Abstract
We present a preliminary ND2 phylogeny of South Asian Cnemaspis, recovering a number of deeply divergent clades within Indian Cnemaspis, endemic to the southern and northern Western Ghats besides the Mysore Plateau and hills of Tamil Nadu. There are a number of unnamed lineages that are >5% divergent on ND2 across the phylogeny, including three from the gracilis clade on an elevation gradient (800–1400 m asl.) around Yercaud in the Shevaroy massif, Salem district, Tamil Nadu. We describe two of these as new species— Cnemaspis shevaroyensis sp. nov. and Cnemaspis thackerayi sp. nov. are both allied to Cnemaspis gracilis and can be diagnosed from all other Indian Cnemaspis by the absence of spine-like scales on flank, heterogeneous dorsal pholidosis, presence of femoral and precloacal pores, tail with enlarged, strongly keeled, conical tubercles forming whorls, a median row of enlarged and smooth sub-caudals. They differ from C. gracilis and each other in body size, the number of tubercles around midbody, the number of tubercles in paravertebral rows, the number of femoral and precloacal pores, the number of poreless scales in-between precloacal pores and between femoral and precloacal pores, and subtle colour pattern differences; besides uncorrected mitochondrial sequence divergence (7.9–16.6 %). We also provide a description of Cnemaspis yercaudensis from its type locality and an additional locality. The discovery of two endemic species and a third unnamed divergent lineage from an isolated massif in peninsular India outside the Western Ghats indicate that many other such understudied hill ranges may harbour high endemic biodiversity.
References
Agarwal, I. (2016) Two new species of ground-dwelling Cyrtodactylus (Geckoella) from the Mysore Plateau, south India. Zootaxa, 4193 (2), 228–244.
https://doi.org/10.11646/zootaxa.4193.2.2
Agarwal, I., Bauer, A.M., Jackman, T.R. & Karanth, K.P. (2014) Insights into Himalayan biogeography from geckos: a molecular phylogeny of Cyrtodactylus (Squamata: Gekkonidae). Molecular Phylogenetics and Evolution, 80, 145–155.
https://doi.org/10.1016/j.ympev.2014.07.018
Agarwal, I., Biswas, S., Bauer, A.M., Greenbaum, E., Jackman, T.R., Silva, A.D. & Batuwita, S. (2017) Cryptic species, taxonomic inflation, or a bit of both? New species phenomenon in Sri Lanka as suggested by a phylogeny of dwarf geckos (Reptilia, Squamata, Gekkonidae, Cnemaspis). Systematics and Biodiversity, 15 (5), 1–13.
https://doi.org/10.1080/14772000.2017.1282553
Agarwal, I., Khandekar, A., Giri, V.B., Ramakrishnan, U. & Karanth, P.K. (2019) The hills are alive with geckos! A radiation of a dozen species on sky islands across peninsular India (Squamata: Gekkonidae, Hemiphyllodactylus). Organisms Diversity & Evolution. 1–21.
https://doi.org/10.1007/s13127-019-00392-5
Bauer, A.M. (2002) Two new species of Cnemaspis (Reptilia: Squamata: Gekkonidae) from Gund, Uttara Kannada, India. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 99, 155–167.
Bauer, A.M., de Silva, A., Greenbaum, E. & Jackman, T.R. (2007) A new species of day gecko from high elevation in Sri Lanka, with a preliminary phylogeny of Sri Lankan Cnemaspis (Reptilia: Squamata: Gekkonidae). Mitteilungen aus dem Museum für Naturkunde in Berlin. Zoologische Reihe, 83 (Supplement), 22–32.
https://doi.org/10.1002/mmnz.200600022
Bauer, A.M., Jackman, T.R., Greenbaum, E., Giri, V. & De Silva, A. (2010) South Asia supports a major endemic radiation of Hemidactylus geckos. Molecular Phylogenetics and Evolution, 57, 343–352.
https://doi.org/10.1016/j.ympev.2010.06.014
Beddome, R.H. (1870) Descriptions of some new lizards from the Madras Presidency. Madras Monthly journal of Medical Science, 1, 30–35.
Cyriac, V.P. & Umesh, P.K. (2013) Current status of Cnemaspis littoralis (Jerdon, 1853) (Sauria: Gekkonidae) with designation of a neotype. Taprobanica: The Journal of Asian Biodiversity, 5, 36–43.
https://doi.org/10.4038/tapro.v5i1.5660
Cyriac, V.P. & Umesh, P.K. (2014) Description of a new ground-dwelling Cnemaspis Strauch, 1887 (Squamata: Gekkonidae), from Kerala, allied to C. wynadensis (Beddome, 1870). Russian Journal of Herpetology, 21 (3), 187–194.
Cyriac, V.P., Johny, A., Umesh, P.K. & Palot, M.J. (2018) Description of two new species of Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from the Western Ghats of Kerala, India, Zootaxa, 4459 (1), 85–100.
https://doi.org/10.11646/zootaxa.4459.1.3
Das, I. (2005) Revision of the genus Cnemaspis Strauch, 1887 (Sauria: Gekkonidae), from the Mentawai and adjacent archipelagos off western Sumatra, Indonesia, with the description of four new species. Journal of Herpetology, 39, 233–247.
https://doi.org/10.1670/61-02A
Das, I. & Bauer, A.M. (2000) Two new species of Cnemaspis (Sauria: Gekkonidae) from Tamil Nadu, southern India. Russian Journal of Herpetology, 7 (1), 17–28.
Das, I. & Sengupta, S. (2000) A new species of Cnemaspis (Sauria: Gekkonidae) from Assam, northeastern India. Journal of South Asian Natural History, 5 (1), 17–23.
Gamble, T., Greenbaum, E., Jackman, T.R., Russell, A.P. & Bauer, A.M. (2012) Repeated origin and loss of adhesive toepads in geckos. PLoS ONE, 7 (6), e39429.
https://doi.org/10.1371/journal.pone.0039429
Gamble, T., Greenbaum, E., Jackman, T.R. & Bauer, A.M. (2015) Into the light: diurnality has evolved multiple times in geckos. Biological Journal of the Linnean Society, 115 (4), 896–910.
https://doi.org/10.1111/bij.12536
Ganesh, S.R., Sreekar, R., Pal, S.P., Ramchandra, G., Srinivasulu, C. & Srinivasulu, B. (2011) Discovery and first description of male Cnemaspis heteropholis Bauer, 2002 (Reptilia: Gekkonidae) from Agumbe, central Western Ghats, India. Journal of Threatened Taxa, 3 (8), 2023–2027.
Ganesh, S.R. & Arumugam, M. (2016) Species richness of montane herpetofauna of southern Eastern Ghats, India: a historical resume and a descriptive checklist. Russian Journal of Herpetology, 23 (1), 7–24.
Ganesh, S.R., Bhupathy, S., David, P., Sathishkumar, N. & Srinivas, G. (2014) Snake fauna of High Wavy Mountains, Western Ghats, India: species richness, status, and distribution pattern. Russian Journal of Herpetology, 21 (1), 53-64.
Giri, V.B., Agarwal, I. & Bauer, A.M. (2009a) Designation of a neotype for Cnemaspis mysoriensis (Jerdon 1853) (Sauria: Gekkonidae), with a redescription and notes on its distribution and habitat. Russian Journal of Herpetology, 16, 256–264.
Giri, V.B., Bauer, A.M. & Gaikwad, K.S. (2009b) A new ground-dwelling species of Cnemaspis Strauch (Squamata: Gekkonidae) from the northern Western Ghats, Maharashtra, India. Zootaxa, 2164, 49–60.
https://doi.org/10.5281/zenodo.189040
Grismer, L.L., Sumontha, M., Cota, M., Grismer, J.L., Wood Jr., P.L., Pauwels, O.S.G. & Kunya, K. (2010) A revision and redescription of the rock gecko Cnemaspis siamensis (Taylor 1925) (Squamata: Gekkonidae) from Peninsular Thailand with descriptions of seven new species. Zootaxa, 2576, 1–55.
Grismer, L.L., Wood Jr., P.L., Anuar, S., Riyanto, A., Ahmad, N., Muin, M.A., Sumontha, M., Grismer, J.L., Onn, C.K., Quah, E.S. & Pauwels, O.S.G. (2014) Systematics and natural history of Southeast asian rock geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia. Zootaxa, 3880 (1), 1–147.
https://doi.org/10.11646/zootaxa.3880.1.1Jerdon, T.C. (1853) Catalogue of reptiles inhabiting the peninsula of India. Journal of the Asiatic Society of Bengal, 22, 462–479.
Kadavul, K. & Parthasarathy, N. (2004) Plant biodiversity and conservation of tropical semi-evergreen forest in the Shervarayan hills of Eastern Ghats, India. Biodiversity & Conservation, 8, 419-437.
https://doi.org/10.1023/A:1008899824399
Khandekar, A. (2019) A new species of rock-dwelling Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from Tamil Nadu, southern India. Zootaxa, 4571 (3): 383–397.
http://doi.org/10.11646/zootaxa.4571.3.6
Macey, J.R., Larson, A., Ananjeva, N.B., Fang, Z. & Papenfuss, T.J. (1997) Two novel gene orders and the role of light-strand replication in rearrangement of the vertebrate mitochondrial genome. Molecular Biology and Evolution, 14, 91–104.
https://doi.org/10.1093/oxfordjournals.molbev.a025706
Manamendra-Arachchi, K., Batuwita, S. & Pethiyagoda, R. (2007) A taxonomic revision of the Sri Lankan day-geckos (Reptilia: Gekkonidae: Cnemaspis), with description of new species from Sri Lanka and southern India. Zeylanica, 7 (1), 9–122.
Mukherjee, D., Bhupathy, S. & Nixon, A.M.A. (2005) A new species of day gecko (Squamata, Gekkonidae, Cnemaspis) from the Anaikatti Hills, Western Ghats, Tamil Nadu, India. Current Science, 89, 1326–1328.
http://www.jstor.org/stable/24110836
Nayaka, S., Upreti, D., Ponmurugan, P. & Ayyappadasan, G. (2013). Two new species of saxicolous Pyxine with yellow medulla from southern India. The Lichenologist, 45 (1), 3–8.
https://doi.org/10.1017/S0024282912000618
Raheem, D., Taylor, H., Ablett, J., Preece, R., Aravind, N. & Naggs, F. (2014). A systematic revision of the land snails of the Western Ghats of India. Tropical Natural History, 4, 1–294.
https://tci-thaijo.org/index.php/tnh/article/view/103091
Ramachandran, A., Radhapriya, P., Jayakumar, S., Dhanya, P. & Geetha, R. (2016) Critical analysis of forest degradation in the southern Eastern Ghats of India: comparison of satellite imagery and soil quality index. PLoS ONE, 11 (1), e0147541.
https://doi.org/10.1371/journal.pone.0147541
Sayyed, A., Pyron, R.A. & Dahanukar, N. (2016) Cnemaspis flaviventralis, a new species of gecko (Squamata: Gekkonidae) from the Western Ghats of Maharashtra, India. Journal of Threatened Taxa, 8, 9619–9629.
https://doi.org/10.11609/jott.2599.8.14.9619-9629
Sayyed, A., Pyron, R.A. & Dileepkumar, R. (2018) Four new species of the genus Cnemaspis Strauch, (Sauria: Gekkonidae) from the northern Western Ghats, India. Amphibian & Reptile Conservation, 12 (2), 1–29.
Silvestro, D. & Michalak, I. (2012) raxmlGUI: A graphical front-end for RAxML. Organisms, Diversity and Evolution, 12, 335–337.
https://doi.org/10.1007/s13127-011-0056-0
Smith, M.A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Vol. II. Sauria. Taylor and Francis, London, xiv + 440 pp., 2 folding maps, 1 pl.
Srinivasulu, C., Kumar, G.C. & Srinivasulu, B. (2015) A new species of Cnemaspis (Sauria: Gekkonidae) from Northern Karnataka, India. Zootaxa, 3947 (1), 85–98.
https://doi.org/10.11646/zootaxa.3947.1.5
Stamatakis, A. (2006) RAxML-VI-HPC: Maximum likelihoodbased phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
https://doi.org/10.1093/bioinformatics/btl446
Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5: Molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution, 28, 2731–2739.
https://doi.org/10.1093/molbev/msr121
Uetz, P. (2018) The Reptile Database. Retrieved from: http://reptile-database.reptarium.cz (accessed-- 2 November 2018)
Wood, P.L. Jr., Heinicke, M.P., Jackman, T.R. & Bauer, A.M. (2012) Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of diversification. Molecular Phylogenetics & Evolution, 65, 992–1003.
https://doi.org/10.1016/j.ympev.2012.08.025