Abstract
A systematic revision based on a morphological and statistical analysis recognizes the following Bridgerian and Uintan brontothere species as valid: Mesatirhinus megarhinus, Metarhinus fluviatilis, Metarhinus diploconus, Sphenocoelus uintensis, Sphenocoelus intermedius, Sphenocoelus hyognathus, Telmatherium validus, Protitanotherium emarginatum, Pseudodiplacodon progressum, Eotitanotherium osborni, Sthenodectes incisivum, and Metatelmatherium ultimum. The valid Bridgerian genus Palaeosyops is not discussed in the present paper. Mesatirhinus, Metarhinus, and Sphenocoelus are grouped within the subfamily Dolichorhininae; Telmatherium, Protitanotherium, Pseudodiplacodon, and Eotitanotherium are grouped within the subfamily Brontotheriinae; and Sthenodectes and Metatelmatherium are listed as Brontotheriidae incertae sedis. It is possible that Metatelmatherium may be a member of the subfamily Brontotheriinae or that it may be closely related to Sthenodectes, justifying the recognition of another North American brontothere subfamily, the Metatelmatheriinae.References
Abdi, H. (2007) Bonferroni and Sidak corrections for multiple comparisons. In: Salkind, N. J. (Ed), Encyclopedia of Measurement and Statistics. Sage Publications, Thousand Oaks, pp. 103–107.
Chow, M.M. (1957) On some Eocene and Oligocene mammals from Kwangsi and Yunnan. Vertebrata Palasiatica, 1(3), 201–214.
Colbert, E.H. (1938) Fossil mammals from Burma in the American Museum of Natural History. Bulletin of the American Museum of Natural History, 74 (6), 435–436.
Cook, H.J. (1926) New Eocene titanotheres from Moffat County, Colorado. Proceedings of the Colorado Museum of Natural History, 6 (3), 12–18.
Cope, E.D. (1872) Second notice of extinct vertebrates from Bitter Creek, Wyoming. Paleontological Bulletin, 7, 1–2.
Cope, E.D. (1873) On the new perissodactyls from the Bridger Eocene. Paleontological Bulletin, 11, 1–2.
Ding, S., Zheng, J., Zhang, Y. & Tong, Y. (1977) The age and characteristics of the Liuniu and the Dongjun faunas, Bose Basin of Guangxi. Vertebrata Palasiatica, 15 (1), 35–45.
Douglass, E. (1909) Preliminary descriptions of some new titanotheres from the Uinta deposits. Annals of the Carnegie Museum, 6(2), 304–313.
Earle, C. (1891) On a new species of Palaeosyops, Palaeosyops megarhinus sp.nov. American Naturalist, 25 (289), 45–47.
Earle, C. (1892) A memoir upon the genus Palaeosyops Leidy and its allies. Journal of the Academy of Natural Sciences of Philadelphia, 2nd series, 9(6), 267–388.
Golz, D.J. (1976) Eocene Artiodactyla of southern California. Natural History Museum of Los Angeles County Bulletin, 26, 1–85.
Golz, D.J. & Lillegraven, J.A. (1977) Summary of known occurrences of terrestrial vertebrates from Eocene strata of southern California. Contributions to Geology, University of Wyoming, 15(1), 43–65.
Granger, W. (1909) Faunal horizons of the Washakie Formation of southern Wyoming. Bulletin of the American Museum of Natural History, 26 (3), 13–24.
Granger, W. & Gregory, W.K. (1938) Addendum. A new titanothere genus from the upper Eocene of Mongolia and North America. In: Colbert, E. H., Fossil mammals from Burma in the American Museum of Natural History. Bulletin of the American Museum of Natural History, 74 (6), 435–436.
Granger, W. & Gregory, W.K. (1943) A revision of the Mongolian titanotheres. Bulletin of the American Museum of Natural History, 80 (10), 349–389.
Gregory, W.K. (1912) Note on the upper Eocene titanotheroid Telmatherium? incisivum Douglass, from the Uinta Basin. Science, new ser., 35 (901), 545.
Gregory, W.K. (1920) Studies in comparative myology and osteology, no. IV. A review of the evolution of the lachrymal bone of vertebrates with special reference to that of mammals. Bulletin of the American Museum of Natural History, 42, 95–263.
Gunnell, G.F. & Yarborough, V.L. (2000) Brontotheriidae (Perissodactyla) from the late Early and middle Eocene (Bridgerian), Wasatch and Bridger Formations, southern Green River Basin, south western Wyoming. Journal of Vertebrate Paleontology, 20 (2), 349–368.
Hatcher, J.B. (1895) On a new species of Diplacodon, with a discussion of the relations of that genus to Telmatotherium. American Naturalist, 29, 1084–1090.
Hay, O.P. (1902) Bibliography and catalogue of the fossil Vertebrata of North America. Bulletin of the United States Geological Survey, 179, 1–868.
Howard, R.W. (1975) The dawnseekers: the first history of American paleontology. Harcourt, Brace, Jovanovich, Inc., New York, 314 pp.
Leidy, J. (1872) On fossils from Wyoming. Proceedings of the Academy of Natural Sciences of Philadelphia, 24, 277.
Li, C. & Ting, S. (1983) The Paleogene mammals of China. Bulletin of the Carnegie Museum of Natural History, 21, 1–93.
Lucas, S.G. (1983) Protitanotherium (Mammalia, Perissodactyla) from the Eocene Baca Formation, west-central New Mexico. New Mexico Journal of Sciences, 23 (2), 39–47.
Mader, B.J. (1989) The Brontotheriidae: a systematic revision and preliminary phylogeny of North American genera. In: Prothero, D. R. & Schoch, R. M. (Eds), The Evolution of Perissodactyls. Oxford University Press, Oxford Monographs on Geology and Geophysics (15), New York, pp. 458–484.
Mader, B.J. (1991) The systematics and phylogeny of North American Eocene brontotheres (Mammalia, Perissodactyla). University of Massachusetts, Amherst, 388 pp.
Mader, B.J. (1998) Brontotheriidae. In: Janis, C. M., Scott, K. M. & Jacobs, L. L. (Eds), Evolution of Tertiary Mammals of North America. Volume I: Terrestrial Carnivores, Ungulates, and Ungulatelike Mammals. Cambridge University Press, New York, pp. 525–536.
Mader, B.J. (2000) Pseudodiplacodon, a new generic name for Diplacodon progressum Peterson (Mammalia, Perissodactyla, Brontotheriidae). Journal of Vertebrate Paleontology, 20(1), 164–166.
Mader, B.J. & Alexander, J.P. (1995) Megacerops kuwagatarhinus n. sp., and unusual brontothere (Mammalia, Perissodactyla) with distally forked horns. Journal of Paleontology, 69(3), 581–587.
Marsh, O.C. (1872) Preliminary description of new Tertiary mammals, Part I. American Journal of Science, 3d series, 4, 122–130.
Marsh, O.C. (1873) Notice of new Tertiary mammals. American Journal of Science, 3d series, 5, 407–411, 485–488.
Marsh, O.C. (1875) Notice of new Tertiary mammals, IV. American Journal of Science, 3d series, 9, 239–250.
Marsh, O.C. (1880) List of genera established by Professor O.C. Marsh, 1862–1879. New Haven, 12 pp.
Martill, D.M. & Turnbull, W.D. (1986) Taphonomy and preservation of a monospecific titanothere assemblage from the Washakie Formation (late Eocene), southern Wyoming. Fourth North American Paleontological Convention, abstracts, A53.
Matthew, W.D. (1899) A provisional classification of the fresh-water Tertiary of the west. Bulletin of the American Museum of Natural History, 12 (3), 19–75.
McCarroll, S.M., Flynn J.J. & Turnbull, W.D. (1996) The mammalian faunas of the Washakie Formation, Eocene age of southern Wyoming. Part III. The perissodactyls. Fieldiana, new series, 33, 1–38.
McKenna, M.C. & Bell, S.K. (1997) Classification of mammals above the species level. Columbia University Press, New York, 631 pp.
Mihlbachler, M.C. (2005) Phylogenetic systematics of the Brontotheriidae (Mammalia, Perissodactyla). Columbia University, New York, 763 pp.
Mihlbachler, M.C., Lucas, S.G. & Emry, R.J. (2004) The holotype specimen of Menodus giganteus, and the “insoluble” problem of Chadronian brontothere taxonomy. In: Lucas, S. G., Zeigler, K. E. & Kondrashov, P. E. (Eds). Paleogene mammals. New Mexico Museum of Natural History and Science Bulletin 26, pp. 129–135.
Osborn, H.F. (1890) The Mammalia of the Uinta formation, Part 3, the Perissodactyla. Transactions of the American Philosophical Society, new series, 16 (9), 461–572.
Osborn, H.F. (1895) Fossil mammals of the Uinta Basin, Expedition of 1894. Bulletin of the American Museum of Natural History, 7 (2), 71–105.
Osborn, H.F. (1908) New or little known titanotheres from the Eocene and Oligocene. Bulletin of the American Museum of Natural History, 24 (32), 599–617.
Osborn, H.F. (1914) Recent results in the phylogeny of the titanotheres. Bulletin of the Geological Society of America, 25, 403–405.
Osborn, H.F. (1929) The titanotheres of ancient Wyoming, Dakota, and Nebraska. U.S. Geological Survey, Monograph 55 (2 volumes), 953pp.
Osborn, H.F., Scott, W.B. & Speir, F. (1878) Paleontological report of the Princeton scientific expedition of 1877. Contributions from the Museum of Geology and Archaeology of Princeton College, 1, 1–106.
Owen, R. (1848) Description of teeth and portions of jaws of two extinct anthracotherioid quadrupeds (Hyopotamus vectianus and Hyop. bovinus) discovered by the Marchioness of Hastings in the Eocene deposits on the N.W.
coast of the Isle of Wight: with an attempt to develop Cuvier's idea of the classification of pachyderms by the number of their toes. Quarterly Journal of the Geological Society of London, 4, 103–141.
Perneger, T.V. (1998) What is wrong with Bonferroni adjustments. British Medical Journal, 316, 1236–1238.
Peterson, O.A. (1914a) A new titanothere from the Uinta Eocene. Annals of the Carnegie Museum, 9 (1–2), 29–52.
Peterson, O.A. (1914b) A small titanothere from the lower Uinta beds. Annals of the Carnegie Museum, 9 (1–2), 53–57.
Peterson, O.A. (1914c) Correction of generic name. Annals of the Carnegie Museum, 9, 220.
Peterson, O.A. (1924) Osteology of Dolichorhinus longiceps Douglass, with a review of the species of Dolichorhinus in the order of their publication. Memoirs of the Carnegie Museum, 9 (4), 405–473.
Peterson, O.A. (1934) New titanotheres from the Uinta Eocene in Utah. Annals of the Carnegie Museum, 22 (3–4), 351–361.
Pilgrim, G.E. (1925) The Perissodactyla of the Eocene of Burma. Memoirs of the Geological Survey of India (new series), 8 (3), 1–28.
Prothero, D.R. & Schoch, R.M. (1989) Origin and evolution of the Perissodactyla: summary and synthesis. In: Prothero, D. R. & Schoch, R. M. (Eds), The Evolution of Perissodactyls. Oxford University Press, Oxford Monographs on Geology and Geophysics (15), New York, 504–529.
Ride, W.D.L., Cogger, H. G., Dupuis, C., Kraus, O., Minelli, A., Thompson, F. C. & Tubbs, P. K. (1999) International Code of Zoological Nomenclature (4th edition), International Trust for Zoological Nomenclature, London. Available from http://www.iczn.org/iczn/ (accessed 22 April 2008).
Riggs, E.S. (1912) New or little known titanotheres from the lower Uinta Formations. Field Museum of Natural History Publication 159, Geological Series, 4 (2), 17–41.
Schoch, R.M. & Lucas, S.G. (1985) The Brontotheriidae a group of Eocene and Oligocene perissodactyls from North America, Asia, and Eastern Europe. In: Fuller, W.A., Nietfeld, M.T. & Harris, M.A. (Eds), Abstracts of papers and posters, Fourth International Theriological Congress, Edmonton, Alberta, Canada, Abstract 0558.
Simpson, G.G. (1945) The principles of classification and a classification of mammals Bulletin of the American Museum of Natural History, 85, 1–350.
Simpson, G.G., Roe, A. & Lewontin, R.C. (1960) Quantitative Zoology. Harcourt, Brace and World, Inc., New York, 440 pp.
Stock, C. (1937) An Eocene titanothere from San Diego County, California, with remarks on the age of the Poway Conglomerate. Proceedings of the National Academy of Sciences, Contribution 219, 23 (2), 48–53.
Tang, X. & Chow, M. (1964) A review of vertebrate-bearing Lower Tertiary of south China. Vertebrata Palasiatica, 8(2), 119–133.
Tang, Y., You, Y., Xu, Q., Qiu, Z. & Hu, Y. (1974) The lower Tertiary of the Bose and Yongle Basins of the Guangxi Zhuang Autonomous Region. Vertebrata Palasiatica, 12(4), 279–292.
Turbull, W.D. & Martill, D.M. (1988) Taphonomy and preservation of a monospecific titanothere assemblage from the Washakie Formation (late Eocene), southern Wyoming. An ecological accident in the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology, 63, 91–108.
Wall, W.P. (1989) The phylogenetic history and adaptive radiation of the Amynodontidae. In: Prothero. D. R. & Schoch, R. M. (Eds), The Evolution of Perissodactyls. Oxford University Press, Oxford Monographs on Geology and Geophysics (15), New York, pp. 341–354.
West, R.M. & Dawson, M.R. (1975) Eocene fossil Mammalia from the Sand Wash Basin, northwestern Moffat County, Colorado. Annals of the Carnegie Museum, 45 (11), 231–253.
Wilson, J.A. (1977) Early Tertiary vertebrate faunas Big Bend area Trans-Pecos Texas: Brontotheriidae. The Pearce - Sellard Series, 25, 1–17.
Woods, S.C. & Stucky, R.K. (1992) Catalogue of the type specimens of fossil vertebrates Denver Museum of Natural History. Proceedings of the Denver Museum of Natural History, 3 (2), 1–12.