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''Daeodon'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
entelodont Entelodontidae, the entelodonts, are an extinct family of pig-like artiodactyls (even-toed ungulates) which inhabited the Northern Hemisphere (Asia, Europe, and North America) from the late Eocene to the Middle Miocene epochs, about 38-19 million ...
even-toed ungulates The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
that inhabited
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 the Car ...
about 23 to 20 million years ago during the latest
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
and earliest
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 ...
. The type species is ''Daeodon shoshonensis'', described by a very questionable holotype by Cope. Some authors synonimize it with ''Dinohyus hollandi'' and several other species (see below), but due to the lack of diagnostic material, this is questionable at best. Another large member of this family, similar in size to ''Daeodon'', is the Asian ''
Paraentelodon ''Paraentelodon'' is an extinct entelodont from the Late Oligocene and Oligocene-Miocene boundary of Asia. The fossils of the type species ''P. intermedium'' were found in Georgia, Kazakhstan and China. An indeterminate species represents in ...
,'' but it is known by very incomplete material.


Taxonomy

The genus ''Daeodon'' was erected by the American anatomist and
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interested ...
in 1878. He classified it as a
perissodactyl Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) o ...
and thought that it was closely related to '' Menodus''. This classification persisted until the description of ''"
Elotherium ''Entelodon'' (meaning "complete teeth", from Ancient Greek ''entelēs'' "complete" and ''odōn'' "tooth", referring to its "complete" eutherian dentition), is an extinct genus of entelodont artiodactyl endemic to Eurasia. Fossils of species a ...
" calkinsi'' in 1905, a very similar and much more complete animal from the same rocks, which was promptly assigned as a species of ''Daeodon'' by Peterson (1909). This led to ''Daeodon''s reclassification as a member of the family
Entelodont Entelodontidae, the entelodonts, are an extinct family of pig-like artiodactyls (even-toed ungulates) which inhabited the Northern Hemisphere (Asia, Europe, and North America) from the late Eocene to the Middle Miocene epochs, about 38-19 million ...
idae. The exact relationships between ''Daeodon'' and other entelodonts are not well understood; some authors (Lucas et al., 1998) consider the greater morphological similarity of ''Daeodon'' to ''
Paraentelodon ''Paraentelodon'' is an extinct entelodont from the Late Oligocene and Oligocene-Miocene boundary of Asia. The fossils of the type species ''P. intermedium'' were found in Georgia, Kazakhstan and China. An indeterminate species represents in ...
'' rather than to earlier North American entelodonts, like ''
Archaeotherium ''Archaeotherium'' ( grc, αρχαιοθήριον, meaning "ancient beast") is an extinct genus of entelodont artiodactyl endemic to North America during the Eocene and Oligocene epochs (35—28  mya), existing for approximately . ''Archa ...
'', as evidence for ''Daeodon'' being a descendant from a
Late Oligocene The Chattian is, in the geologic timescale, the younger of two ages or upper of two stages of the Oligocene Epoch/ Series. It spans the time between . The Chattian is preceded by the Rupelian and is followed by the Aquitanian (the lowest stage ...
immigration of large Asian entelodonts to North America. However, the existence of distinct specimens of ''Archaeotherium'' showing characters reminiscent of those present in both ''Paraentelodon'' and ''Daeodon'' raises the possibility of both genera actually descending from a North American common ancestor. Although not specified in Cope's original description, the name ''Daeodon'' comes from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words , meaning "hostile" or "dreadful" and , meaning "teeth".


Species

The type species of ''Daeodon'' is ''D. shohonensis,'' which is based on a fragment of a lower jaw from the
John Day Formation The John Day Formation is a series of rock strata exposed in the Picture Gorge district of the John Day River basin and elsewhere in north-central Oregon in the United States. The Picture Gorge exposure lies east of the Blue Mountain uplift, whi ...
of
Oregon Oregon () is a U.S. state, state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington (state), Washington, while the Snake River delineates much of it ...
. Several other species were assigned to the genus in the subsequent decades, like ''D. calkinsi'', ''D. mento'' and ''D. minor''. Since 1945, it had been suggested that two other
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
were actually
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
s of ''Daeodon'', but the formalization of this referral didn't take place until the work of Lucas et al. (1998). ''Ammodon leidyanum'', named by Cope's rival,
O. C. Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among h ...
, and ''Dinohyus hollandi,'' a complete skeleton from the Agate Springs quarry of
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 southwe ...
, were found to be indistinguishable from each other and in turn both were indistinguishable from ''D. shoshonensis''. With the exception of ''D. calkinsi'', which was tentatively excluded from ''Daeodon'', the other previously recognized species of ''Daeodon'' were also synonymized to ''D. shoshonensis''. That same year, an obscure entelodont, ''Boochoerus humerosum'', was also synonymized to ''Daeodon'' by Foss and Fremd (1998) and, albeit its status as a distinct species was retained, they note that the differences could still be attributed to individual or population variation or sexual dimorphism.


Description

''Daeodon shoshonensis'' is the largest-known
entelodont Entelodontidae, the entelodonts, are an extinct family of pig-like artiodactyls (even-toed ungulates) which inhabited the Northern Hemisphere (Asia, Europe, and North America) from the late Eocene to the Middle Miocene epochs, about 38-19 million ...
; known adult individuals had skulls about long and were about tall at the shoulders. It's differentiated from other entelodonts by a suite of unique dental characters, the shape and relatively small size of the cheekbone flanges of its skull compared to those of ''
Archaeotherium ''Archaeotherium'' ( grc, αρχαιοθήριον, meaning "ancient beast") is an extinct genus of entelodont artiodactyl endemic to North America during the Eocene and Oligocene epochs (35—28  mya), existing for approximately . ''Archa ...
'', and the small size of its chin
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
, as well as features of its
carpus In human anatomy, the wrist is variously defined as (1) the carpus or carpal bones, the complex of eight bones forming the proximal skeletal segment of the hand; "The wrist contains eight bones, roughly aligned in two rows, known as the carpal ...
and tarsus and the fusion of the bones of the lower leg. Like other entelodonts, its limbs were long and slender with the bones of the foreleg fused together and with only two toes on each foot. It also had a relatively lightly constructed neck for the size of its head, whose weight was mostly supported by muscles and tendons attached to the tall spines of the thoracic
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e, similar to those of modern-day
bison Bison are large bovines in the genus ''Bison'' (Greek: "wild ox" (bison)) within the tribe Bovini. Two extant and numerous extinct species are recognised. Of the two surviving species, the American bison, ''B. bison'', found only in North Ame ...
and
white rhinoceros The white rhinoceros, white rhino or square-lipped rhinoceros (''Ceratotherium simum'') is the largest extant species of rhinoceros. It has a wide mouth used for grazing (behaviour), grazing and is the most social of all rhino species. The white ...
.


Paleoecology


Habitat

''Daeodon'' had a wide range in North America, with many fossils found in Agate Fossil Beds, representing an environment in a transition period between dense forests and expansive prairie, likely a major cause of their extinction in the middle Miocene. It adapted to the grassland with a more cursorial body plan than more basal entelodonts like ''Archaeotherium,'' losing their dewclaws entirely, proximally fused metacarpals, and similar shoulder musculature to bison. The Agate Springs bonebed was a floodplain environment with wet and dry seasons. ''Daeodon'' shared this landscape with small gazelle-like camels, the large browsing chalicothere ''
Moropus ''Moropus'' (meaning "slow foot") is an extinct genus of large perissodactyl ("odd-toed" ungulate) mammal in the chalicothere family. They were endemic to North America during the Miocene from ~20.4—13.6  Mya, existing for approximately ...
'', several species of predatory coyote- to wolf-sized amphicyonids that lived in packs, land beavers (''
Palaeocastor ''Palaeocastor'' ('prehistoric beaver') is an extinct genus of beavers that lived in the North American Badlands during the late Oligocene period to early Miocene. ''Palaeocastor'' was much smaller than modern beavers. There are several specie ...
'') that filled the ecological niche of modern prairie dogs, and thousands of small herd-living rhinoceros. The rhinos suffered massive periodic die-offs in the dry season, but ''Daeodon'' fossils are rare, which suggests they were neither social animals nor especially attracted to carrion.


Diet

''Daeodon'' was omnivorous like all other entelodonts. Enamel patterns suggest eating of nuts, roots, and vines, as well as meat and bones. The superficial similarity to peccaries, hippos, and bears implies a wide range in terms of what plants ''Daeodon'' may have been eating. The dry seasons of North America at the time could get very harsh, so they may have supplemented their water intake by eating grape vines. The extent of its carnivory is debated, but tooth wear suggests they specialized in crushing bone and ripping meat, and bite marks on chalicothere bones suggest they either hunted or scavenged large herbivores. Foss (2001) argues its head was far too heavy to be effective in taking down large prey so it must have relied exclusively on scavenging, but its bison-like adaptations for running, the stereoscopic vision characteristic of predators, and evidence of predation in entelodonts calls this interpretation into question. The uncertainty of their diets suggests they were likely opportunistic omnivores similar to bears, eating whatever they need depending on the circumstance.Foss, S. E., 2001, Systematics and paleobiology of the Entelodontidae (Mammalia, Artiodactyla)
h.D. dissertation Hilda Doolittle (September 10, 1886 – September 27, 1961) was an American modernist poet, novelist, and memoirist who wrote under the name H.D. throughout her life. Her career began in 1911 after she moved to London and co-founded the ...
Dekalb, Northern Illinois University


Behavior

Entelodonts partook in intraspecific face biting, known from tooth marks on their skulls. Males would fight for dominance, possibly using their mandibular tubercles as protection in addition to their function as muscle attachments. Sexual dimorphism of the jugal protections exist in ''Archaeotherium,'' and with a smaller ''Daeodon'' sample size, such dimorphism can't be ruled out for ''Daeodon''. If dimorphic, the function of the expanded jugals was likely for display, supporting large preorbital glands similar to those forest hogs possessed for chemical communication.


In culture

''Daeodon'' appears in the game Ark: Survival Evolved.


References

{{Taxonbar, from=Q133085 Entelodonts Miocene even-toed ungulates Miocene mammals of North America Oligocene mammals of North America Arikareean Hemingfordian White River Fauna Burdigalian genus extinctions Chattian genus first appearances Fossil taxa described in 1878 Taxa named by Edward Drinker Cope Prehistoric even-toed ungulate genera