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Research Article
A multitude of spots! Five new microendemic species of the Cnemaspis gracilis group (Squamata: Gekkonidae) from massifs in the Shevaroy landscape, Tamil Nadu, India
expand article infoIshan Agarwal§, Tejas Thackeray, Akshay Khandekar§|
‡ Thackeray Wildlife Foundation, Mumbai, India
§ National Centre for Biological Sciences, Tata Institute of Fundamental Research, Bengaluru, India
| Shivaji University, Kolhapur, India
Open Access

Abstract

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.

Keywords

Endemic species, integrative taxonomy, microendemism, mountains, southern India, species radiation, taxonomy

Introduction

Diurnality has evolved multiple times within the ancestrally nocturnal Gekkonidae, including numerous reversals (Gamble et al. 2016). Among the most diverse genera of chiefly diurnal gekkonids is the South Asian clade of the paraphyletic Cnemaspis Strauch, 1887 that includes over 100 described species with a disjunct distribution across parts of peninsular India, Sri Lanka and northeast India in the Indian subcontinent, as well as eastern Southeast Asia (Iskandar et al. 2017; Lee et al. 2019; Agarwal et al. 2020a, 2021a; Amarasinghe et al. 2021; Pal et al. 2021; Uetz et al. 2022). Most species of South Asian Cnemaspis are diurnal or crepuscular/ cathemeral, apart from the largely nocturnal wynadensis clade and some species of the beddomei clade (Pal et al. 2021).

South Asian Cnemaspis originated in the Western Ghats in the Paleocene-Eocene and are largely restricted to cool habitats (Agarwal et al. 2020b), 10 of the 13 broad clades within the group endemic to peninsular India and seven of these to the Western Ghats (Pal et al. 2021; Khandekar et al. 2022a). Of the three clades that are distributed outside the Western Ghats, the bangara and mysoriensis clades with four and seven species, respectively, are restricted to the southern edge of the Mysore Plateau and associated hills; and only the C. gracilis clade with six species is distributed in both the Western Ghats and other parts of peninsular India (Agarwal et al. 2021b; Pal et al. 2021; Khandekar et al. 2022a, 2022b).

Cnemaspis gracilis (Beddome) was described by Beddome (1870) from the ‘Palghat Hills’ and the species was thought to be widely distributed in southern India (Smith 1935; Srinivasulu and Srinivasulu 2013). Manamendra-Arachchi et al. (2007) designated a lectotype for C. gracilis and redescribed the species, which is now known with certainty from three localities in Kerala and Tamil Nadu in the vicinity of the Palghat Gap (Khandekar et al. 2022b). The additional five species of the complex have been described in the last three years, each known only from the vicinity of their respective type localities (Khandekar 2019; Khandekar et al. 2019, 2022b; Pal et al. 2021). Cnemaspis jackieii Pal, Mirza, Dsouza & Shanker and C. mundanthuraiensis Khandekar, Thackeray & Agarwal are distributed on the eastern slopes of the Western Ghats, C. shevaroyensis Khandekar, Gaitonde & Agarwal and C. thackerayi Khandekar, Gaitonde & Agarwal in the Shevaroy Hills (Yercaud; along with an additional unnamed lineage C. cf. gracilis), and C. agarwali Khandekar from an isolated hillock just south of the Mysore Plateau, beside C. gracilis from the Palghat Gap (Fig. 1).

Figure 1. 

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 (Khandekar et al. 2019). Yercaud forms a part of the broader Shevaroy Group or Shevaroyan landscape (sensu Raheem et al. 2014), which includes a number of smaller hills and the major ranges with maximum elevations of > 1000 m asl. of Sitteri and Kalrayan Hills to the east of Yercaud, and Kollimalai and Pachaimalai hills to the south and southeast (Fig. 1). We sampled the Shevaroyan landscape as part of an ongoing project on the systematics and taxonomy of peninsular Indian lizards and discovered multiple unidentified Cnemaspis spp. of the bangara clade. In this paper, we describe five new species from the Yercaud-Kollimalai-Pachaimalai Hill complex using morphological data and mitochondrial sequence data to demonstrate the uniqueness of these lineages.

Materials and Methods

Taxon sampling

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).

Molecular data

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 1). The primers L4437 and H5934 (Macey et al. 1997) were used to amplify the protein coding ND2 gene, and L4437 was used to sequence a partial fragment of ND2. Both PCR and sequencing were outsourced to Medauxin (Bangalore, India). We added published sequences of the Cnemaspis gracilis clade and used C. australis Manamendra-Arachchi, Batuwita & Pethiyagoda and C. monticola Manamendra-Arachchi, Batuwita & Pethiyagoda as outgroups (Table 1; after Khandekar et al. 2019, 2022a, 2022c; Pal et al. 2021). We used MEGA 5.2 (Tamura et al. 2011) for sequence alignment using default settings in ClustalW (Thompson et al. 1994), translation to amino acids to check for erroneous stop codons, and to calculate uncorrected pairwise p-distance with the partial deletion option.

Table 1.

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 (Lanfear et al. 2012) which selected partitions by codon position and the HKY + I model for codon position (cp) 1, HKY + G for cp 2 and GTR + G for cp 3. We reconstructed a Maximum Likelihood (ML) phylogeny with the GTR + G model applied for each codon partition in RAxML HPC 8.2.10 (Stamatakis 2014) (since RAxML allows only a single model across partitions) as implemented in raxmlGUI 2.0.9 (Edler et al. 2020) with 10 independent ML runs and support assessed with 500 rapid bootstraps. A partitioned Bayesian analysis was carried out in MrBayes 3.2.7 (Ronquist and Huelsenbeck 2003) with model parameters unlinked across partitions. The final analysis was run for 2,000,000 generations sampling every 200 generations, implementing two parallel runs with four chains each (one cold and three hot) and convergence was determined based on standard deviation of split frequencies (<0.01) and ESS (>>200). A Maximum Clade Credibility tree was built using TreeAnnotator 1.10.4 (Drummond et al. 2012) after discarding the first 25% of trees as burn-in.

Morphological and meristic data

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 1); and comparative morphological data for the most recently described species — C. jackieii was obtained from the original description (Pal et al. 2021). Meristic counts and measurements were taken under a ZEISS Stemi 305 stereo dissecting microscope and on the right side of the body where possible. Colour pattern was recorded from photographs taken in life and specimens in case no photographs were available. We use ocelli to refer to the distinct, dark spots on head and forebody. All measurements were taken with a Mitutoyo digital vernier calliper (to the nearest 0.1 mm). We follow Agarwal et al. (2020a) for body size categories for South Asian Cnemaspis; mensural, meristic and additional morphological character states evaluation is in accordance with Khandekar et al. (2019,): snout vent length (SVL), axilla to groin length (AGL), body height (BH), body width (BW), forearm length (FL), crus length (CL), tail length (TL), tail width (TW), head length (HL), head width (HW), head depth (HD), eye diameter (ED), eye to nares distance (EN), eye to snout distance (ES), eye to ear distance (EE), ear length (EL), internarial distance (IN), interorbital distance (IO); meristic data recorded for all specimens were number of supralabials (SL), infralabials (IL), supralabials at midorbital position (SL M), infralabials at midorbital position (IL M), dorsal tubercle rows including longitudinal rows of spine-like scales on lower flank (DTR), paravertebral tubercles (PVT), ventral scales (VS), mid-body scale rows across the belly (MVSR), precloacal pores (PP), femoral pores (FP), poreless scales between precloacal and femoral pores (SB PP&FP), poreless scales between precloacal pores (SB PP), poreless scales between femoral pores (SB FP), postcloacal tubercles (PCT), transverse subdigital lamellae on finger 1 (LamF1), finger 4 (LamF4), toe 1 (LamT1), toe 4 (LamT4), toe 5 (LamT5).

Results

Phylogenetic relationships

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. 1). These nine lineages fall into two broad clades, the first including C. gracilis as the sister taxon to C. mundanthuraiensis + C. jackieii, these three taxa collectively the sister taxon to Cnemaspis pachaimalaiensis sp. nov. (see description below). The second clade includes one subclade with Cnemaspis agayagangai sp. nov. (see description below) sister to C. shevaroyensis + Cnemaspis fantastica sp. nov. (see description below) and the other subclade with C. agarwali sister to C. rudhira sp. nov. (see description below) (referred to as C. cf. gracilis by Khandekar et al. 2019).

Figure 2. 

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 2). Of the five new species, Cnemaspis salimalii sp. nov. shows ≥ 14.3 % divergence from other members of the clade, Cnemaspis rudhira sp. nov. ≥ 11.0 %, Cnemaspis pachaimalaiensis sp. nov. ≥ 8.5 %, Cnemaspis agayagangai sp. nov. ≥ 7.8 %, and Cnemaspis fantastica sp. nov. ≥ 4.6 % divergence. We describe these five divergent lineages as new species using morphological data.

Table 2.

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

Systematics

Cnemaspis salimalii sp. nov.

Figs 3, 4, 5, 6A, 7A, 8A; Tables 3, 5

Holotype

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.

Paratypes

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.

Etymology

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.

Suggested Common Name

Salim Ali’s dwarf gecko.

Diagnosis

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.

Comparison with members of C. gracilis clade

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.

Table 3.

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
Table 4.

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

Description of the holotype

Adult male in good state of preservation except regenerated portion of tail tip slightly bent towards right (Fig. 3A–E). SVL 35.0 mm, head short (HL/SVL 0.25), wide (HW/HL 0.67), not strongly depressed (HD/HL 0.41), distinct from neck. Loreal region marginally inflated, canthus rostralis not distinct. Snout almost half of head length (ES/HL 0.47), nearly 2.5 times eye diameter (ES/ED 2.33); scales on snout and canthus rostralis subcircular, subequal, smooth anteriorly, becoming weakly keeled, and conical posteriorly; much larger than those on forehead and interorbital region; scales on forehead similar to those on snout and canthus rostralis except smaller and weakly conical; scales on interorbital region even smaller, granular and smooth to weakly keeled; scales on occipital and temporal region heterogeneous, slightly enlarged, weakly keeled, conical tubercles intermixed with smaller, weakly keeled and weakly conical granular scales (Fig. 4A). Eye small (ED/HL 0.20) with round pupil; supraciliaries short, larger anteriorly; seven interorbital scale rows across narrowest point of frontal bone; 30–32 scale rows between left and right supraciliaries at mid-orbit (Fig. 4A, C). Ear-opening deep, oval, small (EL/HL 0.06); eye to ear distance greater than diameter of eye (EE/ED 1.50) (Fig. 4C). Rostral more than 2× wider (1.86 mm) than high (0.80 mm), incompletely divided dorsally by a strongly developed rostral groove and internasal scale for more than half of its height; a single enlarged supranasal on each side, marginally larger than postnasals, separated from each other by a much smaller, elongated internasal scale; two postnasals, upper postnasal marginally larger than lower; rostral in contact with supralabial I, nostril, supranasal, and lower postnasal on either side; nostrils oval, surrounded by two postnasals, supranasal, and rostral on either side; two rows of scales separate orbit from supralabials (Fig. 4C). Mental enlarged, subtriangular, slightly wider (2.01 mm) than high (1.54 mm); two pairs of postmentals, inner pair roughly rectangular, much shorter (0.77 mm) than mental, separated from each other below mental by a single enlarged median chin shield; inner pair bordered by mental, infralabial I, outer postmental, enlarged median chin shield and two enlarged chin shield on either side; outer postmentals roughly subcircular, even smaller (0.63 mm) than inner pair, bordered by inner postmentals, infralabial I and II, and four enlarged chin shields on either side; four enlarged gular scales between left and right outer postmentals; all chin scales bordering postmentals more or less flat, subcircular, smooth, and much smaller than outermost postmentals; scales on rest of throat, even smaller, subequal, and smooth (Fig. 4B). Infralabials bordered below by a row or two of slightly enlarged, much elongated scales, decreasing in size posteriorly. Ten supralabials up to angle of jaw and six at midorbital position on either side; supralabial I largest, rest of the series gradually decreasing in size posteriorly; eight infralabials up to angle of jaw and six at midorbital position on either side; infralabial I largest, gradually decreasing in size posteriorly (Fig. 4C).

Table 5.

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
Figure 3. 

Cnemaspis salimalii sp. nov. (holotype, NRC-AA-1204): A dorsal aspect of body; B ventral aspect of body; C dorsal aspect of tail; D ventral aspect of tail; E lateral aspect of tail. Scale bars 10 mm; photos by Akshay Khandekar.

Figure 4. 

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. 5A–C). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with irregularly arranged rows of enlarged, strongly keeled, conical tubercles; tubercles in approximately 12 longitudinal rows at mid-body including short spine-like scales at lower flank; tubercles in paravertebral rows irregular (Fig. 5A, C). Ventral scales much larger than granular scales on dorsum, smooth, subcircular, subimbricate, subequal from chest to vent; mid-body scale rows across belly 30; 109 scales from mental to anterior border of cloaca (Fig. 5B). Scales on base of neck similar to those on belly; gular region with much smaller, subequal, smooth, flattened scales, those bordering postmentals enlarged, smooth, subcircular, and more or less flattened (Fig. 4B). Five femoral pores on left thigh and four on right, separated by seven poreless scales on left and six on right side from two precloacal pores, precloacal pores separated medially by three poreless scales (Fig. 4D).

Figure 5. 

Cnemaspis salimalii sp. nov. (holotype, NRC-AA-1204): A dorsal aspect of mid-body; B ventral aspect of mid-body; C right side lateral aspect of mid-body. Scale bars 5 mm; photos by Akshay Khandekar.

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. 3A, B). Forelimbs and hindlimbs moderately long, slender (LAL/SVL 0.14; CL/SVL 0.19); digits long, with 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: (1-3-4-4-4 right manus, 1-4-5-8-5 right pes), (1-3-3-4-4 left manus, Fig. 4E; 14585 left pes, Fig. 4F); distal lamellae series: (11-12-13-13-11 right manus, 11-12-14-14-14 right pes), (11-12-13-12-11 left manus, Fig. 4E; 111214–14–14 left pes, Fig. 4F). Relative length of digits (measurements in mm in parentheses): IV (3.8) > III (3.5) > V (3.3) > II (3.1) > I (2.6) (left manus); IV (4.9) > V (4.2) = III (4.2) > II (3.7) > I (2.5) (left pes).

Tail original except tip (15.1 mm) which is regenerated, entire, subcylindrical, slender, slightly longer than snout-vent length (TL/SVL 1.28; Fig. 3C–E). Dorsal scales on tail base weakly keeled, granular, similar in size and shape to granular scales on mid-body dorsum, gradually becoming larger, flattened, imbricate posteriorly, intermixed with enlarged, strongly keeled, distinctly pointed, conical tubercles; enlarged tubercles on the tail forming whorls; six tubercles each on first eight whorls, four in whorls 9–11, only paravertebral tubercles in whorls 12–14, rest of tail tip regenerated (Fig. 3C, E). Scales on ventral aspect of tail much larger than those on dorsal aspect, subimbricate, smooth; median series distinctly larger than rest, roughly pentagonal; scales on tail base slightly smaller than those on mid-body ventrals, smooth, imbricate; a single enlarged, smooth, and conical postcloacal spur on each side (Fig. 3D).

Colouration in life (Fig. 6A)

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.

Figure 6. 

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.

Variation and additional information from type series

Mensural, meristic and additional character state data for the type series is given in Tables 13 respectively. There are four adult and two subadult males and two adult females, ranging in size from 31.3–41.3 mm (Fig. 7A). All paratypes resemble the holotype except as follows: tubercles in paravertebral rows regular in all adult paratypes (condition not discernible in subadults), upper postmentals in contact with each other below mental in NRC-AA-1207, NRC-AA-1208, and NRC-AA-1212; upper postmentals bordered by mental, infralabial I, outer postmental, median chin shield and by a single large chin scale on either side in all paratypes except NRC-AA-1206 in which bordered by two chin scales on left and single on right side. Outer postmental bordered by inner postmental, infralabials I & II in all types except NRC-AA-1206, additionally, bordered by four chin scales on left and three on right side in NRC-AA-1209, four chin scales on left and five on right side in NRC-AA-1212, outer postmental bordered by inner postmental, infralabials II in NRC-AA-1206; outer postmental separated from each other medially by three enlarged chin scales in NRC-AA-1205, NRC-AA-1207, NRC-AA-1209, NRC-AA-1210, and NRC-AA-1211. Five paratypes — NRC-AA-1208, NRC-AA-1209, NRC-AA-1210, NRC-AA-1211 and NRC-AA-1212 with original and complete tails, slightly longer than body except NRC-AA-1210 (TL/SVL 1.32, 1.29, 1.22 and 1.35 respectively), tail marginally longer than body in NRC-AA-1210 (TL/SVL 1.09), tail mostly or completely missing in NRC-AA-1205, NRC-AA-1206, and NRC-AA-1207; original tail faintly banded in all paratypes (Fig. 7A).

Figure 7. 

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.

Distribution and Natural history

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. 1). The new species seems to be diurnal, scansorial, and locally abundant. At collection sites, many individuals (n = >30) were observed active during the daytime (0900–1230 hrs) on old mossy walls and tree trunks below 2 m height in moist evergreen forest patches (Fig. 8A). Individuals of the new species were observed in large numbers inactive during the night, resting on mossy walls and cement culverts along the road inside Semmedu village and Nallathambi Resort. Sympatric geckos encountered on trees and mossy walls at the locality include Cnemaspis yercaudensis Bauer & Das, Hemidactylus parvimaculatus Deraniyagala, Hemidactylus cf. frenatus, and Hemiphyllodactylus kolliensis Agarwal, Khandekar, Giri, Ramakrishnan & Karanth.

Figure 8. 

Habitats of the new species in the vicinity of the type localities: A Cnemaspis salimalii sp. nov.; B Cnemaspis agayagangai sp. nov.; C Cnemaspis fantastica sp. nov.; D Cnemaspis pachaimalaiensis sp. nov. Photos by Akshay Khandekar.

Cnemaspis agayagangai sp. nov.

Figs 9, 10, 11, 6B, 7B, 8B; Tables 6, 7, 8

Holotype

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.

Paratypes

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.

Etymology

The specific epithet is for the type locality of the new species, the Agaya Gangai Waterfalls, and is used as a noun in apposition.

Suggested Common Name

Agaya Gangai dwarf gecko.

Diagnosis

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.

Comparison with members of C. gracilis clade

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.

Description of the holotype

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. 9A–E). SVL 31.2 mm, head short (HL/SVL 0.25), wide (HW/HL 0.67), not strongly depressed (HD/HL 0.47), distinct from neck. Loreal region marginally inflated, canthus rostralis not distinct. Snout half of head length (ES/HL 0.50), almost 2.5 times eye diameter (ES/ED 2.35); scales on snout and canthus rostralis subcircular, subequal, smooth anteriorly, becoming weakly keeled, and conical posteriorly; and much larger than those on forehead and interorbital region; scales on forehead similar to those on snout and canthus rostralis except smaller, elongated, and weakly conical; scales on interorbital region even smaller, granular and weakly keeled; scales on occipital and temporal region heterogeneous, slightly enlarged, weakly keeled, conical tubercles intermixed with smaller, weakly keeled and weakly conical granular scales (Fig. 10A). Eye small (ED/HL 0.21) with round pupil; supraciliaries short, larger anteriorly; six interorbital scale rows across narrowest point of frontal bone; 27 or 28 scale rows between left and right supraciliaries at mid-orbit (Fig. 10A, C). Ear-opening deep, oval, small (EL/HL 0.05); eye to ear distance greater than diameter of eye (EE/ED 1.41) (Fig. 10C). Rostral 2× wider (1.60 mm) than high (0.80 mm), incompletely divided dorsally by a strongly developed rostral groove and internasal scale for more than half of its height; a single enlarged supranasal on each side, slightly larger than postnasals, separated from each other by a much smaller, elongated internasal scale and still smaller scale on snout; two postnasals, upper postnasal marginally larger than lower; rostral in contact with supralabial I, nostril, supranasal, and lower postnasal on either side; nostrils oval, surrounded by two postnasals, supranasal, and rostral on either side; two rows of scales separate orbit from supralabials (Fig. 10C). Mental enlarged, subtriangular, slightly wider (1.90 mm) than high (1.52 mm); two pairs of postmentals, inner pair roughly pentagonal, much shorter (0.80 mm) than mental, separated from each other below mental by a single enlarged median chin shield; inner pair bordered by mental, infralabial I, outer postmental, enlarged median chin shield and an enlarged chin shield on either side; outer postmentals roughly rectangular, even smaller (0.62 mm) than inner pair, bordered by inner postmentals, infralabial I and II, and three enlarged chin shields on either side; three enlarged gular scales between left and right outer postmentals; all chin scales bordering postmentals flat, subcircular, smooth, and smaller than outermost postmentals; scales on rest of throat, small, subequal, and smooth (Fig. 10B). Infralabials bordered below by a row or two of slightly enlarged, much elongated scales, decreasing in size posteriorly. Ten supralabials up to angle of jaw and six at midorbital position on either side; supralabial I largest, rest of the series gradually decreasing in size posteriorly; nine infralabials up to angle of jaw on left and 10 on right, five at midorbital position on either side; infralabial I largest, gradually decreasing in size posteriorly (Fig. 10C).

Figure 9. 

Cnemaspis agayagangai sp. nov. (holotype, NRC-AA-1213): A dorsal aspect of body; B ventral aspect of body; C dorsal aspect of tail; D ventral aspect of tail; E lateral aspect of tail. Scale bars 10 mm; photos by Akshay Khandekar.

Figure 10. 

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. 11A–C). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with a fairly regularly arranged row of enlarged, strongly keeled, conical tubercles; tubercles in approximately 10 longitudinal rows at mid-body including spine-like scales at lower flank; 15 (left) and 14 (right) tubercles in paravertebral row from above forelimb insertion to the hind limb insertion (Fig. 11A, C). Ventral scales much larger than granular scales on dorsum smooth, subcircular, subimbricate, subequal from chest to vent; mid-body scale rows across belly 32; 110 scales from mental to anterior border of cloaca (Fig. 11B). Scales on base of neck similar to those on belly, marginally smaller; gular region with still smaller, subequal, smooth, flattened scales, those bordering postmentals enlarged, smooth, subcircular, and flattened (Fig. 10B). Five femoral pores on either thigh, separated by eight poreless on either side from two precloacal pores, precloacal pores separated medially by two poreless scales (Fig. 10D).

Figure 11. 

Cnemaspis agayagangai sp. nov. (holotype, NRC-AA-1213): A dorsal aspect of mid-body; B ventral aspect of mid-body; C right side lateral aspect of mid-body. Scale bars 5 mm; photos by Akshay Khandekar.

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. 9A, B). Forelimbs and hindlimbs moderately long, slender (LAL/SVL 0.15; CL/SVL 0.17); digits long, with 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: (1-3-3-4-3 right manus, 2-4-4-7-5 right pes), (1-3-3-4-3 left manus, Fig. 10E; 24575 left pes, Fig. 10F); distal lamellae series: (8-9-10-11-9 right manus, 8-10-12-12-11 right pes), (8-10-11-10-9 left manus, Fig. 10E; 8–8*-12-12-12 left pes, Fig. 10F). Relative length of digits (measurements in mm in parentheses): IV (2.8) = III (2.8) > II (2.6) > V (2.3) > I (1.8) (left manus); IV (3.6) > V (3.1) > III (3.0) > II (2.3*) > I (1.9) (left pes).

Tail original except tip (3.1 mm) which is regenerated, entire, subcylindrical, slender, marginally longer than snout-vent length (TL/SVL 1.18; Fig. 9C–E). Dorsal scales on tail base weakly keeled, granular, similar in size and shape to granular scales on mid-body dorsum, gradually becoming larger, flattened, imbricate posteriorly, intermixed with enlarged, strongly keeled, distinctly pointed, conical tubercles; enlarged tubercles on the tail forming whorls; six tubercles each on first six whorls, four in 7–11th whorls, rest of the tail with only paravertebral tubercles (Fig. 9C, E). Scales on ventral aspect of tail much larger than those on dorsal aspect, subimbricate, smooth; median series distinctly larger than rest, roughly rectangular; scales on tail base slightly smaller than those on mid-body ventrals, smooth, imbricate; a single enlarged, weakly keeled and conical postcloacal spur on each side (Fig. 9D).

Colouration in life (Fig. 6B)

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.

Variation and additional information from type series

Mensural, meristic and additional character state data for the type series is given in Tables 68 respectively. There are five adult male and three adult female specimens ranging in size from 29.3–31.8 mm (Fig. 7B). All paratypes resemble holotype except as follows: internasals absent, supranasals in strong contact with each other on snout in NRC-AA-1215, NRC-AA-1216, and NRC-AA-1220. Upper postmentals in contact with each other below mental in NRC-AA-1215, NRC-AA-1214, NRC-AA-1216, NRC-AA-1220, and NRC-AA-1221; upper postmentals bordered by mental, infralabial I, outer postmental, median chin shield, and by a single large chin scale on left side in NRC-AA-1219. Outer postmental bordered by inner postmental, infralabials I & II in all types, additionally, four chin scales on either side in NRC-AA-1216, four chin scales on left side in NRC-AA-1217, NRC-AA-1219, four chin scales on right side in NRC-AA-1218, NRC-AA-1220, and four chin scales on left and five on right side in NRC-AA-1221; outer postmental separated from each other medially by two enlarged chin scales in NRC-AA-1219. Three paratypes — NRC-AA-1217, NRC-AA-1218, and NRC-AA-1220 with original and complete tails, slightly longer than body (TL/SVL 1.16, 1.22, and 1.30 respectively); rest of the paratypes with original but incomplete tails. Original tail distinctly banded in all male paratypes and faintly banded in female paratypes, vertebral blotches not always distinct (Fig. 7B).

Table 6.

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
Table 7.

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
Table 8.

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

Distribution and Natural history

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. 1). The new species was observed to be diurnal, rupicolous, and locally abundant. At collection sites, many individuals (n = >30) were observed active during the daytime (0900–1430 hrs) on rocks and cement walls below 2 m height in moist deciduous to semi evergreen forest patches (Fig. 8B). Individuals of the new species were observed in great numbers across the elevation gradient, along the path to the Agaya Gangai waterfalls. Sympatric geckos encountered at the locality include Cnemaspis yercaudensis, Hemidactylus cf. graniticolus, Hemidactylus leschenaultii Duméril & Bibron, Hemidactylus parvimaculatus, and Hemidactylus cf. frenatus.

Cnemaspis fantastica sp. nov.

Figs 12, 13, 14, 6C, 7C, 8C; Tables 9, 10, 11

Holotype

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.

Paratypes

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.

Etymology

The specific epithet is derived from the Greek phantastikós, alluding to the spectacular colouration of the new species.

Suggested Common Name

Fantastic dwarf gecko.

Diagnosis

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.

Comparison with members of C. gracilis clade

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.

Description of the holotype

Adult male in good state of preservation except tail tip marginally bent towards right (Fig. 12A–E). SVL 31.0 mm, head short (HL/SVL 0.24), wide (HW/HL 0.66), not strongly depressed (HD/HL 0.44), distinct from neck. Loreal region marginally inflated, canthus rostralis not distinct. Snout half of head length (ES/HL 0.50), almost 2.5 times eye diameter (ES/ED 2.43); scales on snout and canthus rostralis subcircular to oval, subequal, smooth anteriorly, becoming weakly keeled, and conical posteriorly; and much larger than those on forehead and interorbital region; scales on forehead similar to those on snout and canthus rostralis except smaller, somewhat elongated, and weakly conical; scales on interorbital region even smaller, granular and weakly keeled; scales on occipital and temporal region heterogeneous, slightly enlarged, weakly keeled, conical tubercles intermixed with smaller, weakly keeled and weakly conical granular scales (Fig. 13A). Eye small (ED/HL 0.20) with round pupil; supraciliaries short, larger anteriorly; five interorbital scale rows across narrowest point of frontal bone; 24 or 25 scale rows between left and right supraciliaries at mid-orbit (Fig. 13A, C). Ear-opening deep, oval, small (EL/HL 0.06); eye to ear distance greater than diameter of eye (EE/ED 1.5) (Fig. 13C). Rostral 2× wider (1.50 mm) than high (0.53 mm), incompletely divided dorsally by a strongly developed rostral groove and internasal scale for more than half of its height; a single enlarged supranasal on each side, slightly larger than upper postnasal, separated from each other by a much smaller, elongated internasal scale and still smaller scale on snout; two postnasals, upper postnasal slightly larger than lower; rostral in contact with supralabial I, nostril, supranasal, and lower postnasal on either side; nostrils oval, surrounded by two postnasals, supranasal, and rostral on either side; two rows of scales separate orbit from supralabials (Fig. 13C). Mental enlarged, subtriangular, slightly wider (1.71 mm) than high (1.34 mm); two pairs of postmentals, inner pair roughly rectangular, much shorter (0.71 mm) than mental, in strong contact with each other below mental; inner pair bordered by mental, infralabial I, outer postmental, enlarged median chin shield and an enlarged chin shield on either side; outer postmentals roughly rectangular, even smaller (0.40 mm) than inner pair, bordered by inner postmentals, infralabial I and II, and four enlarged chin scales on left and three on right side; three enlarged gular scales between left and right outer postmentals; all chin scales bordering postmentals somewhat tubular, subcircular, smooth, and slightly smaller than outermost postmentals; scales on rest of throat, small, subequal, flattened and smooth (Fig. 13B). Infralabials bordered below by a row or two slightly enlarged, much elongated scales, decreasing in size posteriorly. Eight supralabials up to angle of jaw on left and nine on right side, and six at midorbital position on either side; supralabial I largest, rest of the series gradually decreasing in size posteriorly; eight infralabials up to angle of jaw on either side, and six at midorbital position on left and five on right side; infralabial I largest, rest of the series gradually decreasing in size posteriorly (Fig. 13C).

Figure 12. 

Cnemaspis fantastica sp. nov. (holotype, NRC-AA-1222): A dorsal aspect of body; B ventral aspect of body; C dorsal aspect of tail; D ventral aspect of tail; E lateral aspect of tail. Scale bars 10 mm; photos by Akshay Khandekar.

Figure 13. 

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. 14A–C). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with a fairly regularly arranged row of enlarged, strongly keeled, conical tubercles; tubercles in approximately 12 longitudinal rows at mid-body including spine-like scales at lower flank; 17 tubercles in paravertebral rows from above forelimb insertion to the hind limb insertion (Fig. 14A, C). Ventral scales more than twice the size than granular scales on dorsum, smooth, subcircular, subimbricate, subequal from chest to vent; mid-body scale rows across belly 30; 113 scales from mental to anterior border of cloaca (Fig. 14B). Scales on base of neck similar to those on belly, except smaller; gular region with still smaller, subequal, smooth, flattened scales, those bordering postmentals enlarged, smooth, subcircular, and somewhat tubular (Fig. 13B). Five femoral pores on left thigh and four on right, separated by eight poreless on either side from two precloacal pores, precloacal pores separated medially by a single poreless scale (Fig. 13D).

Figure 14. 

Cnemaspis fantastica sp. nov. (holotype, NRC-AA-1222): A dorsal aspect of mid-body; B ventral aspect of mid-body; C right side lateral aspect of mid-body. Scale bars 5 mm; photos by Akshay Khandekar.

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. 12A, B). Forelimbs and hindlimbs moderately long, slender (LAL/SVL 0.14; CL/SVL 0.17); digits long, with 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: (1-3-3-4-4 right manus, 1-4-4-6-5 right pes), (1-4-4-4-4 left manus, Fig. 13E; 14–474 left pes, Fig. 13F); distal lamellae series: (7-9-11-11-10 right manus, 8-10-13-12-12 right pes), (8-9-11-11-10 left manus, Fig. 13E; 81012–12–12 left pes, Fig. 13F). Relative length of digits (measurements in mm in parentheses): IV (2.6) = III (2.6) > II (2.4) > V (2.3) > I (1.7) (left manus); IV (3.5) > V (3.3) = III (3.3) > II (2.9) > I (1.8) (left pes).

Tail half original half regenerated, entire, subcylindrical, slender, marginally longer than snout-vent length (TL/SVL 1.16; Fig. 12C–E). Dorsal scales on tail base weakly keeled, granular, similar in size and shape to granular scales on mid-body dorsum, gradually becoming larger, flattened, imbricate posteriorly, intermixed with enlarged, strongly keeled, distinctly pointed, conical tubercles; enlarged tubercles on the tail forming whorls; six tubercles each on first four whorls, four in 5–8th whorls, only paravertebral tubercles in 9th and 10th whorls, rest of the tail regenerated (Fig. 12C, E). Scales on ventral aspect of original tail much larger than those on dorsal aspect, subimbricate, smooth; median series distinctly larger than rest, roughly pentagonal; scales on tail base slightly smaller than those on mid-body ventrals, smooth, imbricate; a single enlarged, smooth to weakly keeled and conical postcloacal spur on each side (Fig. 12D).

Colouration in life (Fig. 6C)

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.

Variation and additional information from type series

Mensural, meristic and additional character state data for the type series is given in Tables 911 respectively. There are four adult male and three adult female specimens ranging in size from 28.4–32.5 mm (Fig. 7C). All paratypes resemble holotype except as follows: internasal absent, supranasals in strong contact with each other on snout in NRC-AA-1223.

Table 9.

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. 7C).

Table 10.

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

Distribution and Natural history

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. 1). Like most of the other members of its clade, the new species seems to be diurnal, rupicolous, and locally abundant. At collection sites, many individuals (n = >40) were observed active during the daytime (1100–1430 hrs) on rocks, road side rocky cuttings, and building walls below 2 m height, and under cement culverts in moist deciduous to semi-evergreen forest patches (Fig. 8C). Sympatric geckos encountered at the locality include Cyrtodactylus (Geckoella) sp., Hemidactylus cf. graniticolus, Hemidactylus parvimaculatus, Hemidactylus cf. frenatus, and Hemiphyllodactylus kolliensis.

Table 11.

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

Cnemaspis pachaimalaiensis sp. nov.

Figs 15, 16, 17; 6D, 8D, 18A; Tables 12, 13, 14

Holotype

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.

Paratypes

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.

Etymology

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.

Suggested Common Name

Pachaimalai dwarf gecko.

Diagnosis

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.

Comparison with members of C. gracilis clade

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.

Description of the holotype

Adult male in good state of preservation except tail slightly bent towards left (Fig. 15A–E). SVL 28.8 mm, head short (HL/SVL 0.24), wide (HW/HL 0.68), not strongly depressed (HD/HL 0.45), distinct from neck. Loreal region marginally inflated, canthus rostralis not distinct. Snout half of head length (ES/HL 0.50), almost 2.5 times eye diameter (ES/ED 2.33); scales on snout and canthus rostralis subcircular, subequal, weakly keeled, somewhat conical, and much larger than those on forehead and interorbital region; scales on forehead similar to those on snout and canthus rostralis except smaller and elongated, and weakly conical; scales on interorbital region even smaller, granular; scales on occipital and temporal region heterogeneous, enlarged, keeled, conical tubercles intermixed with much smaller, weakly keeled and weakly conical granular scales (Fig. 16A). Eye small (ED/HL 0.21) with round pupil; supraciliaries short, larger anteriorly; five interorbital scale rows across narrowest point of frontal bone; 28–30 scale rows between left and right supraciliaries at mid-orbit (Fig. 16A, C). Ear-opening deep, oval, small (EL/HL 0.05); eye to ear distance greater than diameter of eye (EE/ED 1.60) (Fig. 16C). Rostral more than twice as wide (1.40 mm) as high (0.55 mm), incompletely divided dorsally by a strongly developed rostral groove for more than half of its height; a single enlarged supranasal on each side, slightly larger than upper postnasal, separated from each other by a much smaller, elongated internasal scale and still smaller scale on snout; two postnasals, upper postnasal slightly larger than lower; rostral in contact with supralabial I, nostril, supranasal, and weakly in contact with lower postnasal on either side; nostrils oval, surrounded by two postnasals, supranasal, and rostral on either side; one or two rows of scales separate orbit from supralabials (Fig. 16C). Mental enlarged, subtriangular, marginally wider (1.64 mm) than high (1.30 mm); two pairs of postmentals, inner pair roughly rectangular, much shorter (0.63 mm) than mental, separated from each other below mental by a single enlarged median chin shield; inner pair bordered by mental, infralabial I, outer postmental, enlarged median chin shield and an enlarged chin shield on either side; outer postmentals roughly rectangular, even smaller (0.52 mm) than inner pair, bordered by inner postmentals, infralabial I and II, and four enlarged chin shields on either side; three enlarged gular scales between left and right outer postmentals; all chin scales bordering postmentals flat, subcircular, smooth, and slightly smaller than outermost postmentals; scales on rest of throat granular, small, smooth (Fig. 16B). Infralabials bordered below by a row or two of slightly enlarged, much elongated scales, decreasing in size posteriorly. Eight supralabials up to angle of jaw and six at midorbital position on either side; supralabial I largest, rest of the series gradually decreasing in size posteriorly; seven infralabials up to angle of jaw, five at midorbital position on either side; infralabial I largest, rest of the series gradually decreasing in size posteriorly (Fig. 16C).

Figure 15. 

Cnemaspis pachaimalaiensis sp. nov. (holotype, NRC-AA-1230): A dorsal aspect of body; B ventral aspect of body; C dorsal aspect of tail; D ventral aspect of tail; E lateral aspect of tail. Scale bars 10 mm; photos by Akshay Khandekar.

Figure 16. 

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. 17A–C). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with a fairly regularly arranged row of enlarged, strongly keeled, conical tubercles; tubercles in approximately 11 longitudinal rows at mid-body including short spine-like scales at lower flank; 14 (left) and 16 (right) tubercles in paravertebral row from above forelimb insertion to the hind limb insertion (Fig. 17A, C). Ventral scales much larger than granular scales on dorsum smooth, subcircular, subimbricate, subequal from chest to vent; mid-body scale rows across belly 31; 115 scales from mental to anterior border of cloaca (Fig. 17B). Scales on base of neck similar to those on belly, marginally smaller; gular region with much smaller, smooth, granular scales, those bordering postmentals enlarged, smooth, subcircular, and flattened (Fig. 16B). Five femoral pores on left thigh and four on right, separated by 10 poreless on either side from continuous series of three precloacal pores (Fig. 16D).

Figure 17. 

Cnemaspis pachaimalaiensis sp. nov. (holotype, NRC-AA-1230): A dorsal aspect of mid-body; B ventral aspect of mid-body; C right side lateral aspect of mid-body. Scale bars 5 mm; photos by Akshay Khandekar.

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. 15A, B). Forelimbs and hindlimbs moderately long, slender (LAL/SVL 0.14; CL/SVL 0.17); digits long, with strong, recurved claw, distinctly inflected, distal portions laterally compressed conspicuously. Digits with unpaired lamellae except for a few basal lamellae which are paired, separated into a basal and narrower distal series by single enlarged lamella at inflection; basal lamellae series: (1-3-3-4-3 right manus, 1-4-5-6-5 right pes), (1-3-3-4-3 left manus, Fig. 16E; 14675 left pes, Fig. 16F); distal lamellae series: (9-11-12-11-10 right manus, 8-10-13-12-12 right pes), (8-9-11-11-11 left manus, Fig. 16E; 811131213 left pes, Fig. 16F). Relative length of digits (measurements in mm in parentheses): IV (2.5) > III (2.3) > V (2.0) = II (2.0) > I (1.5) (left manus); IV (3.1) > V (3.0) > III (2.9) > II (2.6) > I (1.9) (left pes).

Tail original, entire, subcylindrical, slender, marginally longer than snout-vent length (TL/SVL 1.11; Fig. 15C–E). Dorsal scales on tail base weakly keeled, granular, similar in size and shape to granular scales on mid-body dorsum, gradually becoming larger, flattened, imbricate posteriorly, intermixed with enlarged, strongly keeled, distinctly pointed, conical tubercles; enlarged tubercles on the tail forming whorls; six tubercles each on first nine whorls, four in 10–13th whorls, rest of the tail with only paravertebral tubercles (Fig. 15A, C). Scales on ventral aspect of tail much larger than those on dorsal aspect, subimbricate, smooth; median series distinctly larger than rest, roughly pentagonal; scales on tail base slightly smaller than those on mid-body ventrals, smooth, imbricate; a single enlarged, conical, and smooth postcloacal spur on each side (Fig. 15B).

Colouration in life (Fig. 6D)

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.

Variation and additional information from type series

Mensural, meristic and additional character state data for the type series is given in Tables 1214 respectively. There are six adult male and a single adult female specimens ranging in size from 28.6–33.6 mm (Fig. 18A). All paratypes resemble holotype except as follows: supranasals in contact with each other behind internasal in NRC-AA-1233 and NRC-AA-1237; upper postmentals in contact with each other below mental in NRC-AA-1231, NRC-AA-1233, NRC-AA-1234, and NRC-AA-1237; upper postmentals separated from each other below mental by two median enlarged chin shields in NRC-AA-1236; upper postmentals bordered by mental, infralabial I, outer postmental, and by a single large chin scale on either side in NRC-AA-1231, NRC-AA-1233, NRC-AA-1236; upper postmentals bordered by mental, infralabial I, outer postmental, median chin shield, and by a single large chin scale on either side in NRC-AA-1234, NRC-AA-1237. Outer postmental bordered by inner postmental, infralabials I & II in all types, additionally, five chin scales on either side in NRC-AA-1231, five chin scales on right in NRC-AA-1233 and NRC-AA-1236, three scales on right in NRC-AA-1232 and NRC-AA-1237; outer postmental separated from each other medially by two enlarged chin scales in NRC-AA-1231, NRC-AA-1233. Three paratypes — NRC-AA-1232, NRC-AA-1233, and NRC-AA-1235 with original and complete tails, slightly longer than body (TL/SVL 1.11, 1.27, and 1.30 respectively); tail entire but partially regenerated in NRC-AA-1234, NRC-AA-1236, and NRC-AA-1237, marginally to slightly longer than body (TL/SVL 1.08, 1.12, and 1.33 respectively); original tail entirely lost, small regenerated portion present in NRC-AA-1231. Ocelli on body between limb insertions are highly variable in all paratypes; original tail banded in all paratypes; regenerated tail orangish in life and yellowish-grey in preservative. (Fig. 18A).

Table 12.

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
Figure 18. 

Paratype series: A Cnemaspis pachaimalaiensis sp. nov., from left to right, NRC-AA-1231–NRC-AA-1237; B Cnemaspis rudhira sp. nov., from left to right, NRC-AA-1239–NRC-AA-1247. Scale bar 10 mm; photos by Akshay Khandekar.

Distribution and Natural history

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. 1). Like most other members of its clade, the new species is diurnal, rupicolous, and fairly abundant locally. At each collection site, many individuals (n = >20) were observed active during the daytime (0900–1630 hrs) on rocks, road side rocky cuttings, and building walls below 2 m height, and under cement culverts in dry deciduous to semi-evergreen forest patches (Fig. 8D). Sympatric geckos encountered at the locality include Cyrtodactylus (Geckoella) sp., Hemidactylus kolliensis Agarwal, Bauer, Giri & Khandekar, Hemidactylus leschenaultii, Hemidactylus whitakeri Mirza, Gowande, Patil, Ambekar & Patel, Hemidactylus parvimaculatus, Hemidactylus cf. frenatus, and Hemiphyllodactylus sp.

Table 13.

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
Table 14.

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 rudhira sp. nov.

Figs 18B–18, 19, 20, 21, 22; Tables 15, 16, 17

Cnemaspis cf. gracilis Khandekar et al. 2019

Holotype

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.

Paratypes

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.

Etymology

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.

Suggested Common Name

Scarlet dwarf gecko.

Diagnosis

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.

Comparison with members of C. gracilis clade

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.

Description of the holotype

Adult male in good state of preservation except tail tip slightly bend towards left, longitudinal skin fold on vertebral region between limb insertions (Fig. 19A–E). SVL 32.3 mm, head short (HL/SVL 0.24), wide (HW/HL 0.66), not strongly depressed (HD/HL 0.47), distinct from neck. Loreal region marginally inflated, canthus rostralis not distinct. Snout half of head length (ES/HL 0.52), marginally more than 2.5 times eye diameter (ES/ED 2.6); scales on snout and canthus rostralis subcircular, subequal, and weakly keeled; much larger than those on forehead and interorbital region; scales on forehead similar to those on snout and canthus rostralis except smaller, elongated, and weakly conical; scales on interorbital region even smaller, granular and weakly keeled; scales on occipital and temporal region heterogeneous, slightly enlarged, weakly keeled, conical tubercles intermixed with smaller, weakly keeled and weakly conical granular scales (Fig. 20A). Eye small (ED/HL 0.20) with round pupil; supraciliaries short, larger anteriorly; six interorbital scale rows across narrowest point of frontal bone; 25 or 26 scale rows between left and right supraciliaries at mid-orbit (Fig. 20A, C). Ear-opening deep, oval, small (EL/HL 0.05); eye to ear distance greater than diameter of eye (EE/ED 1.50) (Fig. 20C). Rostral more than two times wider (1.56 mm) than high (0.72 mm), incompletely divided dorsally by a strongly developed rostral groove and internasal scale for more than half of its height; a single enlarged supranasal on each side, much larger than postnasals, separated from each other by a much smaller, elongated internasal scale and still smaller scale on snout; two postnasals, upper postnasal marginally larger than lower; rostral in contact with supralabial I, nostril, internasal, supranasal, and lower postnasal on either side; nostrils oval, surrounded by two postnasals, supranasal, and rostral on either side; two rows of scales separate orbit from supralabials (Fig. 20C). Mental enlarged, subtriangular, slightly wider (1.97 mm) than high (1.50 mm); two pairs of postmentals, inner pair roughly rectangular, much shorter (0.84 mm) than mental, in strong contact with each other below mental; inner pair bordered by mental, infralabial I, outer postmental, enlarged median chin shield on either side and an enlarged chin shield on left side; outer postmentals roughly rectangular, even smaller (0.63 mm) than inner pair, bordered by inner postmentals, infralabial I and II, and three enlarged chin shields on either side and median chin shield on left side; three enlarged gular scales between left and right outer postmentals; all chin scales bordering postmentals flat, subcircular, smooth, and smaller than outermost postmentals; scales on rest of throat, even smaller, flattened, subequal, and smooth (Fig. 20B). Infralabials bordered below by a row or two of slightly enlarged, much elongated scales, decreasing in size posteriorly. Ten supralabials up to angle of jaw on left, 11 on right side, and six at midorbital position on each side; supralabial I largest, gradually decreasing in size posteriorly; eight infralabials up to angle of jaw on left and nine on right, five at midorbital position on either side; infralabial I largest, gradually decreasing in size posteriorly (Fig. 20C).

Figure 19. 

Cnemaspis rudhira sp. nov. (holotype, NRC-AA-1238): A dorsal aspect of body; B ventral aspect of body; C dorsal aspect of tail; D ventral aspect of tail; E lateral aspect of tail. Scale bars 10 mm; photos by Akshay Khandekar.

Figure 20. 

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. 21A–C). Dorsal pholidosis heterogeneous; weakly keeled granular scales intermixed with a fairly regularly arranged row of enlarged, strongly keeled, conical tubercles; tubercles in approximately 12 longitudinal rows at mid-body including spine-like scales at lower flank; 14 (left) and 15 (right) tubercles in paravertebral row from above forelimb insertion to the hind limb insertion (Fig. 21A, C). Ventral scales much larger than granular scales on dorsum smooth, subcircular, subimbricate, subequal from chest to vent; mid-body scale rows across belly 31; 115 scales from mental to anterior border of cloaca (Fig. 21B). Scales on base of neck similar to those on belly, marginally smaller; gular region with still smaller, subequal, smooth, flattened scales, those bordering postmentals enlarged, smooth, subcircular, and flattened (Fig. 20B). Four femoral pores on either thigh, separated by nine poreless on either side from four precloacal pores, precloacal pores separated medially by a single poreless scale (Fig. 20D).

Figure 21. 

Cnemaspis rudhira sp. nov. (holotype, NRC-AA-1238): A dorsal aspect of mid-body; B ventral aspect of mid-body; C right side lateral aspect of mid-body. Scale bars 5 mm; photos by Akshay Khandekar.

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. 19A, B). Forelimbs and hindlimbs moderately long, slender (LAL/SVL 0.15; CL/SVL 0.18); digits long, with strong, recurved claw, distinctly inflected, distal portions laterally compressed conspicuously. Digits with unpaired lamellae except basal one or two paired on some digits, separated into a basal and narrower distal series by single enlarged lamella at inflection; basal lamellae series: (1-4-4-4-4 right manus, 1-5-5-8-6 right pes), (2-3-4-4-3 left manus, Fig. 20E; 14586 left pes, Fig. 20F); distal lamellae series: (9-10-13-12-10 right manus, 9-11-13-13-11 right pes), (9-10-12-12-10 left manus, Fig. 20E; 9111413–13 left pes, Fig. 20F). Relative length of digits (measurements in mm in parentheses): IV (2.6) > III (2.4) > II (2.3) = V (2.3) > I (1.9) (left manus); IV (3.9) > V (3.3) > III (3.2) > II (2.9) > I (1.8) (left pes).

Tail original except tip (5.1 mm) which is regenerated, entire, subcylindrical, slender, slightly longer than snout-vent length (TL/SVL 1.30; Fig. 19C–E). Dorsal scales on tail base weakly keeled, granular, similar in size and shape to granular scales on mid-body dorsum, gradually becoming larger, flattened, imbricate posteriorly, intermixed with enlarged, strongly keeled, distinctly pointed, conical tubercles; enlarged tubercles on the tail forming whorls; six tubercles each on first eight whorls, four in 9–12th whorls, rest of the tail with only paravertebral tubercles except original and regenerated portion of the tail lacking enlarged tubercles (Fig. 19C, E). Scales on ventral aspect of tail much larger than those on dorsal aspect, subimbricate, smooth; median series distinctly larger than rest, roughly rectangular; scales on tail base slightly larger than those on mid-body ventrals, smooth, imbricate; a single enlarged, weakly keeled and conical postcloacal spur on each side (Fig. 19D).

Colouration in life (Fig. 22)

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.

Figure 22. 

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.

Variation and additional information from type series

Mensural, meristic and additional character state data for the type series is given in Tables 1517 respectively. There are seven adult male and two adult female specimens ranging in size from 27.9–33.8 mm (Fig. 18B). All paratypes resemble holotype except as follows: supranasals in strong contact with each other behind internasal on snout in NRC-AA-1243. Upper postmentals separated from each other below mental by enlarged median chin shield in NRC-AA-1246; upper postmentals bordered by mental, infralabial I, outer postmental, median chin shield, and additionally by a single large chin scale on either side in NRC-AA-1239, NRC-AA-1241, NRC-AA-1245; upper postmental bordered by both infralabial I & II on left and a single large chin scale on either side in NRC-AA-1240, and NRC-AA-1246. Outer postmental bordered by inner postmental, infralabials I & II and additionally, four chin scales on left and three on right side in NRC-AA-1239, NRC-AA-1241, NRC-AA-1244, NRC-AA-1247, four chin scales on either side in NRC-AA-1243; outer postmental bordered by inner postmental, infralabials I (on left), and four chin scales on either side in NRC-AA-1240, NRC-AA-1246; outer postmental separated from each other medially by two enlarged chin scales in NRC-AA-1242, NRC-AA-1243, and NRC-AA-1247. Four paratypes — NRC-AA-1244, NRC-AA-1245, NRC-AA-1246, and NRC-AA-1247 with original and complete tails, slightly longer than body (TL/SVL 1.32, 1.29, 1.29, and 1.19 respectively); NRC-AA-1243 with complete but fully regenerated tail, almost equal to the body (TL/SVL 1.03); NRC-AA-1240 and NRC-AA-1241 with partial but original tail; tail almost entirely lost in NRC-AA-1239 and NRC-AA-1242. Original tail distinctly banded only in two male paratypes — NRC-AA-1240 and NRC-AA-1246 (Fig. 18B).

Table 15.

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

Distribution and Natural history

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. 1). The new species was observed to be diurnal, scansorial, and locally highly abundant. At each collection site, many individuals (n = >30) were observed active during the daytime (0900–1430 hrs) on rocks, cement walls, trees, inside cement culverts etc. all below 2–3 m height in moist deciduous to evergreen forest patches (Fig. 22). Individuals of the new species were observed both daytime and at night in large numbers across the elevation gradient, along the ghat road leading to Yercaud town. Sympatric geckos encountered at the locality include Cnemaspis yercaudensis, Cnemaspis thackerayi, Cyrtodactylus (Geckoella) sp. Hemidactylus cf. graniticolus, Hemidactylus leschenaultii, Hemidactylus parvimaculatus, Hemidactylus cf. frenatus, Hemidactylus whitakeri, and Hemiphyllodactylus aurantiacus (Beddome).

Table 16.

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
Table 17.

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

Discussion

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 bangara clade to 11. Seven divergent lineages of the bangara 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 (Bernstein et al. 2021), numerous radiations in Madagascar (Vences et al. 2009; Uetz et al. 2022), Cnemaspis sensu stricto in Southeast Asia (Grismer et al. 2014) and South Asian Cnemaspis in Sri Lanka (Karunarathna et al. 2023). Additionally, 12 microendemic karst species of Cyrtodactylus Gray are distributed within 90 km of one another in the Salween Basin of Myanmar (Grismer et al. 2018). There is a need for fine scale surveys in unexplored and previously surveyed areas to fully understand patterns of diversity and distribution in this incredibly diverse clade of geckos.

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 (Agarwal et al. 2019b). The timing of initial diversification within the bangara clade as estimated by Agarwal et al. (2020b) was in the middle Miocene, the start of strong seasonality and aridification in peninsular India which is likely to have caused the fragmentation of once relatively contiguous tropical forest (Morley 2007; Pound et al. 2012), likely disproportionately impacting the most cool-adapted, forest-dwelling taxa (as for example high elevation species). These massifs appear to have acted as long term refugia for members of this clade, with complex patterns of dispersal between them and other parts of the range of the bangara clade.

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., Grismer et al. 2013), a finer 3.7 % cut-off was suggested for the Sri Lankan clades of South Asian Cnemaspis based on the lowest divergence between named, morphologically diagnosable species (Agarwal et al. 2017). Additional morphologically distinct Cnemaspis species have been described since show < 5% genetic divergence from their respective sister species (e.g., Batuwita et al. 2019; Karunarathna et al. 2019). Lending support to the specific status of Cnemaspis fantastica sp. nov. is the fact that strong morphological characters separate it from its sister taxon C. shevaroyensis — the presence vs absence of spinelike tubercles and 28–32 scales across the belly versus 21–24 (integration by congruence, sensu Padial et al. 2010). In addition, none of the members of the bangara complex are known from multiple massifs in this landscape, and the low hills separating massifs have additional undescribed species (Khandekar, Thackeray and Agarwal, unpubl. data).

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 (Darwin 1859). Chemical signalling is known in South Asian Cnemaspis, and visual traits have been established in intraspecific communication for at least a few species, though those studies only evaluated gular or gular and brille colouration (Kabir et al. 2019, 2020). Preliminary observations suggest members of the bangara clade are often found in pairs consisting of a single male and female, but quantitative data is needed to ascertain if this is true and what the role of sexual selection, if any, is in the formation of pairs and how this may drive diversification. It is striking that there are two pairs of cryptic, non-sister taxa, C. fantastica sp. nov. + C. agayagangai sp. nov. and C. rudhira sp. nov. + C. gracilis, each pair of which overlaps in practically all aspects of their morphology, differing mainly in male colouration.

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 (Ganesh and Arumugam 2016; Khandekar et al. 2019); Hemidactylus kolliensis is found in both Kollimalai and Pachaimalai while divergent lineages of the Hemidactylus graniticolus complex are found in Yercaud and Kollimalai (Agarwal et al. 2019a); the genus Hemiphyllodactylus has a different endemic species each in Yercaud, Kollimalai and Pachaimalai (Agarwal et al. 2019b; Agarwal, Thackeray and Khandekar unpubl. data). These are all our recent records from the region, as previous records apart from original descriptions of taxa are scanty and many did not include specimens. For example, records of an undescribed Cnemaspis with three precloacal pores and three or four femoral pores from <1350 m the Shevaroys and Kolli Hills, in addition to C. yercaudensis Bauer & Das, the only non bangara clade Cnemaspis known from these hills, may represent C. salimalii sp. nov. from the Kolli Hills and C. fantastica sp. nov. from the Shevaroys, but this cannot be ascertained as no photographs were provided nor specimens collected (Ganesh and Arumugam 2016).

Conservation status

These new species join a large proportion of gekkonid species known only from their type localities (23 %, Meiri et al. 2018; though that analysis considered the type locality to have a maximum latitudinal and longitudinal extent of < 10 km). We surveyed the vicinity of the type localities of all seven species found in the Shevaroy landscape and found that C. rudhira sp. nov. is widely distributed across the elevational gradient in Yercaud, C. thackerayi and C. salimalii sp. nov. are widely distributed on the high elevation plateaus at their respective type localities, while the other three species are extremely local in distribution, each of which is found in only one or a few closely spaced localities on their respective massifs. This implies that each of these species would qualify as critically endangered under IUCN’s criterion B1 of extent of occurrence < 100 km2, meeting subcriterion a. (known from a single location) and b. (iii) (potential for a decline in habitat quality) (IUCN 2012). Cnemaspis shevaroyensis and C. thackerayi are currently listed as critically endangered while C. yercaudensis is endangered (IUCN 2022).

Acknowledgements

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.

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Appendix 1

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.