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Temporal range: Eocene - Oligocene, 35–28 Ma
Arsinoitherium zitteli.jpg
Arsinoitherium zitteli


Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Paenungulata
Order: Embrithopoda
Andrews 1906

Embrithopoda ("heavy-footed") is an order of extinct mammals known from Asia, Africa and eastern Europe. Most of the embrithopod genera are known exclusively from jaws and teeth dated from the late Paleocene to the late Eocene, but the order is best known from its terminal member, the elephantine Arsinoitherium.[2]


While embrithopods bore a superficial resemblance to rhinoceroses, their horns had bony cores covered in keratinized skin and were not made of hair. Not all embrithopods possessed horns, either. Despite their appearance, they were related to elephants, not perissodactyls.[3]

The Embrithopoda are tethytheres[4] and are also believed to be part of the clade Afrotheria. However, a new study of the basal arsinoitheriid, Palaeoamasia, suggests that embrithopods are not tethytheres or even paenungulates, and that they need to be better sampled in an analysis of eutherian relationships to clarify if they are even afrotherians.[5]

Fossils of embrithopods, such as Arsinoitherium, have been found in Egypt, Mongolia, Turkey, Romania, Namibia[6] and Tunisia.[7] Until the 1970s, only Arsinoitherium itself was known, appearing isolated in the fossil record.[3]


McKenna & Manning 1977 and McKenna & Bell 1997 considered Phenacolophus from Mongolia a primitive embrithopod, although this attribution was challenged by several other authors. A 2016 cladistic study, however, confirmed the original embrithopod placement of Phenacolophus.[8][9]

Order Embrithopoda Andrews 1906 sensu Prothero & Schoch 1989 (=Barypoda Andrews 1904)[10]


  1. ^ Rose 2006, pp. 242–3
  2. ^ Rose 2006, p. 265
  3. ^ a b "Introduction to the Embrithopoda". University of California Museum of Paleontology. Retrieved March 2013.  Check date values in: |access-date= (help)
  4. ^ Embrithopoda in the Paleobiology Database. Retrieved March 2013.
  5. ^ Erdal, O., Antoine, P.-O., Sen, S. 2016. New material of Palaeoamasia kansui (Embrithopoda, Mammalia) from the Eocene of Turkey and a phylogenetic analysis of Embrithopoda at the species level. Palaeontology, in press. http://onlinelibrary.wiley.com/doi/10.1111/pala.12247/abstract [Erdal et al.'s inclusion of Embrithopoda in Tabuce et al. (2007) finds them outside of (Ungulata + Afrotheria). Since this clade is not supported by molecular data, it suggests the need to explore the relationships of embrithopods, as they could potentially have affinities with laurasiatheres and "true" ungulates.]
  6. ^ a b M. Pickford; B. Senut; J. Morales; P. Mein; I. M. Sanchez (2008). "Mammalia from the Lutetian of Namibia". Memoir of the Geological Survey of Namibia. 20: 465–514. 
  7. ^ Nicolas Vialle; Gilles Merzeraud; Cyrille Delmer; Monique Feist; Suzanne Jiquel; Laurent Marivaux; Anusha Ramdarshan; Monique Vianey-Liaud; El Mabrouk Essid; Wissem Marzougui; Hayet Khayati Ammar; Rodolphe Tabuce (2013). "Discovery of an embrithopod mammal (Arsinoitherium?) in the late Eocene of Tunisia". Journal of African Earth Sciences. 87: 86–92. doi:10.1016/j.jafrearsci.2013.07.010. 
  8. ^ Radulesco & Sudre 1985; Maas, Thewissen & Kappelman 1998, p. 291; Rose & Archibald 2005, pp. 97–98
  9. ^ Erdal, Ozan; Antoine, Pierre-Olivier; Sen, Sevket; Smith, Andrew (2016). "New material ofPalaeoamasia kansui(Embrithopoda, Mammalia) from the Eocene of Turkey and a phylogenetic analysis of Embrithopoda at the species level". Palaeontology. 59 (5): 631–655. doi:10.1111/pala.12247. ISSN 0031-0239. 
  10. ^ Mikko's Phylogeny Archive [1] Haaramo, Mikko (2007). "†Embrithopoda - arsinoitheres". Retrieved 30 December 2015. 


  • Andrews, C.W. (1904). "Further notes on the mammals of the Eocene of Egypt". Geological Magazine. 1 (4). doi:10.1017/S0016756800119491. OCLC 4668923377. 
  • Andrews, C.W. (1906). A descriptive catalogue of the Tertiary Vertebrata of the Fayûm, Egypt. London: British Museum. OCLC 3675777. 
  • Beadnell, H.J.C. (1902). A preliminary note on Arsinoitherium zitteli Beadnell, from the Upper Eocene strata of Egypt. Cairo: Egyptian Survey Department, Public Works Ministry. OCLC 20609512. 
  • Court, N. (1990). "Periotic anatomy of Arsinoitherium (Mammalia, Embrithopoda) and its phylogenetic implications". Journal of Vertebrate Paleontology. 10 (2): 170–82. doi:10.1080/02724634.1990.10011806. OCLC 4899524631. 
  • Maas, M.C.; Thewissen, J.G.M.; Kappelman, J. (1998). "Hypsamasia seni (Mammalia: Embrithopoda) and other mammals from the Eocene Kartal Formation of Turkey". In Beard, K.C.; Dawson, M.R. Dawn of the Age of Mammals in Asia (PDF). Bulletin of Carnegie Museum of Natural History. 34. pp. 286–297. OCLC 493312921. Retrieved May 2013.  Check date values in: |access-date= (help)
  • McKenna, Malcolm C.; Bell, Susan K. (1997). Classification of Mammals Above the Species Level. New York: Columbia University Press. ISBN 0231110138. OCLC 37345734. 
  • McKenna, M.C.; Manning, E. (1977). "Affinities and palaeobiogeographic significance of the Mongolian Paleogene genus Phenacolophus". Geobios, Memoire special. 1: 61–85. doi:10.1016/S0016-6995(77)80008-9. OCLC 4656767437. 
  • Ozansoy, Fikret (1966). Türkiye Senozoik çağlarında fosil insan formu problemi ve biostratigrafik dayanakları. A.Ü. D.T.C.F. (University of Ankara, Faculty of Languages, History and Geography Publications). 172. Ankara University Press. pp. 1–104. OCLC 16763756. 
  • Radulesco, C.; Iliesco, G.; Iliesco, M. (1976). "Decouverte d'un Embrithopode nouveau (Mammalia) dans la Paléogène de la dépression de Hateg (Roumanie) et considération générales sur la géologie de la région". Neues Jahrbuch für Geologie und Paläontologie Monatshefte. 1 (11): 690–698. 
  • Radulesco, C.; Sudre, J. (1985). "Crivadiatherium iliescui n. sp., nouvel Embrithopode (Mammalia) dans le Paléogène ancien de la depression de Hateg (Roumanie)". Palaeovertebrata. 15 (3): 139–57. 
  • Rose, Kenneth David (2006). The beginning of the age of mammals. Baltimore: JHU Press. ISBN 0801884721. 
  • Rose, Kenneth D.; Archibald, J. David (2005). The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades. JHU Press. ISBN 9780801880223. OCLC 55801049. Retrieved March 2013.  Check date values in: |access-date= (help)
  • Sanders, William J; Kappelman, John; Rasmussen, D Tab (2004). "New large-bodied mammals from the Late Oligocene site of Chilga, Ethiopia" (PDF). Acta palaeontologica polonica. 49 (3). OCLC 716778291. 
  • Şen, Ş.; Heintz, E. (1979). "Palaeoamasia kansui Ozansoy 1966, embrithopode (Mammalia) de l'Eocene de Anatolie". Annales de paléontologie (Vértébres). 65 (1): 73–91.