Abstract
We revise the systematics of Hemidactylus giganteus—an Indian endemic leaf-toed gecko based on recent collections from the type locality and other localities in peninsular India. We provide a comprehensive redescription of H. giganteus sensu stricto and describe three new allied species from the Karnataka and Telangana states in India. We analysed the phylogenetic relationships between the three new species, H. raya sp. nov., H. saxicolus sp. nov., and H. aemulus sp. nov., and found them to be nested within the H. giganteus sensu lato branch. The pairwise genetic divergence between these new cryptic taxa from H. giganteus sensu stricto ranged from 5.88% to 10.95% for the Cyt b gene. The new species also differ from H. giganteus sensu stricto based on number of femoral pores on each thigh and poreless scales separating them (18–23 femoral pores and seven or eight poreless scales versus 17 femoral pores and seven poreless scales in H. raya sp. nov., 26 or 27 femoral pores and nine poreless scales in H. saxicolus sp. nov., and 22–25 femoral pores and nine to ten poreless scales in H. aemulus sp. nov.), corroborated by linear discriminant analysis based on 29 morphometric and meristic characters. Our species delimitation analysis shows the presence of three additional putative new species awaiting formal description within the H. giganteus sensu lato branch.
References
Agarwal, I., Bauer, A.M., Giri, V.B. & Khandekar, A. (2019) An expanded ND2 phylogeny of the brookii and prashadi groups with the description of three new Indian Hemidactylus Oken (Squamata: Gekkonidae). Zootaxa, 4619 (3), 431–458. https://doi.org/10.11646/zootaxa.4619.3.2
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), 1–11. https://doi.org/10.3099/MCZ41.1
Agarwal, I., Thackeray, T. & Khandekar, A. (2020a) Geckos in the granite: two new geckos (Squamata: Gekkonidae) from rocky, scrub habitats in Rishi Valley, Andhra Pradesh, India. Zootaxa, 4838 (4), 451–474. https://doi.org/10.11646/zootaxa.4838.4.1
Agarwal, I., Thackeray, T., Pal, S. & Khandekar, A. (2020b). Granite boulders act as deep-time climate refugia: A Miocene divergent clade of rupicolous Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from the Mysore Plateau, India, with descriptions of three new species. Journal of Zoological Systematics and Evolutionary Research, 58 (4), 1234–1261. https://doi.org/10.1111/jzs.12391
Altschul, S.F., Madden, T.L., Schäffer, A.A., Zhang, J., Zhang, Z., Miller, W. & Lipman, D.J. (1997) Gapped BLAST and PSI-BLAST: a new generation of protein database search programs. Nucleic Acids Research, 25 (17), 3389–3402. https://doi.org/10.1093/nar/25.17.3389
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 (1), 459–465. https://doi.org/10.1016/j.ympev.2010.06.008
Bansal, R. & Karanth, K.P. (2013) Phylogenetic analysis and molecular dating suggest that Hemidactylus anamallensis is not a member of the Hemidactylus radiation and has an ancient late cretaceous origin. Plos One, 8 (5), e60615. https://doi.org/10.1371/journal.pone.0060615
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 (1), 13–28.
Bauer, A.M., Jackman, T.R., Greenbaum, E., Giri, V.B. & De Silva, A. (2010a) South Asia supports a major endemic radiation of Hemidactylus geckos. Molecular Phylogenetics and Evolution, 57 (1), 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., Vya,s 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.
Bruford, M.W., Hanotte, O., Brookfield, J.F.Y. & Burke, T. (1992) Single-locus and multilocus DNA fingerprint. In: Hoelzel, A.R. (Ed.), Molecular Genetic Analysis of Populations: A Practical Approach. IRL Press, Oxford. pp. 225–270.
Ceríaco, L.M.P., Agarwal, I., Marques, M.P. & Bauer, A.M. (2020) A review of the genus Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from Angola, with the description of two new species. Zootaxa, 4746 (1), 1–71. https://doi.org/10.11646/zootaxa.4746.1.1
Chaitanya, R., Agarwal, I., Lajmi, A. & Khandekar, A. (2019) A novel member of the Hemidactylus brookii complex (Squamata: Gekkonidae) from the Western Ghats of Maharashtra, India. Zootaxa, 4646 (2), 236–250. https://doi.org/10.11646/zootaxa.4646.2.2
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
Chandra, K. & Gajbe, P.U. (2005) An inventory of herpetofauna of Madhya Pradesh and Chhattisgarh. Zoo’s Print Journal, 20, 1812–1819. https://doi.org/10.11609/JoTT.ZPJ.1087.1812-9
Chethan Kumar, G., Srinivasulu, C. & Seetharamaraju, M. (2015) On the distribution of the reticulated leaf-toed gecko (Hemidactylus reticulatus). Taprobanica, 7 (4), 272–274.
Chopra, R.N. (1968) On a new locality and habitat of Hemidactylus giganteus. Science and Culture, 38, 376.
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 (3), 233–240.
Daniels, R.J.R. (2000) Reptiles and amphibians of Karnataka. Cobra, 42, 1–11.
Darriba, D., Taboada, G., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772. https://doi.org/10.1038/nmeth.2109
Deepak, V., Khandekar, A., Chaitanya, R. & Karanth, P. (2018) Descriptions of two new endemic and cryptic species of Sitana Cuvier, 1829 from peninsular India. Zootaxa, 4434 (2), 327–365. https://doi.org/10.11646/zootaxa.4434.2.5
Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32 (5), 1792–1797. https://doi.org/10.1093/nar/gkh340
Gaikwad, K.S., Kulkarni, H., Bhambure, R. & Giri, V.B. (2010) Notes on the distribution, natural history and variation of Hemidactylus albofasciatus (Grandison and Soman, 1963) (Squamata: Gekkonidae). Journal of the Bombay Natural History Society, 106 (3), 305–312.
Ganesh, S.R. & Chandramouli, S.R. (2010) Rediscovery of Hemidactylus scabriceps (Annandale, 1906) (Reptilia: Sauria: Gekkonidae) from eastern Tamil Nadu, India. Russian Journal of Herpetology, 17 (1), 70–74. https://doi.org/10.30906/1026-2296-2010-17-1-70-74
Ganesh, S.R., Bhupathy, S., David, P., Kumar, N.S. & 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. https://doi.org/10.30906/1026-2296-2014-21-1-53-64
Ganesh, S.R., Rameshwaran, M., Joseph, N.A. & Jerith, A.M. (2017) On two little-known terrestrial South Asian geckoes Hemidactylus reticulatus and Hemidactylus scabriceps (Reptilia: Gekkonidae). Journal of Threatened Taxa, 9 (5), 10171–10177. https://doi.org/10.11609/jot.3163.9.5.10171-10177
Giri, V.B. (2008) A new rock dwelling Hemidactylus (Squamata: Gekkonidae) from Maharashtra, India. Hamadryad, 32, 25–33.
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 (1), 21–34. https://doi.org/10.11646/zootaxa.1700.1.2
Giri, V.B., Bauer, A.M. & Chaturvedi, N. (2003) Notes on the distribution, natural history and variation of Hemidactylus giganteus Stolickza, 1871. Hamadryad, 27 (2), 217–221.
Giri, V.B., Bauer, A.M. & Gaikwad, K.S. (2009) A new ground-dwelling species of Cnemaspis Strauch (Squamata: Gekkonidae) from the northern Western Ghats, Maharashtra, India. Zootaxa, 2164 (1), 49–60. https://doi.org/10.11646/zootaxa.2164.1.5
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.4347.2.8
Hasegawa, M., Kishino, H. & Yano, T.A. (1985) Dating of the human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution, 22 (2), 160–174. https://doi.org/10.1007/BF02101694
Hutton A.F. & David P. (2009) Notes on a collection of snakes from south India, with emphasis on the snake fauna of the Megamalai Hills (High Wavy Mountains). Journal of Bombay Natural History Society, 105, 299–316.
Khandekar, A., Thackeray, T., Pawar, S. & Agarwal, I. (2020a) A new medium-bodied rupicolous Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from the Sirumalai massif, Tamil Nadu, India. Zootaxa, 4852 (1), 83–100. https://doi.org/10.11646/zootaxa.4852.1.4.
Khandekar, A., Thackeray, T. & Agarwal, I. (2020b) A new cryptic Cnemaspis Strauch (Squamata: Gekkonidae) from an isolated granite hill on the Mysore Plateau, Karnataka, India. Zootaxa, 4845 (4), 509–528. https://doi.org/10.11646/zootaxa.4845.4.3
Kocher, T.D., Thomas, W.K., Meyer, A., Edwards, S.V., Pääbo, S., Villablanca, F.X. & Wilson, A.C. (1989) Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers. Proceedings of National Academy of Sciences, USA, 86 (16), 6196–6200. https://doi.org/10.1073/pnas.86.16.6196
Lajmi, A. & Karanth, P.K. (2020) Eocene-Oligocene cooling and the diversification of Hemidactylus geckos in Peninsular India. Molecular Phylogenetics and Evolution, 142, 106637. https://doi.org/10.1016/j.ympev.2019.106637
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 and Evolution, 16 (3), 659–677. https://doi.org/10.1007/s13127-016-0271-9
Lajmi, A., Giri, V.B., Singh, T. & Agarwal, I. (2020) Two new species of yellow-tailed Hemidactylus Goldfuss, 1820 (Squamata Gekkonidae) from rocky outcrops on the Telangana Plateau, India. Zootaxa, 4895 (4), 483–504. https://doi.org/10.11646/zootaxa.4895.4.2
Lajmi, A., Verma, A. & Karanth, K.P. (2019) Repeated evolution of terrestrial lineages in a continental lizard radiation. Journal of Evolutionary Biology, 33 (1), 57–66. https://doi.org/10.1111/jeb.13544
Mahony, S. (2009) A new species of gecko of the genus Hemidactylus (Reptilia: Gekkonidae) from Andhra Pradesh, India. Russian Journal of Herpetology, 16 (1), 27–34. https://doi.org/10.30906/1026-2296-2009-16-1-27-34
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. (2018) A new cryptic species of ground-dwelling Hemidactylus (Squamata: Gekkonidae) from southern India. Phyllomedusa 17 (2), 169–180. https://doi.org/10.11606/issn.2316-9079.v17i2p169-180
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
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 (1), 51–71.
Mirza, Z.A. & Sanap, R.V. (2014) A new Cryptic species of Gecko of the genus Hemidactylus Oken, 1817 (Reptilia: Gekkonidae) from Southern India. Taprobanica, 6 (1), 12–20.
Murthy, B.H.C.K., Bauer, A.M., Lajmi, A., Agarwal, I. & Giri, V.B. (2015) A new rock dwelling Hemidactylus (Squamata: Gekkonidae) from Chhattisgarh, India. Zootaxa, 4021 (2), 334–350. https://doi.org/10.11646/zootaxa.4021.2.5
Murthy, T.S.N. (1985) A field guide to the lizards of Western Ghats. Records of Zoological Survey of India, Occasional Papers, 72, 1–51, 14 pls.
Pal, S.P., Gaikwad, K.S., Murthy, C., Dutta, S.K. & Giri, V.B. (2013) New locality records of the recently described gecko Hemidactylus aaronbaueri Giri, 2008 with additional notes on natural history. Hamadryad, 36 (2), 162–167.
Purkayastha, J., Madhurima, D., Bauer, A.M., Sengupta, S. & Dutta, S.K. (2010) Notes on the Hemidactylus bowringii complex (Reptilia: Gekkonidae) in India, and a change to the national herpetofaunal list. Hamadryad, 35, 20–27.
R Core Team (2021) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: https://www.R-project.org/ (accessed 23 February 2022)
Rambaut, A., Suchard, M.A., Xie D. & Drummond, A.J. (2014) Tracer v1.6. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 23 February 2022)
Sanyal, D.P. & Dasgupta, G. (1990) On a collection of reptiles from Bastar District, Madhya Pradesh, central India. Hamadryad, 15, 18–20.
Smith, M.A. (1935) The fauna of British India, including Ceylon and Burma. Reptilia and Amphibia. Vol. II. Sauria. Taylor and Francis, London. xiii + 440 pp., 1 pl., 1 folding map.
Soman, P.W. (1966) A new gekkonid in Maharashtra. Science and Culture, 32, 427.
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. & Das, I. (2008) The herpetofauna of Nallamala Hills, Eastern Ghats, India: an annotated checklist, with remarks on nomenclature, taxonomy, habitat use, adaptive types and biogeography. Asiatic Herpetological Research, 11, 110–131.
Srinivasulu, C. & Srinivasulu, B. (2013) Hemidactylus giganteus. The IUCN Red List of Threatened Species, 2013, e.T172701A1369660. https://doi.org/10.2305/IUCN.UK.2013-1.RLTS.T172701A1369660.en
Srinivasulu, C., Chethan Kumar, G. & 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. & Chethan Kumar, G. (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
Srinivasulu, C., Srinivasulu, B. & Molur, S. (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.
Stoliczka, F. (1871) Notes on new or little known Indian lizards. Proceedings of the Asiatic Society of Bengal, 1871, 192–195.
Stoliczka, F. (1872) Notes on various new or little known Indian lizards. Journal of the Asiatic Society of Bengal, 41, 86–135, pls. II–V.
Suchard, M.A., Lemey, P., Baele, G., Ayres, D.L., Drummond, A.J. & Rambaut, A. (2018) Bayesian phylogenetic and phylodynamic data integration using BEAST 1.10. Virus Evolution, 4, vey016. https://doi.org/10.1093/ve/vey016
Tamura, K., Stecher, G., Peterson, D., Filipski, A., & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology & Evolution, 30 (12), 2725–2729. https://doi.org/10.1093/molbev/mst197
Uetz, P., Freed, P, Aguilar, R. & Hošek, J. (Eds.) (2021) The Reptile Database. Available from http://www.reptile-database.org, accessed (accessed 31st May 2021)
Zhang, J., Kapli, P., Pavlidis, P. & Stamatakis, A. (2013) A general species delimitation method with applications to phylogenetic placements, Bioinformatics, 29 (22), 2869–2876. https://doi.org/10.1093/bioinformatics/btt499