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Tremarctinae

The Tremarctinae or short-faced bears is a subfamily of Ursidae that contains one living representative, the spectacled bear of South America, and several extinct species from four genera: the Florida spectacled bear, the North American giant short-faced bears Arctodus, the South American giant short-faced bear Arctotherium as well as Plionarctos (P. edensis and P. harroldorum), which is thought to be ancestral to the other three genera. Of these, the giant short-faced bears may have been the largest ever carnivorans in the Americas. The group is thought to have originated in eastern North America, and then invaded South America as part of the Great American Interchange. Most short-faced bears became extinct at the end of the Pleistocene.

Taxonomy
Tremarctinae was erected by John Merriam and Chester Stock in 1925. Traditionally, analyses of the phylogenetic inner relationships of tremarctines had Plionarctos and Tremarctos as basal groups with respect to a short-faced bear clade of Arctodus and Arctotherium. A study of the affinities of bears belonging to Arctotherium indicates that they were more closely related to the spectacled bear than to Arctodus, implying convergent evolution of large size in the two lineages. Tremarctines are occasionally referred to as arctodonts and/or arctotheres, or tremarctotheres in older scientific literature. Diagnostics Skull Tremarctinae appear to have a disproportionately shorter snouts compared with most modern bears, giving them the name "short-faced." This apparent shortness is an illusion caused by the deep snouts and short nasal bones of tremarctine bears compared with ursine bears; Tremarctinae had shorter and taller skulls, but not a shorter face than most living bears. In addition to being brachycephalic, tremarctine bears' skulls possessed well developed zygomatic arches and glenoid mandibular fossas, Moreover, tremarctine bears' orbits are also bigger, more rounded and lateralized than ursine bears. The features of tremarctine bear's individual morphology, such as dentition, can be quite variable (particularly the M2 molar). Unlike tremarctine bears, ursine bears have only one masseteric fossa on their mandible and more slender and elongated skulls, with generally narrower molars (with the exception of polar bears). Osteological differences between tremarctine and ursine bears also include an extra lateral cusp between the trigonid and talonid on the m1 molar. Being very similar to extant spectacled bears, Ursichnus sudamericanus suggests all tremarctines had long claws and five digits whose tips arrayed in a shallow arc. While all are plantigrade, Tremarctinaes can be distinguished between short-footed (Tremarctos) and long-footed forms (Arctodus). == Evolution ==
Evolution
Tremarctinae originate with their common ancestor, Plionarctos, in the Middle Hemphillian (Late Miocene, ~7Ma) of North America; Plionarctos is last recorded in the early Blancan (Pliocene, ~3Ma).'''' and between Arctotherium and Tremarctos'' at 4.1 Ma. An indeterminate Arctotherium is first recorded from the late Pliocene of El Salvador. In the Early Pleistocene, short-faced bears began to establish themselves more thoroughly in both North and South America. The medium sized Arctodus pristinus inhabited a broad range in the North American continent, with Tremarctos floridanus endemic to the Gulf Coast. The first records of Arctotherium in South America come in the form of the gigantic Arctotherium angustidens, possibly the largest carnivorous land mammal ever, in Argentina circa 1 Ma. What the evolutionary history of Arctotherium in the previous 1.5 million years, and their history in South America, is unclear. In the Middle Pleistocene, both Arctodus and Arctotherium gave way to new forms; Arctodus pristinus gave way to the huge Arctodus simus, which inhabited a pan-continental range, from Alaska to Mexico. Arctotherium angustidens, on the other hand, was replaced by a series of smaller, medium-sized species- Arctotherium vetustum, then shortly thereafter by Arctotherium bonariense, and Arctotherium tarijense. and to reinvade Central America. By the terminal Pleistocene, Arctodus simus, Tremarctos floridanus, Arctotherium tarijense and Arctotherium wingei collectively occupied a range from Alaska to southernmost Patagonia. All of these forms were extinct by the end of the early Holocene. Scholars suggest that the spectacled bear migrated into Central and South America upon the extinction of Arctotherium wingei, if Pleistocene records of Andean Arctotherium sp. aren't confirmed as the spectacled bear. == Paleobiology ==
Paleobiology
Tremarctines inhabited a wide range of niches- from small and mostly herbivorous bears inhabiting more forested habitat, such as Arctotherium wingei and Tremarctos ornatus, to the colossal Arctotherium angustidens and Arctodus simus; plains adapted omnivores with a penchant for large quantities of meat. Although the two giant species appear superficially similar, both species had key, significant differences. While Arctodus simus had a wide range across North America for 800,000 years, Arctotherium angustidens appears to be limited to the Southern Cone, in open plains habitat. Furthermore, whereas Arctodus simus varied its diet between quasi-carnivory in Alaska to classic omnivory, Arctotherium angustidens had similar rates of carnivory across specimens, according to isotope studies. Additionally, the much more gracile form of Arctodus, in contrast with the robust Arctotherium angustidens, has puzzled researchers. ==Systematics==
Systematics
The following taxonomy of the tremarctine bears follow by Mitchell et al. (2016): • Subfamily Tremarctinae Merriam & Stock, 1925 • †Plionarctos Frick, 1926 • †Plionarctos edensis Frick, 1926 • †Plionarctos harroldorum Tedford & Martin, 2001 • †Arctodus Leidy, 1854 • †Arctodus pristinus Leidy, 1854 • †Arctodus simus Cope, 1879 • †Arctotherium Burmeister, 1879 • †Arctotherium angustidens Gervais & Ameghino, 1880 • †Arctotherium bonariense Gervais, 1852 • †Arctotherium tarijense Ameghino, 1902 • †Arctotherium vetustum Ameghino, 1885 • †Arctotherium wingei Ameghino, 1902 • Tremarctos Gervais, 1855 • †Tremarctos floridanus (Gildey, 1928) • Tremarctos ornatus (Cuvier, 1825) – spectacled bear ==See also==
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