Skip to main content Accessibility help

A new species of Cuora (Testudines: Geoemydidae) from the Miocene of Thailand and its evolutionary significance



A new species of Cuora, Cuora chiangmuanensis sp. nov., is described on the basis of a nearly complete shell with limb bones from the late Middle – early Late Miocene Chiang Muan Mine, Phayao Province (Northern Thailand). C. chiangmuanensis is distinguished from other fossil and living Cuora species mainly on the basis of its plastral morphology. Among fossil and extant Cuora taxa, the new species appears to be a missing link between the taxa from Southeast Asia and those from East Asia. It represents the earliest record of the genus and demonstrates that by 11–12 Ma, Asian box turtles were already present in Southeast Asia.


Corresponding author

Author for correspondence:


Hide All
Batsch, A. J. G. C. 1788. Versuch einer Anleitung zur Kenntniss und Geschichte der Thiere und Mineralien. Erster Theil. Allgemeine Geschichte der Natur; besondre der Säugthiere, Vögel, Amphibien und Fische. Jena: Akademischen Buchhandlung, 528 pp.
Bell, T. 1825. A monograph of the tortoise having a moveable sternum, with remarks on their arrangement and affinities. Zoological Journal 2, 299310.
Blanck, T. 2005. Cuora yunnanensis (Boulenger, 1906), the Yunnan box turtle, rediscovered after one-hundred years? Radiata 14, 1033.
Blanck, T., McCord, W. P. & Le, M. 2006. On the Variability of Cuora trifasciata (Bell, 1825). Frankfurt am Main: Edition Chimaira.
Bonin, F., Devaux, B. & Dupre, A. 2006. Turtles of the World. London: Johns Hopkins University Press.
Boulenger, G. A. 1906. Descriptions of new reptiles from Yunnan. Annals and Magazine of Natural History, Series 7 17, 567–8.
Bourret, R. 1939. Notes herpétologiques su l'Indochine française. XVIII. Reptiles et batraciens reçus au Laboratoire des Sciences Naturelles de l'Université au cours de l'année 1939. Descriptions de quatre espèces et d'une variété nouvelles. Bulletin Générale d'Instruction Publique 4 (annexe), 540.
Cantor, T. 1842. General features of Chusan, with remarks on the flora and fauna of that island. Annals and Magazine of Natural History, Series 1 9, 265–78, 361–70, 481–93.
Claude, J. & Tong, H. 2004. Early Eocene turtles from Saint Papoul, France, with the comments on the early evolution of modern Testudinoidea. Oryctos 5, 71113.
Cope, E. D. 1868. On the origin of genera. Proceedings of the Academy of Natural Sciences of Philadelphia 20, 242300.
Deng, T. 2006. Chinese Neogene mammal biochronology. Vertebrata Palasiatica 44, 143–63.
Deng, T. & Qi, G.-Q. 2009. Rhinocerotids (Mammalia, Perissodactyla) from Lufengpithecus site, Lufeng, Yunnan. Vertebrata PalAsiatica 47, 135–52.
Ernst, C. H. 1988. Cuora mccordi, a new Chinese box turtle from Guangxi Province. Proceedings of the Biological Society of Washington 101, 466–70.
Fritz, U. & Havas, P. 2007. Checklist of chelonians of the world. Vertebrate Zoology 57, 149368.
Fong, J., Parham, J. & Fu, J. 2002. A reassessment of the distribution of Cuora flavomarginata Gray, 1868 on mainland China. Russian Journal of Herpetology 9 (1), 914.
Gaffney, E. S. & Meylan, P. A. 1988. A phylogeny of turtles. In The Phylogeny and Classification of the Tetrapods, Vol. 1: Amphibians, Reptiles, Birds (ed. Benton, M. J.), pp. 157219. Oxford: Clarendon Press.
Gray, J. E. 1831. Synopsis Reptilium; or Short Descriptions of the Species of Reptiles. Part I. – Cataphracta. Tortoises, Crocodiles, and Enaliosaurians. London: Treuttel, Wurz and Co., 85 pp.
Gray, J. E. 1855. On some new species of freshwater tortoises from North America, Ceylon and Australia, in the collection of the British Museum. Proceedings of the Zoological Society of London 23, 197202.
Gray, J. E. 1860. On some new species of Mammalia and tortoises from Cambojia. Annals and Magazine of Natural History, Series 3 6, 217–18.
Gray, J. E. 1862. Notice of a new species of Cyclemys from the Lao Mountains, in Siam. Annals and Magazine of Natural History, Series 3 10,157.
Gray, J. E. 1863. Observations on the box tortoises, with the descriptions of three new Asiatic species. Proceedings of the Zoological Society of London 1863, 173–8.
Hasegawa, Y. & Ota, M. 1980. Report on a Pleistocene terrapin, Cyclemys from Katayama Marble Quarry, Mine City, Japan. Science Reports of the Yokohama National University, Section II 27, 4752.
He, T., Zhou, T., Rao, T.-Q. & Zhang, Y.-P. 2007. Molecular identification and phylogenetic position of Cuora yunnanensis . Chinese Science Bulletin 52 (23), 3305–9.
Hirayama, R. 1985. Cladistic analysis of batagurine turtles (Batagurinae: Emydidae: Testudinoidea); a preliminary result. In Studio Palaeocheloniologica 1. Stvdia Geølogica Salmanticensia Volumen Especial 1 (eds. de Broin, F. & Jiménez-Fuentes, E.), pp. 141–57.
Hirayama, R. 2007. Revision of the systematics of the fossil turtles from Japan. Jubilee Publication in Commemoration of Professor Kamei Tadao's 80th Birthday, 145–53.
Honda, M., Yasukawa, Y., Hirayama, R. & Ota, H. 2002. Phylogenetic relationships of the Asian box turtles of the genus Cuora sensu lato (Reptilia: Bataguridae) inferred from mitochondrial DNA sequences. Zoological Science 19, 1305–12.
Iverson, J. B. 2008. A checklist of the turtles of the world.
Iverson, J. B. 1992. A Revised Checklist with Distribution Maps of the Turtles of the World. Richmond, Indiana: Privately Printed.
Iverson, J. B., Brown, R. M., Akre, T. S., Near, T. J., Le, M., Thomson, R. C. & Starkey, D. E. 2007. In search of the tree of life for turtles. In Defining Turtle Diversity: Proceedings of a Workshop on Genetics, Ethics, and Taxonomy of Freshwater Turtles and Tortoises (eds Shaffer, H. B., Fitz Simmons, N. N., Georges, A. & Rhodin, A. G. J.). Chelonian Research Monographs 4, 85106.
Iverson, J. B. & McCord, W. P. 1992. A new subspecies of Cuora galbinifrons (Testudines: Batagurinae) from Hainan Island, China. Proceedings of the Biological Society of Washington 105, 433–9.
Kunimatsu, Y., Ratanasthien, B., Nakaya, H., Saegusa, H. & Nagaoka, S. 2004. Earliest Miocene hominoid from Southeast Asia. American Journal of Physical Anthropology 124, 99108.
Lehr, E., Fritz, U. & Obst, F. J. 1998. Cuora galbinifrons picturata subsp. nov., eine neue Unterart der Hinterindischen Scharnierschildkröte. Herpetofauna 20 (113), 511.
Linnaeus, C. 1758. Systema Naturae, per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio Decima, Reformata. [10th ed.]. Holmiae [Stockholm]: Laurentii Salvii, 824 pp.
Luo, B. & Zong, Y. 1988. A new species of CuoraCuora aurocapitata . Acta Herpetologica Sinica 3, 1315.
McCord, W. P. & Iverson, J. B. 1991. A new box turtle of the genus Cuora Testudines Emydidae with taxonomic notes and a key to the species. Herpetologica 47, 407–20.
Nakaya, H., Saegusa, H., Kunimatsu, Y. & Ratanasthien, B. 2002. Miocene mammalian faunas of Northern Thailand and their geological age. Primate Research 18, 131–41 (in Japanese with English abstract).
Nutaphand, V. 1979. The Turtles of Thailand. Bangkok: Siamfarm Zoological Garden, 222 pp.
Obst, F. J. & Reimann, M. 1994. Bemerkenswerte Variabilität bei Cuora galbinifrons Bourret, 1939, mit Beschreibung einer neuen geographischen Unterart: Cuora galbinifrons bourreti subsp. nov. (Reptilia: Testudines: Criptodira: Bataguridae). Zoologische Abhandlungen, Staatliches Museum für Tierkunde Dresden 48, 125–39.
Parham, J. F. & Li, D. 1999. A new locality for Cuora pani Song 1984 with comments on its known range. Asiatic Herpetological Research 8, 111–13.
Parham, J. F., Simison, W. B., Kozak, K. H., Feldman, C. R. & Shi, H. 2001. New Chinese turtles: endangered or invalid. A reassessment of two species using mitochondrial DNA, allozyme electrophoresis and known-locality specimens. Animal Conservation 4, 357–67.
Parham, J. F., Stuart, B. L., Bour, R. & Fritz, U. 2004. Evolutionary distinctiveness of the extinct Yunnan box turtle (Cuora yunnanensis) revealed by DNA from an old museum specimen. Proceedings of the Royal Society Biological Sciences Series B 271, S391S394.
Pickford, M., Nakaya, H., Kunimatsu, Y., Saegusa, H., Fukuchi, A. & Ratanasthien, B. 2004. Age and taxonomic status of the Chiang Muan (Thailand) hominoids. Comptes Rendus Palevol 3, 6575.
Riche, M. 1801. Testudo amboinensis . In Histoire Naturelle, á Générale et Particulière, des Reptiles. Tome Second (ed. Daudin, F. M.), pp. 309–12. Paris: Dufart.
Schweigger, A. F. 1812. Prodromus monographiae Cheloniorum. Königsberger Archive für Naturwissenschaftliche und Mathematik 1, 271368, 406–58.
Shi, H. 2008. Identification Manual for Trade Turtles in China. Beijing: Encyclopedia of China Publishing House.
Shi, H., Parham, J. F., Simison, W. B., Wang, J., Gong, S. & Fu, B. 2005. A report on the hybridization between two species of threatened Asian box turtles (Testudines: Cuora) in the wild on Hainan Island (China) with comments on the origin of ‘serrata’-like turtles. Amphibia-Reptilia 26 (3): 377–81.
Shikama, T. 1949. The Kuzuü Ossuaries: geological and palaeontological studies of the limestone fissure deposits, in Kuzuü, Totigi Prefecture. Science Report of Tohoku University, 2nd Series (Geology) 23, 1201.
Silaratana, T., Ratanasthien, B., Takayasu, K., Fyfe, W. S., Asnachinda, P., Kandharosa, W. & Kusakabe, M. 2004. Sulfur isotopic implication of Middle Miocene marine incursion in Northern Thailand. Science Asia 30, 4358.
Song, M. T. 1984. A new species of the turtle genus Cuora (Testudoformes: Testudinidae). Acta Zootaxonomica Sinica 9 (3), 330–32.
Spinks, P. Q. & Shaffer, H. B. 2007. Conservation phylogenetics of the Asian box turtles (Geoemydidae, Cuora): mitochondrial introgression, numts, and inferences from multiple nuclear loci. Conservation Genetics 8, 641–57.
Spinks, P. Q., Thomson, R. C., Shaffer, H. B. 2009. A reassessment of Cuora cyclornata Blanck, McCord and Le, 2006 (Testudines, Geoemydidae) and a plea for taxonomic stability. Zootaxa 2018, 5868.
Spinks, P. Q., Thomson, R. C., Zhang, Y., Che, J., Wu, Y. & Shaffer, H. B. 2012. Species boundaries and phylogenetic relationships in the critically endangered Asian box turtle genus Cuora . Molecular Phylogenetics and Evolution 63, 656–67.
Stuart, B. & Parham, J. F. 2004. Molecular phylogeny of the critically endangered Indochinese box turtle (Cuora galbinifrons). Molecular Phylogenetics and Evolution 31, 164–77.
Stuart, B. & Parham, J. F. 2007. Recent hybrid origin of three rare Chinese turtles. Conservation Genetics 8, 169–75.
Suganuma, Y., Hamada, T., Tanaka, S., Okada, M., Nakaya, H., Kunimatsu, Y., Saegusa, H., Nagaoka, S., & Ratanasthien, B. 2006. Magnetostratigraphy of the Miocene Chiang Muan Formation, Northern Thailand: implication for revised chronology of the earliest Miocene hominoid in Southeast Asia. Palaeogeography, Palaeoclimatology, Palaeoecology 239, 7586.
Swofford, D. L. 1998. PAUP*: phylogenetic analysis using parsimony (*and other methods). Sinauer Associates, Sunderland, MA.
Takahashi, A., Otsuka, H. & Ota, H. 2008. Systematic review of Late Pleistocene turtles (Reptilia: Chelonii) from the Ryukyu Archipelago, Japan, with special reference to paleogeographical implication. Pacific Science 62 (3), 395402.
Theobald, W. Jr. 1868. Catalogue of Reptiles in the Museum of the Asiatic Society of Bengal. Journal of the Asiatic Society, Extra Number. Calcutta: Printed at the Baptist Mission Press, 88 pp.
Turtle Taxonomy Working Group [Van Dijk, P. P., Iverson, J. B., Shaffer, H. B., Bour, R. & Rhodin, A. G. J.]. 2011. Turtles of the World, 2011 update: annotated checklist of taxonomy, synonymy, distribution, and conservation status. In Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group (eds A. G. J. Rhodin, P. C. H. Pritchard, P. P. van Dijk, R. A. Saumure, K. A. Buhlmann, J. B. Iverson & R. A. Mittermeier). Chelonian Research Monographs 5, pp. 000.165–000.242, doi:10.3854/crm.5.000.checklist. v4.2011,
Wang, B.-Y. & Qi, G.-Q. 2005. A procupine (Rodentia, Mammalia) from Lufengpithecus site, Lufeng, Yunnan. Vertebrata PalAsiatica 43, 1123.
Yasukawa, Y., Hirayama, R. & Kikida, T. 2001. Phylogenetic relationships of Geoemydine turtles (Reptilia: Bataguridae). Current Herpetology 20, 105–33.
Yeh, H.-K. 1961. The first discovery of a box-turtle in China. Vertebrata PalAsiatica 3, 5864.
Yeh, H.-K. 1981. New fossil Cuora (Emydidae) from Early Pliocene of Yunnan. Vertebrata PalAsiatica 19, 239–46.
Yeh, H.-K. 1983. New remains of Cuora from Zhejiang. Vertebrata PalAsiatica 21: 4953.
Yeh, H.-K. 1985 a. Studies on fossils of Cuora of China and Japan. Acta Hepetologica Sinica 4, 81–7.
Yeh, H.-K. 1985 b. New materials of Cuora pitheca and its generic history. Acta Anthropologiga Sinica 6, 113–17.
Yeh, H.-K. 1994 a. Notes on a fossil Cuora (Testudines) from Tongshan, Anhui. Vertebrata PalAsiatica 32, 66–8.
Yeh, H.-K. 1994 b. Fossil and Recent Turtles of China. Beijing: Science Press.
Yeh, H.-K. & Zhang, J. 1994. Turtle remains from Jiahu site at Wuyang, Henan. Acta Anthropologiga Sinica 13, 32–9.
Zhang, L., Nie, L. W., Cao, C. H. & Zhang, Y. 2008. The complete mitochondrial genome of the keeled box turtle Pyxidea mouhotii and phylogenetic analysis of major turtle groups. Journal of Genetics and Genomics 35, 3340.
Zhao, E. 1990. Cuora zhoui . In A new Chinese box turtle (Testudinata: Emydidae) – Cuora zhoui (eds Zhao, E., Zhou, T. & Ye, P.), pp. 213–16. In From Water Onto Land (ed. E. Zhao). Beijing: Chinese Society for the Study of Amphibians and Reptiles (in Chinese).
Zhou, T., Blanck, T., McCord, W. P. & Li, P.-P. 2008. Tracking Cuora mccordi Ernst, 1988: the first record of its natural habitat; a re-description; with data on captive populations and its vulnerability. Hamadryad 32, 5769.
Zhou, T. & Zhao, E. 2004. On the occurrence of living Cuora yunnanensis since fifty-eight years, and its description. Sichuan Journal of Zoology 23, 325–6.



Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed