Merycoidodontoidea
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Merycoidodontoidea, sometimes called "oreodonts" or "
ruminating Ruminants (suborder Ruminantia) are ungulate, hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principally ...
hogs", is an extinct superfamily of prehistoric
cud Cud is a portion of food that returns from a ruminant's stomach to the mouth to be chewed for the second time. More precisely, it is a bolus of semi-degraded food regurgitated from the reticulorumen of a ruminant. Cud is produced during the phy ...
-chewing
artiodactyl 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 ...
s with short faces and fang-like
canine teeth In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
. As their name implies, some of the better known forms were generally hog-like, and the group has traditionally been placed within the Suina (
pigs The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus ...
,
peccaries A peccary (also javelina or skunk pig) is a medium-sized, pig-like hoofed mammal of the family Tayassuidae (New World pigs). They are found throughout Central and South America, Trinidad in the Caribbean, and in the southwestern area of North ...
and their ancestors), though some recent work suggests they may have been more closely related to camels.Spaulding, M., O'Leary, M.A. & Gatesy, J. (2009): Relationships of Cetacea (Artiodactyla) Among Mammals: Increased Taxon Sampling Alters Interpretations of Key Fossils and Character Evolution. '' PLoS ONE'' no 4(9): e7062.
article
/ref> "Oreodont" means "mountain teeth", referring to the appearance of the molars. Most oreodonts were sheep-sized, though some genera grew to the size of
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult ma ...
. They were heavy-bodied, with short four-toed
hooves The hoof (plural: hooves) is the tip of a toe of an ungulate mammal, which is covered and strengthened with a thick and horny keratin covering. Artiodactyls are even-toed ungulates, species whose feet have an even number of digits, yet the rumi ...
and comparatively long tails. The animals would have looked rather pig- or sheep-like, but features of their teeth indicate they were more closely related to
camelid Camelids are members of the biological family Camelidae, the only currently living family in the suborder Tylopoda. The seven extant members of this group are: dromedary camels, Bactrian camels, wild Bactrian camels, llamas, alpacas, vicuñas, ...
s. They were most likely woodland and grassland
browsers Browse, browser or browsing may refer to: Programs * Web browser, a program used to access the World Wide Web *Code browser, a program for navigating source code * File browser or file manager, a program used to manage files and related objects * ...
, and were widespread in North America during the Oligocene and
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 ...
. Later forms diversified to suit a range of different habitats. For example, ''
Promerycochoerus ''Promerycochoerus'' ("Before '' Merycochoerus''" or "Before Ruminating Hog") is an extinct genus of hippopotamus-like oreodont artiodactyl that lived in Central North America during the Early Miocene. The 1 m (3 ft 4 in) long c ...
'' had adaptations suggesting a semiamphibious lifestyle, similar to that of modern
hippos A hippo or hippopotamus is either of two species of large African mammal which live mainly in and near water: * Hippopotamus * Pygmy hippopotamus Hippo or Hippos may also refer to: Toponymy * The ancient city of Hippo Regius (modern Annaba, Al ...
.


Taxonomy

The two
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
of oreodonts are the Merycoidodontidae (originally known as Oreodontidae) which contains all of the advanced species, and the Agriochoeridae, smaller, primitive oreodonts. Together they form the now-extinct suborder Oreodonta. Oreodonts may have been distantly related to
pigs The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus ...
,
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...
es, and the pig-like
peccaries A peccary (also javelina or skunk pig) is a medium-sized, pig-like hoofed mammal of the family Tayassuidae (New World pigs). They are found throughout Central and South America, Trinidad in the Caribbean, and in the southwestern area of North ...
. Indeed, some scholars place Merycoidodontidae within the pig-related suborder Suina (Suiformes). Other scholars place oreodonts closer to camels in the suborder
Tylopoda Tylopoda (meaning "calloused foot") is a suborder of terrestrial herbivorous even-toed ungulates belonging to the order Artiodactyla. They are found in the wild in their native ranges of South America and Asia, while Australian feral camels ...
. Still, other experts put the oreodonts together with the short-lived cainotheres in the taxonomic suborder
Ancodonta Ancodonta is an infraorder of semiaquatic artiodactyl ungulates including modern hippopotamus and all mammals closer to hippos than to cetaceans (whales). Ancodonts first appeared in the Middle Eocene, with some of the earliest representatives ...
comprising these two groups of extinct ancodonts. All scholars agree, however, that the oreodont was an early form of
even-toed ungulate 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 ...
, belonging to the order Artiodactyla. Today, most evidence points towards the oreodonts being tylopods, along with camels,
xiphodont Xiphodontidae is an extinct family of even-toed ungulates (order Artiodactyla), endemic to Europe during the Eocene 40.4—33.9 million years ago, existing for about 7.5 million years. They were, most likely, all terrestrial herbivores ...
s, and protoceratids. Over 50 genera of Oreodonta have been described in the paleozoological literature. However, oreodonts are widely considered to be taxonomically oversplit, and many of these genera may prove to be synonymous. The last researchers to fully review oreodont taxonomy, C. Bertrand Schultz and Charles H. Falkenbach, have been criticized for erecting excessive numbers of genera, based in part on apparent anatomical differences between different specimens that were actually taphonomic deformations due to postburial forces. Undeformed skulls would be placed in one genus, while skulls crushed from side to side would be placed in a second genus and skulls crushed from front to back would be placed in a third genus. Researchers are beginning to restudy oreodonts and synonymize many genera, but only a few groups have been reviewed.


Natural history

This diverse group of stocky prehistoric mammals grazed amid the
grassland A grassland is an area where the vegetation is dominated by grasses ( Poaceae). However, sedge ( Cyperaceae) and rush ( Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur na ...
s,
prairie Prairies are ecosystems considered part of the temperate grasslands, savannas, and shrublands biome by ecologists, based on similar temperate climates, moderate rainfall, and a composition of grasses, herbs, and shrubs, rather than trees, as the ...
s, or
savanna A savanna or savannah is a mixed woodland- grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
s of
North North is one of the four compass points or cardinal directions. It is the opposite of south and is perpendicular to east and west. ''North'' is a noun, adjective, or adverb indicating direction or geography. Etymology The word ''north ...
and
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
throughout much of the Cenozoic era. First appearing 48 million years ago (Mya) during the warm
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
epoch of the Paleogene period, the oreodonts dominated the American landscape 34 to 23 Mya during the dry Oligocene epoch, but they mysteriously disappeared 4 Mya during the colder
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Neogene period. Today,
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
jaws and teeth of the Oreodonta are commonly found amid the 'Oreodon beds' (
White River Fauna The White River Fauna are fossil animals found in the White River Badlands of South Dakota, Wyoming, Colorado and Nebraska in the United States including Badlands National Park. The fossils have been found in the White River Formation, Chadron F ...
) of the White River
badlands Badlands are a type of dry terrain where softer sedimentary rocks and clay-rich soils have been extensively eroded."Badlands" in ''Chambers's Encyclopædia''. London: George Newnes, 1961, Vol. 2, p. 47. They are characterized by steep slopes, m ...
in
South Dakota South Dakota (; Sioux: , ) is a U.S. state in the North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Lakota and Dakota Sioux Native American tribes, who comprise a large porti ...
,
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 ...
,
Colorado Colorado (, other variants) is a state in the Mountain states, Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the wes ...
and
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to the s ...
. Many oreodont bones have also been reported at the
John Day Fossil Beds National Monument John Day Fossil Beds National Monument is a U.S. national monument in Wheeler and Grant counties in east-central Oregon. Located within the John Day River basin and managed by the National Park Service, the park is known for its well-preserved l ...
in
Oregon Oregon () is a state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington, while the Snake River delineates much of its eastern boundary with Idaho. T ...
. Some oreodonts have been found at
Agate Fossil Beds National Monument Agate Fossil Beds National Monument is a U.S. National Monument near Harrison, Nebraska. The main features of the monument are a valley of the Niobrara River and the fossils found on Carnegie Hill and University Hill. The area largely consists of ...
. In Oligocene/Miocene
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 ...
, oreodonts are surprisingly rare. Instead of the swarms found elsewhere, only six genera of oreodonts are known to have ranged there, and only one, '' Mesoreodon'', is known from a single, good skeleton.


Lifestyle

The majority of oreodonts are presumed to have lived in herds, as suggested by the thousands of individuals in the various mass mortalities seen in the White River
Badlands Badlands are a type of dry terrain where softer sedimentary rocks and clay-rich soils have been extensively eroded."Badlands" in ''Chambers's Encyclopædia''. London: George Newnes, 1961, Vol. 2, p. 47. They are characterized by steep slopes, m ...
, Nebraska Oreodont beds, or Chula Vista, California.


Diversity

Oreodonts underwent a huge diversification during the Oligocene and Miocene, adapting to a number of ecological niches, including: *Semiaquatic – hippo-like ''
Promerycochoerus ''Promerycochoerus'' ("Before '' Merycochoerus''" or "Before Ruminating Hog") is an extinct genus of hippopotamus-like oreodont artiodactyl that lived in Central North America during the Early Miocene. The 1 m (3 ft 4 in) long c ...
'' *Trunked browser – tapir-like ''
Brachycrus ''Brachycrus'' is an extinct genus of oreodont, of the family Merycoidodontidae, endemic to North America. They lived during the Middle Miocene, 16.0—13.6 mya, existing for approximately . The long creature resembled its bigger, earlier ...
'' *Large grazer – cow-sized '' Eporeodon'' *Medium grazer – goat-like ''
Merycoidodon ''Merycoidodon'' ("ruminating teeth") is an extinct genus of herbivorous artiodactyl of the family Merycoidodontidae, more popularly known by the name ''Oreodon'' ("hillock teeth"). It was endemic to North America during the Middle Eocene to Mid ...
'' *Small desert herbivore – goat- to cat-sized ''
Sespia ''Sespia'' ("of Sespe Creek") is an extinct genus of oreodont endemic to North America. They lived during the Late Oligocene 26.3—24.8 Annum, mya, existing for approximately . ''Sespia'' was cat to goat-sized and desert-dwelling. The genus wa ...
'' *Medium desert herbivore – '' Mesoreodon'' and the ubiquitous '' Leptauchenia''


Classification

The family Merycoidodontidae is divided into eleven subfamilies, with four genera not included in any subfamily (''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'') because they are either regarded as basal oreodonts, or their status within the family remains uncertain. *Family †Merycoidodontidae **subfamily ''incertae sedis'' ***†'' Aclistomycter'' ***†''
Merychyus ''Merychyus'' is an extinct genus of oreodont of the family Merycoidodontidae, endemic to North America. It lived during the Miocene, 20.4—10.3 Annum, mya, existing for approximately .Pseudogenetochoerus'' ***†'' Pseudoleptauchenia'' **Subfamily † Oreonetinae ***†'' Bathygenys'' ***†'' Megabathygenys'' ***†'' Oreonetes'' **Subfamily †
Leptaucheniinae Leptaucheniinae ("Delicate Necks") was a taxon of small, goat-like oreodonts with proportionally big heads found throughout North America during the Late Oligocene. Because skeletons of '' Leptauchenia'' and ''Sespia'' have been found by the l ...
***Tribe † Leptaucheniini ****†'' Limnenetes'' ****†'' Leptauchenia'' ***Tribe †
Sespiini ''Sespia'' ("of Sespe Creek") is an extinct genus of oreodont endemic to North America. They lived during the Late Oligocene 26.3—24.8 Annum, mya, existing for approximately . ''Sespia'' was cat to goat-sized and desert-dwelling. The genus wa ...
****†''
Sespia ''Sespia'' ("of Sespe Creek") is an extinct genus of oreodont endemic to North America. They lived during the Late Oligocene 26.3—24.8 Annum, mya, existing for approximately . ''Sespia'' was cat to goat-sized and desert-dwelling. The genus wa ...
'' **Subfamily †
Merycoidodontinae Merycoidodontoidea, sometimes called "oreodonts" or "ruminating hogs", is an extinct superfamily of prehistoric cud-chewing artiodactyls with short faces and fang-like canine teeth. As their name implies, some of the better known forms were gen ...
(syn. Oreodontinae) ***†''
Merycoidodon ''Merycoidodon'' ("ruminating teeth") is an extinct genus of herbivorous artiodactyl of the family Merycoidodontidae, more popularly known by the name ''Oreodon'' ("hillock teeth"). It was endemic to North America during the Middle Eocene to Mid ...
'' (syn. ''Blickohyus'', ''Genetochoerus'', ''Oreodon'', ''Otionohyus'', ''Paramerycoidodon'', ''Prodesmatochoerus'', ''Promesoreodon'', ''Subdesmatochoerus'') ***†'' Mesoreodon'' **Subfamily †
Miniochoerinae ''Miniochoerus'' is an extinct genus of small oreodont endemic to North America. They lived during the Late Eocene to Early Oligocene 38–30.8 mya, existing for approximately . Fossils have been found only in North Dakota, South Dakota, Nebras ...
***†'' Miniochoerus'' (syn. ''Paraminiochoerus'', ''Parastenopsochoerus'', ''Platyochoerus'', ''Pseudostenopsochoerus'', ''Stenopsochoerus'') **Subfamily † Desmatochoerinae ***†'' Desmatochoerus'' ***†'' Eporeodon'' ***†'' Megoreodon'' **Subfamily † Promerycochoerinae ***†''
Promerycochoerus ''Promerycochoerus'' ("Before '' Merycochoerus''" or "Before Ruminating Hog") is an extinct genus of hippopotamus-like oreodont artiodactyl that lived in Central North America during the Early Miocene. The 1 m (3 ft 4 in) long c ...
'' ***†''
Merycoides ''Merycoides'' is an extinct genus of oreodont Merycoidodontoidea, sometimes called "oreodonts" or "ruminating hogs", is an extinct superfamily of prehistoric cud-chewing artiodactyls with short faces and fang-like canine teeth. As their name ...
'' **Subfamily † Merychyinae ***†'' Oreodontoides'' ***†'' Paroreodon'' ***†''
Merycoides ''Merycoides'' is an extinct genus of oreodont Merycoidodontoidea, sometimes called "oreodonts" or "ruminating hogs", is an extinct superfamily of prehistoric cud-chewing artiodactyls with short faces and fang-like canine teeth. As their name ...
'' ***†''
Merychyus ''Merychyus'' is an extinct genus of oreodont of the family Merycoidodontidae, endemic to North America. It lived during the Miocene, 20.4—10.3 Annum, mya, existing for approximately .Eporeodontinae ***†'' Dayohyus'' (syn. ''Eucrotaphus'' deemed '' nomen dubium'') ***†'' Eporeodon'' **Subfamily † Phenacocoelinae ***†'' Phenacocoelus'' ***†''
Hypsiops ''Hypsiops'' is an extinct genus of oreodont of the family Merycoidodontidae endemic to North America. They lived during the Late Oligocene to Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch ...
'' **Subfamily † Ticholeptinae ***†'' Mediochoerus'' ***†'' Ticholeptus'' ***†''Ustatochoerus'' **Subfamily † Merycochoerinae ***†''
Merycochoerus ''Merycochoerus'' (Greek: "ruminant" (merux)-like "swine" (khoiros)) is an extinct genus of oreodont of the family Merycoidodontidae, endemic to North America. They lived during the Early Oligocene 33.9—30.8 mya, existing for approximately ...
'' ***†''
Brachycrus ''Brachycrus'' is an extinct genus of oreodont, of the family Merycoidodontidae, endemic to North America. They lived during the Middle Miocene, 16.0—13.6 mya, existing for approximately . The long creature resembled its bigger, earlier ...
'' In Lander (1998) the classification of Oreodontoidea was as follows: * Family Agriochoeridae Leidy, 1869 (syn. Artionychidae, Eomerycidae, Protoreodontidae) ** Subfamily Agriochoerinae Gill, 1872 (syn. Diplobunopsinae) *** ''Agriochoerus'' Leidy, 1850b (syn. ''Agriomeryx'', ''Artionyx'', ''Coloreodon'', ''Diplobunops'', ''Eomeryx'', ''Merycopater'') *** "Agriochoerus" maximus (Douglass, 1901) ** Subfamily Protoreodontinae Scott, 1890 *** ''Protoreodon'' Scott and Osborn, 1887 (syn. ''Agriotherium'', ''Chorotherium'', ''Hyomeryx'', ''Mesagriochoerus'', ''Protagriochoerus'') *** "Protoreodon" petersoni (Gazin, 1955) *** "Agriochoerus" minimus (Douglass, 1901) *** "Agriochoerus" transmontanus (Stock, 1949) * Family Merycoidodontidae ** Subfamily Bathygeniinae Lander, 1998 *** ''Bathygenys'' Douglass, 1901 (syn. ''Megabathygenys'', ''Parabathygenys'') ** Subfamily Aclistomycterinae Lander, 1998 *** ''Aclistomycter'' Wilson, 1971 ** Subfamily Leptaucheniinae Schultz and Falkenbach, 1940 *** ''Leptauchenia'' Leidy, 1856 (syn. ''Brachymeryx'', ''Cyclopidius'', ''Hadroleptauchenia'', ''Limnenetes'', ''Pithecistes'', ''Pseudocyclopidius'', ''Pseudoleptauchenia'') *** ''Sespia'' Stock, 1930 (syn. ''Megasespia'') ** Subfamily Miniochoerinae Schultz and Falkenbach, 1956 (syn. Oreonetinae, ?Cotylopinae, ?Merycoidodontinae, ?Oreodontinae) ** Subfamily Eucrotaphinae Lander, 1998 ** Subfamily Merycochoerinae Schultz and Falkenbach, 1940 (syn. Desmatochoerinae, Eporeodontinae, Promerycochoerinae) ** Subfamily Phenacocoelinae Schultz and Falkenbach, 1950 ** Subfamily Ticholeptinae Schultz and Falkenbach, 1941 (syn. Merychyinae)


References

{{Taxonbar, from=Q134697 Mammal superfamilies Miocene extinctions White River Fauna Eocene first appearances Tylopoda