A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan

  • Rafaella C. Garbin Department of Geosciences, University of Fribourg, Fribourg; Chemin du Musée 6, 1700 Fribourg
  • Saswati Bandyopadhyay Geological Studies Unit, Indian Statistical Institute, Kolkata; Barrackpore Trunk Road 233, Kolkata
  • Walter G. Joyce Department of Geosciences, University of Fribourg, Fribourg; Chemin du Musée 6, 1700 Fribourg
Keywords: paleontology, Neogene, taxonomy, Siwaliks, turtles

Abstract

Neogene (Siwalik-aged) deposits from India and Pakistan have yielded many vertebrate fossils, of which most were named during the 19th century, including numerous geoemydid turtles. In contrast to many other faunal components from the Siwaliks, geoemydids have not undergone taxonomic revision for more than a century and most fossils have therefore been believed to correspond to recent taxa. In this study, we conduct a taxonomic revision of all previously described geoemydid material from the Siwalik-age. We propose that all specimens of ‘Clemmys’ from the Siwaliks of Punjab, Pakistan should be identified as Melanochelys sivalensis comb. nov.; that Melanochelys tricarinata var. sivalensis represents a valid species, for which we propose the replacement name Melanochelys tapani to avoid homonymy; that specimens originally identified as Batagur cautleyi and Pangshura flaviventer cannot be identified beyond the generic level; and that many fragmentary palatochelydians cannot be identified to any particular species or genus due to the lack of preserved diagnostic osteological characters. With a few exceptions, the Siwalik fauna mostly corresponds in its distribution to that of the recent fauna, indicating a certain amount of geographic stasis. However, as the stratigraphic provenance of most material is poor, it is not possible to discern meaningful temporal patterns.

References

Anquetin J. & Joyce W.G. 2014. A reassessment of the Late Jurassic turtle Eurysternum wagleri (Eucryptodira, Eurysternidae). Journal of Vertebrate Paleontology 34: 1317–1328. https://doi.org/10.1080/02724634.2014.880449

Anderson J.A. 1879. Anatomical and Zoological Researches: Comprising an Account of the Zoological Results of the two Expeditions to Western Yunnan in 1868 and 1875; and a Monograph of the two Cetacean Genera, Platanista and Orcella. Bernard Quaritch, London.

Badam G.L. 1979. Pleistocene Fauna of India: with Special Reference to the Siwaliks. Deccan College, Pune.

Barry J.C., Behrensmeyer A.K., Badgley C.E., Flynn L.J., Peltonen H., Cheema I.U., Pilbeam D., Lindsay E.H., Raza S.M., Rajpar A.R. & Morgan M.E. 2013. The Neogene Siwaliks of the Potwar Plateau, Pakistan. In: Wang X., Flynn L.J. & Fortelius M. (eds) Fossil Mammals of Asia: Neogene biostratigraphy and chronology: 373–399. Columbia University Press, New York.

Boulenger G.A. 1889. Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles in the British Museum (Natural History). Trustees of the British Museum (Natural History), London.

Boulenger G.A. 1890. Reptilia and Batrachia. In: Blanford W.T. (ed.) The Fauna of British India, Including Ceylon and Burma. Taylor and Francis, London.

Chakraborty T., Taral S., More S. & Bera S. 2020. Cenozoic Himalayan foreland basin: an overview and regional perspective of the evolving sedimentary succession. In: Gupta N. & Tandon S.K. (eds) Geodynamics of the Indian Plate: Evolutionary Perspectives. Elsevier Publishing Company. https://doi.org/10.1007/978-3-030-15989-4_11

Chirouze F., Dupont-Nivet G., Huyghe P., van der Beek P., Chakraborty T., Bernet M. & Erens V. 2012. Magnetostratigraphy of the Neogene Siwalik Group in the far eastern Himalaya: Kameng section, Arunachal Pradesh, India. Journal of Asian Earth Sciences 44: 117–135. https://doi.org/10.1016/j.jseaes.2011.05.016

Das I. 1991. The taxonomic status of the Pleistocene turtle Geoclemys sivalensis. Journal of Herpetology 25 (1): 104–107. https://doi.org/10.2307/1564804

Das I. 1994. The identity of the Plio-Pleistocene turtle, Geoemyda pilgrimi Prasad and Satsangi, 1967 (Testudines: Cryptodira: Bataguridae). Hamadryad 19: 41–46.

Das I. & Bhupathy S. 2009a. Hardella thurjii (Gray 1831) – Crowned River Turtle. Chelonian Research Monographs 5: 023.1–023.6. https://doi.org/10.3854/crm.5.023.thurjii.v1.2009

Das I. & Bhupathy S. 2009b. Melanochelys trijuga (Schweigger 1812) – Indian Black Turtle. Chelonian Research Monographs 5: 038.1–038.9. https://doi.org/10.3854/crm.5.038.trijuga.v1.2009

Das I. & Bhupathy S. 2010. Geoclemys hamiltonii (Gray 1830) – Spotted Pond Turtle, Black Pond Turtle. Chelonian Research Monographs 5: 043.1–043.6. https://doi.org/10.3854/crm.5.043.hamiltonii.v1.2010

Duméril A.M.C. & Bibron G. 1835. Erpétologie générale ou Histoire naturelle complète des Reptiles – II. Chéloniens. Librairie encyclopédique de Roret, Paris.

Ernst C.H. & Barbour R.W. 1989. Turtles of the World. Smithsonian Institution Press, Washington D.C.

Falconer H. & Cautley P.T. 1837. On additional fossil species of the order Quadrumana from the Siwalik Hills. Journal of the Asiatic Society of Bengal 6: 354–360.

Falconer H. & Cautley P.T. 1844. Communication on the Colossochelys atlas, a fossil tortoise of enormous size from the Tertiary strata of the Siwalik Hills in the north of India. Proceedings of the Zoological Society of London 1844 (12): 54–84.

Falconer H. & Cautley P.T. 1846. Fauna Antiqua Sivalensis, Being the Fossil Zoology of the Sewalik Hills in the North of India. Proboscidea. Smith, Elder and Co, London. https://doi.org/10.5962/bhl.title.61447

Ferreira G.S., Bandyopadhyay S. & Joyce W.G. 2018. A taxonomic reassessment of Piramys auffenbergi, a neglected turtle from the late Miocene of Piram Island, Gujarat, India. PeerJ 6: e5938. https://doi.org/10.7717/peerj.5938

Fitzinger L.J. 1835. Entwurf einer systematischen Anordnung der Schildkröten nach den Grundsätzen der natürlichen Methode. Annalen des Wiener Museums der Naturgeschichte 1: 105–128.

Flynn L.J, Lindsay E.H., Pilbeam D., Raza S.M., Morgan M.E., Barry J.C., Badgley C.E., Behrensmeyer A.K., Cheema I.U., Rajpar A.R. & Opdyke N.D. 2013. The Siwaliks and Neogene evolutionary biology in South Asia. In: Wang X., Flynn L.J. & Fortelius M. (eds) Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology: 353–372. Columbia University Press, New York. https://doi.org/10.7312/wang15012-014

Gaffney E.S. 1975. A phylogeny and classification of the higher categories of turtles. Bulletin of the American Museum of Natural History 155 (5): 387–436.

Gaffney E.S. & Meylan P.A. 1988. A phylogeny of turtles. In: M.J. Benton (ed.) The Phylogeny and Classification of the Tetrapods Vol I. Amphibians, Reptiles and Birds: 157–219. Clarendon Press, Oxford.

Garbin R.C., Ascarrunz E. & Joyce W.G. 2018. Polymorphic characters in the reconstruction of the phylogeny of geoemydid turtles. Zoological Journal of the Linnean Society 184: 896–918. https://doi.org/10.1093/zoolinnean/zlx106

Garbin R.C., Böhme M. & Joyce W.G. 2019. A new testudinoid turtle from the Middle to Late Eocene of Vietnam. PeerJ 7: e6280. https://doi.org/10.7717/peerj.6280

Gray J.E. 1830. A synopsis of the species of class Reptilia. In: Griffith E. & Pidgeon E. (eds) The Class Reptilia Arranged by the Baron Cuvier, with Specific Descriptions. In: Griffith E. (ed.) The Animal Kingdom Arranged in Conformity with is Organization by the Baron Cuvier, with Additional Descriptions of all the Species Hitherto Named, and of Many not Before noticed. Vol. 9. Reptilia. Whittaker, Treacher and Co, London.

Gray J.E. 1831a. Synopsys Reptilium: or Short Descriptions of the Species of the Reptiles. Part I – Cataphracta. Tortoises, Crocodiles, and Enaliosaurus. Treuttel, Wurz and Co, London.

Gray J.E. 1831b. Illustrations of Indian Zoology, Chiefly Selected from the Collection of Major-General Hardwicke. Vol I, part V. Treuttel, Wurtz, Treuttel jun. and Richter, London. https://doi.org/10.5962/bhl.title.95127

Gray J.E. 1832. Illustrations of Indian Zoology, Chiefly Selected from the Collection of Major-General Hardwicke. Vol II, part XI. Adolphus Richter and Co., London. https://doi.org/10.5962/bhl.title.95127

Gray J.E. 1844. Catalogue of the Tortoises, Crocodiles and Amphisbaenians, in the Collection of the British Museum. Order of the Trustees, London. https://doi.org/10.5962/bhl.title.21639

Gray J.E. 1856 (“1855”). Catalogue of Shield Reptiles in the Collection of the British Museum, Part I – Testudinata (Tortoises). Order of the Trustees, London. https://doi.org/10.5962/bhl.title.5491

Gray J.E. 1862. Notice of a new species of Batagur from North-Western India. Proceedings of the Zoological Society of London 1863: 253.

Gray J.E. 1869. Notes on the families and genera of tortoises (Testudinata), and on the characters afforded by the study of their skulls. Proceedings of the Zoological Society of London 1869: 165–225. https://doi.org/10.1111/j.1469-7998.1869.tb07312.x

Gray J.E. 1870. Supplement to the Catalogue of Shield Reptiles in the Collection of the British Museum, Part I – Testudinata (Tortoises) with figures of the skulls of 36 genera. Order of the Trustees, London.

Günther A.C.L.G. 1864. The Reptiles of British India. Arment Biological Press, Landsville, PA. https://doi.org/10.5962/bhl.title.5012

International Commission on Zoological Nomenclature (ICZN). 1999. International Code of Zoological Nomenclature, Fourth Edition: Adopted by the International Union of Biological Sciences. The International Trust for Zoological Nomenclature, London.

Iverson J.B. 1992. A Revised Checklist with Distribution Maps of the Turtles of the World. Earlham College, privately printed, Richmond.

Joyce W.G. & Lyson T.R. 2010. Pangshura tatrotia, a new species of pond turtle (Testudinoidea) from the Pliocene Siwaliks of Pakistan. Journal of Systematic Palaeontology 8 (3): 449–458. https://doi.org/10.1080/14772019.2010.500879

Joyce W.G., Parham J.F. & Gauthier J.A. 2004. Developing a protocol for the conversion of rank-based taxon names to phylogenetically defined clade names, as exemplified by turtles. Journal of Paleontology 78 (5): 989–1013.

Le M., McCord W.P. & Iverson J.B. 2007. On the paraphyly of the genus Kachuga (Testudines: Geoemydidae). Molecular Phylogenetics and Evolution 45: 398–404. https://doi.org/10.1016/j.ympev.2007.05.002

Lesson R.P. 1831. Catalogue des reptiles qui font partie d`une collection zoologique recueillie dans l`Inde continentale ou en Afrique, et apportée en Franc par M. Lamare-Piquot. Bulletin des Sciences naturelles et Géologie de Paris 25: 119–123.

Lydekker R. 1880. Sketch of the history of the fossil vertebrata of India. Journal of the Asiatic Society of Bengal 49 (2): 8–130.

Lydekker R. 1885a. Indian Tertiary and post-Tertiary Vertebrata, Part III – Siwalik and Narbada Chelonia. Memoirs of the Geological Survey of India, Palaeontologia Indica 10 (3): 155–208.

Lydekker R. 1885b. Catalogue of the Fossil Mammalia in the British Museum (Natural History), Part I – Primates, Chiroptera, Insectivora, Carnivora and Rodentia. Order of the Trustees, London. Available from https://biodiversitylibrary.org/page/17771100 [accessed 11 Feb. 2020].

Lydekker R. 1885c. Catalogue of the Fossil Mammalia in the British Museum (Natural History), Part II – Ungulata: Artiodactyla. Order of the Trustees, London. Available from https://biodiversitylibrary.org/page/17771369 [accessed 11 Feb. 2020].

Lydekker R. 1886a. On a new Emydine chelonian from the Pliocene of India. Quarterly Journal of the Geological Society 42: 540–541. https://doi.org/10.1144/gsl.jgs.1886.042.01-04.54

Lydekker R. 1886b. Catalogue of the Fossil Mammalia in the British Museum (Natural History), Part III – Ungulata: Perissodactyla, Toxodontia, Condylarthra, and Amblypoda. Order of the Trustees, London. Available from https://biodiversitylibrary.org/page/17771723 [accessed 11 Feb. 2020].

Lydekker R. 1886c. Catalogue of the Fossil Mammalia in the British Museum (Natural History), Part IV – Ungulata: Proboscidea. Order of the Trustees, London. Available from https://biodiversitylibrary.org/page/17771933 [accessed 11 Feb. 2020].

Lydekker R. 1887. Indian Tertiary and post-Tertiary Vertebrata, Part I – Siwalik Mammalia. Memoirs of the Geological Survey of India, Palaeontologia Indica 10 (4):1–22.

Lydekker R. 1889a. Notes on Siwalik and Narbada Chelonia. Records of the Geological Survey of India 22 (1): 56–59.

Lydekker R. 1889b. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History). Part III. Chelonia. Order of the Trustees, London.

McDowell S.B. 1964. Partition of the genus Clemmys and related problems in the taxonomy of the aquatic Testudinidae. Proceedings of the Zoological Society of London 143: 239–279. https://doi.org/10.1111/j.1469-7998.1964.tb03860.x

Mertens R. 1969. Eine neue Rasse der Dachschildkröte, Kachuga tecta. Senckenbergiana Biologica 50: 23-30.

Murchinson C. 1868. Biographical sketch. In: Murchinson C. (ed.) Palaeontological Memoirs and Notes of the Late Hugh Falconer, A.M., M.D. Vol I. Fauna Antiqua Sivalensis: XXIII–LIII. Robert Hardwicke, London.

Murray J.A. 1884a. Additions to the reptilian fauna of Sind. The Annals and Magazine of Natural History 14: 106–111. https://doi.org/10.1080/00222938409459777

Murray J.A. 1884b. The Vertebrate Zoology of Sind – a Systematic Account, with Descriptions of all the Known Species of Mammals, Birds, and Reptiles Inhabiting the Province; Observations of their Habits; Tables of their Geographicl Distribution in Persia, Beloochistan, and Afghanistan; Punjab, North-West Provinces, and the Peninsula of India Generally. Education Society’s Press, Bombay. https://doi.org/10.5962/bhl.title.57275

Nanda A.C. 2013. Upper Siwalik mammalian faunas of the Himalayan foothills. Journal of the Palaeontological Society of India 58 (1):75–86.

Nanda A.C., Schleich H.H. & Kotlia B.S. 2016. New fossil reptile records from the Siwalik of North India. Open Journal of Geology 6: 673–691. https://doi.org/10.4236/ojg.2016.68052

Nanda A.C., Sehgal R.K. & Chauhan P.R. 2018. Siwalik-age faunas from the Himalayan Foreland Basin of South Asia. Journal of Asian Earth Sciences 162: 54–68. https://doi.org/10.1016/j.jseaes.2017.10.035

Obst F.J. 1985. Die Welt der Schildkröten. Verlag A. Müller, Rüchslikon-Zürich, Stuttgart, Vienna.

Pereira A.G., Sterli J., Moreira F.R.R. & Schrago C.G. 2017. Multilocus phylogeny and statistical biogeography clarify the evolutionary history of major lineages of turtles. Molecular Phylogenetics and Evolution 113: 59–66. https://doi.org/10.1016/j.ympev.2017.05.008

Pilgrim G.E. 1910. Preliminary note on a revised classification of the Tertiary freshwater deposits of India. Records of the Geological Survey of India 40: 185–205.

Prasad K.N. 1974. The vertebrate fauna from Piram Island, Gujarat, India. Memoirs of the Geological Survey of India, New Series 41: 1–23.

Prasad K.N. & Satsangi P.P. 1967. On a new fossil chelonian from the Siwalik beds of Himachal Pradesh. Records of the Geological Survey of India 95: 533–536.

Praschag P., Hundsdorfer A.K. & Fritz U. 2007. Phylogeny and taxonomy of endangered South and South-east Asian freshwater turtles elucidated by mtDNA sequence variaton (Testudines: Geoemydidae: Batagur, Callagur, Hardella, Kachuga, Pangshura). Zoologica Scripta 36: 429–442. https://doi.org/10.1111/j.1463-6409.2007.00293.x

Praschag P., Sommer R.S., McCarthy C., Gemel R. & Fritz U. 2008. Naming one of the world’s rarest chelonians, the southern Batagur. Zootaxa 1758: 61–68. https://doi.org/10.11646/zootaxa.1758.1.4

Pritchard P.C.H. 1979. Encyclopedia of Turtles. T.F.H. Publications, New Jersey.

Siebenrock F. 1909. Synopsis der rezenten Schildkröten, mit Berücksichtigung der in historischer Zeit ausgestorbenen Arten. Zoologische Jahrbücher Suppl. 10 (3): 427–618. https://doi.org/10.5962/bhl.title.12485

Smith M.A. 1931. The Fauna of British India, Including Ceylon and Burma, Reptilia and Amphibia. Vol. I – Loricata, Testudines. Taylor and Francis, London. https://doi.org/10.2307/1436417

Sonakia A. & Biswas S. 2011. Quaternary geology of the central Narmada valley. Memoirs of the Geological Survey of India, Palaeontologia Indica 53: 3­­­–7.

Strauch A. 1862. Chelonologische Studien, mit besonderer Beziehung auf die Schildkrötensammlung der Kaiserlichen Akademie der Wissenschaften zu St.-Petersburg. Mémoires de l’Académie impériale des Sciences de St.-Pétersbourg, 7ième série 5 (7): 1–197.

Strauch A. 1865. Die Vertheilung der Schildkröten über den Erdball – ein Zoogeographischer Versuch. Mémoires de l’Académie impériale des Sciences de St.-Pétersbourg, 7ième série 8 (13): 1–207. https://doi.org/10.5962/bhl.title.12477

Tewari B.S. & Badam G.L. 1969. A new species of fossil turtle from the Upper Siwaliks of Pinjore, India. Palaeontology 12: 555–558.

Theobald W. 1860. On the tertiary and alluvial deposits of the central portion of the Nerbudda valley. Memoirs of the Geological Survey of India 2: 279–298.

Theobald W. 1868. Catalogue of reptiles in the Museum of the Asiatic Society of Bengal. Journal of the Asiatic Society of Bengal, Extra Number. Baptist Mission Press, Kolkata. https://doi.org/10.5962/bhl.title.5477

Theobald W. 1876. Descriptive Catalogue of the Reptiles of British India. Thacker, Spink and Co, Kolkata. https://doi.org/10.5962/bhl.title.54001

Theobald W. 1877. Description of a new Emydine from the Upper Tertiaries of the Northern Punjab. Records of the Geological Survey of India 10: 43–45.

Tiwari M.P. 2007. Correlation of lithostratigraphy and chronology of the Narmada Valley Quaternary. In: Sankhyan A.R. & Rao V.R. (eds) Human Origins, Genome & People of India: 165–174. Anthropological Survey of India, Kolkata.

Tiwari M.P. & Bhai H.Y.1997. Quaternary stratigraphy of Narmada Valley. Geological Survey of India - Special Publication 46: 33­–63.

Turtle Extinctions Working Group (TEWG). 2015. Turtles and tortoises of the world during the rise and global spread of humanity: first checklist and review of extinct Pleistocene and Holocene chelonians. Chelonian Research Monographs 5: 1–66. https://doi.org/10.3854/crm.5.000e.fossil.checklist.v1.2015

Turtle Taxonomy Working Group (TTWG). 2017. Turtles of the World: Annotated checklist and atlas of taxonomy, synonymy, distribution and conservation status (8th ed.). Chelonian Research Monographs 7: 1–292. https://doi.org/10.3854/crm.7.checklist.atlas.v8.2017

Valdiya K.S. 2010. The Making of India: Geodynamic Evolution. Springer International Publishing, Switzerland.

Wermuth H. & Mertens R. 1977. Testudines, Crocodylia, Rhynchocephalia. Das Tierreich. Walter de Gruyter, Berlin.

Yin A. 2006. Cenozoic tectonic evolution of the Himalayan orogen as constrained by along-strike variation of structural geometry, exhumation history, and foreland sedimentation. Earth-Science Reviews 76: 1–131. https://doi.org/10.1016/j.earscirev.2005.05.004

Zangerl R. 1969. The turtle shell. In: Gans C., D’Abellairs A. & Parsons T.S. (eds) Biology of the Reptilia. Volume 1 – Morphology. Academic Press, New York.

Published
2020-05-26
How to Cite
Garbin, R. C., Bandyopadhyay, S., & Joyce, W. G. (2020). A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan. European Journal of Taxonomy, (652). https://doi.org/10.5852/ejt.2020.652