornata; C. rubraoculus; and C. wallaceii; large-sized Cnemaspis SVL >50 mm: C. anamudiensis Cyriac, Johny, Umesh and Palot, 2018; C. beddomei; C. maculicollis Cyriac, Johny, Umesh and Palot, 2018; and C. smaug); 151–171 longitudinal ventral scales from mental to cloaca (versus 135–140 longitudinal ventral scales from mental to cloaca in C. aaronbaueri, 143–147 in C. nairi, 134–141 in C. nimbus, 148–154 in C. regalis, 122–133 in C. rubraoculus, and 142–150 in C. smaug); 34–44 ventral scales across belly at mid-body (versus 31–33 ventral scales across belly at mid-body in C. aaronbaueri, 30–34 in C. beddomei and C. smaug, 27–31 in C. galaxia, 32–33 in C. nairi, 26–27 in C. nimbus, and 28–29 in C. wallaceii); 24–27 lamellae under digit IV of pes (versus 20–22 lamellae under digit IV of pes in C. anamudiensis, 21–23 in C. beddomei, 23–24 in C. maculicollis, 22 or 23 in C. nimbus and C. wallaceii, 28–31 in C. ornata, 18 or 19 in C. rubraoculus, 20–22 in C. smaug).
Adult male in good state of preservation except head marginally bent towards left and tail tip towards right (Fig.
Cnemaspis azhagu sp. nov. (holotype, NRC-AA-1170): A dorsal aspect of head, B ventral aspect of head, C lateral aspect of right side head, D aspect of cloacal region showing precloacal pores, E ventral aspect of left manus, and F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar and Satpal Gangalmale.
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.
Forelimbs and hindlimbs slightly long, slender (LAL/SVL 0.15); (CL/SVL 0.18); digits long, with a strong, recurved 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 lamellae above the inflection; basal lamellae series: (2-6-6-6-4 right manus, 2-7-7-8-4 right pes), (2-6-6-6-4 left manus, Fig.
Tail original, entire, subcylindrical, slender, slightly longer than snout-vent length (TL/SVL 1.30; Fig.
Dorsal ground colour of body, limbs and tail grey; entire head and region anterior 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 U-shaped light marking and in between uppermost and middle postorbital streaks; a single smaller black ocellus on the right side anterior to forelimb insertions; 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). Original 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.
Additional morphological character states evaluation for the type series of Cnemaspis azhagu sp. nov.. abs. = absent; / = data unavailable.
Type | Holotype | Paratypes | |||||||
Specimen no. | NRC-AA-1170 | NRC-AA-1171 | NRC-AA-1172 | NRC-AA-1173 | NRC-AA-1174 | BNHS 2818 | BNHS 2819 | BNHS 2820 | BNHS 2821 |
Sex | Male | Male | Male | Female | Male | Male | Male | Male | Female |
Anterior extra-brillar fringe scales enlarged (1) or not enlarged (0) | 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 |
Gular scales keeled (1) or smooth (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 |
Precloacal pores continuous (1) or separated (0) | 1 | 1 | 0 | abs. | 0 | 1 | 1 | 1 | abs. |
Precloacal pores elongate (1) or round (0) | 1 | 1 | 1 | abs. | 1 | 0 | 1 | 1 | abs. |
Dorsal pholidosis homogeneous (1) or heterogeneous (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 |
Tubercles linearly arranged (1) or more random (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Spine-like tubercles on flank present (1) or absent (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Lateral caudal furrows present (1) or absent (0) | 0 | 0 | 0 | 0 | / | 0 | 0 | 0 | 0 |
Subcaudals keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | / | 0 | 0 | 0 | 0 |
Single median row of subcaudals keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | / | 0 | 0 | 0 | 0 |
Caudal tubercles encircle tail (1) or not (0) | 0 | 0 | 0 | 0 | / | 0 | 0 | 0 | 0 |
Slightly enlarged median subcaudal scale row (1) or not (0) | 1 | 1 | 1 | 1 | / | 1 | 1 | 1 | 1 |
Enlarged femoral scales present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Subtibial scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Mensural, meristic and additional character state data for the type series is given in Tables
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.
NRC-AA-1175 (AK-R 451), adult male, from Papanasam reserve forest, Mundanthurai forest range (8.6980°N, 77.3561°E; ca. 200 m asl.), Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu state, India; collected by Akshay Khandekar and team on 21 April 2021.
NRC-AA-1176 (AK-R 443), NRC-AA-1177 (AK-R 445), NRC-AA-1178 (AK-R 446), BNHS 2822 (AK-R 448), BNHS 2824 (AK-R 450), and BNHS 2825 (AK-R 452), adult males; BNHS 2823 (AK-R 449), adult female; NRC-AA-1179 (AK-R 447), subadult female. AK-R 443 and AK-R 445 collected from the same locality and other details as holotype; AK-R 446, AK-R 448, and AK-R 450 collected from 8.7038°N, 77.3492°E, ca. 350 m asl.; AK-R 447, AK-R 449, and AK-R 452 collected from 8.7096°N, 77.3622°E; ca. 150 m asl.; all from Papanasam reserve forest, Mundanthurai forest range, Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu state, India; same collectors as holotype except collected on 22 April 2021.
The specific epithet is a toponym for the Mundanthurai forest range of Kalakad-Mundanthurai Tiger Reserve in Tirunelveli district of Tamil Nadu, the type and currently only known locality for this species.
Mundanthurai dwarf gecko.
A small-sized Cnemaspis, snout to vent length less than 33 mm (n=9). 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; 6–8 rows of dorsal tubercles at mid-body, tubercles in paravertebral region irregular; ventral scales subcircular, smooth, subequal from chest to vent, 30–34 scales across belly at mid-body, 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 separated medially by 2–4 poreless scales (n=4/7), precloacal 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 spine-like 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 pentagonal, 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 hindlimbs and margin of belly suffused with yellow).
Cnemaspis mundanthuraiensis sp. nov. is a member of the gracilis clade and can be easily distinguished from all five members of the clade by a combination of the following differing or non-overlapping characters: 6–8 rows of dorsal tubercles at mid-body (versus 9–11 rows of dorsal tubercles at mid-body in C. agarwali Khandekar, 2019; 11–14 in C. gracilis (Beddome, 1870) and C. thackerayi Khandekar, Gaitonde and Agarwal, 2019; 10–14 in C. shevaroyensis Khandekar, Gaitonde and Agarwal, 2019); only a few irregularly arranged tubercles in paravertebral region (versus 12–17 tubercles in paravertebral rows in C. agarwali; 9–12 in C. gracilis; 11–12 in C. jackieii; 13–17 in C. shevaroyensis; and 12–14 in C. thackerayi); spine-like tubercles present on flanks (versus spine-like tubercles absent on flanks in C. agarwali, C. jackieii, C. shevaroyensis, and C. thackerayi); 30–34 ventral scales across belly at mid-body (versus 24–26 ventral scales across belly at mid-body in C. agarwali, 23–25 in C. gracilis, 21–24 in C. shevaroyensis, and 22–25 in C. thackerayi).
Adult male in good state of preservation except tail tip marginally bent towards right (Fig.
Cnemaspis mundanthuraiensis sp. nov. (holotype, NRC-AA-1175): A dorsal aspect of head, B ventral aspect of head, C lateral aspect of right side head, D aspect of cloacal region showing precloacal and femoral pores, E ventral aspect of left manus, and F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar and Satpal Gangalmale.
Body relatively slender (BW/AGL 0.43), trunk less than half of SVL (AGL/SVL 0.41) without ventrolateral folds (Fig.
Mensural (mm) data for the type series of Cnemaspis mundanthuraiensis sp. nov.. Abbreviations are listed in Materials and Methods. * = incomplete tail.
Type | Holotype | Paratypes | |||||||
Specimen no. | NRC-AA-1175 | NRC-AA-1176 | NRC-AA-1177 | NRC-AA-1178 | NRC-AA-1179 | BNHS 2822 | BNHS 2823 | BNHS 2824 | BNHS 2825 |
Sex | Male | Male | Male | Male | Female | Male | Female | Male | Male |
SVL | 31.2 | 31.4 | 30.8 | 32.2 | 22.6 | 32.6 | 29.5 | 27.5 | 29.6 |
TL | 43.5 | 30.7 | 39.7 | 12.9* | 24.5 | 33.0 | 10.8* | 34.5 | 36.9 |
TW | 2.8 | 3.1 | 3.1 | 3.2 | 1.8 | 3.1 | 2.7 | 2.7 | 2.7 |
LAL | 4.8 | 4.8 | 4.9 | 5.1 | 3.4 | 5.2 | 4.6 | 4.4 | 4.5 |
CL | 5.9 | 5.8 | 5.6 | 6.2 | 4.3 | 6.2 | 5.5 | 5.3 | 5.3 |
AGL | 13.1 | 13.2 | 13.3 | 13.4 | 9.2 | 13.6 | 12.1 | 11.6 | 12.2 |
BH | 2.8 | 3.2 | 3.4 | 4.1 | 1.8 | 3.7 | 3.1 | 2.6 | 2.4 |
BW | 5.7 | 5.8 | 5.3 | 6.5 | 3.8 | 6.6 | 5.1 | 4.6 | 4.4 |
HL | 7.6 | 7.8 | 7.9 | 8.1 | 5.7 | 8.6 | 7.0 | 7.1 | 7.0 |
HW | 5.2 | 5.4 | 5.3 | 5.5 | 3.7 | 5.3 | 4.9 | 4.6 | 5.0 |
HD | 3.2 | 3.6 | 3.1 | 3.3 | 2.3 | 3.6 | 2.9 | 2.6 | 2.9 |
ED | 1.5 | 1.6 | 1.5 | 1.6 | 1.3 | 1.8 | 1.5 | 1.4 | 1.5 |
EE | 2.3 | 2.5 | 3.3 | 2.4 | 1.7 | 2.5 | 2.3 | 2.2 | 2.5 |
ES | 4.0 | 4.3 | 3.9 | 4.0 | 2.9 | 4.3 | 3.7 | 3.6 | 3.5 |
EN | 3.2 | 3.2 | 3.2 | 3.3 | 2.4 | 3.5 | 2.9 | 2.9 | 2.9 |
IN | 0.9 | 1.0 | 0.9 | 0.9 | 0.7 | 1.1 | 0.9 | 0.8 | 0.8 |
IO | 1.4 | 1.4 | 1.7 | 1.4 | 1.0 | 1.7 | 1.2 | 1.2 | 1.3 |
EL | 0.5 | 0.4 | 0.5 | 0.5 | 0.3 | 0.6 | 0.4 | 0.4 | 0.4 |
Forelimbs and hindlimbs slightly long, slender (LAL/SVL 0.15); (CL/SVL 0.18); digits long, with a strong, recurved claw, distinctly inflected, distal portions laterally compressed conspicuously. Digits with unpaired lamellae, separated into a basal and narrower distal series by single enlarged lamella at inflection; basal lamellae series: (1-3-4-5-3 right manus, 2-4-5-6-5 right pes), (1-3-3-5-4 left manus, Fig.
Tail original, entire, subcylindrical, slender, slightly longer than snout-vent length (TL/SVL 1.39; Fig.
Dorsal ground colour of head, body, limbs and tail pale brown. Head with indistinct 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 merging with light grey blotches on occiput, middle continuing 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 blotches 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 grey bands. Ventral surfaces white except sternal region, edges of belly and underside of hindlimbs suffused with yellow; gular region with numerous streaks along the posterior edge; underside of tail white. Pupil black, iris golden.
Meristic data for the type series of Cnemaspis mundanthuraiensis sp. nov.. Abbreviations are listed in Materials and Methods except for: L&R = Left & Right; irr = irregular; abs. = absent; * = dorsal tubercle rows and tail tuberculation incomplete.
Type | Holotype | Paratypes | |||||||
Specimen no. | NRC-AA-1175 | NRC-AA-1176 | NRC-AA-1177 | NRC-AA-1178 | NRC-AA-1179 | BNHS 2822 | BNHS 2823 | BNHS 2824 | BNHS 2825 |
Sex | Male | Male | Male | Male | Female | Male | Female | Male | Male |
SL (L&R) | 6&7 | 8&8 | 9&9 | 7&7 | 7&7 | 7&7 | 7&7 | 7&7 | 7&7 |
IL (L&R) | 7&7 | 7&7 | 8&8 | 7&7 | 7&7 | 7&7 | 7&7 | 7&7 | 7&8 |
SL M (L&R) | 6&6 | 6&5 | 5&6 | 6&6 | 7&7 | 5&6 | 6&7 | 6&6 | 6&6 |
IL M (L&R) | 6&6 | 5&5 | 5&6 | 5&6 | 6&6 | 5&5 | 6&6 | 5&5 | 5&6 |
ESFL (L&R) | 4&3 | 2&3 | 3&3 | 6&5 | 4&3 | 6&4 | 5&4 | 3&3 | 3&3 |
PVT (L&R) | irr | irr | irr | irr | irr | irr | irr | irr | irr |
DTR | 6 | 5* | 8 | 6 | 6 | 8 | 6 | 8 | 6 |
MVSR | 30 | 32 | 33 | 34 | 34 | 33 | 30 | 33 | 32 |
VS | 115 | 126 | 120 | 124 | 128 | 121 | 125 | 115 | 122 |
LamF1 (L&R) | 10&10 | 11&11 | 10&10 | 10&11 | 11&11 | 11&11 | 11&11 | 10&11 | 10&10 |
LamF4 (L&R) | 17&18 | 16&15 | 17&16 | 14&14 | 17&17 | 18&18 | 18&18 | 17&17 | 17&16 |
LamT1 (L&R) | 11&11 | 10&11 | 9&10 | 12&12 | 11&11 | 11&11 | 10&10 | 9&9 | 10&10 |
LamT4 (L&R) | 20&19 | 18&19 | 20&20 | 20&19 | 21&22 | 21&20 | 20&19 | 20&20 | 20&20 |
LamT5 (L&R) | 17&17 | 16&16 | 16&17 | 18&19 | 18&19 | 19&19 | 18&18 | 17&17 | 19&18 |
PP (L&R) | 1&1 | abs. | 1&1 | 1&1 | abs. | 1&1 | abs. | abs. | abs. |
SBPP | 2 | abs. | 4 | 2 | abs. | 3 | abs. | abs. | abs. |
SB PP&FP (L&R) | 10&9 | abs. | 10&10 | 11*&10 | abs. | 10&9 | abs. | abs. | abs. |
FP (L&R) | 3&4 | 4&4 | 4&3 | 5&5 | abs. | 5&4 | abs. | 4&4 | 4&4 |
SBFP | abs. | 25 | abs. | abs. | abs. | abs. | abs. | 24 | 23 |
PCT (L&R) | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 |
PVTT (L&R) | 14*&16 | 11*&11* | 13&3 | 6*&5* | 14&14 | / | 6*&6* | 16&16 | 14&14 |
LTT (L&R) | 12&13 | 11*&11* | 6&8 | 6*&7* | 10&9 | / | 6*&6* | 12&12 | 10&12 |
VLTT (L&R) | 8&8 | 6&6 | 6&4 | 6*&6* | 6&6 | / | 7*&7* | 8&8 | 8&7 |
Mensural, meristic and additional character state data for the type series is given in Tables
Cnemaspis mundanthuraiensis sp. nov. is currently known only from around its type locality (Papanasam reserve forest, Mundanthurai forest range, ca. 150–350 m asl.) in Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu (Fig.
Additional morphological character states evaluation for the type series of Cnemaspis mundanthuraiensis sp. nov.. abs. = absent; / = data unavailable.
Type | Holotype | Paratypes | |||||||
Specimen no. | NRC-AA-1175 | NRC-AA-1176 | NRC-AA-1177 | NRC-AA-1178 | NRC-AA-1179 | BNHS 2822 | BNHS 2823 | BNHS 2824 | BNHS 2825 |
Sex | Male | Male | Male | Male | Female | Male | Female | Male | Male |
Anterior extra-brillar fringe scales enlarged (1) or not enlarged (0) | 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 |
Gular scales keeled (1) or smooth (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 |
Precloacal pores continuous (1) or separated (0) | 0 | abs. | 0 | 0 | abs. | 0 | abs. | abs. | abs. |
Precloacal pores elongate (1) or round (0) | 1 | abs. | 1 | 1 | abs. | 1 | abs. | abs. | abs. |
femoral pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | abs. | 1 | abs. | 1 | 1 |
Dorsal pholidosis homogeneous (1) or heterogeneous (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 |
Tubercles linearly arranged (1) or more random (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Spine-like tubercles on flank present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Lateral caudal furrows present (1) or absent (0) | 1 | 1 | 1 | / | 1 | / | 1 | 1 | 1 |
Subcaudals keeled (1) or smooth (0) | 0 | 0 | 0 | / | 0 | / | 0 | 0 | 0 |
Single median row of subcaudals keeled (1) or smooth (0) | 0 | 0 | 0 | / | 0 | / | 0 | 0 | 0 |
Caudal tubercles encircle tail (1) or not (0) | 1 | 1 | 1 | / | 1 | / | 1 | 1 | 1 |
Enlarged median subcaudal scale row (1) or not (0) | 1 | 1 | 1 | / | 1 | / | 1 | 1 | 1 |
Enlarged femoral scales present (1) or absent (0) | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 1 |
Subtibial scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
NRC-AA-1180 (AK-R 648), adult male, from near Sengaltheri forest guesthouse, Kalakad forest range (8.5340°N, 77.4502°E; ca. 960 m asl.), Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu state, India; collected by Akshay Khandekar, and team on 30 April 2021.
NRC-AA-1182 (AK-R 644), BNHS 2826 (AK-R 646), adult males; NRC-AA-1181 (AK-R 643), NRC-AA-1183 (AK-R 645), BNHS 2827 (AK-R 647), BNHS 2828 (AK-R 654), adult females. AK-R 644 and AK-R 646 collected from same locality and other details as holotype; AK-R 643 and AK-R 645 collected from 8.5379°N, 77.4536°E, ca. 910 m asl.; AK-R 647 and AK-R 654 collected from 8.5290°N, 77.4467°E, ca. 1060 m asl; all from Kalakadu reserve forest, Kalakad forest range, Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu state, India, collectors and date same as holotype.
The specific epithet is a toponym for the Kalakad forest range of Kalakad-Mundanthurai Tiger Reserve in Tirunelveli district of Tamil Nadu, the type and currently only known locality for this species.
Kalakad dwarf gecko.
A small-sized Cnemaspis, snout to vent length less than 33 mm (n=7). Dorsal pholidosis heterogeneous; smooth, subcircular, weakly conical granular scales intermixed with irregularly arranged rows of enlarged, smooth, laterally compressed, spine-like scales on flanks; four or five rows of dorsal tubercles at mid-body, enlarged scales or tubercles absent in paravertebral region; ventral scales subcircular, smooth, subimbricate, subequal from chest to vent; 28–34 scales across belly at mid-body, 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 femoral pores on each thigh separated medially by 10 or 11 poreless scales; tail dorsum with enlarged, smooth, flattened 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 grey-brown with indistinct 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.
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 differing or non-overlapping characters: four or five rows of dorsal tubercles at mid-body restricted to flanks, laterally compressed, spine-like (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. palakkadensis; a few scattered spinose tubercles on the flanks in C. palanica Pal, Mirza, Dsouza and Shanker, 2021); 28–34 scales across belly at mid-body (versus 21–23 scales across belly at mid-body 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).
Adult male in good state of preservation except head marginally bent towards right and fully everted hemipenis on left side (Fig.
Cnemaspis kalakadensis sp. nov. (holotype, NRC-AA-1180): A dorsal aspect of head, B ventral aspect of head, C lateral aspect of right side head, D aspect of cloacal region showing femoral pores, E ventral aspect of left manus, and F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar and Satpal Gangalmale.
Body relatively slender (BW/AGL 0.40), trunk less than half of SVL (AGL/SVL 0.42) without ventrolateral folds (Fig.
Forelimbs and hindlimbs slightly long, slender (LAL/SVL 0.13); (CL/SVL 0.16); digits long, with a strong, recurved claw, distinctly inflected, distal portions laterally compressed.
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: (2-3-3-4-4 right manus, 2-4-4-6-5 right pes), (2-3-3-4-4 left manus, Fig.
Tail original, entire, subcylindrical, slender, marginally longer than snout-vent length (TL/SVL 1.12; Fig.
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 longitudinally placed, hourglass-shaped black marking anterior to forelimb insertions, flanked by a poorly defined lighter blotch anteriorly and posteriorly. About four poorly defined 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, elongate mid-dorsal blotches; dorsum of forelimbs, hindlimbs 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 precloacal 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.
Mensural (mm) data for the type series of Cnemaspis kalakadensis sp. nov.. Abbreviations are listed in Materials and Methods. * = incomplete tail.
Type | Holotype | Paratypes | |||||
Specimen no. | NRC-AA-1180 | NRC-AA-1181 | NRC-AA-1182 | NRC-AA-1183 | BNHS 2826 | BNHS 2827 | BNHS 2828 |
Sex | Male | Female | Male | Female | Male | Female | Female |
SVL | 29.6 | 32.8 | 30.5 | 33.1 | 27.4 | 31.6 | 32.5 |
TL | 33.4 | 15.6* | 19.6* | 32.2 | 6.7* | 34.3 | 31.2 |
TW | 2.9 | 2.6 | 2.7 | 2.7 | 2.7 | 2.7 | 3.1 |
LAL | 4.1 | 4.5 | 4.4 | 4.2 | 3.9 | 4.3 | 4.1 |
CL | 5.0 | 5.4 | 5.3 | 5.0 | 4.5 | 5.2 | 4.8 |
AGL | 12.7 | 16.5 | 13.8 | 15.0 | 11.4 | 13.8 | 13.7 |
BH | 3.0 | 4.0 | 2.8 | 3.4 | 2.8 | 3.5 | 4.9 |
BW | 5.2 | 7.8 | 5.2 | 6.7 | 5.0 | 6.1 | 7.3 |
HL | 7.5 | 8.2 | 7.8 | 7.7 | 6.8 | 7.6 | 7.7 |
HW | 4.6 | 5.2 | 4.9 | 4.9 | 4.4 | 4.8 | 5.3 |
HD | 2.9 | 2.9 | 2.8 | 2.8 | 2.5 | 3.0 | 3.6 |
ED | 1.4 | 1.5 | 1.5 | 1.4 | 1.3 | 1.3 | 1.5 |
EE | 2.5 | 2.9 | 2.7 | 2.4 | 2.1 | 2.4 | 2.7 |
ES | 3.5 | 4.0 | 3.6 | 3.9 | 3.5 | 3.7 | 4.0 |
EN | 3.0 | 3.2 | 3.0 | 3.0 | 2.7 | 3.4 | 3.3 |
IN | 0.8 | 0.8 | 0.8 | 0.8 | 0.7 | 0.8 | 0.9 |
IO | 2.0 | 2.1 | 2.0 | 1.9 | 1.7 | 2.1 | 2.4 |
EL | 0.4 | 0.6 | 0.4 | 0.4 | 0.4 | 0.4 | 0.4 |
Meristic data for the type series of Cnemaspis kalakadensis sp. nov.. Abbreviations are listed in Materials and Methods except for: L&R = Left & Right; abs. = absent; * = lamellae and tail tuberculation incomplete.
Type | Holotype | Paratypes | |||||
Specimen no. | NRC-AA-1180 | NRC-AA-1181 | NRC-AA-1182 | NRC-AA-1183 | BNHS 2826 | BNHS 2827 | BNHS 2828 |
Sex | Male | Female | Male | Female | Male | Female | Female |
SL (L&R) | 11&10 | 9&8 | 11&10 | 11&10 | 10&10 | 10&10 | 11&10 |
IL (L&R) | 10&9 | 9&9 | 9&8 | 8&9 | 8&9 | 9&9 | 9&9 |
SL M (L&R) | 8&8 | 8&8 | 10&9 | 9&8 | 8&7 | 8&8 | 7&7 |
IL M (L&R) | 7&6 | 6&7 | 8&8 | 6&6 | 7&6 | 7&6 | 7&6 |
ESFL (L&R) | 3&2 | 4&3 | 4&6 | 5&5 | 3&1 | 4&3 | 2&2 |
PVT | abs. | abs. | abs. | abs. | abs. | abs. | abs. |
DTR | 5 | 4 | 4 | 4 | 4 | 5 | 5 |
MVSR | 30 | 28 | 30 | 33 | 29 | 34 | 28 |
VS | 122 | 129 | 128 | 126 | 125 | 134 | 124 |
LamF1 (L&R) | 11&11 | 11&12 | 12&11 | 12&11 | 12&12 | 12 &12 | 11&11 |
LamF4 (L&R) | 16&16 | 16&16 | 18&17 | 18&18 | 19&18 | 18&6* | 16&16 |
LamT1 (L&R) | 11&11 | 11&11 | 11&12 | 12&11 | 12&12 | 12&12 | 11&11 |
LamT4 (L&R) | 19&19 | 22&21 | 21&20 | 19&20 | 21&21 | 21&22 | 19&19 |
LamT5 (L&R) | 18&18 | 15&19 | 19&18 | 19&19 | 20&18 | 20&18 | 18&17 |
FP (L&R) | 14&14 | abs. | 13&14 | abs. | 13&12 | abs. | abs. |
SBFP (L&R) | 11 | abs. | 10 | abs. | 10 | abs. | abs. |
PCT (L&R) | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 |
PVTT (L&R) | 16&16 | 8*&8* | 13*&13* | 7*&7* | 4*&4* | 18&18 | 4*&4* |
DLTT (L&R) | A | 0 | 3&2 | A | A | A | A |
LTT (L&R) | 2&2 | 0&1 | 1&2 | 2&2 | 1&1 | 4&4 | 2&3 |
Additional morphological character states evaluation for the type series of Cnemaspis kalakadensis sp. nov.. abs. = absent; / = data unavailable.
Type | Holotype | Paratypes | |||||
Specimen no. | NRC-AA-1180 | NRC-AA-1181 | NRC-AA-1182 | NRC-AA-1183 | BNHS 2826 | BNHS 2827 | BNHS 2828 |
Sex | Male | Female | Male | Female | Male | Female | Female |
Anterior extra-brillar fringe scales enlarged (1) or not enlarged (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Ventral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Gular scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Pectoral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
femoral pores elongate (1) or round (0) | 1 | abs. | 0 | abs. | 1 | abs. | abs. |
Dorsal pholidosis homogeneous (1) or heterogeneous (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Dorsal tubercles keeled (1) or not keeled (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Tubercles linearly arranged (1) or more random (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Spine-like scales on flank present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Lateral caudal furrows present (1) or absent (0) | 0 | 0 | 0 | 0 | / | 0 | / |
Subcaudals keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Single median row of subcaudals keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Caudal tubercles encircle tail (1) or not (0) | 0 | 0 | 0 | 0 | / | 0 | / |
Enlarged median subcaudal scale row (1) or not (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Enlarged femoral scales present (1) or absent (0) | 0 | 0 | 1 | 0 | 0 | 0 | 1 |
Subtibial scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Mensural, meristic and additional character state data for the type series is given in Tables
Cnemaspis kalakadensis 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.
South Asian Cnemaspis continue to be named at an unprecedented 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 (
All three new species are from the Kalakad Mundanthurai Tiger Reserve (KMTR), in southern Tamil Nadu; which forms part of the Agasthyamalai Biosphere Reserve along with the contiguous Neyyar, Peppara and Shendurney Wildlife Sanctuaries in Kerala. Spread across 895 km2, KMTR spans an elevational range from <100 m to >1800 m and includes both west-facing and east-facing slopes, receiving rainfall during the summer (southwest) and winter (northeast) monsoons (see
Four clades of South Asian Cnemaspis are distributed 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 14th 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) (
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 granite boulder habitats in southern India outside the Western Ghats (
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 (
We are thankful to Tamil Nadu Forest Department for the necessary permits (permit no. 53/2018). We thank Mr. Teja Bhargava (DFO, Jawadhu) for all his help and co-ordination throughout the fieldwork. At Kalakad Mundanthurai Tiger Reserve, we are grateful to Mr. Yogesh Singh (Field Director), Mr. Gautham (DFO), Forest Range Officers Mr. Karthikeyan Karmegam (Ambasamudram) and Mr. Balaji (Kalakad), Forest guards and watchers Mr. Bharathi Raghu (Thirukurungudi), Mr. Sivasubramaniam for their support and help during the fieldwork at KMTR. The fieldwork wouldn’t have been possible without the help of Satpal Gangalmale, Swapnil Pawar, Vaibhav Patil and Vivek Waghe. Satpal and Vivek helped with data collection and photography. Tarun Karmakar (NCBS field station and museum facility, Bengaluru) and Rahul Khot (BNHS, Mumbai) helped with specimen registrations at respective museums. Krishna Chaitanya helped with the correct formation and etymology of the Tamil specific name. We thank Saunak Pal and Kartik Shanker from CES for the loan of tissue of C. rubraoculus.
Material examined.
Institutional abbreviations are as follows: National Centre for Biological Sciences, Bengaluru (NCBS-AU/ NCBS-BH/ Akshay Khandekar field series [AK/ AK R]); Bombay Natural History Society, Mumbai (BNHS); Centre for Ecological Sciences, Bangalore (CES G).
Cnemaspis agarwali: holotype, NCBS-AU486 (adult male); paratypes, NCBS-AU487, BNHS 2337, NCBS-AU488, NCBSAU490, and BNHS 2338, (adult males), NCBS-AU485, BNHS 2336, and BNHS 2339, (adult females), from Sankari, Salem District, Tamil Nadu, India.
Cnemaspis gracilis: CESG385 from Chittur River, Palakkad District, Kerala, India. AK 133, AK 134, AK 135, AK 136, AK 137, AK 138, AK 139, AK 140, AK 141, AK 142, AK 143, AK 144, from Valparai, Coimbatore District, Tamil Nadu, India.
Cnemaspis shevaroyensis: holotype, NCBS-BH674 (adult male); paratypes, BNHS 2530, BNHS 2531, (adult males), NCBS-BH675, NCBS-BH676, BNHS 2529, (adult females) from the Shevaroy hills, Salem District, Tamil Nadu, India.
Cnemaspis thackerayi: holotype, NCBS-BH670 (adult male); paratypes, NCBS-BH671, BNHS 2527, (adult males), NCBS-BH672, NCBS-BH673, BNHS 2526, BNHS 2528, (adult females) from Yercaud, in Shevaroy hills, Salem District, Tamil Nadu, India.
Cnemaspis littoralis: AK 955, AK 956, AK 957, AK 958, AK 959, AK 960, AK 961, AK 962, from Kozhikode, Kozhikode District, Kerala, India.
Cnemaspis regalis: AK R 453, AK R 454, AK R 455, AK R 456, from Mundanthurai, Tirunelveli District, Tamil Nadu, India.
Cnemaspis beddomei: AK R 523, AK R 524, AK R 525, from Kakachi, Tirunelveli District; AK R 572, AK R 573, AK R 574, Upper Kodiyar, Kanyakumari District, Tamil Nadu, India.