A molecular and morphological study of Otocryptis Wagler, 1830 (Squamata: Agamidae) reveals a new genus from the far south of the Western Ghats, Peninsular India

(original) (raw)

In recent years, several studies have revealed significant unknown and cryptic diversity of agamids in peninsular India, particularly in the Western Ghats. Here, we examine the morphology, anatomy and genetics of the sole Indian representative of the otherwise Sri Lankan agamid genus Otocryptis from the Western Ghats. Our analyses reveal significant distinctions in O. beddomii Boulenger, 1885 with respect to the Sri Lankan members, the type species, O. wiegmanni Wagler, 1830, and O. nigristigma Bahir & de Silva, 2005, warranting a new generic placement. To accommodate the divergent and allopatric O. beddomii from the Western Ghats, we erect a new genus Agasthyagama gen. nov. We re-characterise Agasthyagama beddomii (Boulenger, 1885) comb. nov. based on a syntype (ZSI 15733) and recently collected material. In effect, we restrict the genus Otocryptis, represented by two species, O. wiegmanni and O. nigristigma, to Sri Lanka. We also provide a re-appraisal of the genus Otocryptis sensu stricto, based on data from its type species O. wiegmanni. Our finding adds another endemic agamid genus to the Western Ghats, following Salea Gray, 1845, and the recently described Monilesaurus Pal, Vijayakumar, Shanker, Jayarajan & Deepak, 2018, and Microauris Pal, Vijayakumar, Shanker, Jayarajan & Deepak, 2018_._ In turn, this complements Sri Lankan agamid endemism with Otocryptis, in addition to the accepted endemic radiations of Lyriocephalus Merrem, 1820_, Ceratophora_ Gray, 1835 and Cophotis Peters, 1861. From a systematic perspective, our erection of Agasthyagama gen. nov. likely completes the description of known genus-level diversity in the clade containing Otocryptis Wagler 1830_, Sitana_ Cuvier, 1829 and the recently described Sarada Deepak, Karanth & Giri, 2016.

• 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/10.47605/tapro.v1i1.2
  Arends, J. (2017) Language and Slavery. A Social and Linguistic History of the Suriname Creoles. Creole Language Library 52. John Benjamins Publications, Amsterdam/Philadelphia, 495 pp. https://doi.org/10.1075/cll.52
  Bahir, M.M. & de Silva, A. (2005) Otocryptis nigristigma, a new species of agamid lizard from Sri Lanka. Raffles Bulletin of Zoology, Supplement No. 12, 393–406.
  Bahir, M.M. & Surasinghe, T. (2005) A conservation assessment of the Sri Lankan Agamidae (Reptilia: Sauria). Raffles Bulletin of Zoology, Supplement No. 12, 407–412.
  Bossuyt, F., Meegaskumbura, M., Beenaerts, N., Gower, D.J., Pethiyagoda, R., Roelants, K., Mannaert, A., Wilkinson, M., Bahir, M.M., Manamendra-Arachchi, K. & Ng, P.K. (2004) Local endemism within the Western Ghats-Sri Lanka biodiversity hotspot. Science, 306 (5695), 479–481. https://doi.org/10.1126/science.1100167
  Boulenger, G.A. (1885) Catalogue of the lizards in the British Museum (Nat. Hist.) Vol.-I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. Printed by order of the Trustees, London, xii + 436 pp., 32 pls.
  Boulenger, G.A. (1890) The Fauna of British India, Including Ceylon and Burma. Reptilia and Batrachia. Taylor & Francis, London, xviii + 541 pp. https://doi.org/10.5962/bhl.title.57017
  Chandramouli, S.R. (2009) Status and microhabitat preferences of Otocryptis beddomii Boulenger, 1885 (Reptilia: Agamidae) in Ponmudi hills, Western Ghats, Kerala, India. Taprobanica, 1 (2), 107–110. https://doi.org/10.47605/tapro.v1i2.16
  Daniels, R.J.R. (1991) Ecology and status of little known lizard, Otocryptis beddomii Boulenger. Cobra, 5, 3–4.
  Das, I. (2002) A Photographic guide to Snakes and other Reptiles of India. New Holland Publishers, London, 144 pp.
  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.
  Deepak, V. & Karanth, P. (2018) Aridification driven diversification of fan-throated lizards from the Indian subcontinent. Molecular Phylogenetics and Evolution, 120, 53–62. https://doi.org/10.1016/j.ympev.2017.11.016
  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 the description of one new genus and five new species. Contributions to Zoology, 85 (1), 67–111. https://doi.org/10.1163/18759866-08501004
  Denzer, W., Günther, R. & Manthey, U. (1997) Kommentierter Typenkatalog der Agamen des Museums für Naturkunde der Humbold-Universitt zu Berlin (ehemals Zoologisches Museum Berlin). Mitteilungen aus dem Zoologischen Museum in Berlin, 73, 309–332. https://doi.org/10.1002/mmnz.19970730209
  Edgar, R. (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
  Gray, J.E. (1845) Catalogue of the Specimens of Lizards in the Collection of the British Museum. Printed by the order of the Trustees of the British Museum, London, xxviii + 289 pp.
  Green, E.E. (1902) Bipedal locomotion of a Ceylonese lizard. Nature, 66 (1716), 492–492. https://doi.org/10.1038/066492c0
  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), 1– 11. https://doi.org/10.1186/s12862-016-0611-6
  Gunawardene, N.R., Daniels, A.E., Gunatilleke, I.A.U.N., Gunatilleke, C.V.S., Karunakaran, P.V., Nayak, K.G., Prasad, S., Puyravaud, P., Ramesh, B.R., Subramanian, K.A. & Vasanthy, G. (2007) A brief overview of the Western Ghats-Sri Lanka biodiversity hotspot. Current Science, 93 (11), 1167–1172.
  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 the Bombay Natural History Society, 81 (2), 551–570.
  Jose, J., Ramachandran, K.K. & Nair, P.V. (2007) A rare and little known lizard Otocryptis beddomii from the Myristica swamps of Southern Kerala, India. Herpetological Bulletin, 101, 27–31.
  Kalyaanamoorthy, S., Minh, B., Wong, T., Haeseler, A. & Jermiin, L. (2017) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587–589. https://doi.org/10.1038/nmeth.4285
  Karunarathna, D.S. & Amarasinghe, A.T. (2013) Behavioral ecology and microhabitat use by Lyriocephalus scutatus (Linnaeus, 1758): a monotypic genus in Sri Lanka (Reptilia: Agamidae: Draconinae) with notes on the taxonomy. Russian Journal of Herpetology, 20 (1), 1-15.
  Karunarathna, S., Poyarkov, N.A., Arends, C., Surasinghe, T., Bushuev, A.V., Madawala, M., Gorin, V.A. & de Silva, A. (2020) A new species of the genus Ceratophora Gray, 1835 (Reptilia: Agamidae) from a lowland rainforest in Sri Lanka, with insights on rostral appendage evolution in Sri Lankan agamid lizards. Amphibian & Reptile Conservation, 14 (3), [Taxonomy Section], e259, 103–126.
  Lanfear, R., Calcott, B. & Guindon, S. (2012) Partitionfinder: combined selection of partitioning schemes and substitution model for phylogenetic analyses. Molecular Biology and Evolution, 29 (6), 1695–1701. https://doi.org/10.1093/molbev/mss020
  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
  Maduwage, K. Meegaskumbura, M., de Silva, A. & Pethiyagoda, R. (2008) Phylogenetic implications of hemipenial morphology in Sri Lankan agamid lizards. Current Science, 95 (7), 838–840.
  Maduwage, K. & de Silva, A. (2012) Hemipeneal morphology of Sri Lankan dragon lizards (Sauria: Agamidae). Ceylon Journal of Science, 41 (2), 111–123. https://doi.org/10.4038/cjsbs.v41i2.5381
  Manamendra-Arachchi K & Liyanage S. (1994) Conservation of the agamid lizards of Sri Lanka with illustration of the extant species. Journal of South Asian Natural History, 1, 77–96.
  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 (Supplement 1), 1–8.
  Manthey, U. (2010) Agamid Lizards of Southern Asia. Draconinae 2—Leiolepidinae. Terralog 7b. Edition Chimaira, Frankfurt, 168 pp.
  Minh, B., Nguyen, M. & von Haeseler, A. (2013) Ultrafast Approximation for Phylogenetic Bootstrap. Molecular Biology and Evolution, 30 (5), 1188–1195. https://doi.org/10.1093/molbev/mst024
  Moody, S.M. (1980) Phylogenetic and historical biogeographical relationships of the genera in family Agamidae (Reptilia: Lacertilia). Unpublished PhD Thesis. University Michigan. Michigan.
  Nguyen, L., Schmidt, H., von Haeseler, A. & Minh, B. (2015) IQ-TREE: A fast and stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution, 32, 268–674. https://doi.org/10.1093/molbev/msu300
  Pal, S., Vijayakumar, S.P., Shanker, K., Jayarajan, A. & Deepak, V. (2018) A systematic revision of Calotes Cuvier, 1817 (Squamata: Agamidae) from the Western Ghats adds two genera and reveals two new species. Zootaxa, 4482 (3), 401–450. https://doi.org/10.11646/zootaxa.4482.3.1
  Peters, W. (1864). Über einige neue Säugethiere (Mormops, Macrotus, Vesperus, Molossus, Capromys), Amphibien (Platydactylus, Otocryptis, Euprepes, Ungalia, Dromicus, Tropidonotus, Xenodon, Hylodes), und Fische (Sillago, Sebastes, Channa, Myctophum, Carassius, Barbus, Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin, 1864, 381–399.
  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.
  Rambaut, A., Drummond, A., Xie, D., Baele, G. & Suchard, M. (2018) Posterior Summarization in Bayesian Phylogenetics Using Tracer 1.7. Systematic Biology, 67 (5), 901–904. https://doi.org/10.1093/sysbio/syy032
  Ronquist, F., Teslenko, M., Mark, P., Ayres, D., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M. & Huelsenback, J. (2020) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539–542. https://doi.org/10.1093/sysbio/sys029
  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 (3), 207–214.
  Sambrook, J. & Russel, D. (2001) Molecular Cloning: A Laboratory Manual. 3rd Edition. Cold Spring Harbor Laboratory Press, New York, New York, 2100 pp.
  Smith, M.A. (1935) The fauna of British India, including Ceylon and Burma: Reptilia and Amphibia. Vol. 2. Sauria. Taylor & Francis, London, 440 pp.
  Somaweera, R. & Somaweera, N. (2009) Lizards of Sri Lanka: A Colour Guide with Field Keys. Chimaira, Frankfurt, 304 pp.
  Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution, 30, 2725–2729. https://doi.org/10.1093/molbev/mst197
  Taylor, E.H. (1953) A review of the lizards of Ceylon. University of Kansas Science Bulletin, 35 (12), 1525–1585. https://doi.org/10.5962/bhl.part.26732
  Trifinopoulos, J., Nguyen, L., Haeseler, A. & Minh, B. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acid Research, 44 (W1), W232–W235. https://doi.org/10.1093/nar/gkw256
  Wagler, J.G. (1830) Natürliches System der Amphibien, mit vorangehender Classification der Säugetiere und Vögel. Ein Beitrag zur vergleichenden Zoologie. J.G. Cotta’sche Buchhandlung, München, Stuttgart and Tübingen, 354 pp., 2 foldout pls. https://doi.org/10.5962/bhl.title.108661
  Wang, K., Che, J., Lin, S., Deepak, V., Datta-Roy, A., Jiang, K., Jin, J., Chen, H. & Siler, C. (2018) Multilocus phylogeny and revised classification for mountain dragons of the genus Japalura s.l. (Reptilia: Agamidae: Draconinae) from Asia. Zoological Journal of the Linnean Society, 20, 1–22. https://doi.org/10.1093/zoolinnean/zly034
  Wiegmann, A.F.A. (1831) Zur Amphibienkunde. Isis von Okeni, 24, columns 291–299.