Borophagus
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''Borophagus'' ("gluttonous eater") is an extinct
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 In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...
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 dur ...
, a group of canids endemic to
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
from the Middle
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
epoch through the Early
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
epoch 12—1.8 Mya.


Evolution

''Borophagus'', like other borophagines, are loosely known as "bone-crushing" or "
hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the cl ...
-like" dogs. Though not the most massive borophagine by size or weight, it had a more highly evolved capacity to crunch bone than earlier, larger genera such as ''
Epicyon ''Epicyon'' ("more than a dog") is a large, extinct, canid genus of the subfamily Borophaginae ("bone-crushing dogs"), native to North America. ''Epicyon'' existed for about from the Hemingfordian age of the Early Miocene, to the Hemphillia ...
'', which seems to be an evolutionary trend of the group (Turner, 2004). During the Pliocene epoch, ''Borophagus'' began being displaced by other ''Canid'' species such as '' Canis edwardii'' and later by ''
Aenocyon dirus The dire wolf (''Aenocyon dirus'' ) is an extinct canine. It is one of the most famous prehistoric carnivores in North America, along with its extinct competitor ''Smilodon''. The dire wolf lived in the Americas and eastern Asia during the Lat ...
''. Early species of ''Borophagus'' were placed in the genus ''Osteoborus'' until recently, but the
genera 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 nomenclat ...
are now considered synonyms.


Description

Typical features of this genus are a bulging forehead and powerful jaws; ''Borophagus'' has been considered to be probably a
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feedin ...
by paleontologists in the past. Its crushing
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
teeth and strong jaw muscles would have been used to crack open bone, much like the
hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the cl ...
of the Old World. However, ''Borophagus'' fossils are so abundant and geographically widespread that some paleontologists now argue that ''Borophagus'' must have been both the dominant carnivore of its time, and thus an active predator because carrion feeding alone could not have sustained such a large population.Wang, Xiaoming; and Tedford, Richard H. Dogs: Their Fossil Relatives and Evolutionary History. New York: Columbia University Press, 2008. pp112-3 They note that not all carnivores with bone-cracking ability are scavengers, such as the modern spotted hyena; instead, they interpret the bone-cracking ability as an adaptation to social hunting where complete utilization of a carcass was favored. Coprolites from ''Borophagus'' further vindicate its bone-crushing abilities, while simultaneously indicating it occupied a niche no longer seen in the present-day ecosystems of North America. The discovery of these coprolites also indicates that ''Borophagus'' may have been a social pack-hunter. The adult animal is estimated to have been about 80 cm in length, similar to a
coyote The coyote (''Canis latrans'') is a species of canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecological nich ...
, although it was much more powerfully built.


Species

*'' Borophagus diversidens'' existed for (synonymous with ''Felis hillianus'', ''Hyaenognathus matthewi'', ''Hyaenognathus pachyodon'', ''Hyaenognathus solus'', ''Porthocyon dubius'') *'' Borophagus dudleyi'' existed for *'' Borophagus hilli'' existed for (synonymous with ''Osteoborus crassapineatus'', ''Osteoborus progressus'') *'' Borophagus littoralis'' existed for (syn. ''Osteoborus diabloensis'') *'' Borophagus orc'' existed for *'' Borophagus parvus'' existed for *'' Borophagus pugnator'' existed for (synonymous with ''Osteoborus galushai'') *'' Borophagus secundus'' existed for (synonymous with ''Hyaenognathus cyonoides'', ''Hyaenognathus direptor'', ''Osteoborus secundus'') Existence based on Figure 141 of Wang et al. (1999).


Paleoecology

In North America, in places such as Coffee Ranch in Texas, ''Borophagus'' was contemporary with the bear ''
Agriotherium ''Agriotherium'' is an extinct genus of bears whose fossils are found in Miocene through Pleistocene-aged strata of North America, Europe, Africa, and Asia. This long-lived genus persisted from at least ~11.6–2.5 Mya. Materials from the late- ...
'' as well as the feliform ''
Barbourofelis ''Barbourofelis'' is an extinct genus of large, predatory, feliform carnivoran mammals of the family Barbourofelidae (false saber-tooth cats). The genus was endemic to North America and Eurasia during the Miocene until its extinction during ...
'', the saber-toothed
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 ''Epicyon''. All of these animals were potential competitors that would have occasionally conflicted with ''Borophagus'' for food and territory, though it may also have readily scavenged their kills. Prey for ''Borophagus'' included herbivores like 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 antelope ''
Cosoryx ''Cosoryx'' is an extinct genus of antilocaprid that lived in the Miocene of Nevada. Fossils of this genus have also been found in the Santa Fe Group in New Mexico. ''Cosoryx'' has sometimes been considered synonymous with ''Merycodus ...
'', horses like ''
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 ancient peccary ''
Prosthennops ''Prosthennops'' is a genus of extinct peccaries that lived in North and Central America between the middle Miocene and lower Pliocene (around 15-5 million years ago). Description This animal was very similar to present-day peccaries, both ...
'' and even rhinoceroses like the hippo-like ''
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. *Xiaoming Wang
"The Origin and Evolution of the Dog Family"
Accessed 1/30/06.


Further reading



*Russell Hunt, [https://web.archive.org/web/20110926205607/http://darkwing.uoregon.edu/~rhunt/publications.htm "Ecological Polarities Of the North American Family Canidae: A New Approach to Understanding Forty Million Years of Canid Evolution" (Accessed 1/30/06)].
Wang et al., "Phylogenetic Systematics of the Borophaginae (Carnivora:Canidae)." ''Bulletin of the American Museum of Natural History'', No. 243, Nov. 17 1999. (PDF) (Accessed 4/11/06)
*Xiaoming Wang, Richard H. Tedford, Mauricio Antón, ''Dogs: Their Fossil Relatives and Evolutionary History'', New York : Columbia University Press, 2008; {{Taxonbar, from=Q137418 Borophagines Miocene canids Pliocene carnivorans Neogene mammals of North America Prehistoric carnivoran genera Fossil taxa described in 1892