Abstract
We present an expanded ND2 phylogeny of the brookii and prashadi groups of Indian Hemidactylus. While relationships within the clades are not well resolved, we recover many well supported subclades and a number of deeply divergent lineages within each clade. We use morphological data to describe three of these divergent lineages as new species; Hemidactylus kolliensis sp. nov., a member of the prashadi group from high elevation in the Kollimalai Massif in Tamil Nadu, Hemidactylus chikhaldaraensis sp. nov. and Hemidactylus sankariensis sp. nov., both members of the brookii group, are described from high elevation of the Satpuras in Maharashtra and low elevation of Tamil Nadu, respectively. The three new species exhibit >15% divergence for the ND2 gene from their phylogenetic sisters and differ in a number of morphological characters from all Indian congeners. The discovery of three more endemic Hemidactylus species from rocky habitats suggests that many more remain to be discovered across India.
References
Aengals, R. & Ganesh, S.R. (2013) Rhinophis goweri—A new species of shieldtail snake from the southern Eastern Ghats, India. Russian Journal of Herpetology, 20, 61–65.
Agarwal, I. (2016). Two new species of ground-dwelling Cyrtodactylus (Geckoella) from the Mysore Plateau, South India. Zootaxa, 4193, 228–244.
Agarwal, I., Giri, V.B. & Bauer, A.M. (2011) A new cryptic rock-dwelling Hemidactylus (Squamata: Gekkonidae) from south India. Zootaxa, 2765 (1), 21–37.
https://doi.org/10.11646/zootaxa.2765.1.2
Agarwal, I., Giri, V.B. & Bauer, A.M. (2018). On the Status of Cyrtodactylus malcolmsmithi (Constable, 1949). Breviora, 557: 1–11.
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. [published online]
https://doi.org/10.1007/s13127-019-00392-5
Bansal, R. & Karanth, K.P. (2010) Molecular phylogeny of Hemidactylus geckos (Squamata: Gekkonidae) of the Indian subcontinent reveals a unique Indian radiation and an Indian origin of Asian house geckos. Molecular Phylogenetics and Evolution, 57, 459–465.
https://doi.org/10.1016/j.ympev.2010.06.008
Batuwita, S. & Pethiyagoda, R. (2012) Rediscovery of the Sri Lankan 'house gecko' Hemidactylus pieresii Kelaart (Reptilia: Gekkonidae) with a redescription of Hemidactylus depressus Gray. Zootaxa, 3359, 17–30.
Bauer, A.M. (1999) Evolutionary scenarios in the Pachydactylus group geckos of southern Africa: new hypotheses. African Journal of Herpetology, 48, 53–62.
https://doi.org/10.1080/21564574.1999.9651072
Bauer, A.M., Giri, V.B., Greenbaum, E., Jackman, T.R., Dharne, M.S. & Shouche, Y.S. (2008) On the systematics of the gekkonid genus Teratolepis Günther, 1869: another one bites the dust. Hamadryad, 33, 13–28.
Bauer, A.M., Jackman, T.R., Greenbaum, E., Giri, V. & De Silva, A. (2010a) 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
Bauer, A.M., Jackman, T.R., Greenbaum, E., de Silva, A., Giri, V.B. & Das, I. (2010b) Molecular evidence for the taxonomic status of Hemidactylus brookii group taxa (Squamata: Gekkonidae). The Herpetological Journal, 20, 129–138.
Bauer, A.M., Vyas, R., Jackman, T.R., Lajmi, A. & Giri, V.B. (2012) Hemidactylus porbandarensis Sharma, 1981 is a synonym of Hemidactylus robustus Heyden, 1827. Hamadryad, 36, 46–51.
Carranza, S. & Arnold, E.N. (2006) Systematics, biogeography, and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) elucidated using mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 38, 531–545.
https://doi.org/10.1016/j.ympev.2005.07.012
Chaitanya, R., Lajmi, A. & Giri, V.B. (2018) A new cryptic, rupicolous species of Hemidactylus Oken, 1817 (Squamata: Gekkonidae) from Meghamalai, Tamil Nadu, India. Zootaxa, 4374 (1), 49–70.
https://doi.org/10.11646/zootaxa.4374.1.3
Dandge, P.H. & Tiple, A.D. (2015) A new species of rupicolus gecko of the genus Hemidactylus Oken, 1817 (Reptilia: Squamata: Gekkonidae) from Maharashtra, Central India. Russian Journal of Herpetology, 22, 233–240.
Deso, G., Bauer, A., Probst, J.M. & Sanchez, M. (2013) Hemidactylus parvimaculatus Deraniyagala, 1953 (Squamata: Gekkonidae) à La Réunion: écologie et repartition. Bulletin de la Société Herpétologique de France, 144, 33–49.
Giri, V.B. & Bauer, A.M. (2008) A new ground-dwelling Hemidactylus (Squamata: Gekkonidae) from Maharashtra, with a key to the Hemidactylus of India. Zootaxa, 1700, 21–34.
Giri, V.B., Bauer, A.M., Mohapatra, P.P., Srinivasulu, C. & Agarwal, I. (2017) A new species of large-bodied, tuberculate Hemidactylus Oken (Squamata: Gekkonidae) from the Eastern Ghats, India. Zootaxa, 4347 (2), 331–345.
https://doi.org/10.11646/zootaxa.4347.2.8Grismer, L.L., Wood, P.L., Thura, M.K., Zin, T., Quah, E.S., Murdoch, M.L., Grismer, M.S., Lin, A., Kyaw, H. & Lwin, N. (2017) Twelve new species of Cyrtodactylus Gray (Squamata: Gekkonidae) from isolated limestone habitats in east-central and southern Myanmar demonstrate high localized diversity and unprecedented microendemism. Zoological Journal of the Linnean Society, 45, 251–484.
https://doi.org/10.1093/zoolinnean/zlx057
Grismer, L.L., Wood, P.L. Jr., Anuar, S., Muin, M.A., Quah, E.S.H., McGuire, J.A., Brown, R.M., Van, T.N. & Hong, T.P. (2013) Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zoological Journal of the Linnaean Society, 169, 849–880.
https://doi.org/10.1111/zoj.12064
Kathriner, A., O'shea, M. & Kaiser, H. (2014) Re-examination of Hemidactylus tenkatei van Lidth de Jeude, 1895: Populations from Timor provide insight into the taxonomy of the H. brookii Gray, 1845 complex (Squamata: Gekkonidae). Zootaxa, 3887 (5), 583–599.
https://doi.org/10.11646/Zootaxa.3887.5.5
Khandekar, A. (2019) A new species of rock-dwelling Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from Tamil Nadu, southern India. Zootaxa, 4571 (3), 383–397.
Lajmi, A., Giri, V.B. & Karanth, K.P. (2016) Molecular data in conjunction with morphology help resolve the Hemidactylus brookii complex (Squamata: Gekkonidae). Organisms Diversity & Evolution, 16, 659–677.
https://doi.org/10.1007/s13127-016-0271-9
Lajmi, A., Datta-Roy, A., Vyas, R., Agarwal, I. & Karanth, K.P. (2018a) The curious case of Hemidactylus gujaratensis (Squamata: Gekkonidae). Zootaxa, 4388 (1), 137–142.
https://doi.org/10.11646/zootaxa.4388.1.11
Lajmi, A., Bansal, R., Giri, V. & Karanth, P. (2018b) Phylogeny and biogeography of the endemic Hemidactylus geckos of the Indian subregion suggest multiple dispersals from Peninsular India to Sri Lanka, Zoological Journal of the Linnean Society, zly047.
https://doi.org/10.1093/zoolinnean/zly047
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
Mahony, S. (2011) Taxonomic revision of Hemidactylus brookii Gray: a re-examination of the type series and some Asian synonyms, and a discussion of the obscure species Hemidactylus subtriedrus Jerdon (Reptilia: Gekkonidae). Zootaxa, 3042 (1), 37–67.
https://doi.org/10.11646/zootaxa.3042.1.4
Mirza, Z. A. & Raju, D. (2017). A new rupicolous species of gecko of the genus Hemidactylus Oken, 1817 from the Satpura Hills, Central India. Amphibian & Reptile Conservation, 11 (e137), 51–71.
Mirza, Z.A. & Sanap, R. (2014) New cryptic species of gecko of the genus Hemidactylus Oken, 1817 (Reptilia: Gekkonidae) from Southern India. Taprobanica, 6 (1), 12–20.
https://doi.org/10.4038/tapro.v6i1.7056
Mirza, Z.A., Gowande, G.G., Patil, R., Ambekar, M. & Patel, H. (2018) First appearance deceives many: disentangling the Hemidactylus triedrus species complex using an integrated approach. PeerJ, 6, e5341.
https://doi.org/10.7717/peerj.5341
Oliver, P.M., Laver, R.J., De Mello Martins, F., Pratt, R.C., Hunjan, S. & Moritz, C.C. (2017). A novel hotspot of vertebrate endemism and an evolutionary refugium in tropical Australia. Diversity and Distributions, 23, 53–66.
https://doi.org/10.1111/ddi.12506
Rösler, H. & Glaw, F. (2010) Morphologische Variation und Taxonomie von Hemidactylus brookii Gray, 1845 und Hemidactylus angulatus Hallowell, 1854 sowie phänotypisch ähnlicher Taxa. Spixiana, 33, 139–160.
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
Šmíd, J., Carranza, S., Kratochvíl, L., Gvoždík, V., Nasher, A.K. & Moravec, J. (2013) Out of Arabia: a complex biogeographic history of multiple vicariance and dispersal events in the gecko genus Hemidactylus (Reptilia: Gekkonidae). PLoS One, 8, e64018.[ pmid:23724016]
https://doi.org/10.1371/journal.pone.0064018
Šmíd, J., Moravec, J., Kratochvíl, L., Nasher, A.K., Mazuch, T., Gvoždík, V. & Carranza, S. (2015) Multilocus phylogeny and taxonomic revision of the Hemidactylus robustus species group (Reptilia, Gekkonidae) with descriptions of three new species from Yemen and Ethiopia. Systematics and Biodiversity, 13, 346–368.
https://doi.org/10.1080/14772000.2014.996264
Šmíd, J., Shobrak, M., Wilms, T., Joger, U. & Carranza, S. (2017) Endemic diversification in the mountains: genetic, morphological, and geographical differentiation of the Hemidactylus geckos in southwestern Arabia. Organisms, Diversity and Evolution, 17, 267.
https://doi.org/10.1007/s13127-016-0293-3
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.
Srikanthan, A.N., Swamy, P., Mohan, A.V. & Pal, S. (2018) A distinct new species of riparian rock-dwelling gecko (genus: Hemidactylus) from the southern Western Ghats. Zootaxa, 4434 (1), 141–157.
https://doi.org/10.11646/zootaxa.4434.1.9
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
Srinivasulu, C., Srinivasulu, A. & Kumar, G.C. (2018) A new cryptic rock-dwelling Hemidactylus Oken, 1817 (Squamata: Gekkonidae) from northern Karnataka, India. Zootaxa, 4444 (1), 25–42.
https://doi.org/10.11646/zootaxa.4444.1.2
Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based 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. Available from: http://www.reptile-database.org (accessed 2 November 2018)
Valdecasas, A.G., Williams, D. & Wheeler, Q.D. (2008) 'Integrative taxonomy' then and now: a response to Dayrat 2005. Biological Journal of the Linnean Society, 93, 211–216.
https://doi.org/10.1111/j.1095-8312.2007.00919.x
Zhang, Jiajie, Kapli, P., Pavlidis, P. & Stamatakis, A. (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics, 29, 2869–2876.
https://doi.org/10.1093/bioinformatics/btt499