Daeodon Shoshonensis Skeletal
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''Daeodon'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
entelodont Entelodontidae is 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 early Miocene epochs, about 38-19 million years ago. Their ...
even-toed ungulates Artiodactyls are placental mammals belonging to the order Artiodactyla ( , ). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof). The other t ...
that inhabited
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
about 29 to 15.97 million years ago from the early
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
to late early
Miocene The Miocene ( ) is the first epoch (geology), 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 mea ...
. The type species is ''Daeodon shoshonensis'', described from a very fragmentary holotype by Cope. Some authors synonymize it with ''Dinohyus hollandi'' and several other species (see below), but due to the lack of diagnostic material, this may be questionable. Another large member of this family, possibly larger than ''Daeodon'', is the Asian ''
Paraentelodon ''Paraentelodon'' is an extinct entelodont from the Late Oligocene of Asia. The fossils of the type species ''P. intermedium'' were found in Georgia, Kazakhstan and China. An indeterminate species represents in Bugti Hills which is the late Oli ...
,'' but it is known by very incomplete material.


Taxonomy

The genus ''Daeodon'' was erected by the American anatomist and
paleontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontology, paleontologist, comparative anatomy, comparative anatomist, herpetology, herpetologist, and ichthyology, ichthyologist. Born to a wealthy Quaker fam ...
in 1878. He classified it as a
perissodactyl Perissodactyla (, ), or odd-toed ungulates, is an order of Ungulate, ungulates. The order includes about 17 living species divided into three Family (biology), families: Equidae (wild horse, horses, Asinus, asses, and zebras), Rhinocerotidae ( ...
and thought that it was closely related to ''
Menodus ''Megacerops'' ("large-horned face", from ''wikt:μεγα, méga-'' "large" + ''wikt:κέρας, kéras'' "horn" + ''wikt:ὤψ, ōps'' "face") is an extinct genus of the prehistoric odd-toed ungulate (hoofed mammal) family (biology), family Bro ...
''. This classification persisted until the description of ''" Elotherium" calkinsi'' in 1905, a very similar and much more complete animal from the same rocks, which was promptly assigned as a species of ''Dynohyus'' by Peterson (1909). This led to ''Daeodon''s reclassification as a member of the family
Entelodont Entelodontidae is 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 early Miocene epochs, about 38-19 million years ago. Their ...
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 of Asia. The fossils of the type species ''P. intermedium'' were found in Georgia, Kazakhstan and China. An indeterminate species represents in Bugti Hills which is the late Oli ...
'' rather than to earlier North American entelodonts, like ''
Archaeotherium ''Archaeotherium'' (, meaning "ancient beast") is an extinct genus of entelodont artiodactyl endemic to North America during the Eocene and Oligocene epochs (35–28 mya). ''Archaeotherium'' fossils are most common in the White River Formation ...
'', as evidence for ''Daeodon'' being a descendant from a
Late Oligocene The Chattian is, in the geologic timescale The geologic time scale or geological time scale (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the pro ...
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: 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 of all kno ...
words , meaning "hostile" or "dreadful" and , meaning "teeth".


Species

The type species of ''Daeodon'' is ''D. shoshonensis,'' 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 Mountains (Oregon), ...
of
Oregon Oregon ( , ) is a U.S. state, state in the Pacific Northwest region of the United States. It is a part of the Western U.S., with the Columbia River delineating much of Oregon's northern boundary with Washington (state), Washington, while t ...
. 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''; : 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 name and ...
were actually
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
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. A prolific fossil collector, Marsh was one of the preeminent paleontologists of the nineteenth century. Among his legacies are the discovery or ...
, and ''Dinohyus hollandi,'' a complete skeleton from the Agate Springs quarry of
Nebraska Nebraska ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Ka ...
, 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 is 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 early Miocene epochs, about 38-19 million years ago. Their ...
; 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'' (, meaning "ancient beast") is an extinct genus of entelodont artiodactyl endemic to North America during the Eocene and Oligocene epochs (35–28 mya). ''Archaeotherium'' fossils are most common in the White River Formation ...
'', 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 projectio ...
, 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 Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e, similar to those of modern-day
bison A bison (: bison) is a large bovine in the genus ''Bison'' (from Greek, meaning 'wild ox') within the tribe Bovini. Two extant taxon, extant and numerous extinction, extinct species are recognised. Of the two surviving species, the American ...
and
white rhinoceros The white rhinoceros, also known as the white rhino or square-lipped rhinoceros (''Ceratotherium simum''), is the largest extant species of rhinoceros and the most Sociality, social of all rhino species, characterized by its wide mouth adapted f ...
.


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 early 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 ''
Stenomylus ''Stenomylus'' is an extinct genus of miniature camelid native to North America that is known from the Oligocene and Miocene epochs(23.1~16.3 Ma). Its name is derived from the Greek (, "narrow") and (, "molar"). ''Stenomylus'' was extremely d ...
'', the large browsing
chalicothere Chalicotheriidae (from Ancient Greek ''khálix'', "gravel", and ''theríon'', "beast") is an extinct family of herbivorous, odd-toed ungulate (perissodactyl) mammals that lived in North America, Eurasia, and Africa from the Middle Eocene to the ...
''
Moropus ''Moropus'' (meaning "slow foot") is an extinct genus of large perissodactyl mammal in the chalicothere family. They were endemic to North America during the Miocene from ~20.4–13.6 Mya (unit), Mya, existing for approximately . ''Moropus'' belo ...
'', several species of predatory coyote- to wolf-sized
amphicyonids Amphicyonidae is an extinct family of terrestrial carnivorans belonging to the suborder Caniformia. They first appeared in North America in the middle Eocene (around 45 mya), spread to Europe by the late Eocene (35 mya), and further spread to Asi ...
that lived in packs, land beavers (''
Palaeocastor ''Palaeocastor'' ('ancient beaver') is an extinct genus of beavers that lived in the North American Badlands during the late Oligocene period to early Miocene, 29.5~18.5 million years ago. ''Palaeocastor'' was much smaller than modern beavers. Th ...
'') 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 needed 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 th ...
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.


References

{{Taxonbar, from=Q133085 Entelodonts Miocene Artiodactyla 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 Artiodactyla genera Oligocene Artiodactyla