Epicyon
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''Epicyon'' ("more than a dog") is a large, extinct,
canid Canidae (; from Latin, '' canis'', " dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found withi ...
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 nom ...
of the subfamily
Borophaginae The extinct Borophaginae form one of three subfamilies found within the canid family. The other two canid subfamilies are the extinct Hesperocyoninae and extant Caninae. Borophaginae, called "bone-crushing dogs", were endemic to North America ...
("bone-crushing dogs"), native to North America. ''Epicyon'' existed for about from the
Hemingfordian The Hemingfordian on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 20,600,000 to 16,300,000 years BP. It is usually considered to overlap the lates ...
age of the
Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages: the Aquitanian age, Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 annum, Ma to ...
, to the
Hemphillian The Hemphillian North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 10,300,000 to 4,900,000 years BP. It is usually considered t ...
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-like shape rather than having a skull similar in shape to that of a
wolf The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly un ...
; 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 ''Borophagus'' ("gluttonous eater") is an extinct genus of the subfamily Borophaginae, a group of canids endemic to North America from the Middle Miocene epoch through the Early Pleistocene epoch 12—1.8 Mya. Evolution ''Borophagus'', like o ...
'' (Mya) * ''
Carpocyon ''Carpocyon'' is an extinct genus of the Borophaginae subfamily of canids native to North America. It lived from the Middle to the Late Miocene, 13.6 to 5.3 Ma Mya, existing for approximately . The four species in the genus varied in size, with ...
'' from 20.4 to * ''
Paratomarctus ''Paratomarctus'' is an extinct monospecific genus of the Borophaginae subfamily of canids native to North America. It lived from the Middle to Late Miocene, 16.3—5.3 mya, existing for approximately . It was about the size of a coyote, and was ...
'' from 16.3 to * '' Aelurodon'' from 16.0 to * ''
Canis lepophagus ''Canis lepophagus'' (Latin: : 'dog'; : 'hare' or 'rabbit'; suffix : '-eating'; hence hare-eating dog) is an extinct species of canid which was endemic to much of North America during the Early Pliocene. It is notable because its lineage is prop ...
'' from 10.3 to


Species

Fossil specimens range from
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
to
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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 sout ...
, 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 th ...
,
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
,
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ke ...
, Colorado, Oklahoma, Idaho, Oregon, Arizona within the
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. As well as in
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest Ter ...
,
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 tot ...
. ''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 specime ...
, 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 th ...
.


Taxonomy

''Epicyon'' was first named by
Joseph Leidy Joseph Mellick Leidy (September 9, 1823 – April 30, 1891) was an American paleontologist, parasitologist and anatomist. Leidy was professor of anatomy at the University of Pennsylvania, later was a professor of natural history at Swarthmore ...
in 1858 as a subgenus of ''
Canis ''Canis'' is a genus of the Caninae which includes multiple extant species, such as wolves, dogs, coyotes, and golden jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and de ...
''. 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 The extinct Borophaginae form one of three subfamilies found within the canid family. The other two canid subfamilies are the extinct Hesperocyoninae and extant Caninae. Borophaginae, called "bone-crushing dogs", were endemic to North America ...
.


Paleoecology

In North America, in places such as Coffee Ranch in Texas, ''Epicyon'' shared territory with the bear '' Agriotherium'' and the feliform '' Barbourofelis'',
machairodont Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million until ...
cat '' Amphimachairodus coloradensis'', and fellow canid ''
Borophagus ''Borophagus'' ("gluttonous eater") is an extinct genus of the subfamily Borophaginae, a group of canids endemic to North America from the Middle Miocene epoch through the Early Pleistocene epoch 12—1.8 Mya. Evolution ''Borophagus'', like o ...
''. 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 ''Aepycamelus'' is an extinct genus of camelids that lived during the Miocene 20.6–4.9 million years ago, existing for about . Its name is derived from the Homeric Greek , "high and steep" and κάμηλος – "camel"; thus, "high camel"; ''a ...
'', the pronghorn '' Cosoryx'', horses such as ''
Neohipparion ''Neohipparion'' (Greek: "new" (neos), "pony" (hipparion)) is an extinct genus of equid, from the Neogene (Miocene to Pliocene) of North America and Central America Central America ( es, América Central or ) is a subregion of the Ameri ...
'' and ''
Nannippus ''Nannippus'' is an extinct genus of three-toed horse endemic to North America during the Miocene through Pleistoceneabout 13.3—1.8 million years ago (Mya), living around 11.5 million years. This ancient species of three-toed horse grew up to 3 ...
'', the peccary ''Prosthennops'', and the rhinoceroses such as ''
Teleoceras ''Teleoceras'' (Greek: "perfect" (teleos), "horn" (keratos)) is an extinct genus of grazing rhinoceros. It lived in North America during the Miocene and Pliocene epochs during the Hemingfordian to the end of Hemphillian from around 17.5 to 4. ...
'', 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