Oxyaenidae

Oxyaenidae; Temporal range: Middle Paleocene to late Eocene
	Skull of a Palaeonictis occidentalis
	Reconstruction of Patriofelis ferox
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Mirorder:	Ferae
Clade:	Pan-Carnivora
Order:	†Oxyaenodonta; Van Valen, 1971[1]
Family:	†Oxyaenidae; Cope, 1877[2]
Type genus
	†Oxyaena; Cope, 1874
Subfamilies
	†Machaeroidinae; †Oxyaeninae; †Palaeonictinae; †Tytthaeninae;
Synonyms
	synonyms of order: Oxyaenida (Van Valkenburgh, 2007) ; synonyms of family: Oxyaenoidea (Osborn, 1910) ;

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

Description

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

Taxonomy

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: †Isphanatherium (Lavrov & Averianov, 1998)
      - Isphanatherium ferganensis (Lavrov & Averianov, 1998)
    - Genus: †Machaeroides (Matthew, 1909)
      - †Machaeroides eothen (Matthew, 1909)
      - †Machaeroides simpsoni (Dawson, 1986)
  - Subfamily: †Oxyaeninae (Cope, 1877)
    - Genus: †Argillotherium (Cope, 1877)
      - †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)[5]
      - †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)

Phylogeny

The phylogenetic relationships of family Oxyaenidae are shown in the following cladogram:[6][7][8][9][10][11]

Ferae

Pholidotamorpha

Pan-Carnivora

†Hyaenodonta

†Hyaenodonta (sensu stricto)

	†Altacreodus

	†Tinerhodon

sensu lato

Carnivoramorpha

†Oxyaenodonta

†Oxyaenidae

†Tytthaeninae

†Tytthaena

	†Tytthaena parrisi

	†Tytthaena lichna

†Oxyaeninae

†Oxyaena

	†Oxyaena intermedia

	†Oxyaena forcipata

†Oxyaena gulo

†Oxyaena lupina

†Oxyaena simpsoni

†Oxyaena pardalis

	†Oxyaena woutersi

	†Oxyaena sp. (Europe)

†Argillotherium

†Argillotherium toliapicum

?

†Sarkastodon

	†Sarkastodon henanensis

	†Sarkastodon mongoliensis

†Malfelis

†Malfelis badwaterensis

†Protopsalis

†Protopsalis tigrinus

†Patriofelis

	†Patriofelis ulta

	†Patriofelis ferox

†Dipsalidictis transiens

†Dipsalidictis krausei

†Dipsalidictis platypus

†Dipsalidictis aequidens

†Machaeroidinae

†Apataelurus

	†Apataelurus kayi

	†Apataelurus pishigouensis

†Machaeroidinae sp. (CM 2386)

†Machaeroidinae sp. (FMNH PM 1506)

†Isphanatherium

†Isphanatherium ferganensis

?

†Machaeroides

	†Machaeroides eothen

	†Machaeroides simpsoni

†Palaeonictinae

†Dipsalodon churchillorum

†Dipsalodon sp. (UM 71172)

†Dipsalodon matthewi

†Ambloctonus

	†Ambloctonus major

	†Ambloctonus priscus

	†Ambloctonus sinosus

†Palaeonictis

†Palaeonictis peloria

†Palaeonictis gigantea

	†Palaeonictis wingi

	†Palaeonictis occidentalis

†Altacreodus/Tinerhodon clade

†Dipsalidictis

†Dipsalodon

References

L. Van Valen (1971.) "Adaptive zones and the orders of mammals." Evolution, 25(2), 420–428.
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.
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.
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.
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.