Epicyon
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''Epicyon'' ("more than a dog") is a large, extinct, canid
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
of the subfamily Borophaginae ("bone-crushing dogs"), native to North America. ''Epicyon'' existed for about from the Hemingfordian age of the
Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene Epoch made up of two stages: the Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 Ma to 15.97 ± 0.05 Ma (million years ago). It was p ...
, to the Hemphillian of the Late Miocene. ''Epicyon'' is the largest known canid of all time, with the type species reaching 2.4 m (7.9 ft) in length, 90 cm (35 in) in shoulder height and approximately 100–125 kg (220–276 lb) in body mass. The largest known humerus specimen belonged to an individual weighing up to .


Description

''Epicyon'' had a massive head and powerful jaws that were well adapted for bone-crushing, with enlarged fourth premolars like some hyenas, giving its skull a
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'' native to Africa and India. It has a muscular, broad-chested body; short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphi ...
-like shape rather than having a skull similar in shape to that of a wolf; the adaptation would have allowed ''Epicyon'' to scavenge as well as hunt, giving it access to the nutritious marrow other contemporary carnivores couldn't access. ''Epicyon'' was one of the last of the borophagines, and shared its North American habitat with several other canids, including: * '' Borophagus'' (Mya) * '' Carpocyon'' from 20.4 to * '' Paratomarctus'' from 16.3 to * '' Aelurodon'' from 16.0 to * '' Canis lepophagus'' from 10.3 to


Species

Fossil specimens range from
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to
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the ...
and have been found in
Nebraska Nebraska () is a state in the Midwestern region of the United States. It is bordered by South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Kansas to the south; Colorado to the so ...
, Montana,
Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to ...
,
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,
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe, New Mexico, Santa Fe , LargestCity = Albuquerque, New Mexico, Albuquerque , LargestMetro = Albuquerque metropolitan area, Tiguex , Offi ...
, Colorado, Oklahoma, Idaho, Oregon, Arizona within the
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. As well as in
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,
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tota ...
. ''Epicyon haydeni'', the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...
, existed from 20.6-5.3 million years ago. It is synonymous with ''Aelurodon aphobus, Osteoborus ricardoensis, Osteoborus validus, ''and ''Tephrocyon mortifer,'' and was named by Joseph Leidy as a subgenus. It was recombined as ''Aelurodon haydeni'' by Scott and Osborn in 1890. Further study by Matthew in 1899, Matthew and Gidley in 1904, VanderHoof and Gregory in 1940, McGrew in 1944, Bennett in 1979, (1979) and Becker (1980). It again was recombined as ''Epicyon haydeni'' by Baskin in 1980, Voorhies in 1990, (1990), Baskin (1998), Wang et al. in 1999. Known as the largest species of all canids, it is estimated to have had a body length of , a shoulder height of and a body mass of approximately ,Díaz-Sibaja, R. (2010). "Titanes Vol. 1 Mamíferos." ''Fósil® Revista de Paleontología®''. with the largest known specimen weighing up to . The molars of ''Epicoyn haydeni'' were grindstone-like teeth that allow for a canid diet that includes both meat and plant and insects. The proportional size of an animal's molars is a great measure of the nutritional diversity of its diet. Based on fossilized feces and its robust teeth and jaw muscles it is believed to have consumed large amounts of bone and share a similar digestive tract to modern day hyenas due to their ability to break down bones. They are also believed to be social hunters since ''Epicyon hayden''i is very prevalent in the fossil record as one of the most common meat-eaters in North America during the late Miocene Epoch period. The deadly bite of a ''Epicyon haydeni'' was delivered by the canine teeth, which are placed near the front of the upper and lower jaws, the shortening of the jaws can be an effective method for getting the canines closer to the mandibular condyle, thereby increasing the biting force. ''Epicyon haydeni's'' small clavicle, flexible back, and digitigrade posture are all postcranial features shared with other canids and are likely adaptations designed to increase the animal's stride length. It seems from examinations of the limb proportions and toughness of the skeleton that ''Epicyon haydeni'' was less cursorial than hyaenas or modern wolves but more cursorial than other borophagine species like ''Aelurodon''. Unlike hyenas, ''Epicyon haydeni'' must have used their rearmost lower premolar (p4) and upper carnassial (P4) to crack large bones (ibid.). Smaller bones and bone fragments were likely crushed with the carnassials and postcarnassial molars just as in extant canids. Due to its bigger size and heavier, less gracile skeleton, ''Epicyon haydeni'' was less cursorial and unable to run as long a distance as ''Epicyon saevus''. Instead it relied on bursts of speed and social hunting strategies. ''Epicyon saevus'' existed from 16.3-4.9 million years ago. It is synonymous with ''Aelurodon inflatus'' and was named by Joseph Leidy in 1858 or 1859. In the late 1880s-early 1900s, Scott, Matthew, Cope and Matthew, Troxell recombined the animal as ''Aelurodon saevus''. It was recombined as ''Epicyon saevus'' by Baskin in 1980, Munthe in 1989, Voorhies in 1990, and Wang et al. 1999. Its estimated shoulder height is up to and body mass is up to . ''Epicyon aelurodontoides'' existed from 10.3-4.9 million years ago. It was named by X. Wang and others in 1999. It was found south of the Young Brothers Ranch,
Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to ...
.


Taxonomy

''Epicyon'' was first named by Joseph Leidy in 1858 as a subgenus of '' Canis''. It was also mentioned as belonging to the Aelurodontina by Matthew and Stirton in 1930. Later studies indicates that it was not a species of ''Canis'', but a borophagine.


Paleoecology

In North America, in places such as Coffee Ranch in Texas, ''Epicyon'' shared territory with the bear '' Agriotherium'' and the feliform '' Barbourofelis'', machairodont cat '' Amphimachairodus coloradensis'', and fellow canid '' Borophagus''. All of these animals were potential competitors that would have occasionally conflicted with ''Epicyon ''for food and territory. Prey for ''Epicyon'' included herbivores such as the camel '' Aepycamelus'', the pronghorn '' Cosoryx'', horses such as '' Neohipparion'' and '' Nannippus'', the peccary ''Prosthennops'', and the rhinoceroses such as '' Teleoceras'', all of which could provide a suitable meal through hunting or scavenging.


References

*Alan Turner, "National Geographic: Prehistoric Mammals" (Washington, D.C.: Firecrest Books Ltd., 2004), pp. 112–114.


General references

*Xiaoming Wang, Richard H. Tedford, Mauricio Antón, ''Dogs: Their Fossil Relatives and Evolutionary History'', New York : Columbia University Press, 2008; {{Taxonbar, from=Q2415423 Borophagines Miocene canids Miocene mammals of North America Prehistoric carnivoran genera Fossil taxa described in 1858