New large-bodied mammals from the Late Oligocene site of Chilga, Ethiopia

• Autorzy: Sanders W J , Kappelman J. , Rasmussen D.T.
• Języki publikacji: EN

Abstrakty

EN

Newly recovered fossil proboscideans and embrithopods from Chilga, Ethiopia are described and evaluated taxonomically. They are dated to ca. 28–27 Ma (late Oligocene), temporally intermediate between late Eocene–early Oligocene Afro−Arabian faunas dominated by archaic, endemic taxa, and replacement faunas of the early Miocene marked by a massive influx of Eurasian migrants. The paucity of similar−aged sites in Afro−Arabia makes Chilga critical for delineating the initiation and sequence of this faunal turnover. While most of the genera present at Chilga persist from older Afro−Arabian localities, at higher elevation and farther inland than elsewhere, there are no Eurasian mammals in the fauna. However, the archaic endemics from Chilga differ morphometrically from their older congeners, and include a new embrithopod, Arsinoitherium giganteum sp. nov., and novel species of elephantiform proboscideans, Phiomia major sp. nov., aff. Palaeomastodon sp. nov. A, and aff. Palaeomastodon sp. nov. B. New, primitive deinotheres and gomphotheres also occur at Chilga, extending the fossil records of these proboscideans considerably back in time. The Chilga deinothere, Chilgatherium harrisi sp. nov., differs sufficiently from Prodeinotherium and Deinotherium to be placed in its own subfamily, Chilgatheriinae subfam. nov. The Chilga gomphothere is smaller than Miocene elephantoids, and is referred to cf. Gomphotherium sp. nov. Together, this evidence suggests that indigenous Afro−Arabian taxa had greater ecological versatility than previously suspected and continued to enjoy successful evolutionary trajectories into the late Paleogene. Thus, as they spread into Afro−Arabia, new immigrants from Eurasia may have encountered vibrant local mammalian communities. The demise of many endemic inhabitants followed and remains poorly understood.

Słowa kluczowe

EN

Ethiopia; Gomphotheriidae; Arsinoitheriidae; mammal; fossil; large-bodied mammal; Oligocene; Deinotheriidae; Chilga; paleontology; Palaeomastodontidae

• Wydawca: -
• Czasopismo: Acta Palaeontologica Polonica
• Rocznik: 2004
• Tom: 49
• Numer: 3
• Opis fizyczny: p.365-392,fig.,ref.

Twórcy

autor

Sanders W J

• University of Michigan, 1109 Geddes Avenue, Ann Arbor, Michigan 48109, USA

autor

Kappelman J.

autor

Rasmussen D.T.

Bibliografia

• Adams, C.G., Gentry, A.W., and Whybrow, P.J. 1983. Dating the terminal Tethyan events. In: J. Meulenkamp (ed.), Reconstruction of Marine Environments. Utrecht Micropaleontological Bulletins 30: 273–298.
• Adams, C.G., Bayliss, D.D., and Whittaker, J.E. 1999. The terminal Tethyan event: a critical review of the conflicting age determinations for the disconnection of the Mediterranean from the Indian Ocean.In: P.J. Whybrow and A. Hill (eds.), Fossil Vertebrates of Arabia, 477–484. Yale University Press, New Haven.
• Agustí, J. and Antón, M. 2002. Mammoths, Sabertooths, and Hominids. 65 Million Years of Mammalian Evolution in Europe. 313 pp. Columbia University Press, New York.
• Andrews, C.W. 1901. Fossil vertebrates from Egypt. Zoologist 4: 318–319.
• Andrews, C.W. 1904. Further notes on the mammals of the Eocene of Egypt. Geological Magazine Series 5, 1: 109–115.
• Andrews, C.W. 1905. Note on the species of Palaeomastodon. Geological Magazine Series 5, 2: 512–563.
• Andrews, C.W. 1906. A Descriptive Catalogue of the Tertiary Vertebrata of the Fayum, Egypt. 324 pp. British Museum of Natural History, London.
• Andrews, C.W. 1908. On the skull, mandible, and milk dentition of Palaeomastodon, with some remarks on the tooth change in the Proboscidea in general. Philosophical Transactions of the Royal Society of London B 199: 393–407.
• Andrews, C.W. 1911. On a new species of Dinotherium (Dinotherium hobleyi) from British East Africa. Proceedings of the Zoological Society of London 1911: 943–945.
• Andrews, C.W. and Beadnell, H.J.L. 1902. A preliminary note on some new mammals from the Upper Eocene of Egypt. Survey Department, Public Works Ministry, Cairo 1902: 1–9.
• Andrews, P. and Van Couvering, J.A.H. 1975. Palaeoenvironments in the East African Miocene. In: F. Szalay (ed.), Approaches to Primate Paleobiology, Contributions to Primatology, Vol. 5, 62–103. Karger, Basel.
• Antunes, M.T. 1989. The proboscideans [sic] data, age and paleogeography: evidence from the Miocene of Lisbon. In: E.H. Lindsay, V. Fahlbusch, and P. Mein (eds.), European Neogene Mammal Chronology, 253–262, Plenum Press, New York.
• Arambourg, C. 1961. Note préliminaire sur quelques Vertébrés nouveaux du Burdigalien de Libye. Comptes Rendus Sommaire des Seances de la Société Géologique de France 4: 107–109.
• Arambourg, C. and Burollet, P.F. 1962. Restes de Vertébrés oligocènes en Tunisie centrale. Comptes Rendus Sommaire des Séances de la Société Géologique de France 2: 42–43.
• Arambourg, C., Kikoine, J., and Lavocat, R. 1951. Découverte du genre Moeritherium Andrews dans le Tertiaire continental du Soudan. Comptes rendus des Séances de l’Académie des Sciences 233: 68–70.
• Arambourg, C. and Magnier, P. 1961. Gisements de vertébrés dans le bassin tertaire de Syrte (Libie). Comptes Rendus Hebdomadaires des Séances de l’Academie de Sciences 252: 1181–1183.
• Assefa, G. and Saxena, G.N. 1983. A review of Ethiopian lignites: occurrence, prospects, and possibilities. Energy Exploration and Exploitation 3: 35–42.
• Bachmann, I. 1875. Beschreibung eines Unterkiefers von Dinotherium bavaricum H. v. Meyer. Abhandlungen der Schweizerischen Paläontologischen Gesellschaft 2: 1–20.
• Barry, J.C. and Flynn, L.J. 1989. Key biostratigraphic events in the Siwalik sequence.In: E.H. Lindsay, V. Fahlbusch, and P. Mein (eds.), European Neogene Mammal Chronology, 557–571. Plenum Press, New York.
• Beadnell, H.J.C. 1902. A preliminary note on Arsinoitherium zitteli Beadnell, from the Upper Eocene strata of Egypt. 4 pp. Egyptian Survey Department, Public Works Ministry, Cairo.
• Behrensmeyer, A.K., Potts, R., Plummer, T., Tauxe, L., Opdyke, N., and Jorstad, T. 1995. The Pleistocene locality of Kanjera, Western Kenya: stratigraphy, chronology and paleoenvironments. Journal of Human Evolution 29: 247–274.
• Bergounioux, F.−M. and Crouzel, F. 1959. Nouvelles observations sur un petit Mastodonte du Cartennien de Kabylie.Comptes Rendus Sommaire des Seances de la Société Géologique de France 1959: 101–102.
• Bergounioux, F.−M. and Crouzel, F. 1962. Les Déinothéridés d’Europe. Annales de Paléontologie 48: 1–56.
• Bernor, R.L., Brunet, M., Ginsburg, L., Mein, P., Pickford, M., Rögl, F., Sen, S., Steininger, F., and Thomas, H. 1987. A consideration of some major topics concerning Old World Miocene mammalian chronology, migrations and paleogeography. Geobios 20: 431–439.
• Bishop, W.W., Miller, J.A., and Fitch, F.J. 1969. New potassium−argon age determinations relevant to the Miocene fossil mammal sequence in East Africa. American Journal of Science 267: 669–699.
• Bonaparte, C.L. 1845. Catalogo Metodico dei Mammiferi Europei. 36 pp. Valenciennes, Milan.
• Borissiak, A. 1929. On a new direction in the adaptive radiation of mastodonts. Palaeobiologica 2: 19–33.
• Boschetto, H.B., Brown, F.H., and McDougall, I. 1992. Stratigraphy of the Lothidok Range, northern Kenya, and K/Ar ages of its Miocene primates. Journal of Human Evolution 22: 47–71.
• Cande, S.C. and Kent, D.V. 1995. Revised calibration of the geomagnetic polarity timescale for the Late Cretaceous and Cenozoic. Journal of Geophysical Research 100: 6093–6095.
• Checcia−Rispoli, G. 1914. Sul “Mastodon angustidens” Cuvier dei dintorni di Bugio in provincia di Girgenti. Giornale di Scienze Naturali ed Economiche di Palermo 30: 285–296.
• Coiffait, P.E., Coiffait, B., Jaeger, J.−J., and Mahboubi, M. 1984. Un nouveau gisement à mammifères fossiles d’âge éocène supérieur sur le versant Sud des Nementcha (Algérie orientale); découverte des plus anciens rongeurs d’Afrique. Comptes Rendus de l’Académie des Sciences de Paris 299: 893–898.
• Cooper, C.F. 1922. Miocene Proboscidia [sic] from Baluchistan. Proceedings of the Zoological Society of London 42: 606–626.
• Cope, E.D. 1882. The classification of the ungulate Mammalia. Proceedings of the American Philosophical Society 20: 438–461.
• Coppens, Y., Maglio, V.J., Madden, C.T., and Beden, M. 1978. Proboscidea. In: V.J. Maglio and H.B. S. Cooke (eds.), Evolution of African Mammals, 336–367. Harvard University Press, Cambridge.
• Coryndon, S.C. and Savage, R.J.G. 1973. The origin and affinities of African mammal faunas. Special Papers in Paleontology (The Palaeontological Association, London) 12: 121–135.
• Court, N. 1989. Morphology, Functional Morphology and Phylogeny of Arsinotherium (Mammalia, Embrithopoda). 372 pp. Unpublished thesis, University of Bristol, United Kingdom.
• Court, N. 1990. Periotic anatomy of Arsinotherium (Mammalia, Embrithopoda) and its phylogenetic implications. Journal of Vertebrate Paleontology 10: 170–182.
• Court, N. 1992a. The skull of Arsinoitherium (Mammalia, Embrithopoda) and the higher order interrelationships of ungulates. Palaeovertebrata 22: 1–43.
• Court, N. 1992b. A unique form of dental bilophodonty and a functional interpretation of peculiarities in the masticatory system of Arsinoitherium (Mammalia, Embrithopoda). Historical Biology 6: 91–111.
• Court, N. 1993. Morphology and functional anatomy of the postcranial skeleton in Arsinoitherium (Mammalia, Embrithopoda). Palaeontolographica Abhandlungen A 226: 125–169.
• Court, N. 1995. A new species ofNumidotherium (Mammalia: Proboscidea) from the Eocene of Libya and the early phylogeny of the Proboscidea. Journal of Vertebrate Paleontology 15: 650–671.
• Depéret, C. 1897. Découverte du Mastodon angustidens dans l’étage Cartennien de Kabylie. Bulletin de la Société Géologique de France Series 3, 25: 518–521.
• Drooger, C.W. 1979. Marine connections of the Neogene Mediterranean, deduced from the evolution and distribution of larger foraminifera. Annales Géologiques des Pays Helléniques 1: 361–369.
• Drooger, C.W. 1993. Radial foraminifera; morphometrics and evolution. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afdeeling Natuurkunde, Eerste Reeks 41: 1–242.
• Éhik, J. 1930. Prodinotherium hungaricum n. g., n. sp. Geologica Hungarica, Series Palaeontologica 6: 1–24.
• Eizirik, E., Murphy, W.J., and O’Brien, S.J. 2001. Molecular dating and biogeography of the early placental mammal radiation. The Journal of Heredity 92: 212–219.
• El−Khashab, B. 1977. Approach on genus Arsinoitherium and the diagnosis of its closely related sympatric fossil species, in Lower Oligocene of Fayum Province, Egypt. The Geological Survey of Egypt and Mining Authority, Cairo 62: 29–39.
• El−Khashab, B. 1979. A brief account on Egyptian Paleogene Proboscidea. Annals of the Geological Survey of Egypt 9: 245–260.
• Feseha, M.Y. 2002. Sequence Stratigraphy, Petrography, and Geochronology of the Chilga Rift Basin Sediments, Northwest Ethiopia. 330 pp. Unpublished Ph.D. dissertation, Department of Geological Sciences, University of Texas at Austin.
• Fleagle, J.G. 1986. The fossil record of early catarrhine evolution. In: B. Wood, L. Martin, and P. Andrews (eds.), Major Topics in Primate and Human Evolution ,130–149. Cambridge University Press, Cambridge.
• Flower, B.P. and Kennett, J.P. 1994. The middle Miocene climatic transition: East Antarctic ice sheet development, deep ocean circulation and global carbon cycling. Palaeogeography, Palaeoclimatology, Palaeoecology 108: 537–555.
• Fourtau, R. 1920. Contribution a l’Étude Vertébrés Miocènes de l’Égypte. 121 pp. Survey Department, Ministry of Finance, Egypt, Government Press, Cairo [2nd printing].
• Gagnon, M. 1997. Ecological diversity and community ecology in the Fayum sequence (Egypt). Journal of Human Evolution 32: 133–160.
• Gaziry, A.W. 1976. Jungtertiäre Mastodonten aus Anatolien (Türkei). Geologisches Jahrbuch 22: 3–143.
• Gaziry, A.W. 1987. New mammals from the Jabal Zaltan site, Libya. Senckenbergiana lethaea 68: 69–89.
• Gebo, D.L., MacLatchy, L., Kityo, R., Deino, A., Kingston, J., and Pilbeam, D. 1997. A hominoid genus from the Early Miocene of Uganda. Science 276: 401–404.
• Gentry, A.W. 1987. Mastodons from the Miocene of Saudi Arabia. Bulletin of the British Museum (Natural History) (Geology) 41: 395–407.
• Gheerbrant, E. 1998. The oldest known proboscidean and the role of Africa in the radiation of modern orders of placentals. Bulletin of the Geological Society of Denmark 44: 181–185.
• Gheerbrant, E., Sudre, J., and Cappetta, H. 1996. A Palaeocene proboscidean from Morocco. Nature 383: 68–70.
• Gheerbrant, E., Sudre, J., Cappetta, H., and Bignot, G. 1998. Phosphatherium escuilliei du Thanétien du Bassin des Ouled Abdoun (Maroc), plus ancien Proboscidean (Mammalia) d’Afrique. Geobios 30: 247–269.
• Gheerbrant, E., Sudre, J., Cappetta, H., Iarochène, M., Amaghzaz, M., and Bouya, B. 2002. A new large mammal from the Ypresian of Morocco: evidence of surprising diversity of early proboscideans. Acta Palaeontologica Polonica 47: 493–506.
• Gingerich, P.D. 1993. Oligocene age of the Gebel Qatrani Formation, Fayum, Egypt. Journal of Human Evolution 24: 207–218.
• Ginsburg, L. 1989. The faunas and stratigraphical subdivisions of the Orleanian in the Loire Basin (France). In: E.H. Lindsay, V. Fahlbusch, and P. Mein (eds.), European Neogene Mammal Chronology, 157–176. Plenum Press, New York.
• Göhlich, U.B. 1998. Elephantoidea (Proboscidea, Mammalia) aus dem Mittel− und Obermiozän der Oberen Süßwassermolasse Süddeutschlands: Odontologie und Osteologie. Münchner Geowissenschafliche Abhandlungen A 36: 1–245.
• Göhlich, U.B. 1999. Order Proboscidea. In: G. Rossner and K. Heissig (eds.), The Miocene Land Mammals of Europe, 57–75. F. Pfeil, Munich.
• Gorodisky, H., and Lavocat, R. 1953. Première découverte de mammiferes dans le Tertiaire (Lutétian) du Sénégal. Comptes Rendus Sommaires de la Société Géologique de France 15: 314–316.
• Gräf, I.E. 1957. Die Prinzipien der Artbestimmung bei Dinotherium. Palaeontographica Abhandlungen A 108: 131–185.
• Gray, J.E. 1821. On the natural arrangements of vertebrose animals. London Medical Repository 15: 296–310.
• Gregory, W.K. 1910. The orders of mammals. Bulletin of the American Museum of Natural History 27: 1–524.
• Guerin, C. and Pickford, M. 2003. Ougandatherium napakense nov. gen. nov. sp., le plus ancien Rhinocerotidae Iranotheriinae d’Afrique. Annales de Paléontologie 89: 1–35.
• Gunnell, G.F., Jacobs, B.F., Herendeen, P.S., Head, J.J., Kowalski, E., Msuya, C.P., Mizambwa, F.A., Harrison, T., Habersetzer, J., and Storch, G. 2003. Oldest placental mammal from sub−Saharan Africa: Eocene microbat from Tanzania—evidence for early evolution of sophisticated echolocation. Palaeontologia Electronica 5 (2): 1–10.
• Hamilton, W.R. 1973. A lower Miocene mammalian fauna from Siwa, Egypt. Palaeontology 16: 275–281.
• Haq, B.U., Hardenbol, J., and Vail, P.R. 1987. Chronology of fluctuating sea levels since the Triassic. Science 235: 1156–1167.
• Harris, J.M. 1969. Prodeinotherium from Gebel Zelten, Libya. 303 pp. Unpublished Ph.D. Thesis, University of Bristol, Bristol.
• Harris, J.M. 1973. Prodeinotherium from Gebel Zelten, Libya. Bulletin of the British Museum (Natural History) (Geology) 23: 285–350.
• Harris, J.M. 1975. Evolution of feeding mechanisms in the family Deinotheriidae (Mammalia: Proboscidea). Zoological Journal of the Linnean Society 56: 331–362.
• Harris, J.M. 1977. Deinotheres from South Africa. South African Journal of Science 73: 281–282.
• Harris, J.M. 1978. Deinotherioidea and Barytherioidea. In: V.J. Maglio and H.B.S. Cooke (eds.), Evolution of African Mammals, 315–332. Harvard University Press, Cambridge.
• Harris, J.M. 1983. Family Deinotheriidae. In: J.M. Harris (ed.), Koobi Fora Research Project, Vol. 2, 22–39. Clarendon Press, Oxford.
• Harris, J.M. 1987. Fossil Deinotheriidae from Laetoli. In: M.D. Leakey and J.M. Harris (eds.), Laetoli, A Pliocene Site in Northern Tanzania, 294–297. Clarendon Press, Oxford.
• Harzhauser, M., Piller, W.E., and Steininger, F.F. 2002. Circum−Mediterranean Oligo−Miocene biogeographic evolution—the gastropods’ point of view. Palaeogeography, Palaeoclimatology, Palaeoecology 183: 103–133.
• Hay, O.P. 1922. Further observations on some extinct elephants. Proceedings of the Biological Society of Washington 35: 97–101.
• Hill, A., Drake, R., Tauxe, L., Monaghan, M., Barry, J.C., Behrensmeyer, A.K., Curtis, G., Jacobs, B.F., Jacobs, L., Johnson, N., and Pilbeam, D. 1985. Neogene palaeontology and geochronology of the Baringo Basin, Kenya. Journal of Human Evolution 14: 759–773.
• Hofmann, C., Courtillot, V., Feraud, G. Rochette, P., Yirgus, G., Ketefo, E., and Pik, R. 1997. Timing of the Ethiopian flood basalt event and implications for plume birth and global change. Nature 389: 838–841.
• Holroyd, P.A. 1999. New Pterodontinae (Creodonta: Hyaenodontidae) from the late Eocene–early Oligocene Jebel Qatrani Formation, Fayum province, Egypt. PaleoBios 19: 1–18.
• Holroyd, P.A. and Maas, M.C. 1994. Paleogeography, paleobiogeography, and anthropoid origins.In: J.G. Fleagle and R.F. Kay (eds.), Anthropoid Origins, 297–334. Plenum Press, New York.
• Huttunen, K.J. 2000. Deinotheriidae (Proboscidea, Mammalia) of the Miocene of Lower Austria, Burgendland, and Franzensbad, Czech Republic: Systematics, Odontology and Osteology. 76 pp. Unpublished Dr. Rer. Nat. Thesis, University of Vienna.
• Illiger, C.D. 1811. Prodromus Systematis Mammalium et Avium Additis Terminis Zoographicis Uttriusque Classis. 301 pp. Salfeld, Berlin.
• Janis, C.M. 1993. Evolution in the context of changing climates, vegetation, and tectonic events. Annual Review of Ecology and Systematics 24: 467–500.
• Kalandadze, N.N. and Rautian, S.A. 1992. The system of mammals and historical zoogeography [in Russian]. In: O.L. Rossolimo (ed.), Filogenetika mlekopitaûŝih: issledovaniâ po faune. Sbornik Trudov Zoologičeskogo Muzeâ Moskovskogo Gosudarstvennogo Universiteta 29: 44–152.
• Kappelman, J. 1992. The age of the Fayum primates as determined by paleomagnetic reversal stratigraphy. Journal of Human Evolution 22: 495–503.
• Kappelman, J., Maas, M.C., Sen, S., Alpagut, B., Fortelius, M., and Lunkka, J.−P. 1996. A new early Tertiary mammalian fauna from Turkey and its paleobiogeographic significance. Journal of Vertebrate Paleontology 16: 592–595.
• Kappelman, J., Rasmussen, D.T., Sanders, W.J., Feseha, M., Bown, T., Copeland, P., Crabaugh, J., Fleagle, J., Glantz, M., Gordon, A., Jacobs, B., Maga, M., Muldoon, K., Pan, A., Pyne, L., Richmond, B., Ryan, T., Seiffert, E., Sen, S., Todd, L., Wiemann, M.C., and Winkler, A. 2003. Oligocene mammals from Ethiopia and faunal exchange between Afro−Arabia and Eurasia. Nature 426: 549–552.
• Kappelman, J., Simons, E.L., and Swisher, C. C., III. 1992. New age determinations for the Eocene–Oligocene boundary sediments in the Fayum Depression, northern Egypt. Journal of Geology 100: 647–668.
• Lambert, W.D., and Shoshani, J. 1998. Proboscidea. In: C.M. Janis, K.M. Scott, and L.L. Jacobs (eds.), Evolution of Tertiary Mammals of North America, 606–621, Cambridge University Press, Cambridge.
• Lankester, E.R. 1903. A new extinct monster. Sphere 1903: 238. London.
• Leakey, M.G., Feibel, C.S., Bernor, R.L., Harris, J.M., Cerling, T.E., Stewart, K.M., Storrs, G.W., Walker, A., Werdelin, L., and Winkler, A.J. 1996. Lothagam: A record of faunal change in the late Miocene of East Africa. Journal of Vertebrate Paleontology 16: 556–570.
• Leakey, M.G., Ungar, P.S., and Walker, A. 1995. A new genus of large primate from the Late Oligocene of Lothidok, Turkana District, Kenya. Journal of Human Evolution 28: 519–531.
• Lehmann, U. 1950. Über Mastodontenreste in der Bayerischen Staatssammlung in München. Palaeontographica Abhandlungen A 99: 121–128.
• Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1: Regnum animale. Editio decima, reformata. 824 pp. Laurentii Salvii, Stockholm.
• Liu, F.−G.R. and Miyamoto, M.M. 1999. Phylogenetic assessment of molecular and morphological data for eutherian mammals. Systematic Biology 48: 54–64.
• MacInnes, D.G. 1942. Miocene and post−Miocene Proboscidia [sic] from East Africa. Transactions of the Zoological Society of London 25: 33–106.
• Madden , C.T. 1972. Miocene mammals, stratigraphy and environment of Muruarot Hill, Kenya. Paleobios 14: 1–12.
• Madsen, O., Scally, M., Douady, C.J., Kao, D.J., DeBry, R.W., Adkins, R., Amrine, H.M., Stanhope, M.J., de Jong, W.W., and Springer, M.S. 2001. Parallel adaptive radiations in two major clades of placental mammals. Nature 409: 610–614.
• Maglio, V.J. Patterns of Faunal Evolution. 1978. In: V.J. Maglio and H.B.S. Cooke (eds.), Evolution of African Mammals, 603–619. Harvard University Press, Cambridge.
• Mahboubi, M., Ameur, R., Crochet, J.Y., and Jaeger, J.J. 1984. Earliest known proboscidean from early Eocene of north−west Africa. Nature 308: 543–544.
• Matsumoto, H. 1922. Revision of Palaeomastodon and Moeritherium. Palaeomastodon intermedius, and Phiomia osborni, new species. American Museum Novitates 51: 1–6.
• Matsumoto, H. 1924. A revision of Palaeomastodon dividing it into two genera, and with descriptions of two new species. Bulletin of the American Museum of Natural History 50: 1–58.
• Matsumoto, H. 1926. Contribution to the knowledge of the fossil Hyracoidea of the Fayum, Egypt, with description of several new species. Bulletin of the American Museum of Natural History 56: 253–350.
• Mazo, A.V. 1996. Gomphotheres and mammutids from the Iberian Peninsula. In: J. Shoshani and P. Tassy (eds.), The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives, 136–142. Oxford University Press, Oxford.
• McKenna, M.C. 1975. Toward a phylogenetic classification of the Mammalia. In: W.P. Luckett and F.S. Szalay (eds.), Phylogeny of the Primates, 21–46. Plenum Press, New York.
• McKenna, M.C. and Bell, S.K. 1997. Classification of Mammals Above the Species Level. 631 pp. Columbia University Press, New York.
• Meyer, G.E. 1978. Hyracoidea. In: V.J. Maglio and H.B.S. Cooke (eds.), Evolution of African Mammals, 284–314. Harvard University Press, Cambridge.
• Murphy, W.J., Eizirik, E., Johnson, W.E., Zhang, Y.P., Ryder, O.A., O’Brien, S.J. 2001. Molecular phylogenetics and the origins of placental mammals. Nature 409: 614–618.
• Nakaya, H. 1993. Les Faunes de Mammifères du Miocène Supérieur de Samburu Hills, Kenya, Afrique de l’Est et L’Environnement des Pré−Hominidés. L’Anthropologie (Paris) 97: 9–16.
• Novacek, M.J. 1992. Fossils, topologies, missing data, and the higher level phylogeny of eutherian mammals. Systematic Biology 41: 58–73.
• Nowak, R.M. and Paradiso, J.L. 1983. Walker’s Mammals of the World, 4th Edition. 1362 pp. Johns Hopkins University Press, Baltimore.
• O’Brien, S.J., Eizirik, E., and Murphy, W.J. 2001. On choosing mammalian genomes for sequencing. Science 292: 2264–2266.
• Osborn, H.F. 1919. Palaeomastodon, the ancestor of the long−jawed mastodons only. Proceedings of the National Academy of Sciences 5: 265–266.
• Osborn, H.F. 1921. The evolution, phylogeny, and classification of the Proboscidea. American Museum Novitates 1: 1–15
• Osborn, H.F. 1936. Proboscidea. A Monograph of the Discovery, Evolution, Migration and Extinction of the Mastodonts and Elephants of the World. Volume I. Moerotherioidea, Deinotherioidea, Mastodontoidea. 802 pp. American Museum Press, New York.
• Osborn, H.F. 1942. Proboscidea. A Monograph of the Discovery, Evolution, Migration and Extinction of the Mastodonts and Elephants of the World. Volume II. Stegodontoidea, Elephantoidea. 870 pp. American Museum Press, New York.
• Palmer, R.W. 1924. An incomplete skull of Dinotherium, with notes on the Indian forms. Palaeontologia Indica 7: 1–14.
• Pickford, M. 1981. Preliminary Miocene mammalian biostratigraphy for Western Kenya. Journal of Human Evolution 10: 73–97.
• Pickford, M. 1986a. Première découverte d’une faune mammalienne terrestre paléogène d’Afrique sub−saharienne. Compte Rendus de l’Academie des Sciences, Paris Série II, 302: 1205–1210.
• Pickford, M. 1986b. Cainozoic paleontological sites of Western Kenya. Münchner Geowissenschaftliche Abhandlungen A 8: 1–151.
• Pickford, M. 2001. Afrochoerodon nov. gen. kisumuensis (MacInnes) (Proboscidea, Mammalia) from Cheparawa, Middle Miocene, Kenya.Annales de Paléontologie 87: 99–117.
• Pickford, M. 2002. Ruminants from the early Miocene of Napak, Uganda. Annales de Paléontologie 88: 85–113.
• Pickford, M. and Andrews, P. 1981. The Tinderet Miocene sequence in Kenya. Journal of Human Evolution 10: 11–33.
• Pickford, M., Senut, B., and Gommery, D. 1999. Sexual dimorphism in Morotopithecus bishopi, an early Middle Miocene hominoid from Uganda, and a reassessment of its geological and biological contexts. In: P. Andrews and P. Banham (eds.), Late Cenozoic Environments and Hominid Evolution, 27–38. The Geological Society, London.
• Pickford, M., Senut, B., Hadoto, D., Muisisi, J., and Kariira, C. 1986. Decouvertes recentes dans le sites miocènes de Moroto (Ouganda Oriental): aspects biostratigraphiques et paleoecologiques. Comptes Rendus de l’Academie des Sciences, Paris Série II, 302: 681–686.
• Pickford, M. and Tassy, P. 1980. A new species of Zygolophodon (Mammalia, Proboscidea) from the Miocene hominoid localities of Meswa Bridge and Moroto (East Africa). Neues Jahrbuch für Geologie und Palaeontologie. Monatshefte 4: 235–251.
• Pilgrim, G.E. 1912. The vertebrate fauna of the Gaj Series in the Bugti Hills and the Punjab. Memoirs of the Geological Survey of India, Palaeontologia Indica 4: 1–83.
• Plavcan, J.M. and Cope, D.A. 2001. Metric variation and species recognition in the fossil record. Evolutionary Anthropology 10: 204–222.
• Rădulescu, C. and Sudre, J. 1985. Crivadiatherium iliescui n. sp., nouvel embrithopode (Mammalia) dans le Paléogène ancien de la Dépression de Hateg (Roumanie). Palaeovertebrata 15: 139–157.
• Rasmussen, D.T. 1989. The evolution of the Hyracoidea: a review of the fossil evidence. In: D.R. Prothero and R.M. Schoch (eds.), The Evolution of Perissodactyls, 57–78. Oxford University Press, Oxford.
• Rasmussen, D.T. 1994. The different meanings of a tarsioid−anthropoid clade and a new model of anthropoid origins. In: J.G. Fleagle and R.F. Kay (eds.), Anthropoid Origins, 335–360. Plenum Press, New York.
• Rasmussen, D.T., Gagnon, M., and Simons, E.L. 1990. Taxepody in the carpus and tarsus of Oligocene Pliohyracidae (Mammalia: Hyracoidea) and the phyletic position of hyraxes. Proceedings of the National Academy of Sciences, USA 87: 4688–4691.
• Rasmussen, D.T., Sanders, W.J., and Kappelman, J. (in preparation). New hyracoids, anthracotheres, and creodonts from the late Paleogene site of Chilga, Ethiopia. Journal of Vertebrate Paleontology.
• Raza, S.M. and Meyer, G.E. 1984. Early Miocene geology and paleontology of the Bugti Hills, Pakistan. Memoirs of the Geological Survey of Pakistan 11: 43–63.
• Roger, J., Pickford, M., Thomas, H., de Lapparent de Broin, F., Tassy, P., Van Neer, W., Bourdillon−de−Gissac, C., and Al−Busaidi, S. 1994. Découverte de Vertébrés Fossiles dans le Miocène de la Région du Huqf au Sultanat d’Oman. Annales de Paléontologie, Paris 80: 253–273.
• Roger, O. 1886. Ueber Dinotherium bavaricum, H. v. Meyer. Palaeontographica 32: 215–225.
• Rögl, F. 1998. Palaeogeographic considerations for Mediterranean and Parathethys seaways (Oligocene to Miocene). Annalen des Naturhistorischen Museums in Wien 99A: 279–310.
• Rögl, F. 1999. Oligocene and Miocene palaeogeography and stratigraphy of the circum−Mediterranean region. In: P.J. Whybrow and A. Hill (eds.), Fossil Vertebrates of Arabia, 485–500. Yale University Press, New Haven.
• Rögl, F. and Steininger, F.F. 1983. Vorn Zerfall der Tethys zu Mediterran und Paratethys: Die neogene Paläogeographie und Palinspastik des zirkum−mediterranen Raumes. Annalen des Naturhistorischen Museums, Wien 85/A: 135–163.
• Rögl, F. and Steininger, F.F. 1984. Neogene Paratethys, Mediterranean and Indo−Pacific seaways: implications for the paleobiogeography of marine and terrestrial biotas. In: P. Brenchly (ed.), Fossils and Climate, 171–200. Wiley, London.
• Sach, V.V.J. and Heizmann, E.P.J. 2001. Stratigraphie und Säugetierfaunen der Brackwassermolasse in der Umgebung von Ulm (Südwestdeutschland). Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 310: 1–95.
• Sahni, M.R. and Tripathi, C. 1957. A new classification of the Indian deinotheres and description of D. orlovii sp. nov. Memoirs of the Geological Society of India, Paleontologia India 33: 1–33.
• Sanders, W.J. 2003. Proboscideans from the Sinap Formation, Central Turkey. In: J. Kappelman, M. Fortelius, S. Sen, and R. Bernor (eds.), Geology and Paleontology of the Miocene Sinap Formation, Central Turkey, 202–219. Columbia University Press, New York.
• Sanders, W.J. and Kappelman, J. 2001. A new late Oligocene proboscidean fauna from Chilga, Ethiopia. Scientific Programme and Abstracts, 8th International Theriological Congress 2001, 120. Sun City, South Africa.
• Sanders, W.J. and Miller, E.R. 2002. New proboscideans from the early Miocene of Wadi Moghara, Egypt. Journal of Vertebrate Paleontology 22: 388–404.
• Sarwar, M. 1977. Taxonomy and distribution of the Siwalik Proboscidea. Bulletin of the Department of Zoology, University of the Punjab 10: 1–172.
• Savage, R.J.G. 1969. Early Tertiary mammal locality in southern Libya. Proceedings of the Geological Society, London 1657: 167–171.
• Sen, S. and Heintz, E. 1979. Palaeoamasia kansui Ozansoy 1966, embrithopode (Mammalia) de l’Eocène d’Anatolie. Annales de Paléontologie (Vértébrés) 65: 73–81.
• Shoshani, J. 1992. The controversy continues: an overview of evidence for Hyracoidea−Tethytheria affinity.Israel Journal of Zoology 38: 233–244.
• Shoshani, J. 1996. Para− or monophyly of the gomphotheres and their position within Proboscidea. In: J. Shoshani and P. Tassy (eds.), The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives, 149–177. Oxford University Press, Oxford.
• Shoshani, J. and Tassy, P. 1996. The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives. 472 pp. Oxford University Press, Oxford.
• Shoshani, J., West, R.M., Court, N., Savage, R.J.G., and Harris, J.M.. 1996. The earliest proboscideans: general plan, taxonomy, and palaeoecology. In: J. Shoshani and P. Tassy (eds.), The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives, 57–75. Oxford University Press, Oxford.
• Shoshani, J., Walter, R.C., Libsekal, Y., Abraha, M., and Berhe, S. 2001. The oldest gomphothere in Africa: A late Oligocene proboscidean mandible from the Dogali Formation, eastern Eritrea. Scientific Programme and Abstracts. 8th International Theriological Congress 2001, 128–129. Sun City, South Africa.
• Simons, E.L. 1968. Part I. African Oligocene mammals: introduction, history of study, and faunal succession. Peabody Museum of Natural History, Yale University, Bulletin 28: 1–21.
• Simons, E.L. and Rasmussen, D.T. 1994. A whole new world of ancestors: Eocene anthropoideans from Africa. Evolutionary Anthropology 3: 128–139.
• Simpson, G.G. 1945. The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 85: 1–350.
• Springer, M.S., Cleven, G.C., Madsen, O., de Jong, W.W., Waddell, V.G., Amrine, H.M., and Stanhope, M.J. 1997. Endemic African mammals shake the phylogenetic tree. Nature 388: 61–64.
• Stanhope, M.J., Waddell, V.G., Madsen, O. de Jong, W.W., Hedges, S.B., Cleven, G.C., Kao, D., and Springer, M.S. 1998. Molecular evidence for multiple origins of Insectivora and for a new order of endemic African insectivore mammals. Proceedings of the National Academy of Sciences, USA 95: 9967–9972.
• Steininger, F.F. and Rögl, F. 1979. The Paratethys history—a contribution towards the Neogene geodynamics of the Alpine Orogene (an abstract). Annales Géologiques des Pays Helléniques 3: 1153–1165.
• Symeonidis, N.K. 1970. Ein Dinotherium−Fund in Zentralmakedonien (Griechenland). Annales geologiques des pays Helleniques 21: 334–341.
• Tanner, L.G. 1978. Embrithropoda [sic]. In: V.J. Maglio and H.B.S. Cooke (eds.), Evolution of African Mammals, 279–283. Harvard University Press, Cambridge.
• Tassy, P. 1977a. Présence du genre Choerolophodon Schlesinger (Proboscidea, Mammalia) dans le Miocène est−africain. Comptes Rendus de l’Academie des Sciences, Série D, 284: 2487–2490.
• Tassy, P. 1977b. Découverte de Zygolophodon turicensis (Schinz) (Proboscidea, Mammalia) au Lieu−Dit Malartic a Simorre, Gers (Vindobonien Moyen); Implications paléoécologiques et biostratigraphiques. Geobios 10: 655–659.
• Tassy, P. 1979. Les Proboscidiens (Mammalia) du Miocène d’Afrique orientale: résultats préliminaires. Bulletin de la Société Géologique de France 21: 265–269.
• Tassy, P. 1983a. Les Elephantoidea Miocènes du Plateau du Potwar, Groupe de Siwalik, Pakistan. Ie Partie: Introduction, Cadre chronologique et géographique, Mammutidés, Amébélodontidés. Annales de Paléontologie, Paris 69: 96–136.
• Tassy, P. 1983b. Les Elephantoidea Miocènes du Plateau du Potwar, Groupe de Siwalik, Pakistan. IIe Partie: Choerolophodontes et Gomphothères. Annales de Paléontologie, Paris 69: 235–297.
• Tassy, P. 1985. La place de mastodontes miocènes de l’Ancien Monde dans la phylogénie des Proboscidea (Mammalia): hypothèses et conjectures. 861 pp. Thèse de Doctorat d’Etat ès Sciences Naturelles, Université Pierre et Marie Curie, Paris.
• Tassy, P. 1986. Nouveaux Elephantoidea (Mammalia) dans le Miocène du Kenya. 135 pp. Cahiers de Paléontologie, Éditions du Centre de la Recherche Scientifique, Paris.
• Tassy, P. 1988. Le statut systématique de l’espèce Hemimastodon crepusculi (Pilgrim): L’éternal problème de l’homologie et de la convergence. Annales de Paléontologie 74: 115–127.
• Tassy, P. 1989. The “Proboscidean Datum Event”: how many proboscideans and how many events? In: E.H. Lindsay, V. Fahlbusch, and P. Mein (eds.), European Neogene Mammal Chronology, 237–252. Plenum Press, New York.
• Tassy, P. 1994a. Origin and differentiation of the Elephantiformes (Mammalia, Proboscidea). Verhandlungen Naturwissenschaftlichen Vereins in Hamburg 34: 73–94.
• Tassy, P. 1994b. Gaps, parsimony, and early Miocene elephantoids (Mammalia), with a re−evaluation of Gomphotherium annectens (Matsumoto, 1925). Zoological Journal of the Linnean Society 112: 101–117.
• Tassy, P. 1996a. Dental homologies and nomenclature in the Proboscidea. In: J. Shoshani and P. Tassy (eds.), The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives, 21–25. Oxford University Press, Oxford.
• Tassy, P. 1996b. Who is who among the Proboscidea? In: J. Shoshani and P. Tassy (eds.), The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives, 39–48. Oxford University Press, Oxford.
• Tassy, P. 1996c. The earliest gomphotheres. In: J. Shoshani and P. Tassy (eds.), The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives, 89–91. Oxford University Press, Oxford.
• Tassy, P. and Pickford, M.. 1983. Un nouveau Mastodonte zygolophodonte (Proboscidea, Mammalia) dans le Miocène Inférieur d’Afrique Orientale: Systématique et Paléoenvironnement. Geobios 16: 53–77.
• Tchernov, E., Ginsburg, L., Tassy, P., and Goldsmith, N.F. 1987. Miocene mammals of the Negev (Israel). Journal of Vertebrate Paleontology 7: 284–310.
• Thomas, H. 1985. The early and middle Miocene land connection of the Afro−Arabian plate and Asia: a major event for hominoid dispersal? In: E. Delson (ed.), Ancestors: The Hard Evidence, 42–50. Alan R. Liss, New York.
• Thomas, H., Roger, J. Sen, S., Bourdillon−de−Grissac, C., and Al−Sulaimani, Z. 1989a. Découverte de Vertébrés fossiles dans l’Oligocène inférieur du Dhofar (Sultanat d’Oman). Geobios 22: 101–120.
• Thomas, H., Tassy, P., and Sen, S. 1989b. Paleogene proboscidean remains from the Southern Dhofar (Sultanate of Oman). Scientific Programme and Abstracts. 5th International Theriological Congress 1989, 166. Sydney, Australia.
• Thomas, H., Roger, J., Sen, S., Pickford, M., Gheerbrant, E., Al−Sulaimani, Z., and Al−Busaidi, S. 1999. Oligocene and Miocene terrestrial vertebrates in the Southern Arabian Peninsula (Sultanate of Oman) and their geodynamic and palaeogeographic settings. In: P.J. Whybrow and A. Hill (eds.), Fossil Vertebrates of Arabia, 430–442. Yale University Press, New Haven.
• Tobien, H. 1971. Moeritherium, Palaeomastodon, Phiomia aus dem Paläogen Nordafrikas und die Abstammung der Mastodonten (Proboscidea, Mammalia). Mitteilungen aus dem Geologischen Institut der Technischen Universität Hannover 10: 141–163.
• Tobien, H. 1973. The structure of the mastodont molar (Proboscidea, Mammalia). Part 1: the bunodont pattern. Mainzer Geowissenschaften Mitteilungsblatt 2: 115–147.
• Tobien, H. 1975. The structure of the mastodont molar (Proboscidea, Mammalia). Part 2: the zygodont and zygobunodont patterns. Mainzer Geowissenschaften Mitteilungsblatt 4: 195–233.
• Tobien, H. 1978. The structure of the mastodont molar (Proboscidea, Mammalia). Part 3: The Oligocene mastodont Genera Palaeomastodon, Phiomia and the Eo/Oligocene Paenungulate Moeritherium. Mainzer Geowissenschaften Mitteilungsblatt 6: 177–208.
• Tobien, H. 1988. Les Proboscidiens Deinotheriidae. Palaeovertebrata, Montpellier, Mémoire extraordinaire 1988: 135–175.
• Tobien, H. 1996. Evolution of zygodons with emphasis on dentition. In: J. Shoshani and P. Tassy (eds.), The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives, 76–85. Oxford University Press, Oxford.
• Tsoukala, E.S. and Melentis, J.K. 1994. Deinotherium giganteum Kaup (Proboscidea) from Kassandra Peninsula (Chalkidiki, Macedonia, Greece). Geobios 27: 633–640.
• Vaček, M. 1877. Über österreichische Mastodonten und ihre Beziehungen zu den Mastodon−Arten Europas. Abhandlungen der Kaiserlich−Koniglichen geologischen Reichenstalt 7: 1–45.
• Van Couvering, A.H. and Van Couvering, J.A.. 1976. Early Miocene mammal fossils from East Africa: aspects of geology, faunistics and palaeoecology. In: G.L. Isaac and E.R. McCown (eds.), Human Origins. Louis Leakey and the East African Evidence, 155–207. W.A. Benjamin, Inc., Menlo Park, California.
• Vialli, V. 1966. Sul rinvenimento di Dinoterio (Deinotherium cf. hobleyi Andrews) nelle ligniti di Adi Ugri (Eritrea). Giornale di Geologia (Bologna) 33: 447–458.
• Weinsheimer, O. 1883. Über Dinotherium giganteum Kaup. Palaeontologische Abhandlungen 1: 205–282.
• Welcomme, J.−L. 1994. Le plus ancien crâne de proboscidien d’Europe, Gomphotherium hannibali nov. sp. (Proboscidea, Mammalia), du Miocène inférieur du Languedoc (France). Comptes Rendus de l’Academie des Sciences Serie II, 319: 135–140.
• Welcomme, J.−L., Antoine, P.−O., Duranthon, F., Mein, P., and Ginsburg, L. 1997. Nouvelles découvertes de Vertébrés miocènes dans le synclinal de Dera Bugti (Balouchistan, Pakistan). Comptes Rendus de l’Academie des Sciences 325: 531–536.
• Welcomme, J.−L., Benammi, M., Crochet, J.−Y., Marivaux, L., Métais, G., Antoine, P.−O., and Baloch, I. 2001. Himalayan forelands: paleontological evidence for Oligocene detrital deposits in the Bugti Hills (Balochistan, Pakistan). Geological Magazine 138: 397–405.
• Whybrow, P.J. 1984. Geological and faunal evidence from Arabia for mammal “migrations” between Asia and Africa during the Miocene.Courier Forschungsinstitut Senckenberg 69: 189–198.
• Wight, A.W.R. 1980. Palaeogene vertebrate fauna and regressive sediments of Dur at Talhah, southern Sirt Basin, Libya. In: M.J. Salem and M.T. Busrewil, Geology of Libya, 309–325. Academic Press, London.
• Yemane, K., Bonnefille, R., and Faure, H. 1985. Palaeoclimatic and tectonic implications of Neogene microflora from the northwestern Ethiopian highlands. Nature 318: 653–656.
• Yemane, K., Robert, C., and Bonnefille, R. 1987a. Pollen and clay mineral assemblages of a late Miocene lacustrine sequence from the northwestern Ethiopian highlands. Palaeogeography, Palaeoclimatology, Palaeoecology 60: 123–141.
• Yemane, K., Taieb, M., and Faure, H. 1987b. Limnogeologic studies on an intertrappean continental deposit from the northern Ethiopian Plateau. Journal of African Earth Sciences 6: 91–101.