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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Cladus: Synapsida
Cladus: Eupelycosauria
Cladus: Sphenacodontia
Cladus: Sphenacodontoidea
Cladus: Therapsida
Cladus: Theriodontia
Subordo: Cynodontia
Infraordo: Eucynodontia
Cladus: Probainognathia
Cladus: Prozostrodontia
Cladus: Mammaliaformes
Classis: Mammalia
Subclassis: Trechnotheria
Infraclassis: Zatheria
Supercohors: Theria
Cohors: Eutheria
Infraclassis: Placentalia
Cladus: Boreoeutheria
Superordo: Laurasiatheria
Ordo: †Creodonta
Familia: †Oxyaenidae
Subfamiliae: †Ambloctoninae - †Oxyaeninae - †Tytthaeninae - †Machaeroidinae


Stucky, R.K.; Hardy, T.G. 2007: A new large, hypercarnivorous oxyaenid (Mammalia, Creodonta) from the Middle Eocene of the Wind River Formation, Natrona County, Wyoming. Bulletin of Carnegie Museum of Natural History, 39: 57–65. DOI: 10.2992/0145-9058(2007)39[57:ANLHOM]2.0.CO;2

Vernacular names
русский: Оксиениды
中文: 牛鬣獸科

Oxyaenidae ("sharp hyenas") is a family of extinct carnivorous placental pan-carnivoran mammals.[3] Traditionally classified in order Creodonta, this group is now classified in its own order Oxyaenodonta ("sharp tooth hyenas") within mirorder Ferae. The group contains four subfamilies comprising fourteen genera. Oxyaenids were the first to appear during the late Paleocene in North America, while smaller radiations of oxyaenids in Europe and Asia occurred during the Eocene.[4]


The name of order Oxyaenodonta comes from Ancient Greek ὀξύς- (oxús-) 'sharp', name of hyena genus Hyaena and from Ancient Greek ὀδούς (odoús) 'tooth'.

The name of family Oxyaenidae comes from Ancient Greek ὀξύς- (oxús-) 'sharp', name of hyena genus Hyaena and taxonomic suffix "-idae".[5]

They were superficially cat-like mammals that walked on flat feet, in contrast to modern cats, which walk and run on their toes. Anatomically, characteristic features include a short, broad skull, deep jaws, and teeth designed for crushing rather than shearing, as in the hyaenodonts or modern cats.

Oxyaenids were specialized carnivores that preyed on other terrestrial vertebrates, eggs and insects. They were capable of climbing trees, which is suggested by fossil evidence of their paws.
Classification and phylogeny

Order: †Oxyaenodonta (Van Valen, 1971)
Family: †Oxyaenidae (Cope, 1877)
Subfamily: †Machaeroidinae (Matthew, 1909)
Genus: †Apataelurus (Scott, 1937)
†Apataelurus kayi (Scott, 1937)
†Apataelurus pishigouensis (Tong & Lei, 1986)
Genus: †Diegoaelurus (Zack, Poust & Wagner, 2022)[6]
Diegoaelurus vanvalkenburghae (Zack, Poust & Wagner, 2022)
Genus: †Isphanatherium (Lavrov & Averianov, 1998)
Isphanatherium ferganensis (Lavrov & Averianov, 1998)
Genus: †Machaeroides (Matthew, 1909)
†Machaeroides eothen (Matthew, 1909)
†Machaeroides simpsoni (Dawson, 1986)
Incertae sedis:
†Machaeroidinae sp. (CM 2386) (Zack, 2019)
†Machaeroidinae sp. (FMNH PM 1506) (Tomiya, 2021)
†Machaeroidinae sp. (USNM 173514) (Zack, 2019)
Subfamily: †Oxyaeninae (Cope, 1877)
Genus: †Argillotherium (Davies, 1884)
†Argillotherium toliapicum (Davies, 1884)
Genus: †Dipsalidictis (Matthew & Granger, 1915) <−−−[paraphyletic genus]
†Dipsalidictis aequidens (Matthew & Granger, 1915)
†Dipsalidictis krausei (Gunnell & Gingerich, 1991)
†Dipsalidictis platypus (Matthew & Granger, 1915)
†Dipsalidictis transiens (Matthew & Granger, 1915)
Genus: †Malfelis (Stucky & Hardy, 2007)[7]
†Malfelis badwaterensis (Stucky & Hardy, 2007)
Genus: †Oxyaena (Cope, 1874)
†Oxyaena forcipata (Cope, 1874)
†Oxyaena gulo (Matthew & Granger, 1915)
†Oxyaena intermedia (Denison, 1938)
†Oxyaena lupina (Cope, 1874)
†Oxyaena pardalis (Matthew & Granger, 1915)
†Oxyaena simpsoni (Van Valen, 1966)
†Oxyaena woutersi (Lange-Badré & Godinot, 1982)
Genus: †Patriofelis (Leidy, 1870)
†Patriofelis ferox (Marsh, 1872)
†Patriofelis ulta (Leidy, 1870)
Genus: †Protopsalis (Cope, 1880)
†Protopsalis tigrinus (Cope, 1880)
Genus: †Sarkastodon (Granger, 1938)
†Sarkastodon henanensis (Tong & Lei, 1986)
†Sarkastodon mongoliensis (Granger, 1938)
Subfamily: †Palaeonictinae (Denison, 1938) [synonym: Ambloctoninae (Cope, 1877)]
Genus: †Ambloctonus (Cope, 1875)
†Ambloctonus major (Denison, 1938)
†Ambloctonus priscus (Matthew & Granger, 1915)
†Ambloctonus sinosus (Cope, 1875)
Genus: †Dipsalodon (Jepsen, 1930) <−−−[paraphyletic genus]
†Dipsalodon churchillorum (Rose, 1981)
†Dipsalodon matthewi (Jepsen, 1930)
Genus: †Palaeonictis (de Blainville, 1842)
†Palaeonictis gigantea (de Blainville, 1842)
†Palaeonictis occidentalis (Osborn, 1892)
†Palaeonictis peloria (Rose, 1981)
†Palaeonictis wingi (Chester, 2010)
Subfamily: †Tytthaeninae (Gunnell & Gingerich, 1991)
Genus: †Tytthaena (Gingerich, 1980)
†Tytthaena lichna (Rose, 1981)
†Tytthaena parrisi (Gingerich, 1980)


The phylogenetic relationships of family Oxyaenidae are shown in the following cladogram:[8][9][10][11][12][13]






Hyaenodonta (sensu stricto)

 sensu lato 




Tytthaena lichna

Tytthaena parrisi


Patriofelis ferox

Patriofelis ulta


Protopsalis tigrinus


Malfelis badwaterensis


Sarkastodon henanensis

Sarkastodon mongoliensis


Argillotherium toliapicum


Oxyaena woutersi

Oxyaena sp. (Europe)

Oxyaena lupina

Oxyaena simpsoni

Oxyaena pardalis

Oxyaena gulo

Oxyaena forcipata

Oxyaena intermedia

Dipsalidictis transiens

Dipsalidictis krausei

Dipsalidictis platypus

Dipsalidictis aequidens


Apataelurus kayi

Apataelurus pishigouensis

†Machaeroidinae sp. (CM 2386)

†Machaeroidinae sp. (FMNH PM 1506)

†Machaeroidinae sp. (USNM 173514)


Diegoaelurus vanvalkenburghae


Isphanatherium ferganensis


Machaeroides eothen

Machaeroides simpsoni


Dipsalodon churchillorum

Dipsalodon sp. (UM 71172)

Dipsalodon matthewi


Ambloctonus major

Ambloctonus priscus

Ambloctonus sinosus


Palaeonictis peloria

Palaeonictis gigantea

Palaeonictis occidentalis

Palaeonictis wingi

Altacreodus/Tinerhodon clade


E. D. Cope (1877.) "Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874." Report upon United States Geographical Surveys West of the One Hundredth Meridian, in charge of First Lieut. G.M. Wheeler, Corps of Engineers, U.S. Army, Vol. IV Paleontology, Part II, pp. 1-365. Government Printing Office, Washington, D.C.
L. Van Valen (1971.) "Adaptive zones and the orders of mammals." Evolution, 25(2), 420–428.
Halliday, Thomas J. D.; Upchurch, Paul; Goswami, Anjali (2015). "Resolving the relationships of Paleocene placental mammals" (PDF). Biological Reviews. 92 (1): 521–550. doi:10.1111/brv.12242. ISSN 1464-7931. PMC 6849585. PMID 28075073.
Gunnel, Gregg F.; Gingerich, Philip D. (30 Sep 1991). "Systematics and evolution of late Paleocene and early Eocene Oxyaenidae (Mammalia, Creodonta) in the Clarks Fork Basin, Wyoming" (PDF). Contributions from the Museum of Paleontology. The University of Michigan. 28 (7): 141–180. Retrieved 2010-01-03.
Dixon, Dougal (2008). World Encyclopedia of Dinosaurs & Prehistoric Creatures. Lorenz Books. ISBN 978-0754817307.
Zack, Shawn P.; Poust, Ashley W.; Wagner, Hugh (2022-03-15). "Diegoaelurus, a new machaeroidine (Oxyaenidae) from the Santiago Formation (late Uintan) of southern California and the relationships of Machaeroidinae, the oldest group of sabertooth mammals". PeerJ. 10: e13032. doi:10.7717/peerj.13032. ISSN 2167-8359. PMID 35310159.
Stucky, R. K.; Hardy, T. G. (2007). "A new large hypercarnivorous oxyaenid (Mammalia, Creodonta) from the Middle Eocene of the Wind River Formation, Natrona County, Wyoming". Bulletin of Carnegie Museum of Natural History. 39: 57–65. doi:10.2992/0145-9058(2007)39[57:anlhom];2.
Gunnel, Gregg F.; Gingerich, Philip D. (1991). "Systematics and evolution of late Paleocene and early Eocene Oxyaenidae (Mammalia, Creodonta) in the Clarks Fork Basin, Wyoming" (PDF). Contributions from the Museum of Paleontology. The University of Michigan. 28 (7): 141–180.
Stephen G. B. Chester; Jonathan I. Bloch; Ross Secord; Doug M. Boyer (2010). "A new small bodied species of Palaeonictis (Creodonta, Oxyaenidae) from the Paleocene-Eocene thermal maximum". Journal of Mammalian Evolution. 17 (4): 227–243. doi:10.1007/s10914-010-9141-y. S2CID 15058311.
F. Solé, E. Gheerbrant, and M. Godinot. (2011.) "New data on the Oxyaenidae from the Early Eocene of Europe; biostratigraphic, paleobiogeographic and paleoecologic implications." Palaeontologia Electronica 14(2):13A:1-41
Solé, Floréal; Ladevèze, Sandrine (2017). "Evolution of the hypercarnivorous dentition in mammals (Metatheria,Eutheria) and its bearing on the development of tribosphenic molars". Evolution & Development. 19 (2): 56–68. doi:10.1111/ede.12219. PMID 28181377. S2CID 46774007.
Prevosti, F. J., & Forasiepi, A. M. (2018). "Introduction. Evolution of South American Mammalian Predators During the Cenozoic: Paleobiogeographic and Paleoenvironmental Contingencies"

Shawn P. Zack (2019). "A skeleton of a Uintan machaeroidine 'creodont' and the phylogeny of carnivorous eutherian mammals". Journal of Systematic Palaeontology. 17 (8): 653–689. doi:10.1080/14772019.2018.1466374. S2CID 89934728.

Further reading

David Lambert and the Diagram Group. The Field Guide to Prehistoric Life. New York: Facts on File Publications, 1985. ISBN 0-8160-1125-7

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