Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2020-11-06
Page range: 1–62
Abstract views: 580
PDF downloaded: 9

Disentangling vines: a study of morphological crypsis and genetic divergence in vine snakes (Squamata: Colubridae: Ahaetulla) with the description of five new species from Peninsular India

Center for ecological Sciences, Indian Institute of Science, Bangalore 560012, Karnataka, India.
Center for ecological Sciences, Indian Institute of Science, Bangalore 560012, Karnataka, India.
Center for ecological Sciences, Indian Institute of Science, Bangalore 560012, Karnataka, India. Bombay Natural History Society, Hornbill House, Opp. Lion Gate, S. B. S. Road, Fort, Mumbai, India, 400001
Center for ecological Sciences, Indian Institute of Science, Bangalore 560012, Karnataka, India.
Center for ecological Sciences, Indian Institute of Science, Bangalore 560012, Karnataka, India.
Chennai Snake Park, Rajbhavan post, Chennai 600 022, Tamil Nadu, India.
Reptilia cryptic species morphology Sri Lanka taxonomy vine snakes Western Ghats

Abstract

We carried out a taxonomic revision of Ahaetulla species inhabiting Peninsular India, using a multiple criteria approach (including genetics, morphology, and geography). Our work included populations of the A. nasuta complex (widespread across the entire region, including the Western Ghats), the A. pulverulenta complex (in the Western Ghats, within Peninsular India) and the A. dispar complex (endemic to the Southern Western Ghats) which all revealed undocumented cryptic diversity. Here, we describe five new species and effect nomenclatural changes to some recognised taxa. In the A. nasuta complex, we describe four species from several latitudinal blocks of the Western Ghats and make nomenclatural emendations to the plains populations in the Indian peninsula. We effect nomenclatural change in the A. pulverulenta population of the Western Ghats and describe a new species from the A. dispar group. Our study highlights the use of a multi-criteria approach in unraveling cryptic diversity. This study also reveals a striking case of discordance between morphological and genetic divergence, and the way this is reflected in previous taxonomic and nomenclatural treatments of these populations.

 

References

  1. 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.

    Annandale, N. (1906) Notes on the fauna of a desert tract in southern India. Part I. Batrachians and reptiles, with remarks on the reptiles of the desert region of the North–West Frontier. Memoirs of the Asiatic Society of Bengal, 1(10), 183–222.

    Arévalo, E., Davis, S.K. & Sites, J.W. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in Central Mexico. Systematic Biology, 43, 387–418.

    https://doi.org/10.1093/sysbio/43.3.387

    Bauer, A.M. (2015) Patrick Russell’s snakes and their role as type specimens. Hamadryad, 37 (1&2), 18–65.

    Bauer, A.M., Vogel, G. & Campbell, P. (2015) A preliminary consideration of the dry snake skin specimens of Patrick Russell. Hamadryad, 37(1&2), 73–84.

    Beddome, R.H. (1862) Notes upon the land and freshwater snakes of the Madras Presidency. Madras Quarterly Journal of Medical Sciences, 5: 1–31.

    Bell, T. (1825) On Leptophina, a group of serpents comprising the Genus Drynus of Merrem, and a newly formed Genus proposed to be named Leptophis. The Zoological Journal, 2, 328. London.

    Bleeker, P. de. (1856) Reis door de Minahassa en den Moluskschen Archipel gedaan in de maanden September en Oktober 1855 in het gevolg van den gouverneur generaal Mr. A.J. Duymaer van Twist. Eerste deel. Lange & Co., Batavia, viii + 288 pp.

    Boie, F. (1826) Bulletin des sciences naturelles et de géologie. Paris, (Ferussac) 9: 237.

    Boie, F. (1827) Bemerkungen über Merrem’s Versuch eines Systems der Amphibien, 1. Lieferung: Ophidier. Isis van Oken 20: 508–566.

    Bonnaterre, P.J. (1790) Ophiologie, in Tableau encyclopédique et méthodique des trois règnes de la nature. Panconoke, Paris. 53 p.

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

    https://doi.org/10.5962/bhl.title.57017

    Boulenger, G.A. (1896) Catalogue of the snakes in the British Museum. 3: Taylor & Francis, London. 193 p.

    Burbrink, F.T., Lawson, R. & Slowinski, J.B. (2000) Mitochondrial DNA phylogeography of the polytypic north American rat snake (Elaphe obsoleta): a critique of the subspecies concept. Evolution, 54, 2107–2118.

    https://doi.org/10.1111/j.0014-3820.2000.tb01253.x

    Chanard, T., Grossmann, W., Gumprecht, A. & Schulz, K.D. (1999) Amphibians and reptiles of Peninsular Malaysia and Thailand – an illustrated checklist [bilingual English and German]. Bushmaster Publications, Würselen, Germany. 240 Pp.

    Chandramouli, S.R. & Ganesh, S.R. (2010) Herpetofauna of Southern Western Ghats, India − reinvestigated after decades. Taprobanica, 2 (2), 72–85.

    https://doi.org/10.4038/tapro.v2i2.3145

    Cherian, P.T., Rema Devi, K. & Ravichandran, M.S. (2000) Ichthyo and Herpetofaunal diversity of Kalakkad Wildlife Sanctuary. Zoo’s Print Journal, 15 (2), 203–206.

    https://doi.org/10.11609/JoTT.ZPJ.15.2.203-6

    Cope, E.D. (1886) An Analytical Table of the Genera of Snakes. Proceedings of the American Philosophical Society 23 (124), 479–499.

    Dalman, J. W. (1823) Analecta Entomologica. Holmiæ. 7 p.

    https://doi.org/10.5962/bhl.title.66069

    Darwin, C. (1871). The Descent of Man and Selection in Relation to Sex. Vol. II, John Murray, UK.

    https://doi.org/10.5962/bhl.title.24784

    Das, I. & Chaturvedi, N. (1998) Catalogue of the herpetological types in the collection of the Bombay Natural History Society. Hamadryad, 23 (2): 150-156.

    Das, I. (2002) A photographic guide to snakes and other reptiles of India. New Holland Publisher, UK Ltd. 144 pp.

    Das, I. (2010) A Field Guide to the Reptiles of South-East Asia. New Holland Publishers, UK Ltd. 260 pp.

    Das, I. & de Silva, A. (2005) A photographic guide to snakes and other reptiles of Sri Lanka. New Holland Publication, UK, 144 pp.

    David, P. & Dubois, A. (2005) Découverte et redescription de l’holotype d’Ahaetulla pulverulenta (Dumérl, Bibron & Duméril, 1854) (Reptilia, Serpentes, Colubridae), avec une remarque sur le statut de Dryinus fuscus Dumérl, Bibron & Duméril, 1854. Zoosystema, 27 (1), 163–178.

    Deepak, V., Harikrishnan, S., Vasudevan, K. & Smith, E.N. (2010) Redescription of Bibron’s coral snake, Calliophis bibroni (Jan 1858) with notes and new records from south of the Palghat and Shencottah gaps of the Western Ghats, India. Hamadryad, 35 (1), 1–10.

    Deepak, V. Narayanan, S., Sarkar, V., Dutta S K. & Mohapatra, P.P. (2019) A New Species of Ahaetulla Link, 1807 (Serpentes: Colubridae: Ahaetullinae) from India. Journal of Natural History, 53(9–10), 497–516.

    https://doi.org/10.1080/00222933.2019.1589591

    De Silva, P.H.D.H. 1969. Taxonomic studies on Ceylon snakes of the family Colubridae. Spolia Zeylanica, 31 (2), 431-546

    Deraniyagala, P.E.P. (1955) A colored atlas of some vertebrates from Ceylon. Vol.3. Serpentoid Reptilia. Colombo.121 pp.

    https://doi.org/10.2307/1439475

    Duméril, A.M.C., Bibron, G. & Dumeril, A.H.A. (1854) Erpétologie générale ou histoire naturelle complète des reptiles. Tome septième. Deuxième partie, comprenant l’histoire des serpents venimeux. Paris, Librairie Encyclopédique de Roret: i-xii + 781-1536. https://doi.org/10.5962/bhl.title.118790

    Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797.

    https://doi.org/10.1093/nar/gkh340

    Figueroa, A., McKelvy, A.D., Grismer, L.L., Bell, C.D. & Lailvaux S.P. (2016) A Species-Level Phylogeny of Extant Snakes with Description of a New Colubrid Subfamily and Genus. PLoS ONE, 11(9), e0161070.

    https://doi.org/10.1371/journal.pone.0161070

    Fitzinger, L.J.F.J. (1843) Systema Reptilium. Fasciculus primus. Amblyglossae. Braumüller et Seidel, Wien. 27 p.

    Fujisawa, T. & Barraclough T.G. (2013) Delimiting species using single-locus data and the generalized mixed Yule coalescent approach: a revised method and evaluation on simulated data sets. Systematic Biology, 62, 707–724.

    https://doi.org/10.1093/sysbio/syt033

    Ganesh, S. R. & Chandramouli, S.R. (2011) On the nomenclature and taxonomy of the south Indian colubrid snake Ahaetulla perroteti (Duméril, Bibron & Duméril, 1854). Herpetological Bulletin, (117), 19–24.

    Gaulke, M. (1994) Eine neue Unterart des Malaysischen Baumschnüfflers, Ahaetulla prasina suluensis n. subsp. (Reptilia: Serpentes: Colubridae). Senckenbergiana Biologica, 73 (1&2), 45–47.

    Gower, D.J. & Winkler, J.D. (2007) Taxonomy of the Indian snake Xylophis Beddome (Serpentes: Caenophidia), with description of a new species. Hamadryad, 31 (2), 315–329.

    Gower, D.J. & Wickramasinghe L.J.M. (2016) Recharacterization of Rhinophis dorsimaculatus Deraniyagala, 1941 (Serpentes: Uropeltidae), including description of new material. Zootaxa, 4158 (2), 203–212.

    https://doi.org/10.11646/zootaxa.4158.2.3

    Gray, J. E. (1825) A synopsis of the genera of reptiles and Amphibia, with a description of some new species. Annals of Philosophy, 10, 193–217.

    Günther, A.C.L.G. (1858) Catalogue of Colubrine snakes of the British Museum. London, I – XVI, 1 – 281.

    Günther, A.C.L.G. (1864) The Reptiles of British India. London (Taylor & Francis), xxvii + 452 pp.

    Gutell, R.R. (1994) Collection of small subunit (16S- and 16S-like) ribosomal RNA structures. Nucleic Acids Research, 22, 3502–3507.

    https://doi.org/10.1093/nar/22.17.3502

    Hutton, A.F. & David, P. (2009) Notes on a collection of snakes from South India, with emphasis on the snake fauna of Meghamalai hills (High Wavy Mountains). Journal of the Bombay Natural History Society 105 (3), 299–316.

    Ishwar, N.M., Chellam, R. & Kumar, A. (2001) Distribution of forest floor reptiles in the rainforest of Kalakad-Mundanthurai Tiger Reserve, South India. Current Science, 80 (3), 413–418.

    Jan, G. (1863) Elenco sistematico degli ofidi descritti e disegnati per l’Iconografia générale. A. Lombardi, Milano, vii + 143 pp. https://doi.org/10.5962/bhl.title.106683

    Jerdon, T.C. (1854) Catalogue of the Reptiles inhabiting the Peninsula of India. Part 2. Journal of the Asiatic Society of Bengal, XXII: 522–534.

    Jins, V.J., Bhupathy, S. & Panigrahi, M. (2014) New record of Beddome’s Coral Snake, Calliophis beddomei Smith 1943 from the Southern Western Ghats, India. Herpetology Notes, 7, 555–557.

    Kjer, K.M. (1995) Use of rRNA secondary structure in phylogenetic studies to identify homologous positions: an example of alignment and data presentation from the frogs. Molecular Phylogenetics and Evolution, 4, 314–330.

    https://doi.org/10.1006/mpev.1995.1028

    Lacepède, B.G.E. (1789) Histoire Naturelle des Quadrupèdes Ovipares et de Serpens. Vol.2. lmprimerie du Roi, Hôtel de Thou, Paris, 671 pp. https://doi.org/10.5962/bhl.title.50642

    Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29(6), 1695–1701.

    https://doi.org/10.1093/molbev/mss020

    Lazell, J.D. (2002) The herpetofauna of Shek Kwu Chau, South Chinese Sea, with descriptions of two new colubrid snakes. Memoirs of the Hong Kong Natural History Society, 25, 82.

    Link, H.F. (1807) Beschreibung der Naturalien–Sammlung der Universität zu Rostock, Vol. 2. Rostock: Adlers Erben.

    https://doi.org/10.5962/bhl.title.69815

    Linnaeus, C. (1758) Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiæ. 225 pp.

    https://doi.org/10.5962/bhl.title.542

    Luo, A., Lan, H., Ling, C., Zhang, A., Shi, L., Ho, S.Y.W. & Zhu, C. (2018) A simulation study of sample size for DNA barcoding. Ecology and Evolution, 5, 5869–5879.

    https://doi.org/10.1002/ece3.1846

    Malhotra, A & Davis, K. (1991) A report on the herpetological survey of the Srivilliputhur Reserve Forests, Tamil Nadu. Journal of the Bombay Natural History Society, 88, 2, 157–166.

    Mallik, A.K., Achyuthan N.S., Ganesh, S.R., Pal, S.P., Vijayakumar, S.P. & Shanker, K. (2019) Discovery of a deeply divergent new lineage of vine snake (Colubridae: Ahaetuliinae: Proahaetulla gen. nov.) from the Southern Western Ghats of Peninsular India with a revised key for Ahaetuliinae. PLoS ONE, 14(7), e0218851.

    https://doi.org/10.1371/journal.pone.0218851

    Meise, W. & Hennig, W. (1932) Die Schlangengattung Dendrophis. Zoologischer Anzeiger, 99 (11/12), 273-297.

    Mell, R. (1931) List of Chinese snakes. Lingnan Science Journal, Canton (1929) 8, 119–219.

    McCann, C. (1940). Extension of the range of the brown whip snake (Dryophis pulverulentus Jan.). Journal of the Bombay Natural History Society, 42, 200.

    Merrem, B. (1820) Versuch eines Systems der Amphibien we (Tentamen Systematis Amphibiorum). J. C. Kriegeri, Marburg. 191 pp.

    https://doi.org/10.5962/bhl.title.5037

    Miralles, A. & David, P. (2010) First record of Ahaetulla mycterizans (Linnaeus, 1758) (Reptilia, Squamata, Colubridae) from Sumatra, Indonesia, with an expanded definition. Zoosystema, 32 (3), 449–456.

    https://doi.org/10.5252/z2010n3a6

    Mohapatra, P.P., Dutta, S.K., Kar, N.B., Das, A., Murthy, B.H.C.K., & Deepak, V. (2017) Ahaetulla nasuta anomala (Annandale, 1906) (Squamata: Colubridae), resurrected as a valid species with marked sexual dichromatism. Zootaxa, 4263 (2), 318–332.

    https://doi.org/10.11646/zootaxa.4290.3.13

    Mukherjee, D. & Bhupathy, S. (2007) A new species of wolf snake (Serpentes: Colubridae: Lycodon) from Anaikatti Hills, Western Ghats, Tamil Nadu, India. Russian Journal of Herpetology, 14 (1), 21–26.

    Palumbi, S.R. (1996) Nucleic acids II: the polymerase chain reaction. In: Hillis, D.M., Moritz, C., Mable, B.K. (Eds.). Molecular Systematics. USA: Massachusetts, Sunderland; Sinauer Associates, Inc.

    Parkinson, C.L. (1999) Molecular systematics and biogeographical history of the pitvipers as determined by mitochondrial ribosomal sequences. Copeia 1999 (3), 576–586

    https://doi.org/10.2307/1447591.

    Puillandre, N., Lambert, A., Brouillet, S. & Achaz, G. (2012) ABGD, automatic barcode gap discovery for primary species delimitation. Molecular Ecology, 21, 1864–1877.

    https://doi.org/10.1111/j.1365-294X.2011.05239.x

    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

    Rao, K.T., Ghate, H.V., Sudhakar, A.M., Javed, S.M. & Krishna, I.S.R. (2005) Herpetofauna of Nallamalai Hills with eleven new records for the region including ten new records for Andhra Pradesh. Zoos’ Print Journal, 20(1), 1737-1740. https://doi.org/10.11609/JoTT.ZPJ.1232.1737-40.

    Sambrook, J., Fritsch, E.F., & Maniatis, T. (1989) Molecular Cloning: A Laboratory Manual. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York, USA.

    Savage, J.M. (1952) Two centuries of confusion: the history of the snake name Ahaetulla. Bulletin of the Chicago Academy of Sciences, 9 (11), 203–216.

    https://doi.org/10.1093/nq/s9-I.11.216h

    Savage, J.M. & Oliver, J.A. (1956) Proposed addition to the” Official List of Generic Names in Zoology” of” Ahaetulla” Link, 1807, with” Ahaetulla mycterizans” Link, 1807, as the type species (Class Reptilia). Bulletin of Zoological Nomenclature, 12, 147-152.

    Schinz, H.R. (1833) Naturgeschichte und Abbildungen der Reptilien. Brodtmann’s lithographischer Anstalt, Schasshausen, iv + 240 pp., 102 pls.

    Shanker, K., Ganeshaiah, K.N. and Vijayakumar, S.P. (2017) Unpacking the species conundrum: philosophy, practice and ways forward. Journal of Genetics 96, 413-430.

    Sharma, R.C. (1976) Records of the reptiles of Goa. Records of the Zoological Survey of India, 71, 149–167.

    Shaw, G. (1802) General Zoology, or Systematic Natural History. Vol.3, part 2. G. Kearsley, Thomas Davison, London, UK.

    Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML. Organisms Diversity & Evolution, 12 (4), 335–337.

    https://doi.org/10.1007/s13127-011-0056-0

    Smith, M.A. (1943) The Fauna of British India, Ceylon and Burma, including the whole of the Indo–Chinese Subregion. Reptilia and Amphibia. Vol. 3. Serpentes. Taylor. & Francis, London. 584 pp.

    Smith, E. N., Ogale, H., Deepak, V. & Giri, V.B. (2012) A new species of coralsnake of the genus Calliophis (Squamata: Elapidae) from the west coast of Peninsular India. Zootaxa, 3437, 51–68.

    Somaweera, R. (2006) Sri Lankawe Sarpayin (‘The snakes of Sri Lanka’). Colombo, Wildlife Heritage Trust of Sri Lanka. (in Sinhala).

    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: 1069 molecular evolutionary genetics analysis using maximum likelihood, 1070 evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution, 28 (1071), 2731–2739.

    https://doi.org/10.1093/molbev/msr121

    Taylor, E.H. (1965) The serpents of Thailand and adjacent waters. Kansas University Science Bulletin, 45(9), 609-1096.

    Theobald, W. (1868) Catalogue of reptiles in the Museum of the Asiatic Society of Bengal, 1866. Journal of the Asiatic Society of Bengal (Extra number), 37, i–vi + 1–88 pp. + i–iii.

    Uetz, P., Cherikh, S., Shea, G., Ineich, I., Campbell, P.D., Doronin, I.V., Rosado, J., Wynn, A., Tighe, K.A., Mcdiarmid, R., Lee, J.L., Köhler, G., Ellis, R., Doughty, P., Raxworthy, C.J., Scheinberg, L., Resetar, A., Sabaj, M., Schneider, G., Franzen, M., Glaw, F., Böhme, W., Schweiger, S., Gemel, R., Couper, P., Amey, A., Dondorp, E., Ofer, G., Meiri, S. & Wallach, V. (2019) A global catalog of primary reptile type specimens. Zootaxa, 4695(5), 438-450.

    Varma, V. & Gharat, G. (2019) An unusual morph in Green Vine Snake, Ahaetulla nasuta spp. (Lacépède, 1789) (Serpentes: Colubridae: Ahaetullinae) from Matheran, Maharashtra, India. Records of the Zoological Survey of India, 119(1), 88–90.

    Vogel, G. & Rooijen, J.V. (2011a) A new species of Dendrelaphis (Serpentes: Colubridae) from the Western Ghats – India. Taprobanica, 3 (2), 77–85.

    https://doi.org/10.4038/tapro.v3i2.3963

    Vogel, G. and Rooijen, J.V. (2011b) Contributions to a Review of the Dendrelaphis pictus (Gmelin, 1789) Complex (Serpentes: Colubridae)–3. The Indian Forms, with the description of a new species from the Western Ghats. Journal of Herpetology, 45, (1), 100–110.

    https://doi.org/10.1670/09-035.1

    Vogel, G. & Ganesh, S.R. (2013) A new species of cat snake (Reptilia: Serpentes: Colubridae: Boiga) from dry forests of eastern peninsular India. Zootaxa, 3637 (2), 158–168.

    https://doi.org/10.11646/zootaxa.3637.2.6

    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, 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, 283 (1836), RSBP 20161011.

    https://doi.org/10.1098/rspb.2016.1011

    Wagler, J.G. (1830) Natürliches System der Amphibien, mit vorangehender Classification der Säugethiere und Vögel. Ein Beitrag zur vergleichenden Zoologie. J. G. Cotta Buchhandlung, München. 184 pp.

    https://doi.org/10.5962/bhl.title.108661

    Wall, F. (1908a) Remarks on some recently acquired snakes. Journal of the Bombay Natural History Society, 18, 778–784.

    Wall, F. (1908b) A new colour variety of the common green whip-snake (Dryophis mycterizans). Journal of the Bombay Natural History Society, 18, 919.

    Wall, F. (1910a) Remarks on the varieties and distribution of the common green whip snake (Dryophis mycterizans). Journal of the Bombay Natural History Society, 20 (1), 229–230.

    Wall, F. (1910b) Varieties of the common green whip snake (Dryophis mycterizans). Journal of the Bombay Natural History Society, 20, 524.

    Wall, F. (1919) Notes on a collection of snakes made in the Nilgiri hills and adjacent Wynaad. Journal of the Bombay Natural History Society, 26, 552–584.

    Wall, F. (1921) Ophidia Taprobanica or the Snakes of Ceylon, Colombo Museum, Colombo. 291 pp.

    https://doi.org/10.5962/bhl.title.54947

    Wallach, V., Williams, K.L. & Boundy, J. (2014) Snakes of the World: A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, Florida, 1237 pp.

    https://doi.org/10.1201/b16901

    Wang, C., Agrawal, S., Laudien, J., Häussermann, V. & Held, C. (2016) Discrete phenotypes are not underpinned by genome–wide genetic differentiation in the squat lobster Munida gregaria (Crustacea: Decapoda: Munididae): a multimarker study covering the Patagonian shelf. BMC Evolutionary Biology, 16: 258.

    https://doi.org/10.1186/s12862-016-0836-4

    Whitaker, R. & Captain, A. (2004) Snakes of India–the Field Guide. Draco Books, Chengalpattu, India. 495 pp.

    Wiens, J.J. (2007) Species Delimitation: New Approaches for Discovering Diversity. Systematic Biology, 56 (6), 875–878.

    https://doi.org/10.1186/s12862-016-0836-4

    Yang, Z. (2015) The BPP program for species tree estimation and species delimitation. Current Zoology, 61, 854–865.

    https://doi.org/10.1093/czoolo/61.5.854

    Zhang, J., 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

    Zheng, Y. & Wiens, J.J. (2016) Combining phylogenomic and supermatrix approaches, and a time–calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species. Molecular Phylogenetics and Evolution, 94, 537–547.

    https://doi.org/10.1016/j.ympev.2015.10.009