PBDB Taxon

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Borophagus

Mammalia - Carnivora - Canidae

Taxonomy

Borophagus was named by Cope (1892) [also, but incorrectly said to be 1893]. It is not extant.

It was reranked as Dinocyon (Borophagus) by Matthew (1902); it was synonymized subjectively with Dinocyon by Matthew (1902); it was considered a nomen dubium by Matthew (1918).

It was assigned to Hyaenidae by Hay (1902); to Dinocyon by Matthew (1902); to Simocyoninae by Kurten and Anderson (1980); to Canidae by Cope (1892), Matthew and Stirton (1930), Stirton and VanderHoof (1933), VanderHoof and Gregory (1940) and Carroll (1988); to Borophagina by Wang et al. (1999), Wang et al. (2004) and Wang et al. (2008); and to Borophaginae by Bjork (1970), Tseng and Geisler (2016), Albright et al. (2019) and Bōgner and Samuels (2022).

Species

B. diversidens (syn. Hyaenognathus solus, Porthocyon dubius, Hyaenognathus pachyodon, Hyaenognathus matthewi, Felis hillanus), B. dudleyi, B. hilli (syn. Osteoborus crassapineatus, Osteoborus progressus), B. littoralis (syn. Osteoborus diabloensis), B. orc, B. parvus, B. pugnator (syn. Osteoborus galushai), B. secundus (syn. Hyaenognathus direptor, Hyaenognathus cyonoides), Dinocyon (Borophagus) meandrinus, Osteoborus crassipinaetus, Porthocyon matthewi

Synonyms

Cynogulo is a genus.

It was synonymized subjectively with Borophagus by Wang et al. (1999).
Hyaenognathus was named by Merriam (1903).

It was synonymized subjectively with Borophagus by VanderHoof (1936), Schultz (1937), Johnston (1939), Stirton (1939), VanderHoof and Gregory (1940), Dalquest (1969), Richey (1979) and Wang et al. (1999).
Osteoborus was named by Stirton and VanderHoof (1933). Its type is Hyaenognathus cyonoides.

It was synonymized subjectively with Borophagus by Wang et al. (1999).

It was assigned to Canidae by Stirton and VanderHoof (1933).
Pliogulo was named by White (1941) [preoccupied].

It was replaced with Cynogulo by Kretzoi (1968); it was synonymized subjectively with Osteoborus by Webb (1969); it was synonymized subjectively with Borophagus by Richey (1979), Tedford et al. (1987) and Wang et al. (1999).
Porthocyon was named by Merriam (1903).

It was synonymized subjectively with Hyaenognathus by Matthew and Stirton (1930) and VanderHoof (1931); it was synonymized subjectively with Osteoborus by VanderHoof and Gregory (1940) and Richey (1979); it was synonymized subjectively with Borophagus by Stirton (1939) and Wang et al. (1999).

Synonymy list

Year	Name and author
1892	Borophagus Cope
1902	Borophagus Hay p. 776
1902	Dinocyon (Borophagus) Matthew p. 289
1903	Hyaenognathus Merriam
1903	Porthocyon Merriam
1903	Hyaenognathus Merriam p. 278
1903	Porthocyon Merriam p. 283 figs. Pl. 29 and P. 30, Fig 1
1930	Borophagus Matthew and Stirton
1933	Borophagus Stirton and VanderHoof
1933	Osteoborus Stirton and VanderHoof p. 175
1940	Borophagus VanderHoof and Gregory
1941	Pliogulo White
1970	Borophagus Bjork p. 14
1980	Borophagus Kurten and Anderson p. 165
1988	Borophagus Carroll
1999	Borophagus Wang et al. p. 266
2004	Borophagus Wang et al. p. 451
2008	Borophagus Wang et al.
2016	Borophagus Tseng and Geisler p. 1
2019	Borophagus Albright et al. p. 172
2022	Borophagus Bōgner and Samuels

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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	Borophaginae
genus	Borophagus

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.

G. †Borophagus Cope 1892

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†Borophagus diversidens Cope 1892

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Invalid names: Felis hillanus Cope 1893 [synonym], Hyaenognathus matthewi Freudenberg 1910 [synonym], Hyaenognathus pachyodon Merriam 1903 [synonym], Hyaenognathus solus Stock 1932 [synonym], Porthocyon dubius Merriam 1903 [synonym]

†Borophagus dudleyi White 1941

†Borophagus hilli Johnston 1939

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Invalid names: Osteoborus crassapineatus Olsen 1956 [synonym], Osteoborus progressus Hibbard 1944 [synonym]

†Borophagus littoralis VanderHoof 1931

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Invalid names: Osteoborus diabloensis Richey 1938 [synonym]

†Borophagus orc Webb 1969

†Borophagus parvus Wang et al. 1999

†Borophagus pugnator Cook 1922

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Invalid names: Osteoborus galushai Webb 1969 [synonym]

†Borophagus secundus VanderHoof 1931

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Invalid names: Hyaenognathus cyonoides Martin 1928 [synonym], Hyaenognathus direptor Matthew 1924 [synonym]

†Dinocyon (Borophagus) meandrinus Hatcher 1894

Invalid names: Cynogulo [synonym], Hyaenognathus Merriam 1903 [synonym], Osteoborus Stirton and VanderHoof 1933 [synonym], Pliogulo White 1941 [synonym], Porthocyon Merriam 1903 [synonym]

Diagnosis

Reference	Diagnosis
R. A. Stirton and V. L. VanderHoof 1933 (Osteoborus)	First three premolars greatly reduced in size; anterior pre- molars (1-3 / 1 or 2-3) with prominent central cusp and small or absent anterior and posterior cusps; p4 with posterior accessory cusp above talonid; parastyle on P!; protocone on P! vestigial or absent; M;: with metaconid; m1 with wide inner heel; m2-3 intermediate in structure between Hyaenognathus and Aelurodon; superior branch of premaxilla strong, and meets anterior extension of frontals, excluding maxillaries from contact with nasals.

Measurements

No measurements are available

Composition:

phosphaticsubp

Environment:

terrestrialsubc

Locomotion:

actively mobilec

Life habit:

ground dwellingf

Diet:

carnivoreg

Reproduction:

viviparoussubc

Comments:

"meat/bone group" (Van Valkenburgh 1988: based on "Osteoborus" cyonoides)g

Created:

2009-01-04 14:34:14

Modified:

2009-01-04 16:34:14

Source: g = genus, f = family, subc = subclass, c = class, subp = subphylum

References: Van Valkenburgh 1988, Carroll 1988, Nowak 1991, Hendy et al. 2009, Ji et al. 2002, Lillegraven 1979

Age range: base of the early Early Hemphillian to the top of the Gelasian or 10.30000 to 1.80600 Ma

Collections (142 total)

Time interval	Ma	Country or state	Original ID and collection number
Miocene	23.03 - 5.333	Mexico (Chihuahua)	Osteoborus sp. (18714)
Clarendonian	12.5 - 9.4	USA (Idaho)	B. pugnator (19023)
Clarendonian	12.5 - 9.4	USA (California)	B. littoralis (19380) Osteoborus cyonoides (19318) Osteoborus diabloensis (19185 19322 19323 19324) Osteoborus littoralis (19553)
Clarendonian	12.5 - 9.4	USA (Texas)	B. secundus (18091)
Late/Upper Clarendonian	12.5 - 9.4	USA (Oregon)	B. pugnator (19036)
Clarendonian - Hemphillian	12.5 - 4.9	USA (Oklahoma)	B. sp. (70119)
Tortonian	11.62 - 7.246	USA (California)	B. littoralis (19466)
Late/Upper Miocene	11.608 - 5.333	USA (Oregon)	Osteoborus cyonoides (154947)
Late/Upper Miocene	11.608 - 5.333	USA (California)	B. littoralis (19308)
Late/Upper Miocene	11.608 - 5.333	El Salvador	Osteoborus cyonoides (18720)
Hemphillian	10.3 - 4.9	USA (Oklahoma)	B. secundus (170201) Osteoborus cyonoides (18304)
Hemphillian	10.3 - 4.9	USA (Texas)	Osteoborus cyonoides (18097 18175 18198) Osteoborus pugnator, Osteoborus sp. (18072) Osteoborus sp. (18073)
Hemphillian	10.3 - 4.9	USA (Colorado)	Osteoborus pugnator (18495)
Hemphillian	10.3 - 4.9	Mexico (Hidalgo)	B. secundus (212307)
Hemphillian	10.3 - 4.9	Honduras	Osteoborus cyonoides (18732 18736 18739 18750)
Hemphillian	10.3 - 4.9	USA (Kansas)	B. pugnator (18111) Hyaenognathus cyonoides, Osteoborus cyonoides (18140) Osteoborus cyonoides (18166 18241 18342) Osteoborus sp. (18046 18170 18263 18415)
Hemphillian	10.3 - 4.9	Mexico (Jalisco)	B. secundus (18745 189788 189790) Osteoborus cyonoides (18747)
Hemphillian	10.3 - 4.9	USA (Nebraska)	B. pugnator (18193 18254) B. secundus (18033) B. sp. (18086) Osteoborus sp. (18441)
Hemphillian	10.3 - 4.9	USA (California)	B. sp. (19479) Osteoborus sp. (19342 19419)
Hemphillian	10.3 - 4.9	USA (Nevada)	Osteoborus sp. (18975)
Hemphillian	10.3 - 4.9	USA (Florida)	B. dudleyi, B. pugnator (18565) B. hilli (18527 200705) B. pugnator (18504 18602) B. pugnator, Osteoborus orc (18601) Osteoborus dudleyi (18502) Osteoborus galushai (18545 18560 18563) Osteoborus orc (18564)
Hemphillian	10.3 - 4.9	USA (New Mexico)	Osteoborus sp. (19348 19351)
Hemphillian	10.3 - 4.9	USA (Oregon)	B. secundus (18970)
Late/Upper Hemphillian	10.3 - 4.9	USA (Arizona)	B. parvus (19514 19612)
Late/Upper Hemphillian	10.3 - 4.9	Mexico (Jalisco)	Osteoborus cyonoides (18715)
Late/Upper Hemphillian	10.3 - 4.9	USA (Texas)	B. hilli (18036 18093) Osteoborus sp. (18110)
Late/Upper Hemphillian	10.3 - 4.9	USA (California)	B. parvus, B. secundus (19582) B. secundus (19445) Osteoborus cyonoides (19604)
Late/Upper Hemphillian	10.3 - 4.9	USA (New Mexico)	Osteoborus secundus (19473)
Late/Upper Hemphillian	10.3 - 4.9	USA (Florida)	B. hilli (18533 18595) B. hilli, B. pugnator (18513 18532) B. pugnator (18573)
Late/Upper Hemphillian	10.3 - 4.9	USA (Nebraska)	B. secundus (18432) Osteoborus direptor (18499) Osteoborus sp. (18126 18359)
Late/Upper Hemphillian	10.3 - 4.9	Mexico (Guanajuato)	B. secundus (18713) Osteoborus cyonoides (18737 18740)
Late/Upper Hemphillian	10.3 - 4.9	Mexico (Chihuahua)	Osteoborus sp. (18755)
Late/Upper Hemphillian	10.3 - 4.9	Mexico (Zacatecas)	B. secundus (157349)
early Early Hemphillian - late Early Hemphillian	10.3 - 4.9	USA (Idaho)	B. sp. (218109 218111)
early Early Hemphillian - late Early Hemphillian	10.3 - 4.9	USA (Florida)	Osteoborus galushai (18539)
Late/Upper Hemphillian - Zanclean	10.3 - 3.6	USA (Florida)	B. hilli (18535)
Late/Upper Hemphillian - Blancan	10.3 - 1.8	USA (Indiana)	B. sp. (18320)
Late/Upper Hemphillian - Blancan	10.3 - 1.8	USA (Tennessee)	B. sp. (26574)
Zanclean	5.333 - 3.6	USA (North Carolina)	B. dudleyi, B. orc (52582)
Blancan	4.9 - 1.8	USA (Idaho)	Borophaginae indet. (19808) B. sp. (19867 19913)
Blancan	4.9 - 1.8	USA (Kansas)	B. diversidens (19932) B. sp. (19936) Osteoborus progressus (19922)
Blancan	4.9 - 1.8	Mexico (Baja California Sur)	B. diversidens (19673 20075)
Blancan	4.9 - 1.8	USA (Washington)	B. diversidens (20062) B. sp. (20063)
Blancan	4.9 - 1.8	USA (Nevada)	B. diversidens (19965)
Blancan	4.9 - 1.8	USA (New Mexico)	B. diversidens (19974) B. sp. (19981)
Blancan	4.9 - 1.8	USA (California)	B. diversidens (20117 20119) B. pachyodon (19663 19701) B. solus (19664 19665) Hyaenognathus pachydon (19668)
Blancan	4.9 - 1.8	El Salvador	B. hilli (107827)
Blancan	4.9 - 1.8	USA (Florida)	B. diversidens (19757)
Blancan	4.9 - 1.8	USA (South Carolina)	B. hilli (178479)
Blancan	4.9 - 1.8	Mexico (Guanajuato)	Canidae indet. (20077)
Blancan	4.9 - 1.8	USA (Nebraska)	B. diversidens (19957 19963) B. sp. (19954 19955 19958 19959 19960)
Blancan	4.9 - 1.8	USA (Texas)	B. diversidens (20023 20026 20032 20034 20060) B. sp. (20033 20050)
Blancan	4.9 - 1.8	USA (Arizona)	B. diversidens (19662) B. sp. (19625 19628 19631 19657 19658)
Piacenzian	3.6 - 2.588	USA (North Carolina)	Osteoborus dudleyi (18550)
Gelasian	2.588 - 1.806	USA (Florida)	B. diversidens (19758)