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Type: Article
Published: 2018-09-19
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A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species

Centre for Ecological Sciences, Indian Institute of Science, Bangalore, 560012, India. Bombay Natural History Society, Hornbill House, Dr Salim Ali Chowk, Mumbai 400001, India.
Centre for Ecological Sciences, Indian Institute of Science, Bangalore, 560012, India.
Centre for Ecological Sciences, Indian Institute of Science, Bangalore, 560012, India.
Centre for Ecological Sciences, Indian Institute of Science, Bangalore, 560012, India.
Centre for Ecological Sciences, Indian Institute of Science, Bangalore, 560012, India. Department of Life Sciences, The Natural History Museum, London SW7 5BD, United Kingdom.
Reptilia Agamidae Calotes new genus Microauris Monilesaurus Psammophilus Western Ghats

Abstract

Lizards of the genus Calotes are geographically restricted to South Asia, Indo-China and parts of Southeast Asia. The greatest diversity of the genus is from the biodiversity hotspots in South Asia: Western Ghats (Peninsular India), Sri Lanka and Indo-Burma. Here, we present a systematic revision of members of the genus Calotes from Peninsular India using a combination of molecular phylogeny, geographical distribution and morphological characters. We show that Calotes from the Western Ghats is paraphyletic and consists of three major clades, one of which is widely distributed in South and Southeast (SE) Asia, while the others are restricted to Peninsular India. The Peninsular Indian clade is composed of two sister clades: Psammophilus, with a wider distribution and a second clade, composed of two extant species, Calotes rouxii and Calotes ellioti and two new species, all restricted to the Western Ghats region. Based on morphological differences, we retain the generic status of Psammophilus and assign its sister clade to a new genus Monilesaurus gen. nov. and transfer the following species, C. rouxii and C. ellioti, to this new genus. We also provide diagnoses and descriptions for two new species recognized within Monilesaurus gen. nov. In addition, Calotes aurantolabium from the Western Ghats was observed to be deeply divergent and to share a sister-relationship with the clade composed of Calotes, Monilesaurus gen. nov., and Psammophilus. Based on its phylogenetic position and morphological attributes, we assign this species to a new genus Microauris gen. nov. These new discoveries highlight the evolutionary significance of the Western Ghats in housing novel lizard diversity.

 

References

  1. Amarasinghe, A.A.T., Karunarathna, D.M.S.S. & Fujinuma, J. (2014a) A new Calotes species from Sri Lanka with a redescription of Calotes liolepis Boulenger, 1885. Herpetologica, 70 (3), 323–338.
    https://doi.org/10.1655/HERPETOLOGICA-D-13-00087

    Amarasinghe, A.A.T., Karunarathna, D.M.S.S., Hallermann, J., Fujinuma, J., Grillitsch, H. & Campbell, P.D. (2014b) A new species of the genus Calotes (Squamata: Agamidae) from high elevations of the Knuckles Massif of Sri Lanka. Zootaxa, 3785 (1), 059–078.
    http://doi.org/10.11646/zootaxa.3785.1.5

    Amarasinghe, A.A.T., Manthey, U., Stöckli, E., Ineich, I., Kullander, S.O., Tiedemann, F., McCarthy, C. & Gabadage, D.E. (2009) The original descriptions and figures of Sri Lankan Agamid lizards (Squamata: Agamidae) of the 18th and 19th centuries. Taprobanica, 1 (1), 2–15.
    https://doi.org/10.4038/tapro.v1i1.2771

    Ananjeva, N.B. & Stuart, B.L. (2001) The agamid lizard Ptyctolaemus phuwuanensis Manthey and Nabhitabhata, 1991 from Thailand and Laos represents a new genus. Russian Journal of Herpetology, 8 (3), 165–170.

    Ananjeva, N.B., Orlov, N.L., Nguyen, Q.T. & Nazarov, R.A. (2007) A new species of Pseudocophotis (Agamidae: Acrodonta: Lacertilia: Reptilia) from central Vietnam. Russian Journal of Herpetology, 14 (2), 153–160.

    Anderson, J. (1879) Reptilia and amphibia. In: Anderson, J. (Ed.), Anatomical and Zoological Researches: Comprising an Account of the Zoological Results of the Two Expeditions to Western Yunnan in 1868 and 1875. Vol. I. Bernard Quaritch, London, pp. 705– 860.

    Bahir, M.M. & Silva, A. (2005) Otocryptis nigristigma, a new species of agamid lizard from Sri Lanka. In: Yeo, C.J., Ng, P.K.L. & Pethiyagoda, R. (Eds.), Contributions to biodiversity exploration and research in Sri Lanka. The Raffles Bulletin of Zoology, 12 (Supplement), 393–406.

    Bhupathy, S. & Kannan, P. (1997) Status of Agamid lizards in the Western Ghats of Tamil Nadu, India. Technical Report, No. 5. Salim Ali Centre for Ornithology and Natural History, Coimbatore, 27 pp.

    Biju, S.D., Garg, S., Gururaja, K.V, Shouche, Y. & Walujkar, S.A. (2014 a) DNA barcoding reveals unprecedented diversity in Dancing Frogs of India (Micrixalidae, Micrixalus): a taxonomic revision with description of 14 new species. Ceylon Journal of Science, 43 (1), 1–87.
    https://doi.org/10.4038/cjsbs.v43i1.6850

    Biju, S.D., Garg, S., Mahony, S., Wijayathilaka, N., Senevirathne, G. & Meegaskumbura, M.(2014b) DNA barcoding, phylogeny and systematics of Golden-backed frogs (Hylarana, Ranidae) of the Western Ghats-Sri Lanka biodiversity hotspot with the description of seven new species. Contributions to Zoology, 83, 269–335.

    Biswas, S. (1975) Reptilia from Bhutan with description of a new species of Calotes Rafinesque. Journal of Bombay Natural History Society, 72, 774–777.

    Boulenger, G.A. (1885) Catalogue of lizards in the British museum. Vol. 1. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae, Second edition. British Museum (Natural History), London, 436 pp.

    Boulenger, G.A. (1890) The Fauna of British India, including Ceylon and Burma: Reptilia and Batrachia. Taylor and Francis, London, 541 pp.

    Chaitanya, R., Giri, V.B. & Deepak, V. (2017) On the recent designation of a neotype for the taxon Calotes versicolor (Daudin, 1802) (Reptilia: Agamidae). Zootaxa, 4317 (3), 585–587.
    https://doi.org/10.11646/zootaxa.4317.3.10

    Daniels, R.J.R. & Ishwar, N.M. (1994) Rarity and the herpetofauna of the Southern Eastern Ghats, India. Cobra, 16, 2–14.

    Das, I., Dattagupta, B. & Gayen, N.C. (1998) History and catalogue of reptile types in the collection of the Zoological Survey of India. Journal of South Asian natural History, 3 (2), 121–172.

    Das, I., Charles, J.K. & Edwards, D.S. (2008) Calotes versicolor (Squamata: Agamidae)—A new invasive squamate from Borneo. Current Herpetology, 27 (2), 109–112.
    https://doi.org/10.3105/1881-1019-27.2.109

    Das, I. & Lakim, M. (2008) A second specimen of Complicitus nigrigularis (Ota & Hikida, 1991). Hamadryad, 32 (1), 82–84.

    Dahanukar N., Modak N., Krutha K., Nameer P.O., Padhye A.D. & Molur S. (2016) Leaping frogs (Anura: Ranixalidae) of the Western Ghats of India: An integrated taxonomic review. Journal of Threatened Taxa, 8 (10), 9221–9288. https://doi.org/10.11609/jott.2532.8.10.9221-9288

    Deepak, V., Vyas, R., Giri, V.B. & Karanth, K.P. (2015) A taxonomic mystery for more than 180 years: the identity and systematic position of Brachysaura minor (Hardwicke & Gray, 1827) Vertebrate Zoology, 65 (3), 371–381.

    Deepak, V., Giri, V.B., Asif, M., Dutta, S.K., Vyas, R., Zambre, A.M., Bhosale, H. & Karanth, K.P. (2016) Systematics and phylogeny of Sitana (Reptilia: Agamidae) of Peninsular India, with description of one new genus and five new species. Contributions to Zoology, 85, 67–111.

    Denzer, W., Günther, R. & Manthey, U. (1997) Annotated type catalogue of the Agamid lizards (Reptilia: Squamata: Agamidae) in the Museum für Naturkunde der Humbolt-Universität zu Berlin (former Zoological Museum Berlin). Mitteilungen aus dem Zoologischen Museum in Berlin, 73 (2), 309–332.
    https://doi.org/10.1002/mmnz.19970730209

    Denzer, W., Manthey, U., Mahlow, K. & Böhme W. (2015) The systematic status of Gonocephalus robinsonii Boulenger, 1908 (Squamata: Agamidae: Draconinae). Zootaxa, 4039 (1), 129–144.
    https://doi.org/10.11646/zootaxa.4039.1.5

    Dumril, A.M.C. & Bibron, G. (1837) Erpétologie Générale ou Histoire Naturelle Complete des Reptiles. Vol. 4. Librairie Encyclopédique Roret, Paris, 570 pp.

    Ganesh, S.R. & Chandramouli, S.R. (2013) Identification of two similar Indian Agamid lizards Calotes nemoricola Jerdon, 1853 and C. grandisquamis Günther, 1875. Russian Journal of Herpetology, 20, 33–35.

    Ganesh, S.R., Chadramouli, S.R., Sreekar, R. & Shankar, P.G. (2013) Reptiles of the central Western Ghats, India—a reappraisal and revised checklist, with emphasis on the Agumbe Plateau. Russian Journal of Herpetology, 20, 81–89.

    Gowande, G. (2016) Molecular phylogeography of Calotes versicolor from the Western Ghats of India. Msc dissertation, Savitribai Phule Pune University, Pune, 36 pp.

    Gowande, G., Mishra, A. & Mirza, Z.A. (2016) Neotype designation for Calotes versicolor Daudin, 1802 (Sauria: Agamidae) with notes on its systematics. Zootaxa, 4126 (2), 271–279.
    https://doi.org/10.11646/zootaxa.4126.2.7

    Gray, E. (1845) Catalogue of the specimens of lizards in the collection British museum. Trustees of die British Museum/ Edward Newman, London, 289 pp.

    Grismer, J.L., Schulte, J.A., Alexander, A., Wagner, P., Travers, S.L., Buehler, M.D., Welton, L.J. & Brown, R.M. (2016) The Eurasian invasion: phylogenomic data reveal multiple Southeast Asian origins for Indian Dragon Lizards. BMC evolutionary biology, 16, 1–11.
    https://doi.org/10.1186/s12862-016-0611-6

    Grismer, L.L., Quah, E.S.H., Wood Junior, P.L., Anuar, S., Muin, A., Davis, H.R., Murdoch, M.L., Grismer, J.L., Cota, M. & Cobos, A.J. (2016) Dragons in the mist: three new species of Pseudocalotes Fitzinger (Squamata: Agamidae) from the sky island archipelago of Peninsular Malaysia. Zootaxa, 4136 (3), 461–490.
    https://doi.org/10.11646/zootaxa.4136.3.3

    Günther, A. (1864) The Reptiles of British India. Taylor & Francis, London, 452 pp.

    Günther, A. (1870) Descriptions of a new Indian lizard of the genus Calotes. Proceedings of the Zoological Society of London, 778–779.

    Günther, A. (1872) Descriptions of some Ceylonese Reptiles and Batrachians. Annals Magazine of natural History, 4, 85–88.
    https://doi.org/10.1080/002229372011951778

    Günther, A. (1875) Second report on collections of Indian Reptiles obtained by the British Museum. Proceedings of the Zoological Society of London, 1875, 224–234.

    Hallermann, J. (2000) A new species of Calotes from the Moluccas (Indonesia) with notes on the biogeography of the genus (Sauria: Agamidae). Bonner Zoologische Beiträge, 49, 155–163.

    Hallermann, J. & Böhme, W. (2000) A review of the genus Pseudocalotes (Squamata: Agamidae), with description of a new species from West Malaysia. Amphibia-Reptilia, 21 (2), 193–210.
    https://doi.org/10.1163/156853800507372

    Hallermann, J. & McGuire, J.A. (2001) A new species of Pseudocalotes (Squamata: Agamidae) from Bukit Larut, West Malaysia. Herpetologica, 57 (3), 255–265.

    Hallermann, J., Orlov, N. & Ananyeva, N. (2004) Notes on distribution and colour pattern of the rare agamid lizard Lophocalotes ludekingii (Bleeker 1860) in Sumatra (Indonesia). Salamandra, 40, 303–306.

    Hallermann, J. & Böhme, W. (2007) On the validity of two recently described species of the endemic Sri Lankan genus Cophotis Peters, 1861 (Squamata: Agamidae) from the Knuckles Forest Range. Salamandra, 43, 187–190.

    Hallermann, J., Truong, N.Q., Orlov, N. & Ananjeva, N. (2010) A new species of the genus Pseudocalotes (Squamata: Agamidae) from Vietnam. Russian Journal of Herpetology, 17 (1), 31–40.

    Hardwicke, T. & Gray, J.E. (1827) A synopsis of the species of saurian reptiles, collected in India by Major-General Hardwicke. Zoological Journal, 3, 213–229.

    Hartmann, T., Geissler, P., Nikolay, A.J., Ihlow, P.F., Galoyan, E.A., Rödder, D. & Böhme, W. (2013) A new species of the genus Calotes Cuvier, 1817 (Squamata: Agamidae) from southern Vietnam. Zootaxa, 3599 (3), 246–260.
    https://doi.org/10.11646/zootaxa.3599.3.3

    Harikrishnan, S., Vasudevan, K. & Choudhury, B.C. (2010) A review of herpetofaunal descriptions and studies from Andaman and Nicobar Islands, with an updated checklist. In: Ramakrishna, Raghunathan, C. & Sivaperuman, C. (Eds.), Recent Trends in Biodiversity of Andaman and Nicobar Islands. Zoological Survey of India, Kolkata, pp 387–398.

    Harikrishnan, S. & Vasudevan, K. (2013) Rediscovery of Calotes andamanensis Boulenger, 1891, and assessment of its generic allocation (squamata: sauria: Agamidae). Herpetozoa, 26, 3–13.

    Harvey, M.B., Hamidy, A., Kurniawan, N., Shaney, K. & Smith, E.N. (2014) Three new species of Pseudocalotes (Squamata: Agamidae) from southern Sumatra, Indonesia. Zootaxa, 3841 (2), 211–238.
    https://doi.org/10.11646/zootaxa.3841.2.3

    Harvey, M. B., Shaney, K., Hamidy, A., Kurniawan, N. & Smith, E. N. (2017) A new species of Pseudocalotes (Squamata: Agamidae) from the Bukit Barisan Range of Sumatra with an Estimation of its phylogeny. Zootaxa, 4276 (2), 215–232.
    https://doi.org/10.11646/zootaxa.4276.2.4

    Huang, Y., Guo, X., Ho, S., Shi, H., Li, J., Li, J., Cai, B. & Wang, Y. (2013) Diversification and Demography of the Oriental Garden Lizard (Calotes versicolor) on Hainan Island and the Adjacent Mainland. PloS One, 8 (6), e64754.
    https://doi.org/10.1371/journal.pone.0064754

    Honda, M., Ota, H., Kobayashi, M., Nabhitabhata, J., Yong, H., Sengoku, S. & Hikida, T. (2000) Phylogenetic Relationships of the Family Agamidae (Reptilia: Iguania) Inferred from Mitochondrial DNA Sequences. Zoological Science, 17 (4), 527–537.
    https://doi.org/10.2108/0289-0003(2000)17[527:PROTFA]2.0.CO;2

    Inger, R.F. (1960) A review of the agamid lizards of the genus Phoxophrys Hubrecht. Copeia, 3, 221–225.
    https://doi.org/10.2307/1439661

    Inger, R.F. & Stuebing, R.B. (1994) First record of the lizard genus Pseudocalotes (Lacertilia: Agamidae) in Borneo, with description of a new species. Raffles Bulletin of Zoology, 42 (4), 961–965.

    Inger, R. F., Shaffer, H. B, Koshy, M. & Bakde, R. (1984) A report on a collection of amphibians and reptiles from the Ponmudi, Kerala, South India. Journal of Bombay Natural History Society, 81, 551–570.

    Ishwar, N.M. & Das, I. (1998) Rediscovery of Calotes andamanensis Boulenger 1891 and a reassessment of the type locality. Journal of Bombay Natural History Society, 95, 513–514.

    Jerdon, T.C. (1854) Catalogue of the Reptiles inhabiting the Peninsula of India. Part 1. Journal of Asiatic Society of Bengal, 22, 462–479.

    Krishnan, S. (2008) New Species of Calotes (Reptilia: Squamata: Agamidae) from the Southern Western Ghats, India. Journal of Herpetology, 42 (3), 530–535.
    https://doi.org/10.1670/06-198.1

    Leong, T.M. (2001) The first female topotype of the female agamid Pseudocalotes flavigula (Smith, 1924) from Peninsular Malaysia, with description of colouration in life. Hamadryad, 26, 336–338.

    Macey, J.R., Schulte, J.A., Larson, A., Ananjeva, N.B., Wang, Y., Pethiyagoda, R., Rastegar-Pouyani, N. & Papenfuss, T.J. (2000) Evaluating trans-tethys migration: an example using acrodont lizard phylogenetics. Systematic Biology, 49 (2), 233–256.
    https://doi.org/10.1093/sysbio/49.2.233

    Mahony, S. (2010) Systematic and taxomonic revaluation of four little known Asian agamid species, Calotes kingdonwardi Smith, 1935, Japalura kaulbacki Smith, 1937, Salea kakhienensis Anderson, 1879 and the monotypic genus Mictopholis Smith, 1935 (Reptilia: Agamidae). Zootaxa, 2514, 1–23.

    Manamendra-Arachchi, K., de Silva, A. & Amarasinghe, T. (2006) Description of a second species of Cophotis (Reptilia: Agamidae) from the highlands of Sri Lanka. Lyriocephalus, 6, 1–8.

    Manthey, U. (2008) Agamid Lizards of Southern Asia, Draconinae 1. Edition Chimaira, Frankfurt, 160 pp.

    Manthey, U. & Denzer, W. (1992) The true Angle-head agamas of the genus Gonocephalus Kaup (Sauria: Agamidae) IV. Gonocephalus mjoebergi Smith, 1925 and Gonocephalus robinsonii Boulenger, 1908. Sauria, 1, 15–19.

    Manthey, U. & Denzer, W. (2000) Description of a new genus Hypsicalotes gen. nov (Sauria: Agamidae) from Mount Kinabalu, north Borneo, with remarks on the generic identity of Gonocephalus schultzewestrumi Urban, 1999. Hamadryad, 25, 13–20.

    Manthey, U. & Nabhitabhata, J. (1991) Eine neue Agame, Ptyctolaemus phuwuanensis sp. n. (Sauria: Agamidae), aus Nordost-Thailand. Sauria, 13, 3–6.

    Moody, S.M. (1980) Phylogenetic and historical biogeographical relationships of the genera in the family Agamidae (Reptilia; Lacertilia). Unpublished thesis, The University of Michigan, Ann Arbor, Michigan, 373 pp.

    Naniwadekar, R. & Deepak, V. (2008) New distribution record for Calotes nemoricola Jerdon, 1853 from the Kudremukh hills, Karnataka, India. Journal of Bombay Natural History Society, 105 (1), 99.

    Ota, H. & Hikida, T. (1991) Taxonomic review of the lizards of the genus Calotes Cuvier, 1817 (Agamidae Squamata) from Sabah, Malaysia. Tropical Zoology, 4 (2), 179–192.
    https://doi.org/10.1080/03946975.1991.10539488

    Ota, H. & Hikida, T. (1996) The second specimen of Calotes kinabaluensis de Grijs (Squamata: Agamidae) from Sabah, Malaysia, with comments on the taxonomic status of the species. Journal of Herpetology, 30 (2), 288–291.
    https://doi.org/10.2307/1565529

    Pethiyagoda, R. & Manamendra-Arachchi, K. (1998) A revision of the endemic Sri Lankan agamid lizard genus Ceratophora Gray, 1835, with description of two new species. Journal of South Asian natural History, 3 (1), 1–50.

    Pyron, R.A., Burbrink, F.T. & Wiens, J.J. (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology, 13, 93.
    https://doi.org/10.1186/1471-2148-13-93

    Radder, R.S. (2006) An overview of geographic variation in the life history traits of the tropical agamid lizard, Calotes versicolor. Current Science, 91, 1354–1363.

    Radder, R.S., Saidapur, S.K. & Shanbhag, B.A. (2005) Population density, microhabitat use and activity pattern of the Indian rock lizard, Psammophilus dorsalis (Agamidae). Current Science, 89 (3), 560–566.

    Rambaut, A., Suchard, M., Xie, D. & Drummond, A. (2014) Tracer. Version 1. 6. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 12 August 2014)

    Robin, V. V, Sinha, A. & Ramakrishnan, U. (2010) Ancient geographical gaps and paleo-climate shape the phylogeography of an endemic bird in the sky islands of southern India. PloS One, 5 (10), e13321.
    https://doi.org/10.1371/journal.pone.0013321

    Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.
    https://doi.org/10.1093/sysbio/sys029

    Schulte II, J.A., Vindum, J.V., Win, H., Thin, T., Lwin, K.S. & Shein, A.K. (2004) Phylogenetic relationships of the genus Ptyctolaemus (Squamata: Agamidae), with a description of a new species from Chin Hills of Western Myanmar. Proceedings of the California Academy of Sciences, 55, 222–247.

    Samarawickrama, V.A.M.P.K., Ranawana, K.B., Rajapaksha, D.R.N.S., Ananjeva, N.B., Orlov, N.L., Ranasinghe, J.M.A.S. & Samarawickrama, V.A.P. (2006) A new species of the genus Cophotis (Squamata: Agamidae) from Sri Lanka. Russian Journal of Herpetology, 13, 207–214.

    Smith, M.A. (1935) The Fauna of British India, Including Ceylon and Burma; Reptilia and Amphibia. Vol. II. Sauria. Taylor and Francis, London, 440 pp.

    Simon, C., Franke, A. & Martin, A. (1991) The polymerase chain reaction: DNA extraction and amplification. In: Hewitt, G.M., Johnson, A.W.B. & Young, J.P.W. (Eds.), Molecular Techniques in Taxonomy. NATO ASI Series. Series H: Cell Biology. Vol. 57. Springer, Berlin, Heidelberg, pp. 329–355.
    https://doi.org/10.1007/978-3-642-83962-7_22

    Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML. Organism Diversity and Evolution, 12, 335–337.
    https://doi.org/10.1007/s13127-011-0056-0

    Srinivasulu, C., Srinivasulu, B. & Molur, S. (Compilers) (2014) The Status and Distribution of Reptiles in the Western Ghats, India. Conservation Assessment and Management Plan (CAMP). Wildlife Information Liaison Development Society, Coimbatore, Tamil Nadu, 148 pp.

    Stamatakis, A., Ludwig, T. & Meier, H. (2005) RAxML-III: a fast program for maximum likelihood-based inference of large phylogenetic trees. Bioinformatics, 21, 456–463.
    https://doi.org/10.1093/bioinformatics/bti191

    Stoliczka, F. (1871) Notes on new or little known Indian lizards. Proceedings of the Asiatic Society of Bengal, 9, 192–195.

    Stoliczka, F. (1872) Notes on various new or little known Indian lizards. Proceedings of the Asiatic Society of Bengal, 41, 86–135.

    Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA 5: Molecular Evolutionary Genetics Analysis version 5.1. Molecular Biology and Evolution, 30, 2725–2729.

    Taylor, E.H. (1963) The lizards of Thailand. University of Kansas Science Bulletin, 44, 687–1077.

    Uetz, P. & Hošek, J. (2016) The Reptile Database. Available from: http://reptile-database.reptarium.cz (accessed 6 May 2016)

    Van Bocxlaer, I., Biju, S.D., Willaert, B., Giri, V.B., Shouche, Y.S. & Bossuyt, F. (2012) Mountain-associated clade endemism in an ancient frog family (Nyctibatrachidae) on the Indian subcontinent. Molecular Phylogenetics and Evolution, 62, 839–847.
    https://doi.org/10.1016/j.ympev.2011.11.027

    Vijayakumar, S.P., Dinesh, K.P., Prabhu, M.V. & Shanker, K. (2014) Lineage delimitation and description of nine new species of bush frogs (Anura: Raorchestes, Rhacophoridae) from the Western Ghats Escarpment. Zootaxa, 3893 (4), 451–488.
    https://doi.org/10.11646/zootaxa.3893.4.1

    Vijayakumar, S.P., Menezes, R.C., Jayarajan, A. & Shanker, K. (2016) Glaciations, gradients, and geography: multiple drivers of diversification of bush frogs in the Western Ghats Escarpment. Proceedings of the Royal Society B: Biological Sciences, 283, 20161011.
    https://doi.org/10.1098/rspb.2016.1011

    Vindum, J.V., Win, H., Thin, T., Lwin, K.S., Shein, A.K. & Tun, H. (2003) A new Calotes (Squamata: Agamidae) from the Indo-Burman range of western Myanmar (Burma). Proceedings of the California Academy of Science, 54 (1), 1–16.

    Zhao, E. & Li, S. (1984) A new species of Calotes (Lacertilia: Agamidae) from Xizang (Tibet) [in Chinese]. Acta Herpetologica Sinica, 3 (4), 77–78.

    Zug, G.R., Brown, H.H., Schulte, J.A. & Vindum, J.V. (2006) Systematics of the Garden Lizards, Calotes versicolor Group (Reptilia, Squamata, Agamidae), in Myanmar: Central Dry Zone Populations. Proceedings of the California Academy of Sciences, 57 (2), 35–68.