PBDB Taxon

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Morphology	Ecology and taphonomy	External Literature Search	Age range and collections

Aenocyon dirus (dire wolf)

Mammalia - Carnivora - Canidae

Taxonomy

Canis dirus was named by Leidy (1858). Its type specimen is ANSP 11614, a maxilla, and it is a 3D body fossil. Its type locality is Ohio River at Evansville, which is in a Pleistocene terrestrial horizon in Indiana.

It was recombined as Canis (Aenocyon) dirus by Stock et al. (1946); it was recombined as Aenocyon dirus by Merriam (1918), Hibbard (1949), Hibbard and Taylor (1960), Ruiz-Ramoni et al. (2022), Prevosti (2023).

Synonyms

Canis primaevus was named by Hodgson (1833).

It was replaced with Canis indianensis by Leidy (1873); it was recombined as Cuon primaevus by Bourguignat (1875); it was synonymized subjectively with Canis dirus by Merriam (1912); it was synonymized subjectively with Aenocyon dirus by Ruiz-Ramoni et al. (2022).
Canis ayersi was named by Sellards (1916) [species named after Frank Ayers, who collected the holotype]. Its type specimen is UF/FGS 280, a skull, and it is a 3D body fossil. Its type locality is Vero, which is in a Pleistocene channel sandstone/sandstone in Florida.

It was recombined as Aenocyon ayersi by Merriam (1918); it was synonymized subjectively with Canis dirus by Lundelius (1972), Martin (1974), Nowak (1979), Kurten and Anderson (1980), Kurten (1984); it was synonymized subjectively with Aenocyon dirus by Ruiz-Ramoni et al. (2022), Prevosti (2023).
Canis indianensis was named by Leidy (1869).

It was synonymized subjectively with Canis dirus by Merriam (1912), Troxell (1915), Nowak (1979), Kurten and Anderson (1980), Kurten (1984); it was synonymized subjectively with Aenocyon dirus by Ruiz-Ramoni et al. (2022), Prevosti (2023).
Canis mississippiensis was named by Allen (1876). It is a 3D body fossil.

It was synonymized subjectively with Canis dirus by Merriam (1912), Nowak (1979), Kurten and Anderson (1980), Kurten (1984); it was synonymized subjectively with Aenocyon dirus by Ruiz-Ramoni et al. (2022), Prevosti (2023).
Canis himalaicus was named by Ogilby (1836).

It was synonymized subjectively with Cuon primaevus by Bourguignat (1875).
Cuon buansu was named by Gervais (1855).

It was synonymized subjectively with Cuon primaevus by Bourguignat (1875).
Cuon grayformis was named by Hodgson (1863).

It was synonymized subjectively with Cuon primaevus by Bourguignat (1875).

Synonymy list

Year	Name and author
1833	Canis primaevus Hodgson
1836	Canis himalaicus Ogilby
1854	Canis primaevus Leidy
1855	Cuon buansu Gervais
1858	Canis dirus Leidy p. 21
1863	Cuon grayformis Hodgson
1869	Canis indianensis Leidy
1873	Canis indianensis Leidy p. 230
1875	Cuon primaevus Bourguignat
1876	Canis mississippiensis Allen p. 49
1902	Canis dirus Hay p. 775
1902	Canis indianensis Hay p. 775
1903	Canis indianensis Merriam p. 288 figs. Pl. 30, Fig. 2
1916	Canis ayersi Sellards p. 121
1918	Aenocyon ayersi Merriam
1918	Aenocyon dirus Merriam
1929	Canis ayersi Simpson
1931	Canis dirus Elftman p. 5
1940	Canis dirus Olson p. 42
1940	Canis ayersi Olson p. 44
1946	Canis (Aenocyon) dirus Stock et al. p. 109
1949	Aenocyon dirus Hibbard
1960	Aenocyon dirus Hibbard and Taylor
1966	Canis dirus Slaughter p. 481
1972	Canis dirus Kurten and Anderson
1972	Canis dirus Lundelius, Jr.
1974	Canis dirus Martin
1979	Canis dirus Nowak
1980	Canis dirus Kurten and Anderson p. 171
1984	Canis dirus Kurten
1988	Canis dirus Berta
2008	Canis dirus Wang et al.
2009	Canis dirus Tedford et al. p. 144 figs. 40, 43, 44, 56A–F; appendices 2, 3
2019	Canis dirus Albright et al. p. 173
2021	Canis dirus Lu et al.
2022	Aenocyon dirus Ruiz-Ramoni et al. p. 103
2023	Aenocyon dirus Prevosti

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Rank	Name	Author
kingdom	Animalia	()
—	Bilateria
—	Eubilateria	Ax 1987
—	Deuterostomia
phylum	Chordata	Haeckel 1874
subphylum	Vertebrata
superclass	Gnathostomata
—	Osteichthyes	()
subclass	Sarcopterygii	()
subclass	Dipnotetrapodomorpha	(Nelson 2006)
subclass	Tetrapodomorpha	()
—	Tetrapoda
—	Reptiliomorpha
—	Anthracosauria
subclass	Amphibiosauria	Kuhn 1967
—	Cotylosauria	()
—	Amniota
subclass	Synapsida
—	Therapsida	()
infraorder	Cynodontia	()

Rank	Name	Author
—	Mammaliamorpha
—	Mammaliaformes
class	Mammalia
—	Cladotheria
—	Zatheria
subclass	Tribosphenida	()
subclass	Theria
—	Eutheria	()
—	Placentalia
—	Boreoeutheria
—	Laurasiatheria
—	Scrotifera
—	Ferae	()
—	Carnivoramorpha	Wyss and Flynn 1993
—	Carnivoraformes	Flynn et al.
order	Carnivora
family	Canidae
subfamily	Caninae	Gill 1872
genus	Aenocyon
species	dirus	()

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

†Aenocyon dirus Leidy 1858 [dire wolf]

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Invalid names: Canis ayersi Sellards 1916 [synonym], Canis indianensis Leidy 1869 [synonym], Canis mississippiensis Allen 1876 [synonym], Canis nehringi Ameghino 1902 [synonym], Canis primaevus Hodgson 1833 [synonym]

Diagnosis

Reference	Diagnosis
R. H. Tedford et al. 2009	Canis dirus is distinguished by a number of autapomorphies related to hypercarnivory: loss of m2 anterolabial cingulum; wide palate; greatly reduced m1–m2 metaconids and entoconids; greatly reduced M1–M2 hypocones; very weak or discontinuous anterolingual cingu- lum across M1 protocone. It shares with C. armbrusteri two synapomorphies: short and little recurved canine, and a reduced P4 protocone. With C. lupus it shares four additional synapomorphies: paroccipital pro- cess strongly expanded posteriorly; frontal sinus reaching the frontoparietal suture; long incisive foramina that extend to or behind canine alveoli; M1 parastyle very weak, preparacrista directed anteriorly, not to parastyle.