Research Article |
Corresponding author: Ishan Agarwal ( ishan.agarwal@gmail.com ) Corresponding author: Akshay Khandekar ( akshaykhandekar555@gmail.com ) Academic editor: Uwe Fritz
© 2022 Ishan Agarwal, Tejas Thackeray, Akshay Khandekar.
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:
Agarwal I, Thackeray T, Khandekar A (2022) A multitude of spots! Five new microendemic species of the Cnemaspis gracilis group (Squamata: Gekkonidae) from massifs in the Shevaroy landscape, Tamil Nadu, India. Vertebrate Zoology 72: 1137-1186. https://doi.org/10.3897/vz.72.e94799
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South Asian Cnemaspis are one of the most diverse clades of gekkonids in South Asia with their highest diversity in the Western Ghats and Sri Lanka. These geckos include only a few nocturnal species and are largely diurnal or cathemeral and restricted to relatively cool habitats. One of the prominently diurnal subgroups in South Asian Cnemaspis is the bangara clade, which includes six species distributed in southern India on the eastern slopes of the Western Ghats, the southern Eastern Ghats and Palghat Gap. In this paper, we describe five more species of the bangara clade from the Shevaroyan landscape, including three from Kollimalai and one each from Yercaud and Pachaimalai, all in Tamil Nadu. These new species show 4.6–19.7 % uncorrected sequence divergence on the mitochondrial ND2 gene from each other and known species of the bangara clade and are morphologically diagnosable in body size, the number of paravertebral tubercles between limb insertions, the number of dorsal tubercle rows, the number of ventral scale rows across the belly, the number of femoral and precloacal pores and poreless scales separating these series, and aspects of colouration. The discovery of these five new species adds to the growing discoveries of cool-adapted species in southern India outside the Western Ghats and highlights the role of sky-islands in diversification. The Shevaroyan landscape shows high levels of microendemism with eight species distributed in an area of < 2000 km2, and all these species restricted to much smaller areas of actual distribution. With an area of < 500 km2 respectively, the massif of Pachaimalai has a single endemic and the massifs of Yercaud and Kollimalai have three endemic Cnemaspis species each.
Endemic species, integrative taxonomy, microendemism, mountains, southern India, species radiation, taxonomy
Diurnality has evolved multiple times within the ancestrally nocturnal Gekkonidae, including numerous reversals (
South Asian Cnemaspis originated in the Western Ghats in the Paleocene-Eocene and are largely restricted to cool habitats (
Cnemaspis gracilis (Beddome) was described by
Elevation map showing the distribution of members of the Cnemaspis gracilis clade in peninsular India. Major hill ranges are marked by bold text, within the Western Ghats: AG, Agasthyamalai; AN, Anaimalai; DV, Devarmalai; N, Nilgiris; PL, Palani; and outside the Western Ghats: BR, Biligirangan; J, Jawadhu; K, Kollimalai; KR, Kalrayan; MM, Male Mahadeshwara; P, Pachaimalai; S, Sitteri; SR, Sirumalai; Y, Yercaud; YL, Yelagiri. The Mysore Plateau and Palghat Gap are also marked.
The highest diversity within the bangara clade is in Yercaud, an isolated massif with a plateau above 1000 m asl. and a maximum elevation of 1623 m asl., on which two named species and one unnamed species occur (
Surveys were conducted in both day and night time, specimens were spotted on rocks, building walls, sometimes on tree trunks, and collected by hand, followed by euthanasia using isoflurane after taking colour photos in life. Liver 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 ~12–24 hours, washed and kept in tap water for ~24 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 (NCBS/ NRC).
Total genomic DNA was extracted from tail/ liver tissues using the Qiagen DNeasy Blood & Tissue extraction kit for the new species and some individuals of Cnemaspis gracilis (Table
List of Cnemaspis sequences used in this study. Museum and voucher abbreviations as follows: AK, Akshay Khandekar field series; CES G (Karanth lab field series) and CES L (Centre for Ecological Sciences, Bangalore); NCBS and NRC (National Centre for Biological Sciences, Bangalore); ZM, Zeeshan Mirza field series.
Species | Voucher | Locality | GenBank Accession number |
Cnemaspis agarwali | BNHS 2336 (AK 107) | India, Tamil Nadu, Salem District, Sankari | MK792466 |
Cnemaspis agarwali | NCBS-AU485 (AK 108) | India, Tamil Nadu, Salem District, Sankari | MK792467 |
Cnemaspis agayagangai sp. nov. | NRC-AA-1215 (AK 267) | India, Tamil Nadu, Namakkal District, Agaya Gangai Waterfalls | OP709694 |
Cnemaspis agayagangai sp. nov. | NRC-AA-1214 (AK 268) | India, Tamil Nadu, Namakkal District, Agaya Gangai Waterfalls | OP709695 |
Cnemaspis australis | ZM003 | India, Kerala, Peppara | MZ701834 |
Cnemaspis gracilis | AK 135 | India, Tamil Nadu, Coimbatore District, Valparai | MK792470 |
Cnemaspis gracilis | CES L 606 | India, Tamil Nadu, Palakkad District, Chennathanair RF | OP709696 |
Cnemaspis gracilis | CES L 607 | India, Tamil Nadu, Palakkad District, Chennathanair RF | OP709697 |
Cnemaspis gracilis | CES G 385 | India, Kerala, Palakkad District, near Chittur river | MK7924 |
Cnemaspis jackieii | CES L 192 | India, Tamil Nadu, Vairavankulam RF, near Karuppanadhi dam | MZ701804 |
Cnemaspis fantastica sp. nov. | NRC-AA-1223 (AK 284) | India, Tamil Nadu, Namakkal District, Kollimalai ghat | OP709698 |
Cnemaspis fantastica sp. nov. | NRC-AA-1224 (AK 285) | India, Tamil Nadu, Namakkal District, Kollimalai ghat | OP709699 |
Cnemaspis fantastica sp. nov. | CES G 131 | India, Tamil Nadu, Namakkal District, Kollimalai ghat | OP709700 |
Cnemaspis monticola | CES L 044 | India, Kerala, Wayanad District, Manikunjmalai | MZ701803 |
Cnemaspis mundanthuraiensis | NRC-AA-1176 (AKR 443) | India, Tamil Nadu, Tirunelveli District, Mundanthurai forest range | ON494557 |
Cnemaspis mundanthuraiensis | NRC-AA-1177 (AKR 445) | India, Tamil Nadu, Tirunelveli District, Mundanthurai forest range | ON494558 |
Cnemaspis pachaimalaiensis sp. nov. | NRC-AA-1231 (AK 708) | India, Tamil Nadu, Tiruchirapalli District, Pachaimalai | OP709701 |
Cnemaspis pachaimalaiensis sp. nov. | NRC-AA-1232 (AK 709) | India, Tamil Nadu, Tiruchirapalli District, Pachaimalai | OP709702 |
Cnemaspis rudhira sp. nov. | NRC-AA-1239 (AK 208) | India, Tamil Nadu, Salem District, Yercaud | MK792461 |
Cnemaspis rudhira sp. nov. | NRC-AA-1240 (AK 209) | India, Tamil Nadu, Salem District, Yercaud | MK792462 |
Cnemaspis rudhira sp. nov. | NRC-AA-1241 (AK 212) | India, Tamil Nadu, Salem District, Yercaud | MK792463 |
Cnemaspis rudhira sp. nov. | NRC-AA-1242 (AK 213) | India, Tamil Nadu, Salem District, Yercaud | MK792464 |
Cnemaspis salimalii sp. nov. | NRC-AA-1205 (AK 257) | India, Tamil Nadu, Namakkal District, Kollimalai | OP709703 |
Cnemaspis salimalii sp. nov. | NRC-AA-1206 (AK 258) | India, Tamil Nadu, Namakkal District, Kollimalai | OP709704 |
Cnemaspis shevaroyensis | NCBS-BH675 (AK 205) | India, Tamil Nadu, Salem District, Yercaud | MK792468 |
Cnemaspis shevaroyensis | NCBS-BH674 (AK 204) | India, Tamil Nadu, Salem District, Yercaud | MK792469 |
Cnemaspis thackerayi | CES G 143 | India, Tamil Nadu, Salem District, Yercaud | MK792471 |
The best fit models of sequence evolution and partitions were selected using the Bayesian Information Criteria in Partitionfinder 2 (
Morphological data were collected from a total of 44 specimens of the five new species. We restricted morphological comparisons to the bangara clade (see Results). Comparative morphological data of four of six members of the bangara clade included the type series; topotypic as well as additional specimens were used for C. gracilis (Beddome) (all listed in Appendix
The partial ND2 sequences for the new species ranged from 325–851 nucleotides. The monophyly of the bangara clade is well supported (bootstrap support 100, Posterior Probability 1.0), within which a basal split separates Cnemaspis salimalii sp. nov. (see description below) from the remaining lineages in the clade, with a subsequent divergence separating C. thackerayi from the remaining lineages (Fig.
Maximum likelihood tree of the bangara clade of South Asian Cnemaspis based on a partial fragment of ND2 with photographs of the species in life (not to scale) and sketches showing dorsal colour pattern of head and forebody (not to scale); note that C. salimalii sp. nov. and C. thackerayi have the same pattern. Bootstrap support and Posterior Probability (only values ≥ 65 % and 0.99 shown) depicted at nodes, outgroups not shown.
Uncorrected pairwise sequence divergence within previously named species of the bangara clade ranges from 6.1–19.7 % (Table
Pairwise uncorrected ND2 sequence divergence between members of the Cnemaspis gracilis clade of South Asian Cnemaspis, numbers in bold along diagonal represent intraspecific diversity.
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | |
1 | C. agarwali | 0.0 | |||||||||
2 | C. agayagangai sp. nov. | 7.8 | 1.0 | ||||||||
3 | C. fantastica sp. nov. | 8.2 | 9.8 | 0.4 | |||||||
4 | C. gracilis | 8.7 | 10.4 | 11.3 | 0.2 | ||||||
5 | C. jackieii | 9.0 | 9.8 | 10.4 | 7.8 | - | |||||
6 | C. mundanthuraiensis | 8.8 | 10.7 | 10.6 | 7.6 | 6.1 | 0.4 | ||||
7 | C. pachaimalaiensis sp. nov. | 9.9 | 10.5 | 12.0 | 9.1 | 8.5 | 8.9 | 1.8 | |||
8 | C. rudhira sp. nov. | 11.0 | 12.8 | 13.2 | 13.8 | 11.1 | 13.6 | 14.2 | 1.0 | ||
9 | C. salimalii sp. nov. | 17.2 | 16.4 | 17.3 | 15.8 | 18.2 | 16.8 | 14.3 | 19.7 | 1.1 | |
10 | C. shevaroyensis | 7.9 | 9.5 | 4.6 | 10.2 | 9.7 | 10.3 | 10.6 | 13.3 | 17.8 | 0.7 |
11 | C. thackerayi | 13.2 | 14.4 | 14.1 | 12.9 | 12.3 | 13.3 | 13.5 | 16.6 | 14.3 | 13.6 |
NRC-AA-1204 (AK 683), adult male, from the vicinity of Nallathambi resort, (11.2865°N, 78.3381°E; ca. 1150 m asl.), Semmedu, Kolli hills, Namakkal district, Tamil Nadu state, India; collected by Akshay Khandekar, Swapnil Pawar and Tejas Thackeray on 28th May 2019.
NRC-AA-1205 (AK 257), NRC-AA-1206 (AK 258), subadult males, NRC-AA-1207 (AK 259), NRC-AA-1208 (AK 261), NRC-AA-1209 (AK 263), adult males, NRC-AA-1210 (AK 265), adult female, same locality as holotype except collected by Akshay Khandekar, Ishan Agarwal, Nikhil Gaitonde, Varad Giri, Chaitanya R, and Aniruddha Dutta-Roy on 20th December 2018; NRC-AA-1212 (AK690), adult male, NRC-AA-1211 (AK 689), adult female, same data as holotype.
The specific epithet is a patronym honouring the eminent ornithologist Dr. Salim Ali (1896–1987) for his immense contributions to field research and conservation in India.
Salim Ali’s dwarf gecko.
A medium-sized Cnemaspis, snout to vent length up to 41.3 mm (n = 9). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with irregularly arranged rows of enlarged, strongly keeled, conical tubercles; last one or two rows of enlarged tubercles on flank weakly keeled, short and spine-like; 11–13 rows of dorsal tubercles at mid-body, 16–18 tubercles in paravertebral rows, paravertebral rows rarely irregular (n = 1/7); ventral scales smooth, subcircular, subimbricate, subequal from chest to vent, 30–33 scales across belly at mid-body, 109–128 longitudinal scales from mental to cloaca; subdigital scansors smooth, unpaired, unnotched; 10–12 lamellae under digit I of manus and pes, 15–18 lamellae under digit IV of manus and 20–24 lamellae under digit IV of pes; males with 3–5 femoral pores on each thigh separated by 5–7 poreless scales from series of 2–4 precloacal pores, precloacal pores separated medially by three or four poreless scales (n = 7/9); tail with enlarged, strongly keeled, pointed, and spine-like tubercles forming whorls; median row of subcaudals smooth, roughly pentagonal, and distinctly enlarged. Dorsum with diffuse light tan blotches including some in a vertebral row and numerous smaller orange blotches; a single black dorsal ocellus on neck, venter off-white with black speckles; original tail in males with eight or nine faint bands, regenerated tail brown.
Cnemaspis salimalii sp. nov. is a member of the bangara clade and can be easily distinguished from all six members of the clade by a combination of the following differing or non-overlapping characters: medium sized Cnemaspis, SVL up to 41 mm (versus small Cnemaspis SVL < 35 mm in C. agarwali, C. gracilis, C. jackieii, C. mundanthuraiensis, and C. shevaroyensis); 16–18 tubercles in paravertebral rows (versus only a few irregularly arranged tubercles in paravertebral region in C. mundanthuraiensis, 10–14 in C. gracilis; 11 or 12 in C. jackieii, 12–14 in C. thackerayi); 11–13 rows of dorsal tubercles at mid-body (versus eight or nine rows of dorsal tubercles at mid-body in C. jackieii, 6–8 rows of dorsal tubercles at mid-body in C. mundanthuraiensis); short spine-like tubercles present on flanks (versus spine-like tubercles absent on flanks in C. agarwali, C. jackieii, C. shevaroyensis, and C. thackerayi); 30–33 ventral scales across belly at mid-body (versus 24–26 ventral scales across belly at mid-body in C. agarwali, 26–29 (rarely 30) in C. gracilis, 21–24 in C. shevaroyensis, and 22–25 in C. thackerayi); single dorsal ocellus on occiput absent, single dorsal ocellus on neck present (versus a single dorsal ocellus each on occiput and neck, two pairs on either side of neck and just posterior to forelimb insertions in C. shevaroyensis; a single dorsal ocellus present on occiput and neck, two pairs on either side just anterior and sometimes posterior to forelimb insertions in C. agarwali). Cnemaspis salimalii sp. nov. is diagnosed against Cnemaspis agayagangai sp. nov., Cnemaspis fantastica sp. nov., Cnemaspis pachaimalaiensis sp. nov., and Cnemaspis rudhira sp. nov. as part of their respective descriptions below.
Mensural (mm) data for the type series of Cnemaspis salimalii sp. nov.. Abbreviations are listed in Materials and Methods. * = incomplete tail.
Type | Holotype | Paratypes | |||||||
museum number | NRC-AA-1204 | NRC-AA-1205 | NRC-AA-1206 | NRC-AA-1207 | NRC-AA-1208 | NRC-AA-1209 | NRC-AA-1210 | NRC-AA-1211 | NRC-AA-1212 |
Sex | Male | Male | Male | Male | Male | Male | Female | Female | Male |
SVL | 35.0 | 31.3 | 31.6 | 36.8 | 36.5 | 37.6 | 41.1 | 41.3 | 36.4 |
TL | 45.0 | 2.7* | 6.7* | 14.0* | 48.5 | 48.6 | 45.2 | 50.6 | 49.2 |
TW | 3.5 | 3.0 | 2.7 | 3.5 | 3.8 | 3.7 | 3.5 | 4.0 | 3.4 |
LAL | 5.1 | 4.6 | 4.6 | 5.6 | 5.2 | 5.6 | 6.1 | 5.8 | 5.6 |
CL | 6.7 | 5.5 | 5.2 | 7.1 | 6.4 | 6.6 | 7.5 | 7.0 | 6.7 |
AGL | 13.3 | 12.6 | 12.9 | 15.7 | 15.0 | 15.6 | 18.8 | 17.8 | 14.3 |
BH | 3.3 | 2.8 | 2.6 | 3.4 | 3.7 | 4.3 | 3.5 | 4.1 | 3.6 |
BW | 5.8 | 5.5 | 4.6 | 6.7 | 6.3 | 7.4 | 7.0 | 8.3 | 6.4 |
HL | 8.9 | 7.7 | 7.5 | 9.1 | 8.8 | 8.9 | 9.8 | 9.7 | 9.2 |
HW | 6.0 | 5.4 | 5.1 | 6.0 | 6.0 | 6.4 | 6.6 | 6.6 | 6.2 |
HD | 3.7 | 2.7 | 3.0 | 3.4 | 4.2 | 3.5 | 3.4 | 4.0 | 3.8 |
ED | 1.8 | 1.6 | 1.6 | 2.1 | 1.8 | 1.9 | 2.0 | 1.9 | 1.9 |
EE | 2.7 | 2.5 | 2.5 | 2.6 | 2.9 | 2.8 | 3.0 | 3.0 | 2.8 |
ES | 4.2 | 3.9 | 3.7 | 4.5 | 4.4 | 4.5 | 4.9 | 4.9 | 4.7 |
EN | 3.4 | 3.1 | 2.9 | 3.4 | 3.5 | 3.5 | 4.1 | 3.9 | 3.5 |
IN | 1.2 | 1.0 | 0.9 | 1.1 | 1.1 | 1.1 | 1.1 | 1.3 | 1.2 |
IO | 1.5 | 1.3 | 0.8 | 1.6 | 1.4 | 1.4 | 1.6 | 1.6 | 1.5 |
EL | 0.6 | 0.5 | 0.4 | 0.7 | 0.6 | 0.6 | 0.8 | 0.7 | 0.5 |
Meristic data for the type series of Cnemaspis salimalii sp. nov.. Abbreviations are listed in Materials and Methods except for: L&R = Left & Right; irr = irregular; * = paravertebral tubercles and lamellae incomplete; / = not available; abs. = absent;.
Type | Holotype | Paratypes | |||||||
Museum number | NRC-AA-1204 | NRC-AA-1205 | NRC-AA-1206 | NRC-AA-1207 | NRC-AA-1208 | NRC-AA-1209 | NRC-AA-1210 | NRC-AA-1211 | NRC-AA-1212 |
Sex | Male | Male | Male | Male | Male | Male | Female | Female | Male |
SL (L&R) | 10&10 | 8&9 | 8&8 | 9&9 | 9&8 | 9&9 | 9&10 | 8&7 | 9&9 |
IL (L&R) | 8&8 | 7&8 | 7&7 | 8&9 | 9&9 | 9&8 | 9&9 | 8&8 | 9&8 |
SL M (L&R) | 6&6 | 6&6 | 7&6 | 5&6 | 7&7 | 6&6 | 5&5 | 5&5 | 5&5 |
IL M (L&R) | 6&6 | 5&6 | 6&5 | 5&6 | 6&6 | 6&5 | 5&5 | 5&6 | 5&6 |
PVT (L&R) | irr | / | / | 18&18 | 18&18 | *&17 | 18&18 | 16&17 | 18&17 |
DTR | 12 | 11 | / | 11 | 12 | 12 | 12 | 13 | 12 |
MVSR | 30 | 33 | 30 | 30 | 33 | 32 | 33 | 32 | 31 |
VS | 109 | 114 | 113 | 118 | 116 | 128 | 117 | 114 | 117 |
LamF1 (L&R) | 12&12 | 11&12 | 10&10 | 11&11 | 11&11 | 11&11 | 11&11 | 10&11 | 11&11 |
LamF4 (L&R) | 16&17 | 15&16 | 18&16 | 18&18 | 16&17 | 17&17 | 16&16 | 18&18 | 18&18 |
LamT1 (L&R) | 12&12 | 11&11 | 10&10 | 10&11 | 10&10 | 11&10 | 10&10 | 11&11 | 10&11 |
LamT4 (L&R) | 22&22 | 21&22 | 22&22 | 21&22 | 21&21 | 21&20 | 22&22 | 15*&22 | 24&23 |
LamT5 (L&R) | 19&19 | 15*&19 | 18&18 | 19&20 | 19&18 | 19&19 | 18&18 | 18&19 | 20&21 |
PP L&R | 1&1 | 1&1 | 1&1 | 1&1 | 2&2 | 1&1 | abs. | abs. | 1&2 |
SBPP | 3 | 3 | 3 | 3 | 3 | 4 | abs. | abs. | 3 |
SB PP&FP (L&R) | 7&6 | 6&6 | 7&6 | 7&6 | 7&6 | 6&6 | abs. | abs. | 6&5 |
FP (L&R) | 5&4 | 4&4 | 4&4 | 4&3 | 3&4 | 4&3 | abs. | abs. | 5&5 |
SBFP | abs. | abs. | abs. | abs. | abs. | 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 |
Adult male in good state of preservation except regenerated portion of tail tip slightly bent towards right (Fig.
Additional morphological character states evaluation for the type series of Cnemaspis salimalii sp. nov.. abs. = absent; / = data unavailable.
Types | Holotype | Paratypes | |||||||
Museum number | NRC-AA-1204 | NRC-AA-1205 | NRC-AA-1206 | NRC-AA-1207 | NRC-AA-1208 | NRC-AA-1209 | NRC-AA-1210 | NRC-AA-1211 | NRC-AA-1212 |
Sex | Male | Male | Male | Male | Male | Male | Female | Female | 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 | 0 | 0 | 0 | 0 | 0 | abs. | abs. | 0 |
Precloacal pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | 1 | 1 | abs. | abs. | 1 |
femoral pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | 0 | 1 | abs. | abs. | 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 scales 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 keeled subcaudals (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) | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 1 |
Subtibial scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Occipital ocellus present (1) or absent (0) | 0 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 1 |
Ocelli anterior of the shoulder present (1) or absent (0) & number | 1 (1) | 1 (1) | 1 (1) | 0 | 1 (1) | 1 (1) | 1 (1) | 1 (1) | 1 (1) |
Ocelli posterior of the shoulder present (1) or absent (0) & number | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Original tail banded (1) or not (0) | 1 | / | / | 1 | 1 | 1 | 1 | 1 | 1 |
Cnemaspis salimalii sp. nov. (holotype, NRC-AA-1204): 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; F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar.
Body relatively slender (BW/AGL 0.43), trunk less than half of SVL (AGL/SVL 0.38) without ventrolateral folds; short spine-like scales on flank present (Fig.
Scales on palms and soles small, smooth, rounded, and flattened; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small granular, weakly keeled, imbricate scales that are twice the size of granules on body dorsum, largest on anterolateral aspect of hands and feet; scales on upper arm larger than lower; posterolateral aspect of limbs with small weakly keeled to smooth granular scales; ventral aspect of forelimbs with small, smooth, subimbricate scales, larger on lower arm than upper arm; ventral aspect of hindlimb with enlarged, smooth, flattened, subimbricate scales, slightly larger than body ventrals (Fig.
Tail original except tip (15.1 mm) which is regenerated, entire, subcylindrical, slender, slightly longer than snout-vent length (TL/SVL 1.28; Fig.
Dorsum of head, body, limbs and tail base mottled light brown. Head with some orange blotches and alternating yellow and dark bands on labials. Two orangish-brown postorbital streaks terminating anterior to forelimb insertions and one suborbital streak extending onto throat. A single black ocellus with a margin of orangish scales on neck. Dorsum with five light tan vertebral blotches from neck to tail base, orangish-brown blotches interspersed with smaller yellowish-grey spots on rest of dorsum and flank. Dorsum of limbs more muted than back, digits with alternating dark and light bands. Tail with five indistinct dark brown bands with a brown regenerated tail tip. Venter off-white, with black speckles under limbs and throat.
Colouration in life: A Cnemaspis salimalii sp. nov., adult male (holotype, NRC-AA-1204); B Cnemaspis agayagangai sp. nov., adult male (holotype, NRC-AA-1213); C Cnemaspis fantastica sp. nov., adult male (holotype, NRC-AA-1222); D Cnemaspis pachaimalaiensis sp. nov., adult male (holotype, NRC-AA-1230). Photos by Tejas Thackeray.
Mensural, meristic and additional character state data for the type series is given in Tables
Paratype series: A Cnemaspis salimalii sp. nov., from left to right, NRC-AA-1205–NRC-AA-1212; B Cnemaspis agayagangai sp. nov., from left to right, NRC-AA-1214–NRC-AA-1221; C Cnemaspis fantastica sp. nov., from left to right, NRC-AA-1223– NRC-AA-1229. Scale bar 10 mm; photos by Akshay Khandekar.
Cnemaspis salimalii sp. nov. is currently known only from around its type locality (from vicinity of Nallathambi resort, Semmedu, Kolli Hills, ca. 1100–1300 m asl.) in Namakkal district, Tamil Nadu (Fig.
NRC-AA-1213 (AK 699), adult male, near Agaya Gangai waterfalls (11.2640°N, 78.3925°E; ca. 860 m asl.), Kolli hills, Namakkal district, Tamil Nadu state, India; collected by Akshay Khandekar, Swapnil Pawar, and Tejas Thackeray on 29th May 2019.
NRC-AA-1215 (AK 267), NRC-AA-1214 (AK 268), adult males, same collection data as holotype; NRC-AA-1216 (AK 269), NRC-AA-1217 (AK 270) adult males, NRC-AA-1218 (AK 700), adult female, from near Arappaleeswarar temple (11.2645°N, 78.3906°E; ca. 940 m asl.); and NRC-AA-1220 (AK 702), adult male, NRC-AA-1219 (AK 701), NRC-AA-1221 (AK 703) adult females, from near Agaya Gangai waterfalls (11.2656°N, 78.3943°E; ca. 780 m asl.), collected by Akshay Khandekar, Ishan Agarwal, Nikhil Gaitonde on 20th December 2018.
The specific epithet is for the type locality of the new species, the Agaya Gangai Waterfalls, and is used as a noun in apposition.
Agaya Gangai dwarf gecko.
A small-sized Cnemaspis, snout to vent length up to 31.8 mm (n = 9). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with a fairly regularly arranged rows of enlarged, strongly keeled, conical tubercles; last one or two rows of enlarged tubercles on flank weakly keeled, spine-like; 10–12 rows of dorsal tubercles at mid-body, 14–18 tubercles in paravertebral rows; ventral scales smooth, subcircular, subimbricate, subequal from chest to vent, 28–34 scales across belly at mid-body, 106–120 longitudinal scales from mental to cloaca; subdigital scansors smooth, unpaired, unnotched; 9–11 lamellae under digit I of manus and 9–12 lamellae under digit I of pes; 14–16 lamellae under digit IV of manus and 17–20 lamellae under digit IV of pes; males (n = 6/9) with four or five femoral pores on each thigh separated by 8–10 poreless scales from series of two precloacal pores, precloacal pores separated medially by single (rarely 2, n = 1/6) poreless scales; tail with enlarged, strongly keeled, pointed, and spine-like tubercles forming whorls; median row of subcaudals smooth, roughly rectangular, and distinctly enlarged. Dorsum orange, mottled with numerous small light grey spots and fine black spots, light grey vertebral blotches sometimes distinct; a single central black dorsal ocellus each on neck and occiput separated by a light grey blotch, ocellus on neck flanked anteriorly on each side by a slightly larger ocellus, ocellus on occiput sometimes flanked on each side by smaller, indistinct ocellus; venter off-white with black speckles, two indistinct pairs of streaks on throat; original tail in males with about 8–10 alternating dark and light grey bands, regenerated tail orange-brown.
Cnemaspis agayagangai sp. nov. is a member of the bangara clade and can be easily distinguished from all members of the clade by a combination of the following differing or non-overlapping characters: small-sized Cnemaspis with maximum SVL 32 mm (versus medium-sized Cnemaspis, SVL up to 41 mm in C. thackerayi, and C. salimalii sp. nov.); 14–18 tubercles in paravertebral rows (versus only a few irregularly arranged tubercles in paravertebral region in C. mundanthuraiensis, 10–14 in C. gracilis; 11 or 12 in C. jackieii);
10–12 rows of dorsal tubercles at mid-body (versus eight or nine rows of dorsal tubercles at mid-body in C. jackieii, 6–8 rows of dorsal tubercles at mid-body in C. mundanthuraiensis); spine-like tubercles present on flanks (versus spine-like tubercles absent on flanks in C. agarwali, C. jackieii, C. shevaroyensis, and C. thackerayi); 28–34 ventral scales across belly at mid-body (versus 24–26 ventral scales across belly at mid-body in C. agarwali, 21–24 in C. shevaroyensis, and 22–25 in C. thackerayi); a single central dorsal ocellus each on occiput and neck, ocellus on neck flanked anteriorly on each side by a slightly larger ocellus (versus a single central dorsal ocellus each on occiput and neck in C. gracilis, C. mundanthuraiensis, C. thackerayi; single dorsal ocellus on occiput absent, single dorsal ocellus on neck present in C. salimalii sp. nov.; a single dorsal ocellus each on occiput and neck, a smaller pair on either side just anterior to forelimb insertion in C. jackieii; a single dorsal ocellus each on occiput and neck, two pairs on either side just anterior and posterior to forelimb insertions in C. shevaroyensis). Cnemaspis agayagangai sp. nov. is diagnosed against Cnemaspis fantastica sp. nov., Cnemaspis pachaimalaiensis sp. nov., and Cnemaspis rudhira sp. nov. as part of their respective descriptions below.
Adult male in good state of preservation except regenerated portion of the tail tip slightly bent towards right, small skin injury left of sternum, and claw on 2nd digit of left foot missing (Fig.
Cnemaspis agayagangai sp. nov. (holotype, NRC-AA-1213): 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; F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar.
Body relatively slender (BW/AGL 0.51), trunk less than half of SVL (AGL/SVL 0.40) without ventrolateral folds; spine-like scales on flank present (Fig.
Scales on palms and soles granular, smooth, rounded, and flattened; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small granular, weakly keeled, imbricate scales which are twice the size of granules on the body dorsum, largest on anterolateral aspect of the hands and feet; posterolateral aspect of limbs with small weakly keeled to smooth granular scales; ventral aspect of forelimbs with small, smooth, subimbricate scales, larger on lower arm than upper arm; ventral aspect of hindlimb with enlarged, smooth, flattened, subimbricate scales, slightly larger than body ventrals (Fig.
Tail original except tip (3.1 mm) which is regenerated, entire, subcylindrical, slender, marginally longer than snout-vent length (TL/SVL 1.18; Fig.
Dorsum of head, body, and tail base orange, limbs brown. Head with numerous light grey blotches and some black spots, light grey and dark bands on labials. Two dark postorbital streaks flanked by slightly broader light grey streaks terminating anterior to forelimb insertion. A single central black dorsal ocellus on neck and one on occiput separated by a larger light grey blotch, ocellus on neck flanked anteriorly on each side by a slightly larger ocellus, ocellus on occiput flanked one each side by a smaller ocellus; all ocelli with a fine orange margin. Dorsum with six light grey vertebral blotches from forelimb insertions to tail base, interspersed with smaller light-grey spots (sometimes forming streaks) and fine black spots on rest of dorsum and flank. Dorsum of limbs more muted than back, digits with alternating dark and light bands. Tail with seven or eight alternating light grey and black bands with an orange regenerated tip. Venter off-white with black speckles, two indistinct pairs of streaks on throat.
Mensural, meristic and additional character state data for the type series is given in Tables
Mensural (mm) data for the type series of Cnemaspis agayagangai sp. nov.. Abbreviations are listed in Materials and Methods. * = incomplete tail.
Type | Holotype | Paratypes | |||||||
museum number | NRC-AA-1213 | NRC-AA-1215 | NRC-AA-1214 | NRC-AA-1216 | NRC-AA-1217 | NRC-AA-1218 | NRC-AA-1219 | NRC-AA-1220 | NRC-AA-1221 |
Sex | M | M | M | M | M | F | F | M | F |
SVL | 31.2 | 30.0 | 30.2 | 31.8 | 29.3 | 29.8 | 31.7 | 31.5 | 30.4 |
TL | 37.0 | 14.9* | 12.2* | 20.6* | 34.0 | 36.5 | 19.0* | 41.1 | 21.3 |
TW | 3.0 | 3.2 | 3.1 | 2.7 | 2.7 | 2.9 | 2.4 | 3.0 | 2.6 |
LAL | 4.7 | 4.4 | 4.6 | 5.0 | 4.0 | 3.8 | 4.2 | 4.3 | 4.2 |
CL | 5.6 | 5.5 | 5.6 | 5.7 | 5.1 | 4.9 | 4.9 | 5.4 | 4.8 |
AGL | 12.6 | 12.4 | 12 | 12.7 | 11.8 | 12.4 | 12.8 | 12.2 | 12.3 |
BH | 3.2 | 3.4 | 3.2 | 3.2 | 3.2 | 3.3 | 3.2 | 3.2 | 3.6 |
BW | 6.5 | 5.5 | 6.0 | 5.8 | 5.1 | 5.6 | 7.1 | 5.3 | 6.2 |
HL | 8.0 | 7.3 | 7.7 | 8.0 | 7.3 | 7.0 | 7.5 | 7.6 | 7.6 |
HW | 5.4 | 4.9 | 5.1 | 5.0 | 4.8 | 4.8 | 4.9 | 5.0 | 4.8 |
HD | 3.8 | 3.4 | 3.2 | 3.3 | 3.0 | 3.3 | 3.4 | 4.0 | 3.1 |
ED | 1.7 | 1.6 | 1.6 | 1.8 | 1.6 | 1.5 | 1.7 | 1.8 | 1.6 |
EE | 2.4 | 2.4 | 2.5 | 2.5 | 2.1 | 2.3 | 2.7 | 2.5 | 2.5 |
ES | 4.0 | 3.8 | 3.9 | 3.9 | 3.6 | 3.6 | 3.6 | 4.0 | 3.9 |
EN | 3.2 | 3.0 | 3.2 | 3.2 | 3.6 | 3.0 | 3.1 | 3.2 | 3.1 |
IN | 1.0 | 1.0 | 0.9 | 1.0 | 0.9 | 0.9 | 1.0 | 0.9 | 1.0 |
IO | 1.1 | 1.2 | 1.2 | 1.1 | 1.0 | 1.0 | 1.0 | 1.3 | 1.1 |
EL | 0.4 | 0.4 | 0.4 | 0.5 | 0.5 | 0.5 | 0.6 | 0.6 | 0.4 |
Meristic data for the type series of Cnemaspis agayagangai sp. nov.. Abbreviations are listed in Materials and Methods except for: L&R = Left & Right; abs. = absent; * = paravertebral tubercles and lamellae incomplete; / = not available.
Type | Holotype | Paratypes | |||||||
Museum number | NRC-AA-1213 | NRC-AA-1215 | NRC-AA-1214 | NRC-AA-1216 | NRC-AA-1217 | NRC-AA-1218 | NRC-AA-1219 | NRC-AA-1220 | NRC-AA-1221 |
Sex | M | M | M | M | M | F | F | M | F |
SL (L&R) | 10&10 | 9&8 | 8&9 | 9&9 | 8&8 | 8&9 | 9&8 | 8&9 | 9&10 |
IL (L&R) | 9&10 | 8&8 | 6&7 | 8&9 | 7&7 | 7&8 | 8&7 | 8&8 | 7&8 |
SL M (L&R) | 6&6 | 5&5 | 6&6 | 6&6 | 6&6 | 6&6 | 5&5 | 5&5 | 6&6 |
IL M (L&R) | 5&5 | 5&5 | 5&5 | 6&6 | 5&5 | 5&6 | 4&5 | 6&6 | 5&6 |
PVT (L&R) | 15&14 | 15&16 | 14&15 | 14*&14* | 16&16 | 16&16 | / | 14&14 | 18&18 |
DTR | 10 | 12 | 11 | 11 | 11 | 10 | 10 | 12 | 11 |
MVSR | 32 | 28 | 33 | 31 | 32 | 31 | 30 | 30 | 34 |
VS | 110 | 115 | 110 | 114 | 118 | 110 | 106 | 120 | 113 |
LamF1 (L&R) | 9&9 | 9&9 | 10&6* | 9&9 | 10&10 | 11&11 | 10&10 | 11&11 | 11&10 |
LamF4 (L&R) | 14&15 | 16&15 | 17&16 | 15&14 | 16&15 | 16&16 | 15&15 | 16&16 | 15&15 |
LamT1 (L&R) | 10&10 | 9&10 | 10&10 | 9&9 | 9&10 | 12&12 | 10&10 | 10&10 | 9&10 |
LamT4 (L&R) | 19&19 | 20&20 | 19&20 | 17&17 | 20&20 | 20&20 | 14*&18 | 18&19 | 19&19 |
LamT5 (L&R) | 17&16 | 17&19 | 18&17 | 16&16 | 17&17 | 18&18 | 17&15 | 18&18 | 17&17 |
PP L&R | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | abs. | abs. | 1&1 | abs. |
SBPP | 2 | 1 | 1 | 1 | 1 | abs. | abs. | 1 | abs. |
SB PP&FP (L&R) | 8&8 | 10&10 | 9&8 | 9&9 | 8&8 | abs. | abs. | 9&8 | abs. |
FP (L&R) | 5&5 | 4&4 | 5&5 | 5&4 | 4&4 | abs. | abs. | 5&5 | abs. |
SBFP | abs. | abs. | abs. | abs. | abs. | 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 |
Additional morphological character states evaluation for the type series of Cnemaspis agayagangai sp. nov.. abs. = absent; / = data unavailable.
Types | Holotype | Paratypes | |||||||
Museum number | NRC-AA-1213 | NRC-AA-1215 | NRC-AA-1214 | NRC-AA-1216 | NRC-AA-1217 | NRC-AA-1218 | NRC-AA-1219 | NRC-AA-1220 | NRC-AA-1221 |
Sex | M | M | M | M | M | F | F | M | F |
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 | 0 | 0 | 0 | 1 | abs. | abs. | 0 | abs. |
Precloacal pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | 1 | abs. | abs. | 1 | abs. |
femoral pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | 1 | abs. | abs. | 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 scales 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 | 1 | 1 |
Subcaudals keeled (1) or smooth (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 |
Caudal tubercles encircle tail (1) or not (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Enlarged median subcaudal scale row (1) or not (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Enlarged femoral scales present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 |
Subtibial scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Occipital ocellus present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Ocelli anterior of the shoulder present (1) or absent (0) & number | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) |
Ocelli posterior of the shoulder present (1) or absent (0) & number | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Original tail banded (1) or not (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Cnemaspis agayagangai sp. nov. is currently known only from around its type locality (from Agaya Gangai waterfalls, Kolli hills, ca. 700–1000 m asl.) in Namakkal district, Tamil Nadu (Fig.
NRC-AA-1222 (AK 688), adult male, near Tree view point, (11.3192°N, 78.3460°E; ca. 1060 m asl.), Kolli Hills, Solakkadu, Namakkal district, Tamil Nadu state, India; collected by Akshay Khandekar, Swapnil Pawar, and Tejas Thackeray on 28th May 2019.
NRC-AA-1224 (AK 285), NRC-AA-1225 (AK 286), adult males, same locality data as holotype; NRC-AA-1223 (AK 284), NRC-AA-1226 (AK 684), adult females (11.3240°N, 78.3419°E; ca. 800 m asl.), Kolli Hills; and NRC-AA-1227 (AK 685), NRC-AA-1229 (AK 687), adult males, NRC-AA-1228 (AK 686), adult female (11.3270°N, 78.3392°E; ca. 600 m asl.) Kolli Hills collected by Akshay Khandekar, Ishan Agarwal, Nikhil Gaitonde, on 20th December 2018.
The specific epithet is derived from the Greek phantastikós, alluding to the spectacular colouration of the new species.
Fantastic dwarf gecko.
A small-sized Cnemaspis, snout to vent length up to 32.5 mm (n = 8). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with a fairly regularly arranged rows of enlarged, strongly keeled, conical tubercles; last one or two rows of enlarged tubercles on flank weakly keeled, spine-like; 11–13 rows of dorsal tubercles at mid-body, 15–17 tubercles in paravertebral rows; ventral scales smooth, subcircular, subimbricate, subequal from chest to vent, 28–32 scales across belly at mid-body, 110–120 longitudinal scales from mental to cloaca; subdigital scansors smooth, unpaired, unnotched; 8–10 lamellae under digit I of manus and pes, 14–16 lamellae under digit IV of manus and 16–20 lamellae under digit IV of pes; males with four or five femoral pores on each thigh separated by 7–9 poreless scales from series of 2–4 precloacal pores, precloacal pores separated medially by one or two poreless scales; tail with enlarged, strongly keeled, pointed, and spine-like tubercles forming whorls; median row of subcaudals smooth, roughly pentagonal, and distinctly enlarged. Dorsum reddish, mottled with numerous small yellow spots some of which form an indistinct vertebral line; a single central ocellus on neck, flanked posteriorly by a pair of much larger squarish blotches and anteriorly by a pair of subequal squarish blotches, indistinct spot on occiput; venter off-white with black speckles, two distinct pairs of black streaks on throat; throat off-white with two pairs of black streaks; original tail in males with 8–9 alternating dark and light grey bands, regenerated tail orange.
Cnemaspis fantastica sp. nov. is a member of the bangara clade and can be easily distinguished from all members of the clade by a combination of the following differing or non-overlapping characters: small-sized Cnemaspis with maximum SVL 32 mm (versus medium-sized Cnemaspis, SVL up to 41 mm in C. thackerayi and C. salimalii sp. nov.); 15–17 tubercles in paravertebral rows (versus only a few irregularly arranged tubercles in paravertebral region in C. mundanthuraiensis, 10–14 in C. gracilis; 11 or 12 in C. jackieii);
11–13 rows of dorsal tubercles at mid-body (versus eight or nine rows of dorsal tubercles at mid-body in C. jackieii, 6–8 rows of dorsal tubercles at mid-body in C. mundanthuraiensis); spine-like tubercles present on flanks (versus spine-like tubercles absent on flanks in C. agarwali, C. jackieii, C. shevaroyensis, and C. thackerayi); 28–32 ventral scales across belly at mid-body (versus 24–26 ventral scales across belly at mid-body in C. agarwali, 21–24 in C. shevaroyensis, and 22–25 in C. thackerayi); a single central ocellus on neck, flanked posteriorly by a pair of much larger squarish blotches and anteriorly by a pair of subequal squarish blotches, indistinct spot on occiput (versus a single central dorsal ocellus each on occiput and neck, ocellus on neck flanked anteriorly on each side by a slightly larger ocellus in C. agayagangai sp. nov., a single central dorsal ocellus each on occiput and neck in C. gracilis, C. mundanthuraiensis, C. thackerayi; single dorsal ocellus on occiput absent, single dorsal ocellus on neck present in C. salimalii sp. nov.; a single dorsal ocellus each on occiput and neck, a smaller pair on either side just anterior to forelimb insertion in C. jackieii). Cnemaspis fantastica sp. nov. overlaps in all morphological and meristic characters to C. agayagangai sp. nov. apart from the condition of the ventral scales in the original tail, which are roughly pentagonal and in a relatively regular series, size more than half tail width (versus irregular in shape and arrangement, size less than half tail width in C. agayagangai sp. nov.); and colour pattern, with the dorsal ocelli on the new species relatively larger and squarish with the central ocellus smallest and forming an X with five ocelli (versus four smaller, rounded subequal ocelli forming a diamond in C. agayagangai sp. nov.). Cnemaspis fantastica sp. nov. is diagnosed against Cnemaspis pachaimalaiensis sp. nov. and Cnemaspis rudhira sp. nov. as part of their respective descriptions below.
Adult male in good state of preservation except tail tip marginally bent towards right (Fig.
Cnemaspis fantastica sp. nov. (holotype, NRC-AA-1222): 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; F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar.
Body relatively slender (BW/AGL 0.45), trunk less than half of SVL (AGL/SVL 0.39) without ventrolateral folds; short spine-like scales on flank present (Fig.
Scales on palms and soles granular, smooth, subcircular, and flattened; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small granular, weakly keeled, imbricate scales which are twice the size of granules on the body dorsum, largest on anterolateral aspect of the hands and feet; posterolateral aspect of limbs with small weakly keeled to smooth granular scales; scales on upper hand and thigh larger than lower hand and shank respectively; ventral aspect of forelimbs with small, smooth, subimbricate scales, larger on lower arm than upper arm; ventral aspect of hindlimb with enlarged, smooth, flattened, subimbricate scales, slightly larger than body ventrals (Fig.
Tail half original half regenerated, entire, subcylindrical, slender, marginally longer than snout-vent length (TL/SVL 1.16; Fig.
Dorsum of head, body, limbs and tail base reddish. Head with numerous yellow spots, yellow and dark bands on labials, postorbital streaks indistinct. A single central ocellus on neck, flanked posteriorly by a pair of much larger squarish blotches and anteriorly by a pair of subequal squarish blotches, indistinct spot on occiput; all ocelli black with a fine orange and diffuse yellow margin. Dorsum mottled with numerous small yellow spots some of which form an indistinct vertebral line and fine black spots. Dorsum of limbs more muted than back with indistinct yellow bands, digits with alternating dark and light bands. Tail with four black and three light grey bands with an orange regenerated tip. Venter off-white with black speckles, two distinct pairs of black streaks on throat.
Mensural, meristic and additional character state data for the type series is given in Tables
Mensural (mm) data for the type series of Cnemaspis fantastica sp. nov.. Abbreviations are listed in Materials and Methods. * = incomplete tail.
Type | Holotype | Paratypes | ||||||
museum number | NRC-AA-1222 | NRC-AA-1223 | NRC-AA-1224 | NRC-AA-1225 | NRC-AA-1226 | NRC-AA-1227 | NRC-AA-1228 | NRC-AA-1229 |
Sex | M | F | M | M | F | M | F | M |
SVL | 31.0 | 31.9 | 32.5 | 30.5 | 32.1 | 28.4 | 31.9 | 31.4 |
TL | 36.0 | 12.9* | 10.3* | 9.5* | 19.5* | 28.5 | 36.1 | 38.4 |
TW | 3.1 | 2.5 | 2.8 | 2.5 | 2.9 | 2.8 | 2.8 | 3.3 |
LAL | 4.4 | 4.4 | 4.5 | 4.4 | 4.2 | 4.0 | 4.4 | 4.6 |
CL | 5.4 | 4.9 | 5.5 | 5.2 | 5.0 | 5.0 | 5.2 | 5.2 |
AGL | 12.3 | 13.6 | 12.9 | 12.9 | 13.6 | 12.1 | 13.8 | 12.8 |
BH | 3.1 | 4.0 | 3.3 | 3.0 | 3.7 | 3.0 | 3.8 | 3.5 |
BW | 5.6 | 6.8 | 5.8 | 5.1 | 6.7 | 6.1 | 6.7 | 6.5 |
HL | 7.7 | 7.8 | 7.9 | 7.6 | 7.4 | 7.0 | 7.8 | 7.6 |
HW | 5.1 | 5.0 | 5.0 | 4.9 | 5.0 | 5.0 | 5.4 | 5.1 |
HD | 3.4 | 3.4 | 3.4 | 3.0 | 3.2 | 3.3 | 3.2 | 3.5 |
ED | 1.6 | 1.5 | 1.6 | 1.5 | 1.6 | 1.4 | 1.7 | 1.7 |
EE | 2.4 | 2.5 | 2.4 | 2.0 | 2.6 | 2.2 | 2.5 | 2.5 |
ES | 3.9 | 3.8 | 3.8 | 3.7 | 3.9 | 3.5 | 3.7 | 3.9 |
EN | 3.1 | 3.3 | 3.1 | 3.1 | 3.1 | 2.8 | 3.1 | 3.0 |
IN | 1.0 | 1.1 | 1.0 | 0.8 | 1.0 | 1.0 | 1.1 | 1.0 |
IO | 1.4 | 1.2 | 0.9 | 1.2 | 1.2 | 1.2 | 1.0 | 1.2 |
EL | 0.5 | 0.5 | 0.4 | 0.5 | 0.5 | 0.4 | 0.4 | 0.6 |
Upper postmentals marginally in contact with each other below mental in NRC-AA-1224; upper postmentals separated from each other below mental by a single median enlarged chin shields in NRC-AA-1227 and NRC-AA-1228; upper postmentals bordered by mental, infralabial I, outer postmental, and by a single large chin scale on either side in NRC-AA-1223, NRC-AA-1225, NRC-AA-1226, and NRC-AA-1229. Outer postmental bordered by inner postmental, infralabials I & II in all types, additionally, four chin scales on either side in NRC-AA-1223, NRC-AA-1226, NRC-AA-1228, five chin scales on right in NRC-AA-1227, and five chin scales on left and four on right side in NRC-AA-1229; outer postmental separated from each other medially by two enlarged chin scales in NRC-AA-1223, NRC-AA-1225, NRC-AA-1226, and NRC-AA-1229. Two paratypes — NRC-AA-1228 and NRC-AA-1229 with original and complete tails, slightly longer than body (TL/SVL 1.13 and 1.22 respectively); tail entire but mostly regenerated in NRC-AA-1227, equal to body length (TL/SVL 1.00); tail entire but incomplete in NRC-AA-1225 and NRC-AA-1226 (TL = 9.5 and 19.5 mm respectively); tail almost entirely regenerated and largely lost in NRC-AA-1223 and NRC-AA-1224; original tail distinctly banded in males and faintly in female paratypes; regenerated tail orangish in life and yellowish-grey in preservative. (Fig.
Meristic data for the type series of Cnemaspis fantastica sp. nov.. Abbreviations are listed in Materials and Methods except for: L&R = Left & Right; abs. = absent; / = not available.
Type | Holotype | Paratypes | ||||||
Museum number | NRC-AA-1222 | NRC-AA-1223 | NRC-AA-1224 | NRC-AA-1225 | NRC-AA-1226 | NRC-AA-1227 | NRC-AA-1228 | NRC-AA-1229 |
Sex | M | F | M | M | F | M | F | M |
SL (L&R) | 8&9 | 9&8 | 8&8 | 8&8 | 9&8 | 9&10 | 8&9 | 9&9 |
IL (L&R) | 8&8 | 9&8 | 7&7 | 8&8 | 8&7 | 8&9 | 7&8 | 7&7 |
SL M (L&R) | 6&6 | 6&6 | 6&6 | 6&6 | 5&5 | 6&6 | 6&6 | 6&6 |
IL M (L&R) | 6&5 | 5&5 | 5&5 | 6&6 | 5&5 | 5&5 | 5&5 | 5&6 |
PVT (L&R) | 17&17 | 16&16 | 16&16 | 16&17 | 16&16 | / | 15&15 | 17&17 |
DTR | 12 | 13 | 12 | 12 | 11 | 12 | 11 | 11 |
MVSR | 30 | 31 | 32 | 30 | 32 | 31 | 29 | 28 |
VS | 113 | 110 | 116 | 113 | 117 | 120 | 114 | 112 |
LamF1 (L&R) | 9&8 | 8&8 | 9&9 | 8&8 | 9&10 | 9&9 | 9&9 | 9&9 |
LamF4 (L&R) | 15&15 | 15&14 | 14&15 | 14&14 | 16&16 | 16&16 | 14&15 | 15&15 |
LamT1 (L&R) | 9&9 | 8&9 | 9&9 | 9&9 | 9&9 | 10&10 | 10&10 | 9&10 |
LamT4 (L&R) | 19&18 | 17&16 | 19&19 | 19&19 | 18&18 | 20&19 | 18&18 | 19&17 |
LamT5 (L&R) | 16&17 | 15&15 | 16&16 | 16&16 | 16&16 | 18&18 | 17&17 | 16&17 |
PP L&R | 1&1 | abs. | 2&2 | 1&1 | abs. | 1&1 | abs. | 1&1 |
SBPP | 1 | abs. | 1 | 2 | abs. | 2 | abs. | 2 |
SB PP&FP (L&R) | 8&8 | abs. | 7&8 | 8&8 | abs. | 9&9 | abs. | 8&9 |
FP (L&R) | 5&4 | abs. | 5&5 | 5&5 | abs. | 5&5 | abs. | 4&4 |
SBFP | abs. | abs. | abs. | abs. | abs. | abs. | abs. | abs. |
PCT (L&R) | 1&1 | 0&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 |
Cnemaspis fantastica sp. nov. is currently known only from around its type locality (Karavallicombai reserve forest, Kolli hills, between an elevational gradient of ca. 600–1100 m asl.) in Namakkal district, Tamil Nadu (Fig.
Additional morphological character states evaluation for the type series of Cnemaspis fantastica sp. nov.. abs. = absent; / = data unavailable.
Types | Holotype | Paratypes | ||||||
museum number | NRC-AA-1222 | NRC-AA-1223 | NRC-AA-1224 | NRC-AA-1225 | NRC-AA-1226 | NRC-AA-1227 | NRC-AA-1228 | NRC-AA-1229 |
Sex | M | F | M | M | F | M | F | M |
Anterior extra-brillar fringe scales enlarged (1) or not enlarged (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Ventral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Gular scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Pectoral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Precloacal pores continuous (1) or separated (0) | 0 | abs. | 0 | 0 | abs. | 0 | abs. | 0 |
Precloacal pores elongate (1) or round (0) | 1 | abs. | 1 | 0 | abs. | 0 | abs. | 1 |
femoral pores elongate (1) or round (0) | 1 | abs. | 1 | 1 | abs. | 1 | abs. | 1 |
Dorsal pholidosis homogeneous (1) or heterogeneous (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 |
Tubercles linearly arranged (1) or more random (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Spine-like scales on flank present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Lateral caudal furrows present (1) or absent (0) | 1 | / | / | / | 1 | / | 0 | 1 |
Subcaudals keeled (1) or smooth (0) | 0 | / | / | / | 0 | / | 0 | 0 |
Single median row of keeled subcaudals (1) or smooth (0) | 0 | / | / | / | 0 | / | 0 | 0 |
Caudal tubercles encircle tail (1) or not (0) | 1 | / | / | / | 1 | / | 1 | 1 |
Enlarged median subcaudal scale row (1) or not (0) | 1 | / | / | / | 1 | / | 1 | 1 |
Enlarged femoral scales present (1) or absent (0) | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 1 |
Subtibial scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Occipital ocellus (present or absent) | P | P | A | A | P | P | A | P |
Ocelli anterior of the shoulder present (1) or absent (0) & number | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) |
Ocelli posterior of the shoulder present (1) or absent (0) & number | 1 (2) | 1 (2) | 1 (2) | 1 (2) | 1 (2) | 1 (2) | 1 (2) | 1 (2) |
Original tail banded (1) or not (0) | 1 | / | / | 1 | 1 | 1 | 1 | 1 |
NRC-AA-1230 (AK 711), adult male, from near Mangalam waterfalls, (11.3422°N, 78.6047°E; ca. 650 m asl.), Pachaimalai hills, Trichy district, Tamil Nadu state, India; collected by Akshay Khandekar, Ishan Agarwal, Swapnil Pawar and Tejas Thackeray on 30th May 2019.
NRC-AA-1231 (AK 708), NRC-AA-1232 (AK 709), adult males, same data as holotype; NRC-AA-1233 (AK 712), adult male, NRC-AA-1234 (AK 713), adult female, from near Shri Kaliyamman temple (11.3642°N, 78.5910°E; ca. 960 m asl.); NRC-AA-1235 (AK 730), NRC-AA-1236 (AK 731), NRC-AA-1237 (AK 753), adult males, from Pachaimalai RF (11.3167°N, 78.6018°E; ca. 840 m asl.), same data as holotype.
The specific epithet is a toponym for the Pachaimalai hills in Trichy district of Tamil Nadu, the type and currently only known locality for this species.
Pachaimalai dwarf gecko.
A small-sized Cnemaspis, snout to vent length up to 33.6 mm (n = 8). Dorsal pholidosis heterogeneous; weakly keeled, weakly conical, granular scales intermixed with a fairly regularly arranged rows of enlarged, strongly keeled, conical tubercles; last one or two rows of enlarged tubercles on flank short spine-like; 10–12 rows of dorsal tubercles at mid-body, 13–17 tubercles in paravertebral rows; ventral scales subcircular, smooth, subequal from chest to vent, 30–33 scales across belly at mid-body, 112–125 longitudinal scales from mental to cloaca; subdigital scansors smooth, unpaired, unnotched; 9–11 lamellae under digit I of manus and 9–12 lamellae under digit I of pes, 15–18 lamellae under digit IV of manus and 18–22 lamellae under digit IV of pes; males with 4–7 femoral pores on each thigh separated by 7–11 poreless scales from continuous series of 2–5 precloacal pores (n = 7/8); tail with enlarged, strongly keeled, pointed, and spine-like tubercles forming whorls; median row of subcaudals smooth, roughly pentagonal, and distinctly enlarged. Dorsum orange, mottled with numerous light grey spots and fine black spots; a large central black dorsal ocellus on neck flanked anteriorly and posteriorly on each side by elongate dark ocelli, smaller ocellus on occiput flanked on each side by a smaller ocellus; indistinct rows of smaller dark ocelli may be present; venter off-white with black speckles, two distinct pairs of black streaks on throat; original tail in males with 9–11 alternating dark and light grey bands, regenerated tail orange.
Cnemaspis pachaimalaiensis sp. nov. is a member of the bangara clade and can be easily distinguished from all members of the clade by a combination of the following differing or non-overlapping characters: males with continuous series of precloacal pores (versus precloacal pore series medially separated by at least one poreless scale in males of C. agarwali, C. agayagangai sp. nov., C. gracilis, C. jackieii, C. fantastica sp. nov., C. salimalii sp. nov. C. thackerayi, C. shevaroyensis; precloacal pores either absent or medially separated by 2–4 poreless scales in C. mundanthuraiensis); small-sized Cnemaspis with maximum SVL 32 mm (versus medium-sized Cnemaspis, SVL up to 41 mm in C. thackerayi, and C. salimalii sp. nov.); 13–17 tubercles in paravertebral rows (versus only a few irregularly arranged tubercles in paravertebral region in C. mundanthuraiensis, 11 or 12 in C. jackieii); 10–12 rows of dorsal tubercles at mid-body (versus eight or nine rows of dorsal tubercles at mid-body in C. jackieii, 6–8 rows of dorsal tubercles at mid-body in C. mundanthuraiensis); short spine-like tubercles present on flanks (versus spine-like tubercles absent on flanks in C. agarwali, C. jackieii, C. shevaroyensis, and C. thackerayi); 30–33 ventral scales across belly at mid-body (versus 24–26 ventral scales across belly at mid-body in C. agarwali, 21–24 in C. shevaroyensis, and 22–25 in C. thackerayi); a large central black dorsal ocellus on neck flanked anteriorly and posteriorly on each side by elongate dark ocelli, smaller ocellus on occiput flanked on each side by a smaller ocellus; indistinct rows of smaller dark ocelli may be present (versus a single central dorsal ocellus each on occiput and neck in C. gracilis, C. mundanthuraiensis, C. thackerayi; single dorsal ocellus on occiput absent, single dorsal ocellus on neck present in C. salimalii sp. nov.; a single dorsal ocellus each on occiput and neck, a smaller pair on either side just anterior to forelimb insertion in C. jackieii). Cnemaspis pachaimalaiensis sp. nov. is diagnosed against Cnemaspis rudhira sp. nov. as part of its descriptions below.
Adult male in good state of preservation except tail slightly bent towards left (Fig.
Cnemaspis pachaimalaiensis sp. nov. (holotype, NRC-AA-1230): 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; F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar.
Body relatively slender (BW/AGL 0.46), trunk less than half of SVL (AGL/SVL 0.39) without ventrolateral folds; short spine-like scales on flank present (Fig.
Scales on palms and soles granular, smooth, rounded, and flattened; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small granular, weakly keeled, imbricate scales which are twice the size of granules on the body dorsum, largest on anterolateral aspect of the hands and feet; posterolateral aspect of limbs with small weakly keeled to smooth granular scales; ventral aspect of forelimbs with small, smooth, subimbricate scales, larger on lower arm than upper arm; ventral aspect of hindlimb with enlarged, smooth, flattened, subimbricate scales, slightly larger than body ventrals (Fig.
Tail original, entire, subcylindrical, slender, marginally longer than snout-vent length (TL/SVL 1.11; Fig.
Dorsum of head, body, limbs and tail base orange-brown. Head with numerous yellow blotches and some black spots, yellow and dark bands on labials. Three dark postorbital streaks, all terminating anterior to forelimb insertions, suborbital streak continues onto throat. A large central black dorsal ocellus on neck flanked anteriorly and posteriorly on each side by elongate dark ocelli, smaller ocellus on occiput flanked on each side by a smaller ocellus; approximately three rows of three smaller dark ocelli; all ocelli with a diffuse orange margin. Dorsum mottled with smaller light-grey spots and fine black spots. Dorsum of limbs more muted than back, digits with alternating dark and light bands. Tail with seven or eight alternating light grey and black bands with an orange regenerated tip. Venter off-white with black speckles, two distinct pairs of black streaks on throat.
Mensural, meristic and additional character state data for the type series is given in Tables
Mensural (mm) data for the type series of Cnemaspis pachaimalaiensis sp. nov.. Abbreviations are listed in Materials and Methods. * = incomplete tail.
Type | Holotype | Paratypes | ||||||
museum number | NRC-AA-1230 | NRC-AA-1231 | NRC-AA-1232 | NRC-AA-1233 | NRC-AA-1234 | NRC-AA-1235 | NRC-AA-1236 | NRC-AA-1237 |
Sex | Male | Male | Male | Male | Female | Male | Male | Male |
SVL | 28.8 | 30.7 | 28.6 | 31.5 | 33.6 | 30.6 | 30.3 | 31.9 |
TL | 32.2 | 8.5* | 32.0 | 40.7 | 36.3 | 40.0 | 34.0 | 42.6 |
TW | 2.7 | 3.0 | 2.4 | 3.1 | 2.8 | 2.9 | 2.9 | 3.3 |
LAL | 4.3 | 4.5 | 3.7 | 4.5 | 4.8 | 4.6 | 4.5 | 4.6 |
CL | 5.1 | 5.5 | 4.8 | 5.6 | 5.3 | 5.3 | 5.6 | 5.4 |
AGL | 11.5 | 11.7 | 10.8 | 12.7 | 14.4 | 13 | 12 | 13.3 |
BH | 3.0 | 3.3 | 2.7 | 3.4 | 3.8 | 3.1 | 3.3 | 3.5 |
BW | 5.4 | 5.5 | 4.7 | 6.4 | 6.4 | 5.6 | 6.3 | 6.4 |
HL | 7.0 | 7.5 | 7.2 | 8.0 | 7.8 | 7.6 | 7.7 | 8.0 |
HW | 4.8 | 4.9 | 4.6 | 5.5 | 5.2 | 5.3 | 5.1 | 5.2 |
HD | 3.2 | 3.5 | 3.0 | 3.7 | 3.4 | 3.4 | 3.2 | 3.3 |
ED | 1.5 | 1.6 | 1.6 | 1.8 | 1.9 | 1.5 | 1.6 | 1.8 |
EE | 2.4 | 2.3 | 2.2 | 2.5 | 2.5 | 2.4 | 2.3 | 2.6 |
ES | 3.5 | 3.8 | 3.5 | 3.7 | 4.1 | 3.9 | 3.9 | 4.0 |
EN | 2.7 | 3.0 | 2.8 | 3.0 | 3.4 | 3.0 | 3.1 | 3.2 |
IN | 0.8 | 1.0 | 0.8 | 1.1 | 1.1 | 1.0 | 1.0 | 0.9 |
IO | 1.1 | 1.0 | 1.0 | 1.1 | 1.1 | 1.3 | 1.3 | 1.5 |
EL | 0.4 | 0.4 | 0.4 | 0.4 | 0.6 | 0.4 | 0.6 | 0.6 |
Cnemaspis pachaimalaiensis sp. nov. is currently known only from around its type locality (near Mangalam waterfalls, Pachaimalai Hills, between an elevational gradient of ca. 600–1000 m asl.) in Trichy district, Tamil Nadu (Fig.
Meristic data for the type series of Cnemaspis pachaimalaiensis sp. nov.. Abbreviations are listed in Materials and Methods except for: L&R = Left & Right; abs. = absent; * = paravertebral tubercles and lamellae incomplete.
Type | Holotype | Paratypes | ||||||
Museum number | NRC-AA-1230 | NRC-AA-1231 | NRC-AA-1232 | NRC-AA-1233 | NRC-AA-1234 | NRC-AA-1235 | NRC-AA-1236 | NRC-AA-1237 |
Sex | Male | Male | Male | Male | Female | Male | Male | Male |
SL (L&R) | 8&8 | 9&8 | 8&8 | 8&8 | 9&8 | 8&8 | 8&8 | 8&8 |
IL (L&R) | 7&7 | 7&6 | 7&8 | 8&7 | 8&7 | 7&7 | 7&8 | 7&7 |
SL M (L&R) | 6&6 | 6&6 | 6&7 | 6&6 | 6&6 | 6&5 | 6&6 | 6&6 |
IL M (L&R) | 5&5 | 5&5 | 5&6 | 5&5 | 5&5 | 5&5 | 5&5 | 5&5 |
PVT (L&R) | 14&16 | 15&17 | 13*&15 | 15&14 | 13&13 | 16&14 | 16&16 | 15&14 |
DTR | 11 | 12 | 11 | 10 | 10 | 11 | 12 | 10 |
MVSR | 31 | 30 | 30 | 32 | 32 | 32 | 31 | 33 |
VS | 115 | 112 | 118 | 113 | 125 | 117 | 124 | 118 |
LamF1 (L&R) | 9&10 | 10&10 | 10&10 | 10&10 | 9&10 | 10&10 | 10&10 | 11&10 |
LamF4 (L&R) | 15&15 | 17&18 | 16&15 | 16&16 | 15&16 | 17&17 | 17&18 | 17&16 |
LamT1 (L&R) | 9&9 | 10&10 | 9&10 | 10&10 | 10&9 | 10&10 | 10&8* | 12&11 |
LamT4 (L&R) | 19&18 | 22&22 | 19&19 | 18&18 | 20&* | 21&21 | 22&21 | 21&22 |
LamT5 (L&R) | 18&17 | 18&19 | 18&19 | 18&17 | 18&18 | 20&19 | 18&17 | 18&19 |
PP L&R | 3 | 4 | 5 | 3 | 0 | 4 | 3 | 4 |
SBPP | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
SB PP&FP (L&R) | 10&10 | 9&9 | 7&7 | 8&7 | 0 | 8&8 | 10&10 | 9&11 |
FP (L&R) | 5&4 | 5&4 | 7&7 | 6&6 | 0 | 7&7 | 5&6 | 7&6 |
SBFP | A | A | A | A | 0 | A | A | A |
PCT (L&R) | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 | 1&1 |
Additional morphological character states evaluation for the type series of Cnemaspis pachaimalaiensis sp. nov.. abs. = absent; / = data unavailable.
Types | Holotype | Paratypes | ||||||
Museum number | NRC-AA-1230 | NRC-AA-1231 | NRC-AA-1232 | NRC-AA-1233 | NRC-AA-1234 | NRC-AA-1235 | NRC-AA-1236 | NRC-AA-1237 |
Sex | Male | Male | Male | Male | Female | Male | Male | Male |
Anterior extra-brillar fringe scales enlarged (1) or not enlarged (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Ventral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Gular scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Pectoral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Precloacal pores continuous (1) or separated (0) | 1 | 1 | 1 | 1 | abs. | 1 | 1 | 1 |
Precloacal pores elongate (1) or round (0) | 1 | 1 | 0 | 1 | abs. | 1 | 1 | 1 |
femoral pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | abs. | 1 | 1 | 1 |
Dorsal pholidosis homogeneous (1) or heterogeneous (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 |
Tubercles linearly arranged (1) or more random (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Spine-like scales on flank present (1) or absent (0) | 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 keeled subcaudals (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 | 0 | 1 | 0 | 1 |
Subtibial scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Occipital ocellus present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Ocelli anterior of the shoulder present (1) or absent (0) & number | 1 (3) | 1 (2*) | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) | 1 (3) |
Ocelli posterior of the shoulder present (1) or absent (0) & number | 1 (+3) | 1 (2) | 1 (2) | 0 | 1 (2) | 0 | 1 (2) | 1 (2) |
Original tail banded (1) or not (0) | 1 | / | 1 | 1 | 1 | 1 | 1 | 1 |
Cnemaspis cf. gracilis
NRC-AA-1238 (AK 566), adult male, from near Sri Salaipaarai Muniappan Temple, Yercaud, in the Shevaroy hill range (11.7761°N, 78.1900°E; 1060 m asl.), Salem district, Tamil Nadu state, India, collected by Akshay Khandekar, Swapnil Pawar, and Tejas Thackeray on 2nd January 2019.
NRC-AA-1246 (AK 567), adult male, NRC-AA-1247 (AK 568), adult female, same data as holotype; NRC-AA-1239 (AK 208), NRC-AA-1240 (AK 209), NRC-AA-1241 (AK 212) adult males, from Yercaud Ghat, in the Shevaroy hill range (11.7796°N, 78.1911°E; 1200 m asl.), and NRC-AA-1242 (AK 213) adult female (11.7655°N, 78.1884°E; 800 m asl.), collected by Akshay Khandekar, Ishan Agarwal, Nikhil Gaitonde, on 18th December 2018; NRC-AA-1243 (AK 539), NRC-AA-1244 (AK 541), NRC-AA-1245 (AK 543), adult males, from near Botanical garden, Yercaud, in the Shevaroy hill range (11.7810°N, 78.2035°E; 1400 m asl.), same collection data as holotype.
The specific epithet is from the Sanskrit rudhira which means blood, alluding to the blood-red colouration of this beautiful species, and is used as a noun in apposition.
Scarlet dwarf gecko.
A small-sized Cnemaspis, snout to vent length up to 33.8 mm (n = 10). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with a fairly regularly arranged rows of enlarged, strongly keeled, conical tubercles; last one or two rows of enlarged tubercles on flank weakly keeled, spine-like; 10–12 rows of dorsal tubercles at mid-body, 13–17 tubercles in paravertebral rows; ventral scales smooth, subcircular, subimbricate, subequal from chest to vent, 30–32 scales across belly at mid-body, 101–121 longitudinal scales from mental to cloaca; subdigital scansors smooth, mostly unpaired, unnotched; 8–11 lamellae under digit I of manus and pes; 14–17 lamellae under digit IV of manus and 17–21 lamellae under digit IV of pes; males with four or five femoral pores on each thigh separated by 6–9 poreless scales from series of 4–6 precloacal pores, precloacal pores separated medially by single (rarely 2, n = 1/8) poreless scales; tail with enlarged, strongly keeled, pointed, and spine-like tubercles forming whorls; median row of subcaudals smooth, roughly rectangular, and distinctly enlarged. Dorsum orange, mottled with numerous small light grey spots and fine black spots with an indistinct series of light grey vertebral blotches extending from neck to tail base; single central black dorsal ocellus on neck and smaller ocellus on occiput, separated by a light grey blotch; venter off-white with black speckles, margin of throat strongly marked; original tail in males grey or with 9–11 alternating dark and light grey bands, regenerated tail orange.
Cnemaspis rudhira sp. nov. is a member of the bangara clade and can be easily distinguished from all members of the clade by a combination of the following differing or non-overlapping characters: small-sized Cnemaspis with maximum SVL 34 mm (versus medium-sized Cnemaspis, SVL up to 41 mm in C. thackerayi, and C. salimalii sp. nov.); 13–17 tubercles in paravertebral rows (versus only a few irregularly arranged tubercles in paravertebral region in C. mundanthuraiensis, 11 or 12 in C. jackieii); 10–12 rows of dorsal tubercles at mid-body (versus eight or nine rows of dorsal tubercles at mid-body in C. jackieii, 6–8 rows of dorsal tubercles at mid-body in C. mundanthuraiensis); spine-like tubercles present on flanks (versus spine-like tubercles absent on flanks in C. agarwali, C. jackieii, C. shevaroyensis, and C. thackerayi); 30–32 ventral scales across belly at mid-body (versus 24–26 ventral scales across belly at mid-body in C. agarwali, 26–29 (rarely 30) in C. gracilis, 21–24 in C. shevaroyensis, and 22–25 in C. thackerayi); males with two (rarely 3) precloacal pore on each side which are separated medially by single (rarely 2) poreless scales (versus males with single (rarely 2) precloacal pore on each side which are separated medially by 2–4 poreless scales in C. gracilis; precloacal pores either absent or single precloacal pores on each side which are separated medially by 2–4 poreless scales in C. mundanthuraiensis; males with continuous series of precloacal pores in C. pachaimalaiensis sp. nov.; single central dorsal ocellus each on occiput and neck (versus a single dorsal ocellus present on occiput and neck, two pairs on either side just anterior and sometimes posterior to forelimb insertions in C. agarwali; a single central dorsal ocellus each on occiput and neck, ocellus on neck flanked anteriorly on each side by a slightly larger ocellus in C. agayagangai sp. nov.; a single central ocellus on neck, flanked posteriorly by a pair of much larger squarish blotches and anteriorly by a pair of subequal squarish blotches, indistinct spot on occiput in C. fantastica sp. nov.; a large central black dorsal ocellus on neck flanked anteriorly and posteriorly on each side by elongate dark ocelli, smaller ocellus on occiput flanked on each side by a smaller ocellus; indistinct rows of smaller dark ocelli may be present in C. pachaimalaiensis sp. nov.; a single dorsal ocellus present on occiput and neck, two pairs on either side just anterior and posterior to forelimb insertions in C. shevaroyensis.
Adult male in good state of preservation except tail tip slightly bend towards left, longitudinal skin fold on vertebral region between limb insertions (Fig.
Cnemaspis rudhira sp. nov. (holotype, NRC-AA-1238): 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; F ventral aspect of left pes. Scale bars 5 mm; photos by Akshay Khandekar.
Body relatively slender (BW/AGL 0.51), trunk less than half of SVL (AGL/SVL 0.38) without ventrolateral folds; spine-like scales on flank present (Fig.
Scales on palm and soles granular, smooth, subcircular, subimbricate and flattened; scales on dorsal aspects of limbs heterogeneous in shape and size; mixture of small granular, weakly keeled, imbricate scales which are twice the size of granules on the body dorsum, largest on anterolateral aspect of the hands and feet; posterolateral aspect of limbs with small weakly keeled to smooth granular scales; scales on lower arm and shank small, subimbricate, and keeled; ventral aspect of forelimbs with small, smooth, subimbricate scales, larger on lower arm than upper arm; ventral aspect of hindlimb with enlarged, smooth, flattened, subimbricate scales, slightly larger than body ventrals (Fig.
Tail original except tip (5.1 mm) which is regenerated, entire, subcylindrical, slender, slightly longer than snout-vent length (TL/SVL 1.30; Fig.
Dorsum of head, body, limbs and tail base orange. Head with numerous light grey and yellow blotches and fine black spots, light grey and dark bands on labials; indistinct grey postorbital streaks. A single central black dorsal ocellus on neck and a smaller one on occiput separated by a larger light grey blotch, both ocelli with an orange margin. Dorsum with numerous light-grey spots and fine black spots and six light grey vertebral blotches from forelimb insertions to tail base. Dorsum of limbs with yellow reticulation, digits with alternating dark and light bands. Tail grey with an orange regenerated tip. Venter off-white with black speckles, margin of throat strongly marked.
A Colouration in life of Cnemaspis rudhira sp. nov. showing sexual dichromatism: top adult female (paratype, NRC-AA-1247), bottom adult male (holotype, NRC-AA-1238), photo by Tejas Thackeray; B Habitat at the type locality of Cnemaspis rudhira sp. nov. showing general habitat, rocky outcrop in evergreen forest. Photo by Akshay Khandekar.
Mensural, meristic and additional character state data for the type series is given in Tables
Mensural (mm) data for the type series of Cnemaspis rudhira sp. nov.. Abbreviations are listed in Materials and Methods. * = incomplete tail.
Type | Holotype | Paratypes | ||||||||
Museum number | NRC-AA-1238 | NRC-AA-1239 | NRC-AA-1240 | NRC-AA-1241 | NRC-AA-1242 | NRC-AA-1243 | NRC-AA-1244 | NRC-AA-1245 | NRC-AA-1246 | NRC-AA-1247 |
Sex | Male | Male | Male | Male | Female | Male | Male | Male | Male | Female |
SVL | 32.3 | 32.9 | 32.1 | 30.5 | 27.9 | 33.8 | 31.5 | 33.5 | 33.7 | 29.8 |
TL | 42.3 | 2.2* | 12.1* | 19.7* | 16.7* | 35.1 | 41.8 | 43.5 | 43.8 | 35.6 |
TW | 3.3 | 3.3 | 3.3 | 3.0 | 2.3 | 3.2 | 3.0 | 3.1 | 3.0 | 2.7 |
LAL | 4.9 | 4.9 | 4.7 | 4.5 | 3.7 | 4.9 | 4.4 | 4.9 | 5.0 | 4.1 |
CL | 5.9 | 6.1 | 5.7 | 5.7 | 4.6 | 6.0 | 5.5 | 5.9 | 6.0 | 5.1 |
AGL | 12.5 | 12.6 | 12.9 | 12.6 | 12.6 | 14.3 | 12.5 | 14.3 | 15.0 | 12.4 |
BH | 3.5 | 3.4 | 3.4 | 2.6 | 2.2 | 3.1 | 2.4 | 3.7 | 3.1 | 3.0 |
BW | 6.4 | 5.8 | 5.9 | 5.4 | 5.0 | 5.4 | 5.1 | 5.4 | 5.6 | 5.7 |
HL | 8.0 | 8.6 | 8.1 | 7.6 | 7.0 | 8.3 | 8.0 | 8.1 | 7.4 | 7.3 |
HW | 5.3 | 5.6 | 5.5 | 5.0 | 4.6 | 5.2 | 5.1 | 5.2 | 5.0 | 4.9 |
HD | 3.8 | 3.7 | 3.6 | 3.2 | 2.5 | 3.4 | 3.1 | 3.5 | 3.7 | 3.5 |
ED | 1.6 | 1.8 | 1.8 | 1.6 | 1.5 | 1.8 | 1.7 | 1.6 | 1.7 | 1.5 |
EE | 2.5 | 2.6 | 2.5 | 2.4 | 2.1 | 2.5 | 2.5 | 2.4 | 2.3 | 2.4 |
ES | 4.2 | 4.3 | 4.2 | 3.5 | 3.2 | 4.1 | 4.1 | 4.1 | 3.8 | 3.5 |
EN | 3.3 | 3.2 | 3.4 | 3.0 | 2.7 | 3.4 | 3.3 | 3.3 | 3 | 2.9 |
IN | 1.0 | 1.1 | 1.0 | 1.0 | 0.8 | 1.1 | 0.9 | 1 | 0.9 | 0.9 |
IO | 1.3 | 1.1 | 1.1 | 1.0 | 0.7 | 1.1 | 1.4 | 1.5 | 1.3 | 1.1 |
EL | 0.4 | 0.5 | 0.5 | 0.5 | 0.4 | 0.4 | 0.6 | 0.5 | 0.4 | 0.4 |
Cnemaspis rudhira sp. nov. is known from a broad elevation gradient of ca. 800–1400 m asl. around its type locality, Yercaud, in the Shevaroy hills, Salem district, Tamil Nadu (Fig.
Meristic data for the type series of Cnemaspis rudhira sp. nov.. Abbreviations are listed in Materials and Methods except for: L&R = Left & Right; abs. = absent; * = lamellae incomplete.
Type | Holotype | Paratypes | ||||||||
Museum number | NRC-AA-1238 | NRC-AA-1239 | NRC-AA-1240 | NRC-AA-1241 | NRC-AA-1242 | NRC-AA-1243 | NRC-AA-1244 | NRC-AA-1245 | NRC-AA-1246 | NRC-AA-1247 |
Sex | Male | Male | Male | Male | Female | Male | Male | Male | Male | Female |
SL (L&R) | 10&11 | 7&9 | 9&9 | 9&10 | 7&7 | 10&10 | 9&8 | 10&10 | 10&9 | 8&8 |
IL (L&R) | 8&9 | 7&7 | 9&8 | 8&8 | 6&7 | 8&10 | 7&7 | 10&7 | 8&9 | 7&6 |
SL M (L&R) | 6&6 | 6&6 | 6&6 | 6&6 | 6&6 | 6&6 | 6&6 | 6&6 | 7&7 | 6&6 |
IL M (L&R) | 5&5 | 5&5 | 5&5 | 5&6 | 6&5 | 5&6 | 5&6 | 6&6 | 5&5 | 5&5 |
PVT (L&R) | 14&15 | 13&14 | 14&14 | 13&14 | 15&14 | 14&14 | 13&14 | 14&14 | 17&15 | 14&15 |
DTR | 12 | 10 | 11 | 11 | 12 | 11 | 11 | 10 | 11 | 12 |
MVSR | 31 | 32 | 31 | 32 | 31 | 32 | 30 | 31 | 31 | 30 |
VS | 115 | 112 | 101 | 118 | 110 | 121 | 116 | 116 | 118 | 115 |
LamF1 (L&R) | 11&10 | 9&9 | 9&9 | 10&10 | 9&9 | 11&11 | 9&9 | 10&10 | 8&8 | 10&10 |
LamF4 (L&R) | 16&16 | 15&15 | 15&4* | 17&16 | 15&15 | 17&16 | 14&15 | 15&16 | 16&15 | 16&16 |
LamT1 (L&R) | 10&10 | 2*&11 | 10&10 | 10&9 | 9&9 | 11&10 | 9&10 | 10&11 | 8&8 | 10&10 |
LamT4 (L&R) | 21&21 | 19&20 | 10*&19 | 19&19 | 19&20 | 19&20 | 17&17 | 19&20 | 17&18 | 20&20 |
LamT5 (L&R) | 19&17 | 18&19 | 18&18 | 19&0* | 17&18 | 20&19 | 18&17 | 19&19 | 16&17 | 18&17 |
PP L&R | 2&2 | 2&2 | 2&2 | 3&3 | abs. | 2&2 | 2&2 | 2&2 | 2&2 | abs. |
SBPP | 1 | 1 | 1 | 1 | abs. | 1 | 1 | 1 | 2 | abs. |
SB PP&FP (L&R) | 9&9 | 8&8 | 6&6 | 7&7 | abs. | 8&8 | 7&7 | 8&7 | 7&7 | abs. |
FP (L&R) | 4&4 | 4&4 | 5&4 | 5&5 | abs. | 5&4 | 5&4 | 5&5 | 5&5 | abs. |
SBFP | abs. | abs. | abs. | abs. | abs. | abs. | 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 | 1&1 |
Additional morphological character states evaluation for the type series of Cnemaspis rudhira sp. nov.. abs. = absent; / = data unavailable.
Types | Holotype | Paratypes | ||||||||
Museum number | NRC-AA-1238 | NRC-AA-1239 | NRC-AA-1240 | NRC-AA-1241 | NRC-AA-1242 | NRC-AA-1243 | NRC-AA-1244 | NRC-AA-1245 | NRC-AA-1246 | NRC-AA-1247 |
Sex | Male | 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 | 1 |
Ventral scales keeled (1) or smooth (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 |
Pectoral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Precloacal pores continuous (1) or separated (0) | 0 | 0 | 0 | 0 | abs. | 0 | 0 | 0 | 0 | abs. |
Precloacal pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | abs. | 1 | 1 | 1 | 1 | abs. |
femoral pores elongate (1) or round (0) | 1 | 1 | 1 | 1 | abs. | 1 | 1 | 1 | 1 | abs. |
Dorsal pholidosis homogeneous (1) or heterogeneous (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 |
Tubercles linearly arranged (1) or more random (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Spine-like scales on flank present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Lateral caudal furrows present (1) or absent (0) | 1 | / | / | 1 | / | / | 1 | 1 | 1 | 1 |
Subcaudals keeled (1) or smooth (0) | 0 | / | / | 0 | / | / | 0 | 0 | 0 | 0 |
Single median row of keeled subcaudals (1) or smooth (0) | 0 | / | / | 0 | / | / | 0 | 0 | 0 | 0 |
Caudal tubercles encircle tail (1) or not (0) | 1 | / | / | 1 | / | / | 1 | 1 | 1 | 1 |
Enlarged median subcaudal scale row (1) or not (0) | 1 | / | / | 1 | / | / | 1 | 1 | 1 | 1 |
Enlarged femoral scales present (1) or absent (0) | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 1 |
Subtibial scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Occipital ocellus present (1) or absent (0) | 1 | 0 | 0 | 1 | 1 | 0 | 0 | P | P | P |
Ocelli anterior of the shoulder present (1) or absent (0) & number | 1 (1) | 1 (1) | 1 (1) | 1 (1) | 1 (1) | 1 (1) | 1 (1) | 1 (1) | 1 (1) | 1 (1) |
Ocelli posterior of the shoulder present (1) or absent (0) & number | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Original tail banded (1) or not (0) | 0 | / | 1 | 0 | / | 0 | 0 | 0 | 1 | 0 |
The description of these five new species from the Shevaroy Group of hills in southern India brings the number of Cnemaspis species known from peninsular India outside the Western Ghats to 23 and from the gracilis clade to 11. Seven divergent lineages of the gracilis clade are now known from three hill blocks that are < 70 km from one another, with three lineages each in Yercaud and Kollimalai, the two largest massifs. The massif of Yercaud rises from < 350–1623 m asl, Kollimalai from < 200–1400 m asl, and Pachaimalai from < 200–1000 m, each < 500 km2 in area. Though at a relatively small spatial scale, the incredibly high microendemism within a single clade in a continental setting is perhaps comparable in sheer numbers of species per unit area to only a few squamate radiations on large continental islands such as Bavayia Roux, Dierogekko Bauer, Jackman, Sadlier & Whitaker and Nannoscincus Günther on Grand Terre in New Caledonia (
Potential explanations for the high level of diversity and microendemism seen in the bangara clade in the Shevaroy landscape include the role of elevation and intrinsic habitat heterogeneity, dispersal, and sexual selection. These mountains are known to form part of a sky-island complex for geckos of the genus Hemiphyllodactylus with very different forests and climate from the surrounding lowland habitats (
Cnemaspis fantastica sp. nov. and its sister taxon C. shevaroyensis have the lowest genetic divergence within known species of Indian Cnemaspis at 4.6%. Though a 5 % p-distance cut-off has been proposed as indicative of species divergence in gekkonids (e.g.,
All the species of the bangara clade are sexually dichromatic, males in general with much brighter dorsal colouration, dark ocelli on the forebody, strong markings on the throat and a strongly banded tail, apart from the two medium-bodied species that are restricted to high elevations, C. salimalii sp. nov. and C. thackerayi which are less striking in the differences between sexes. As these species are conspicuously diurnal, there is a potential role for sexual selection based on visual traits to drive diversification within the group (
The ancestral state for colour pattern of the forebody (in males) appears to be a single, central ocellus on the neck (seen in C. salimalii sp. nov. and C. thackerayi), and the most common colour pattern is a single, central ocellus each on the neck and occiput (seen in C. gracilis, C. mundanthuraiensis and C. rudhira sp. nov.; C. jackieii has two additional small lateral ocelli). The remaining species have multiple ocelli on the neck (C. agarwali and C. agayagangai sp. nov.) or multiple ocelli on the neck and between forelimbs (C. fantastica sp. nov., C. pachaimalaiensis sp. nov. and C. shevaroyensis). Though our phylogeny does not receive high support at more basal nodes and is based on a partial fragment of one mitochondrial gene, both multi-ocellate and two single central ocelli-bearing morphs occur in multiple, well-supported clades within the phylogeny. Understanding the evolution of colour pattern in this diverse clade needs more sampling of genes and taxa.
Other geckos within this landscape show contrasting patterns of distribution — Cnemaspis yercaudensis from the mysoriensis clade of South Asian Cnemaspis is a truly widely distributed mid to high elevation species, with little genetic structure or morphological differentiation between populations in the Shevaroy landscape (
These new species join a large proportion of gekkonid species known only from their type localities (23 %,
We are thankful to the Tamil Nadu Forest Department for the necessary permits to carry out this study (permit no. 53/2018). We thank Teja Bhargava (former DFO, Jawadhu) for his help and co-ordination throughout fieldwork. Fieldwork assistance was provided by Swapnil Pawar, Vaibhav Patil, Nikhil Gaitonde, Anuradha Batabyal, Varad Giri, Aniruddha Dutta-Roy, Chaitanya R., Satpal Gangalmale and Vivek Waghe. Tarun Karmakar (NCBS field station and museum facility, Bengaluru) helped with the specimen registrations. We are thankful to Uma Ramakrishnan for lab support at NCBS, and Lee Grismer and two anonymous reviewers for their comments.
Material examined.
Institutional abbreviations are as follows: National Centre for Biological Sciences, Bengaluru (NCBS-AU/ NCBS-BH/NRC-AA/ 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 mundanthuraiensis: holotype, NRC-AA-1175 (adult male); paratypes, NRC-AA-1176, NRC-AA-1177, NRC-AA-1178, BNHS 2822, BNHS 2824, and BNHS 2825, (adult males), BNHS 2823, (adult female), NRC-AA-1179, (subadult female) from Papanasam reserve forest, Mundanthurai forest range, Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu state, India.