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A new box turtle from the Miocene/Pliocene boundary (latest Hemphillian) of Oklahoma and a refined chronology of box turtle diversification

  • Walter G. Joyce (a1) (a2), Andrea Petričević (a1), Tyler R. Lyson (a3) (a4) and Nicholas J. Czaplewski (a5)


A near complete shell from the Hemphillian 4 (Miocene/Pliocene boundary) Buis Ranch local fauna of Beaver County, Oklahoma, represents a fossil box turtle. An anterior contact of neural III and neural V with costal III and costal V only, respectively, presence of a small contact between the suprapygal and eleventh peripherals, development of a thin peripheral lip for articulation with the posterior plastral lobe, placement of the vertebral III/IV sulcus on neural VII, presence of two anterior musk duct glands, a rounded posterior plastral lobe, an elongate shell outline, and a complete neural series diagnose the fossil as a new species, Terrapene parornata n. sp. A phylogenetic analysis of fossil box turtles places T. parornata along the phylogenetic stem of the extant taxon T. ornata. The holotype of ‘Terrapene longinsulae’ cannot be distinguished from Terrapene ornata and is therefore synonymized. Finally, ‘Terrapenecorneri lacks characters of crown group Terrapene and may therefore represent a stem box turtle. The provenance of the holotype of ‘Terrapene longinsulae’ is more poorly known than previously recognized and this specimen may originate from Kansas or Nebraska and be early Miocene to late Pleistocene in age. Terrapene parornata is therefore the oldest demonstrable representative of crown group Terrapene (ca. 5.3–4.6 Ma). ‘Terrapenecorneri from the late Barstovian of Nebraska and fragmentary material from the middle Barstovian of Nebraska by contrast are the oldest representative of the Terrapene lineage (ca. 14.5–11.5 Ma). A review of morphological characters related to shell kinesis reveals that most are highly correlated. The results of the phylogenetic analysis converge upon those of molecular data when these correlated characters are omitted from the analysis.



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Agassiz, L. 1857. Contributions to the Natural History of the United States of America. Little, Brown, and Company, Boston, 452 p.
Angielczyk, K. D., Feldman, C. R., and Miller, G. R. 2011. Adaptive evolution of plastron shape in emydine turtles. Evolution, 65:377394.
Auffenberg, W. 1958. Fossil turtles of the genus Terrapene in Florida. Bulletin of the Florida State Museum Biological Sciences, 3:5392.
Auffenberg, W. 1967. Further notes on fossil box turtles of Florida. Copeia, 1967:319325.
Barbour, T., and Stetson, H. C. 1931. A revision of the Pleistocene species of Terrapene of Florida. Bulletin of the Museum of Comparative Zoology, 72:295299.
Batsch, A. J. G. C. 1788. Versuch einer Anleitung, zur Kenntniß und Geschichte der Thiere und Mineralien. Akademische Buchhandlung, Jena, 528 p.
Bell, T. 1825. A monograph of the tortoises having a moveable sternum with remarks on their arrangement and affinities. Zoological Journal, 2:299310.
Berggren, W. A., Hilgen, F. J., Langereis, C. G., Kent, D. V., Obradovich, J. D., Raffi, I., Raymo, M. E., and Shackleton, N. J. 1995. Late Neogene chronology: New perspectives in high-resolution stratigraphy. Geological Society of America Bulletin, 107:12721287.
Bickham, J. W., Lamb, T., Minx, P., and Patton, J. C. 1996. Molecular systematics of the genus Clemmys and the intergeneric relationships of emydid turtles. Herpetologica, 52:8997.
Bramble, D. M. 1974. Emydid shell kinesis: Biomechanics and evolution. Copeia, 1974:707727.
Brattstrom, B. H. 1967. A succession of Pliocene and Pleistocene snake faunas from the High Plains of the United States. Copeia, 1967:188202.
Burke, R. L., Leuteritz, T. E., and Wolf, A. J. 1996. Phylogenetic relationships of emydine turtles. Herpetologica, 52:572584.
Coker, R. E. 1910. Diversity in the scutes of Chelonia. Journal of Morphology, 21:175, pls. 1–14.
Cope, E. D. 1868. On the origin of genera. Proceedings of the Academy of Natural Sciences of Philadelphia, 1868:242300.
Cope, E. D. 1870. Synopsis of the extinct Batrachia, Reptilia and Aves of North America. Transactions of the American Philosophical Society, 14:1252.
Cope, E. D. 1878. Descriptions of new Vertebrata from the upper Tertiary formations of the West. Proceedings of the American Philosophical Society, 17:219231.
Davy, C. M., and Murphy, R. W. 2009. Explaining patterns of deformity in freshwater turtles using MacCulloch's hypothesis. Canadian Journal of Zoology, 87, 5:433439.
Dolliver, P. N. 1984. Cenozoic evolution of the Canadian River Basin. Baylor Geological Studies Bulletin, 42:196.
Ernst, C. H., Lovich, J. E., and Barbour, R. W. 1994. Turtles of the United States and Canada. Smithsonian Institution Press, Washington D.C., 578 p.
Feldman, C. R., and Parham, J. F. 2002. Molecular phylogenetics of emydine turtles: Taxonomic revision and the evolution of shell kinesis. Molecular Phylogenetics and Evolution, 22:388398.
Felsenstein, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39:783791.
Fitzinger, L. 1826. Neue Classification der Reptilien nach ihren natürlichen Verwandtschaften. Verlag J. G. Heubner, Wien, 66 p.
Gaffney, E. S., and Meylan, P. A. 1988. A phylogeny of turtles, p. 157219InBenton, M. J.(ed.), The Phylogeny and Classification of the Tetrapods, Vol. 1: Amphibians, Reptiles, Birds. The Systematics Association Special Vol. 35A. Clarendon Press, Oxford.
Gilmore, C. W. 1927. On fossil turtles from the Pleistocene of Florida. Proceedings of the United States National Museum, 71:110.
Hay, O. P. 1906. Descriptions of two new genera (Echmatemys and Xenochelys) and two new species (Xenochelys formosa and Terrapene putnami) of fossil turtles. Bulletin of the American Museum of Natural History, 22:2731.
Hay, O. P. 1907. Descriptions of seven new species of turtles from the Tertiary of the United States. Bulletin of the American Museum of Natural History, 23:847863.
Hay, O. P. 1908. Descriptions of five species of North American fossil turtles, four of which are new. Proceedings of the United States National Museum, 35:161169.
Hay, O. P. 1916. Descriptions of some Floridian fossil vertebrates. Annual Report of the Florida State Geological Survey, 8:3976.
Hay, O. P. 1917. Vertebrata mostly from stratum No. 3, at Vero, Florida, together with descriptions of new species. Annual Report of the Florida State Geological Survey, 9:4368.
Hay, O. P. 1921. Descriptions of some Pleistocene vertebrates found in the United States. Proceedings of the United States National Museum, 58:83146.
Hazard, E. B. 1961. The subgeneric status and distribution in time of Citellus rexroadensis. Journal of Mammalogy, 42:477483.
Hibbard, C. W. 1954. A new Pliocene vertebrate fauna from Oklahoma. Papers of the Michigan Academy of Sciences, Arts, and Letters, 39:339359.
Hibbard, C. W. 1963. Tanupolama vera (Matthew) from the late Hemphillian of Beaver County, Oklahoma. Transactions of the Kansas Academy of Sciences, 66:267269.
Hillis, D. M., and Bull, J. J. 1993. An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic Biology, 42:182192
Holman, J. A. 1973. A new Pliocene snake, genus Elaphe, from Oklahoma. Copeia, 1973:574580.
Holman, J. A. 1975. Herpetofauna of the WaKeeney Local Fauna of Trego County Kansas. University of Michigan Papers on Paleontology, 12:4966.
Holman, J. A., and Corner, R. G. 1985. A Miocene Terrapene (Testudines: Emydidae) and other Barstovian turtles from south-central Nebraska. Herpetologica, 41:8893.
Holman, J. A. 1987. Herpetofauna of the Egelhoff site (Miocene: Barstovian) of North-Central Nebraska. Journal of Vertebrate Paleontology, 7:109120.
Holman, J. A. and Fritz, U. 2005. The box turtle genus Terrapene (Testudines: Emydidae) in the Miocene of the U.S.A. The Herpetological Journal, 15:8190.
Honey, J. G., Martin, R. A., and Peláez-Campomanes, P. 2005. Stratigraphic framework of early Pliocene fossil localities along the north bank of the Cimarron River, Meade County, Kansas. Ameghiniana, 42:461472.
Hulbert, R. C. Jr, Czaplewski, N. J., and Webb, S. D. 2005. New records of Pseudhipparion simpsoni (Mammalia, Equidae) from the late Hemphillian of Oklahoma and Florida. Journal of Vertebrate Paleontology, 25:737740.
Hutchison, J. H., and Bramble, D. M. 1981. Homology of the plastral scales of the Kinosternidae and related turtles. Herpetologica, 37:7385.
Iverson, J. B. 1992. A Revised Checklist with Distribution Maps of the Turtles of the World. Privately printed, Richmond, Virginia, 363 p.
Izett, G. A., and Honey, J. G. 1995. Geological map of the Irish Flats NE quadrangle, Meade County, Kansas. United States Geological Survey Miscellaneous Series Investigations Map I–2498.
Joyce, W. G., and Bell, C. J. 2004 A review of the comparative morphology of extant testudinoid turtles (Reptilia: Testudines). Asiatic Herpetological Research, 10:53109.
Legler, J. M. 1960. Natural history of the ornate box turtle, Terrapene ornata ornata Agassiz. University of Kansas Publications Museum of Natural History, 11:527669.
Linnaeus, C. 1758. Systema Naturae, Vol. 1 (tenth edition). Laurentius Salvius, Holmia, 824 p.
Lynn, W. G. 1937. Variation in scutes and plates in the boxturtle, Terrapene carolina. The American Naturalist, 71:421426.
Martin, R. A., Honey, J. G., and Peláez-Campomanes, P. 2000. The Meade Basin rodent project: A progress report. Paludicola, 3:132.
Martin, R. A., Honey, J. G., Peláez-Campomanes, P., Goodwin, H. T., Baskin, J. A., and Zakrzewski, R. J. 2002. Blancan lagomorphs and rodents of the Deer Park assemblages, Meade County, Kansas. Journal of Paleontology, 76:10721090.
Martin, R. A., Hurt, R. T., Honey, J. G., and Peláez-Campomanes, P. 2003. Late Pliocene and early Pleistocene rodents from the northern Borchers Badlands (Meade County, Kansas), with comments on the Blancan–Irvingtonian boundary in the Meade Basin. Journal of Paleontology, 77:9851001.
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, 143:239279.
Merrem, B. 1820. Versuch eines Systems der Amphibien. Johann Christian Krieger, Marburg, 191p.
Milstead, W. M. 1956. Fossil turtles of Friesenhahn Cave, Texas, with the description of a new species of Testudo. Copeia, 1956:162171.
Milstead, W. W. 1967. Fossil box turtles (Terrapene) from central North America and box turtles of eastern Mexico. Copeia, 1967:168179.
Milstead, W. W. 1969. Studies on the evolution of the box turtles (genus Terrapene). Bulletin of the Florida State Museum Biological Sciences, 14:1113.
Minx, P. 1996. Phylogenetic relationships among the box turtles, genus Terrapene. Herpetologica, 52:584597.
Moodie, K. B., and Van Devender, T. R. 1978. Fossil box turtles (Genus Terrapene) from southern Arizona. Herpetologica, 34:172174.
Oelrich, T. 1953. A new boxturtle from the Pleistocene of southwestern Kansas. Copeia, 1953:3338.
Parker, G. H. 1901. Correlated abnormalities in the scutes and bony plates of the carapace of the sculptured tortoise. The American Naturalist, 35:1724.
Parmley, D., and Holman, J. A. 1995. Hemphillian (late Miocene) snakes from Nebraska, with comments on Arikareean through Blancan snakes of midcontinental North America. Journal of Vertebrate Paleontology, 15:7995.
Schmidt, K. P., and Owens, D. W. 1944. Amphibians and reptiles of northern Coahuila, Mexico. Field Museum of Natural History, Zoological Series, 29:97115.
Smith, H. M., and Smith, R. B. 1979. Synopsis of the Herpetofauna of Mexico, Vol. VI. Guide to Mexican Turtles. Lundberg, Augusta, West Virginia.
Spinks, P. Q., and Shaffer, H. B. 2009. Conflicting mitochondrial and nuclear phylogenies for the widely disjunct Emys (Testudines: Emydidae) species complex, and what they tell us about biogeography and hybridization. Systematic Biology, 58:120.
Stejneger, L. 1925. New species and subspecies of American turtles. Journal of the Washington Academy of Sciences, 15:462463.
Stevens, M. S. 1966. The osteology and relationships of the Pliocene ground squirrel, Citellus dotti Hibbard, from the Ogallala Formation of Beaver County, Oklahoma. Pearce-Sellards Series, 4:124.
Swofford, D. L. 2002. PAUP∗. Phylogenetic Analysis Using Parsimony (∗and other methods). Version 4b10. Sinauer Associates, Sunderland, Massachusetts.
Tedford, R. H., Albright, L. B. III, Barnosky, A. D., Ferrusquía-Villafranca, I., Hunt, R. M. Jr, Storer, J. E., Swisher, C. C. III, Voorhies, M. R., Webb, S. D., and Whistler, D. P. 2004. Mammalian biochronology of the Arikareean through Hemphillian interval (late Oligocene through early Pliocene epochs), p. 169231. InWoodburne, M. O.(ed.), Late Cretaceous and Cenozoic Mammals of North America: Biostratigraphy and Geochronology. Columbia University Press, New York.
Tihen, J. A. 1955. A new Pliocene species of Ambystoma, with remarks on other fossil ambystomids. Contributions to Paleontology, Museum of Paleontology of the University of Michigan, 12:229244.
Velo-Antón, G., Becker, C. G., and Cordero-Rivera, A. 2011. Turtle carapace anomalies: The roles of genetic diversity and environment. PLoS ONE, 6:e18714.
Waagen, G. N. 1972. Musk glands in recent turtles. M.S. thesis, University of Utah, 64 p.
Werdelin, L. 1985. Small Pleistocene felines of North America. Journal of Vertebrate Paleontology, 5:194210.
Wright, D. B. 1989. Phylogenetic relationships of Catagonus wagneri: Sister taxa from the Tertiary of North America, p. 281308. InRedford, K. H. and Eisenberg, J. F.(eds.), Advances in Neotropical Mammalogy. Sandhill Crane Press, Gainesville, Florida.
Zangerl, R. and Johnson, R. G. 1957. The nature of shield abnormalities in the turtle shell. Fieldiana: Geology, 10:341362.
Zug, G. R. 1969. Fossil Chelonians, Chrysemys and Clemmys, from the upper Pliocene of Idaho. Great Basin Naturalist, 29:8287.

A new box turtle from the Miocene/Pliocene boundary (latest Hemphillian) of Oklahoma and a refined chronology of box turtle diversification

  • Walter G. Joyce (a1) (a2), Andrea Petričević (a1), Tyler R. Lyson (a3) (a4) and Nicholas J. Czaplewski (a5)


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