The Tremarctinae or short-faced bears is a

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...

of Ursidae that contains one living representative, the spectacled bear (''Tremarctos ornatus'') of

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

, and several extinct species from four genera: the Florida spectacled bear (''Tremarctos floridanus''), the North American giant short-faced bears '' Arctodus'' (''A. pristinus'' and ''A. simus''), the South American giant short-faced bear ''

Arctotherium ''Arctotherium'' ("bear beast") is an extinct genus of the Pleistocene Tremarctinae, short-faced bears endemic to Central America, Central and South America. ''Arctotherium'' migrated from North America to South America during the Great American In ...

'' (including ''A. angustidens'', ''A. vetustum'', ''A. bonariense'', ''A. wingei'', and ''A. tarijense)'' 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 ('' Arctodus simus'' and '' Arctotherium angustidens'') may have been the largest ever carnivorans in the

Americas The Americas, sometimes collectively called America, are a landmass comprising the totality of North America and South America.''Webster's New World College Dictionary'', 2010 by Wiley Publishing, Inc., Cleveland, Ohio. When viewed as a sing ...

. 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 The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...

Evolution

Tremarctinae originate with their common ancestor, '' Plionarctos,'' in the Middle Hemphillian (earliest

Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

, ~10Ma) of North America; '' Plionarctos'' is last recorded in the early Blancan (Early Pliocene, ~3.3Ma). Around the

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 ...

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58ursids, experienced an explosive radiation in diversity, as C4 vegetation ( grasses) and open habitats dominated, the world experienced a major temperature drop and increased seasonality, and a faunal turnover which extinguished 60–70% of all Eurasian faunal genera, and 70–80% of North American genera. Correspondingly, a 2016 study suggested that the mean divergence dates for ''Arctotherium'', ''Arctodus'' and ''Tremarctos'' were 4.8 Ma, and between ''Arctotherium'' and ''Tremarctos'' at 4.1 Ma. A further study calculated the divergence date between ''Arctodus'' and ''Tremarctos'' at 5.5 Ma. All three genera are first recorded from the Blancan (Pliocene/Pleistocene boundary) of North America. An indeterminate ''

'' is first recorded from the late Pliocene of

El Salvador El Salvador, officially the Republic of El Salvador, is a country in Central America. It is bordered on the northeast by Honduras, on the northwest by Guatemala, and on the south by the Pacific Ocean. El Salvador's capital and largest city is S ...

. 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 ''

'' gave way to new forms; ''Arctodus pristinus'' gave way to the huge ''Arctodus simus'', which inhabited a pan-continental range, from

Alaska Alaska ( ) is a non-contiguous U.S. state on the northwest extremity of North America. Part of the Western United States region, it is one of the two non-contiguous U.S. states, alongside Hawaii. Alaska is also considered to be the north ...

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. ''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''. Although the smaller ''Arctotherium wingei'' is only known from Late Pleistocene records, the species' more archaic position in the ''Arctotherium'' family tree also suggests an origin in the Middle Pleistocene. ''Arctotherium wingei'' was the only known species of ''Arctotherium'' to principally inhabit a range north of the Southern Cone, and to reinvade

Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...

. 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. Around this time, '' Tremarctos ornatus'', otherwise known as the spectacled bear, starts appearing in the South American fossil record. 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.

Anatomy

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 a deeper but not a shorter face than most living bears. Osteological differences between tremarctine and ursine bears include an extra lateral cusp between the trigonid and talonid on the m1 molar, a premasseteric fossa on the mandible, and often an entepicondylar foramen on the humerus of tremarctine bears. Additionally, tremarctine bears' skulls are deeper and more brachycephalic, their zygomatic arches and glenoid fossas are well developed, and they have larger molars in comparison with ursines. Moreover, tremarctine bears'

orbits In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...

are also bigger, more rounded and lateralized. The ichnotaxon ''Ursichnus sudamericanus'' is referred to ''Arctotherium tarijense''; unlike the footprints of ursine bears, the heel impression is absent in the manus, and the digit III imprint is longer than the remaining digits. 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.

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. However, it has been posited that the

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58procyonid '' Chapalmalania'' in South America, and both '' Borophagus'' and '' Agriotherium'' in North America, was a shared impetus for gigantism in '' Arctodus'' and ''

''. '' Arctodus'' and '' Tremarctos'' share characteristics common to herbivorous bears. This includes cheek teeth with large surface areas, a deep mandible, and large mandibular muscle attachments. Because herbivorous carnivorans lack an efficient digestive tract for breaking down plant matter via microbial action, they must break down plant matter via extensive chewing or grinding, and thus possess features to create a high mechanical advantage of the jaw. This presents the possibility that these traits may be an ancestral condition of the group, if not an indication of their preferred dietary habits.

Systematics

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 ''

''. 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 Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...

of large size in the two lineages. Tremarctines are occasionally referred to as ''arctodonts'' or ''arctotheres'' in older scientific literature, although this has fallen out of fashion.

Taxonomy

The following taxonomy of the tremarctine bears follow by Mitchell et al. (2016): * Subfamily Tremarctinae Merriam & Stock, 1925 ** †'' Plionarctos'' Frick, 1926 *** †''Plionarctos harroldorum'' Tedfored & Martin, 2001 *** †''Plionarctos edensis'' Frick, 1926 ** †'' Arctodus'' Leidy, 1854 *** †''Arctodus simus'' Cope, 1879 *** †''Arctodus pristinus'' Leidy, 1854 ** †''

'' Burmeister, 1879 *** †''Arctotherium angustidens'' Gervais & Ameghino, 1880 *** †''Arctotherium vetustum'' Ameghino, 188) *** †''Arctotherium wingei'' Ameghino, 1902 *** †''Arctotherium bonariense'' (Gervais, 1852) *** †''Arctotherium tarijense'' Ameghino, 1902 ** '' Tremarctos'' Gervais, 1855 *** †'' Tremarctos floridanus'' (Gildey, 1928) *** '' Tremarctos ornatus'' (Cuvier, 1825) – spectacled bear

References

External links

Yukon Beringia Interpretive Centre - North American Short-Faced Bear
{{Taxonbar, from=Q337197 Bears Mammal subfamilies Fossil taxa described in 1925