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Embrithopoda Temporal range: Paleocene - Oligocene, 56–28 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Arsinoitherium zitteli
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Paenungulatomorpha
Grandorder:	Paenungulata
Mirorder:	Tethytheria
Order:	†Embrithopoda Andrews 1906
Families
†Stylolophus †Arsinoitheriidae †Palaeoamasiidae

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; however, the order is best known from its terminal member, the elephantine Arsinoitherium.

Description

While embrithopods bore a superficial resemblance to rhinoceroses, their horns had bony cores covered in keratinized skin. Not all embrithopods possessed horns, either. Despite their appearance, they have been regarded as related to elephants, not perissodactyls.

As tethytheres, the Embrithopoda have been believed to be part of the clade Afrotheria. However, a 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. It is also not clear if embrithopods originated in Africa or Eurasia. However, recent findings demonstrate an African origin for embrithopods and furthermore a relationship with other paenungulates, albeit having diverged earlier than previously thought.

Fossils of embrithopods, such as Arsinoitherium, have been found in Egypt, Ethiopia, Kenya, Morocco, Mongolia, Turkey, Romania, Namibia, Tunisia and Croatia. Until the 1970s, only Arsinoitherium itself was known, appearing isolated in the fossil record.

Classification

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 found Phenacolophus as a stem-perissodactyl and the embrithopods at the base of Altungulata. More recently, an afrothere identity has been vindicated, albeit more basal than previously assumed.

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

• Genus †Stylolophus [de] Gheerbrant et al, 2018
• Family †Arsinoitheriidae Andrews 1904
  • Genus †Namatherium [de] Pickford et al., 2008
  • Genus †Arsinoitherium Beadnell 1902
• Family †Palaeoamasiidae Şen & Heintz 1979
  • Genus †Hypsamasia Maas, Thewissen & Kappelman 1998
  • Genus †Palaeoamasia Ozansoy 1966
  • Genus †Crivadiatherium Radulesco, Iliesco & Iliesco 1976

Notes

1. Emmanuel Gheerbrant; Arnaud Schmitt; László Kocsis (2018). "Early African fossils elucidate the origin of embrithopod mammals". Current Biology. Online edition. doi:10.1016/j.cub.2018.05.032.
2. Rose 2006, pp. 242–3
3. Rose 2006, p. 265
4. ^ "Introduction to the Embrithopoda". University of California Museum of Paleontology. Retrieved 7 March 2013.
5. Embrithopoda in the Paleobiology Database. Retrieved March 2013.
6. ^ Erdal, O.; Antoine, P.-O.; Sen, S.; Smith, A. (2016). "New material of Palaeoamasia 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.
7. ^ Gheerbrant, E.; Schmitt, A.; Kocsis, L. (July 2018). "Early African fossils elucidate the origin of embrithopod mammals". Current Biology. 28 (13): 2167-2173.e2. Retrieved 28 September 2024.
8. ^ 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.
9. 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.
10. Rab danas
11. Koenigswald, W. v. (2012). "Unique differentiation of radial enamel in Arsinoitherium (Embrithopoda, Tethytheria)". Historical Biology. 25 (2): 183–192. doi:10.1080/08912963.2012.714658.
12. Radulesco & Sudre 1985; Maas, Thewissen & Kappelman 1998, p. 291; Rose & Archibald 2005, pp. 97–98
13. Mikko's Phylogeny Archive Haaramo, Mikko (2007). "†Embrithopoda - arsinoitheres". Retrieved 30 December 2015.

References

• Andrews, C.W. (1904). "Further notes on the mammals of the Eocene of Egypt". Geological Magazine. 1 (4): 157–162. Bibcode:1904GeoM....1..157A. 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" (PDF). In Beard, K.C.; Dawson, M.R. (eds.). Dawn of the Age of Mammals in Asia. Bulletin of Carnegie Museum of Natural History. Vol. 34. pp. 286–297. OCLC 493312921. Retrieved 9 May 2013.
• 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. 10 (Suppl 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). Vol. 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 28 April 2013.
• 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.

v t e Paenungulatomorpha
Kingdom: Animalia Phylum: Chordata Class: Mammalia Infraclass: Placentalia Superorder: Afrotheria
Paenungulatomorpha †Abdounodus †Hadrogeneios †Ocepeia Hyracoidea †Dimaitherium †Helioseus †Megalohyrax †Microhyrax †Namahyrax †Regubahyrax †Rukwalorax †Saghatherium †Seggeurius †Selenohyrax †Thyrohyrax †Geniohyidae Abdahyrax Brachyhyrax Bunohyrax Geniohyus Nengohyrax Pachyhyrax †Titanohyracidae Afrohyrax Antilohyrax Rupestrohyrax Titanohyrax †Pliohyracidae Hengduanshanhyrax Kvabebihyrax Meroehyrax Parapliohyrax Pliohyrax Postschizotherium Prohyrax Procaviidae Dendrohyrax †Gigantohyrax Heterohyrax Procavia Tethytheria †Desmostylia? Paleoparadoxiidae Ashoroa Paleoparadoxia Neoparadoxia Desmostylidae Behemotops Cornwallius Desmostylus Kronokotherium Ounalashkastylus †Embrithopoda Stylolophus Arsinoitheriidae Arsinoitherium Namatherium Palaeoamasiidae Crivadiatherium Hypsamasia Palaeoamasia Proboscidea see Proboscidea includes elephants Sirenia see Sirenia includes dugongs and manatees

• Embrithopods
• Eocene mammals
• Oligocene mammals
• Rupelian extinctions