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Cranial morphology and systematics of an extraordinary sample of the Late Neogene dwarf tapir, Tapirus polkensis (Olsen)

  • Richard C. Hulbert (a1), Steven C. Wallace (a2), Walter E. Klippel (a3) and Paul W. Parmalee (a4)


The previously poorly known “Tapiravus” polkensis Olsen, 1960 (Mammalia, Perissodactyla, Tapiridae) is now known from abundant, well preserved specimens from both the type area in central Florida and from the Gray Fossil Site (GFS) in eastern Tennessee. The latter has produced over 75 individuals, the greatest number of tapirids from a single fossil site, including many articulated skeletons. Almost all linear measurements taken on skulls, mandibles, and cheek teeth from GFS have coefficients of variation less than 10 (most between 3 and 7), indicating the presence of a single species. However, the sample reveals considerable intraspecific variation for a few key morphologic features, including development of the sagittal crest, outline shape of the nasals, and number and relative strength of lingual cusps on the P1. The Florida sample of T. polkensis is more limited, but has the same state as the GFS sample for all preserved characters of systematic significance, and while the Florida teeth are on average smaller (especially narrower lower cheek teeth), they fall either within or just below the observed range of the Gray Fossil Site population. The new material supports a reassignment of “Tapiravus” polkensis to the genus Tapirus, and demonstrates that the geologic age of the species is significantly younger than previously thought, Hemphillian rather than Barstovian. Tapirus polkensis is the smallest known North American Tapirus, and smaller than any of the extant species in the genus, with an estimated average mass of 125 kg.



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Benson, R. N. 1998. Radiolarians and diatoms from the Pollack Farm Site, Delaware: marine-terrestrial correlation of Miocene vertebrate assemblages of the middle Atlantic Coastal Plain, pp. 519. In Benson, R. N. (ed.), Geology and Paleontology of the Lower Miocene Pollack Farm Fossil Site, Delaware. Delaware Geological Survey Special Publication 21.
Boeuf, O. 1991. Tapirus jeanpiveteaui nov. sp., nouvelle espèce de Tapiridae (Perissodactyla) du Pliocène de Charente (France). Palaeontographica, Abteilung A, 217:177194.
Colbert, M. W. 2005. The facial skeleton of the early Oligocene Colodon (Perissodactyla, Tapiroidea). Paleontologica Electronica, 8:12A <>.
Colbert, M. W. and Schoch, R. M. 1998. Tapiroidea and other moropomorphs, pp. 569582. In Janis, C. M., Scott, K. S., and Jacobs, L. L. (eds.), Evolution of Tertiary Mammals of North America, Volume 1: Terrestrial Carnivores, Ungulates, and Ungulatelike Mammals. Cambridge University Press, Cambridge.
Cook, H. J. 1927. A new rhinoceros of the genus Teleoceras from Colorado. Proceedings of the Colorado Museum of Natural History, 7:15.
Croizet, J.-B. and Jobert, A. C. G. 1828. Reserches sur les Ossements Fossiles du Department du Puy-de-Dôme. Paris, 224 p.
Desantis, L. R. G. and Wallace, S. C. 2008. Neogene forest refugium from the Appalachians of Tennessee, USA: Geochemical evidence from fossil mammal teeth. Palaeogeography, Palaeoclimatology, Palaeoecology, 266:5968.
Desmarest, A. G. 1819. Tapirs fossiles, pp. 458461. In Nouveau Dictionnaire d'Histoire Naturelle (ed.), Nouvelle Edition. Chez Deterville, Paris, vol. 32.
Fejar, O. 1964. The lower-Villafranchian vertebrates from Hajnáōka near Filákovo in southern Slovakia. Ústrední Ústav Geologický, Praha, 116 pp.
Ferrero, B. S. and Noriega, J. I. 2007. A new upper Pleistocene tapir from Argentina: remarks on the phylogenetics and diversification of Neotropical Tapiridae. Journal of Vertebrate Paleontology, 27:504511.
Florida Geological Survey. 1959. Known active mineral producers of Florida 1956-1957. Biennial Report, Florida Geological Survey, 13:7084.
Frick, C. 1921. Extinct vertebrate faunas of the badlands of Bautista Creek and San Timoteo Cañon, southern California. University of California Publications, Bulletin of the Department of Geology, 12:277424.
Frick, C. 1926. The Hemicyoninae and an American Tertiary bear. Bulletin of the American Museum of Natural History, 56:1119.
Gazin, C. L. and Collins, R. L. 1950. Remains of land mammals from the Miocene of the Chesapeake Bay Region. Smithsonian Miscellaneous Collections, 116(2): 121.
Gill, T. N. 1865. Description of the genus Elasmognathus. Proceedings of the Academy of Natural Sciences Philadelphia, 17:183.
Guérin, C. and Eisenmann, V. 1994. Les tapirs (Mammalia, Perissodactyla) du Miocène supérieur d'Europe occidentale. Geobios, 27:113127.
Hatcher, J. B. 1894. A median horned rhinoceros from the Loup Fork Beds of Nebraska. American Geologist, 13:149150.
Hershkovitz, P. 1954. Mammals of northern Colombia, preliminary report no. 7: tapirs (genus Tapirus), with a systematic review of American species. Proceedings of the United States National Museum, 103:465496.
Holbrook, L. T. 2002. The unusual development of the sagittal crest in the Brazilian tapir (Tapirus terrestris). Journal of Zoology, London, 256:215219.
Hooijer, D. A. 1947. On fossil and prehistoric remains of Tapirus from Java, Sumatra, and China. Zoologische Mededeelingen Museum Leiden, 27:253299.
Huang, W. and Qiren, F. 1991. Wushan Homind Site. China Ocean Press, Beijing, 205 p. (In Chinese)
Hulbert, R. C. 1995. The giant tapir, Tapirus haysii, from Leisey Shell Pit 1A and other Florida Irvingtonian localities. Bulletin of the Florida Museum of Natural History, 37:515551.
Hulbert, R. C. 1999. Nine million years of Tapirus (Mammalia, Perissodactyla) from Florida. Journal of Vertebrate Paleontology, 19(3, supplement):53A.
Hulbert, R. C. 2005. Late Miocene Tapirus (Mammalia, Perissodactyla) from Florida, with description of a new species, Tapirus webbi. Bulletin of the Florida Museum of Natural History, 45:465494.
Hulbert, R. C., MacFadden, B. J., and Waldrop, J. S. 2001. Perissodactyla, pp. 280306. In Hulbert, R. C. (ed.), The Fossil Vertebrates of Florida. University Press of Florida, Gainesville.
Hulbert, R. C. and Wallace, S. C. 2005. Phylogenetic analysis of late Cenozoic Tapirus (Mammalia, Perissodactyla). Journal of Vertebrate Paleontology, 25(3, supplement):72A.
Janis, C. M. 1990. Correlation of cranial and dental variables with body size in ungulates and macropodids, pp. 255291. In Damuth, J. and MacFadden, B. J. (eds.), Body Size in Mammalian Paleobiology, Estimation and Biological Implications. Cambridge University Press, Cambridge.
Jefferson, G. T. 1989. Late Cenozoic tapirs (Mammalia: Perissodactyla) of western North America. Los Angeles County Museum Contributions in Science, Number 406, 21 p.
Kaup, J. J. 1833. Description d'ossements fossils de Mammifères inconnus jusqu'à présent, qui se trouvent au Muséum grand-ducal de Darmstadt. 2ème cahier (Chalicotherium, Tapir, Porcs, Carnivores). Darmstadt, 31 pp.
Kretzoi, M. 1951. The Hipparion-fauna von Csakvar. Földtany Közlöny, 81: 402417.
Leidy, J. 1859. Descriptions of vertebrate fossils, pp. 99122. In Holmes, F. S. (ed.), Post-Pleiocene Fossils of South Carolina. Russell and Jones, Charleston.
Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonymis, locis. Tenth Edition. Laurentii Salvii, Stockholm, 824 p.
Lundelius, E. L. and Slaughter, B. H. 1976. Notes on American Pleistocene tapirs, pp. 226243. In Churcher, C. S. (ed.), Athlon: Essays in Paleobiology in Honour of Loris Shano Russell. Royal Ontario Museum, Toronto.
Marsh, O. C. 1871. Description of Lophiodon validus. Proceedings of the Academy of Natural Sciences, Philadelphia, 23:910.
Marsh, O. C. 1877. Notice of some new vertebrate fossils. American Journal of Science, series 3, 14:249256.
Martin, H. T. 1928. Two new carnivores from the Pliocene of Kansas. Journal of Mammalogy, 9:233236.
Matthew, W. D. and Granger, W. 1923. New fossil mammals from the Pliocene of Szechuan, China. Bulletin of the American Museum of Natural History, 48:563598.
Merriam, J. C. 1911. Tertiary mammal beds of Virgin Valley and Thousand Creek in northwestern Nevada. University of California Publications, Bulletin of the Department of Geological Sciences, 6:199304.
Merriam, J. C. 1913. Tapir remains from late Cenozoic beds of the Pacific coast region. University of California Publications, Bulletin of the Department of Geology, 7:169175.
Olsen, S. J. 1959. Fossil mammals of Florida. Florida Geological Survey Special Publication Number 6, 75 p.
Olsen, S. J. 1960. Age and faunal relationships of Tapiravus remains from Florida. Journal of Paleontology, 34:164167.
Owen, R. 1870. On fossil remains of mammals found in China. Journal of the Geological Society of London, 26:417434.
Padilla, M. and Dowler, R. C. 1994. Tapirus terrestris. Mammalian Species, Number 481, 8 p.
Parmalee, P. W., Klippel, W. E., Meylan, P. A., and Holman, J. A. 2002. A Late Miocene-early Pliocene population of Trachemys (Testudines: Emydidae) from East Tennessee. Annals of the Carnegie Museum of Natural History, 71:233239.
Ray, C. and Sanders, A. E. 1984. Pleistocene tapirs in the eastern United States, pp. 283315. In Genoways, H. H. and Dawson, M. R. (eds.), Contributions in Quaternary Vertebrate Paleontology: A Volume in Memorial to John E. Guilday. Carnegie Museum of Natural History, Pittsburgh, Special Publication 8.
Robertson, J. S. 1976. Latest Pliocene mammals from Haile XVA, Alachua County, Florida. Bulletin of the Florida State Museum, 20:111186.
Roulin, X. 1829. Memoir pour servir a l'histoire du tapir; et description dune espece nouvelle appartenant aux hautes regions de la Cordillere des Andes. Annales des Sciences Naturelle Zoologie Paris, 17:2655.
Rustioni, M. 1992. On Pliocene tapirs from France and Italy. Bollettino della Societá Paleontologica Italiana, 31:269294.
Savage, D. E. and Russell, D. E. 1983. Mammalian Paleofaunas of the World. Addison-Wesley, Reading, Massachusetts, 432 p.
Schlaikjer, E. M. 1937. A new tapir from the lower Miocene of Wyoming. Bulletin of the Museum of Comparative Zoölogy, 80:231251.
Schoch, R. M. 1984. The type specimens of Tapiravus validus and ?Tapiravus rarus (Mammalia, Perissodactyla), with a review of the genus, and a new report of Miotapirus (Miotapirus marslandensis Schoch and Prins, new species) from Nebraska. Postilla, Number 195, 12 p.
Schubert, B. W. and Wallace, S. C. 2006. Amphibians and reptiles of the Mio-Pliocene Gray Fossil Site and their paleoecologic implications. Journal of Vertebrate Paleontology, 26(supplement to 3:122A.
Schultz, C. B., Martin, L. D., and Corner, R. G. 1975. Middle and late Cenozoic tapirs from Nebraska. Bulletin of the Nebraska State Museum, 10:121.
Scott, K. M. 1990. Postcranial dimensions of ungulates as predictors of body size, pp. 301335. In Damuth, J. and MacFadden, B. J. (eds.), Body Size in Mammalian Paleobiology, Estimation and Biological Implications. Cambridge University Press, Cambridge.
Sellards, E. H. 1916. Fossil vertebrates from Florida: a new Miocene fauna; new Pliocene species; the Pleistocene fauna. Annual Report, Florida Geological Survey, 8:77119.
Sellards, E. H. 1918. The skull of a Pleistocene tapir including description of a new species and a note on the associated fauna and flora. Annual Report, Florida Geological Survey, 10:5770.
Simpson, G. G. 1945. Notes on Pleistocene and Recent tapirs. Bulletin of the American Museum of Natural History, 86:3382.
Shunk, A., Driese, S. G., and Clark, G. M. 2006. Latest Miocene to earliest Pliocene sedimentation and climate record derived from paleosinkhole fill deposits, Gray Fossil Site, northeastern Tennessee, U.S.A. Palaeogeography, Palaeoclimatology, Palaeoecology, 231:265278.
Spassov, N. and Ginsburg, L. 1999. Tapirus balkanicus nov. sp., nouveau tapir (Perissodactyla, Mammalia) du Turolien de Bulgarie. Annales de Paléontologie, 85:265276.
Sugarman, P. J., Miller, K. G., Owens, J. P., and Feigenson, M. D. 1993. Strontium-isotope and sequence stratigraphy of the Miocene Kirkwood Formation, southern New Jersey. Geological Society of America Bulletin, 105: 423436.
Tedford, R. H. and Hunter, M. E. 1984. Miocene marine-nonmarine correlations, Atlantic and Gulf Coastal Plains, North America. Palaeogeography, Palaeoclimatology, Palaeoecology, 47:126151.
Tedford, R. H., Albright, L. B., Barnosky, A. D., Ferrusquila-Villafranca, I., Hunt, R. M., Storer, J. E., Swisher, C. C., Voorhies, M. R., Webb, S. D., and Whistler, D. P. 2004. Mammalian biochronology of the Arikareean through Hemphillian interval (late Oligocene through earliest Pliocene epochs) in North America, pp. 169231, In Woodburne, M. O. (ed.), Late Cretaceous and Cenozoic Mammals of North America: Biostratigraphy and Geochronology. New York, Columbia University Press.
Tong, H. 2004. Tapiridae, pp. 233250. In Zheng, S. (ed.), Jianshi Hominid Site. Science Press, Beijing. (In Chinese)
Tong, H. 2005. Dental characters of the Quaternary tapirs in China, their significance in classification and phylogenetic assessment. Geobios, 38: 139150.
Tong, H., Liu, J., and Han, L. 2002. On fossil remains of early Pleistocene tapir (Perissodactyla, Mammalia) from Fanchang, Anhui. Chinese Science Bulletin, 47:586590.
Wallace, S. C. and Hulbert, R. C. 2005. Morphology and systematics of the Hemphillian (Miocene) Gray Fossil Site tapir. Journal of Vertebrate Paleontology, 25(3, supplement): 127A.
Wallace, S. C., Nave, J., and Burdick, K. 2002. Preliminary report on the recently discovered Gray Fossil Site (Miocene), Washington Co., Tennessee: with comments on observed paleopathologies - the advantages of a large sample. Journal of Vertebrate Paleontology, 22(3, supplement): 117A.
Wallace, S. C. and Wang, X. 2004. Two new carnivores from an unusual late Tertiary forest biota in eastern North America. Nature, 431:556559.
Webb, S. D. 1969. The Pliocene Canidae of Florida. Bulletin of the Florida State Museum, 14:273308.
Webb, S. D. 1973. Pliocene pronghorns of Florida. Journal of Mammalogy, 54:203221.
Webb, S. D. 2000. Evolutionary history of New World Cervidae, pp. 3864. In Vrba, E. and Schaller, G. (eds.), Antelopes, Deer, and Relatives: Fossil Record Behavioral Ecology, Systematics and Conservation. Yale University Press, New Haven.
Webb, S. D. and Hulbert, R. C. 1986. Systematics and evolution of Pseudhipparion (Mammalia, Equidae) from the Late Neogene of the Gulf Coastal Plain and the Great Plains, pp. 237272. In Flanagan, K. M. and Lillegraven, J. A. (eds.), Vertebrates, Phylogeny, and Philosophy. University of Wyoming Contributions to Geology, Special Paper 3.
Webb, S. D., Hulbert, R. C., Morgan, G. S., and Evans, H. F. 2008. Terrestrial mammals of the Palmetto Fauna (early Pliocene, latest Hemphillian) from the Central Florida Phosphate District, pp. 293312. In Wang, X. and Barnes, L. G. (eds.), Geology and Vertebrate Paleontology of Western and Southern North America, Contributions in Honor of David P. Whistler. Natural History Museum of Los Angeles County Science Series, Number 41.
Webb, S. D. and Tessman, N. 1968. A Pliocene vertebrate fauna from low elevation in Manatee County, Florida. American Journal of Science, 266: 777811.
White, T. E. 1942. Additions to the fauna of the Florida phosphates. Proceedings of the New England Zoology Club, 21:8791.
Yarnell, K. L. 1980. Systematics of late Miocene Tapiridae (Mammalia, Perissodactyla) from Florida and Nebraska. Unpublished , , Gainesville, 125 p.

Cranial morphology and systematics of an extraordinary sample of the Late Neogene dwarf tapir, Tapirus polkensis (Olsen)

  • Richard C. Hulbert (a1), Steven C. Wallace (a2), Walter E. Klippel (a3) and Paul W. Parmalee (a4)


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