Arctodus

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''Arctodus'' belongs to the subfamily Tremarctinae, which appeared in North America during the earliest parts of the late Miocene epoch in the form of ''[[Plionarctos]]''. The medium-sized ''Arctodus pristinus,'' ''[[Tremarctos floridanus]]'' and ''Arctotherium sp.'' evolved from ''Plionarctos'' in the [[Blancan|Blancan age]] of North America.<ref name="Emslie-1995">{{cite journal |last1=Emslie |first1=Steven D. |year=1995 |title=The fossil record of Arctodus pristinus (Ursidae: Tremarctinae) in Florida |journal=Bulletin of the Florida Museum of Natural History |volume=37 |issue=15 |pages=501–514 |doi=10.58782/flmnh.hduf9651 |url=https://www.floridamuseum.ufl.edu/wp-content/uploads/sites/35/2017/03/Vol-37-No-15.pdf |s2cid=168164209 }}</ref><ref>{{Cite journal |last1=Soibelzon |first1=Leopoldo H. |last2=Romero |first2=M.R. Aguilar |date=2008-10-14 |title=A Blancan (Pliocene) short-faced bear from El Salvador and its implications for Tremarctines in South America |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |volume=250 |issue=1 |pages=1–8 |doi=10.1127/0077-7749/2008/0250-0001 |url=http://sedici.unlp.edu.ar/handle/10915/5361 }}</ref><ref name="Schubert-2010a">{{Cite journal |last1=Schubert |first1=Blaine |last2=Hulbert |first2=Richard |last3=MacFadden |first3=Bruce |last4=Searle |first4=Michael |last5=Searle |first5=Seina |date=2010-01-01 |title=Giant Short-faced Bears (Arctodus simus) in Pleistocene Florida USA, a Substantial Range Extension |url=https://www.researchgate.net/publication/250071137 |journal=Journal of Paleontology |volume=84 |issue=1 |pages=79–87 |doi=10.1666/09-113.1 |bibcode=2010JPal...84...79S |s2cid=131532424}}</ref> The genetic divergence date for ''Arctodus'' is ~5.5 million years ago,<ref name="Pedersen 2728–2736.e8" /> around the [[Late Miocene|Miocene]]-[[Zanclean|Pliocene]] boundary (~5 Mya), when tremarctine bears, along with other ursids, experienced an explosive radiation in diversity, as [[C4 carbon fixation|C<sub>4</sub> vegetation]] ([[Poaceae|grasses]]) and open habitats dominated. The world experienced a major temperature drop and increased seasonality, and a faunal turnover which extinguished 70–80% of North American genera.<ref>{{Cite journal |last1=Krause |first1=Johannes |last2=Unger |first2=Tina |last3=Noçon |first3=Aline |last4=Malaspinas |first4=Anna-Sapfo |last5=Kolokotronis |first5=Sergios-Orestis |last6=Stiller |first6=Mathias |last7=Soibelzon |first7=Leopoldo |last8=Spriggs |first8=Helen |last9=Dear |first9=Paul H |last10=Briggs |first10=Adrian W |last11=Bray |first11=Sarah CE |last12=O'Brien |first12=Stephen J |last13=Rabeder |first13=Gernot |last14=Matheus |first14=Paul |last15=Cooper |first15=Alan |date=December 2008 |title=Mitochondrial genomes reveal an explosive radiation of extinct and extant bears near the Miocene-Pliocene boundary |journal=BMC Evolutionary Biology |language=en |volume=8 |issue=1 |page=220 |doi=10.1186/1471-2148-8-220 |issn=1471-2148 |pmc=2518930 |pmid=18662376 |bibcode=2008BMCEE...8..220K |doi-access=free }}</ref><ref name="Donohue2" />	''Arctodus'' belongs to the subfamily Tremarctinae, which appeared in North America during the earliest parts of the late Miocene epoch in the form of ''[[Plionarctos]]''. The medium-sized ''Arctodus pristinus,'' ''[[Tremarctos floridanus]]'' and ''Arctotherium sp.'' evolved from ''Plionarctos'' in the [[Blancan|Blancan age]] of North America.<ref name="Emslie-1995">{{cite journal |last1=Emslie |first1=Steven D. |year=1995 |title=The fossil record of Arctodus pristinus (Ursidae: Tremarctinae) in Florida |journal=Bulletin of the Florida Museum of Natural History |volume=37 |issue=15 |pages=501–514 |doi=10.58782/flmnh.hduf9651 |url=https://www.floridamuseum.ufl.edu/wp-content/uploads/sites/35/2017/03/Vol-37-No-15.pdf |s2cid=168164209 }}</ref><ref>{{Cite journal |last1=Soibelzon |first1=Leopoldo H. |last2=Romero |first2=M.R. Aguilar |date=2008-10-14 |title=A Blancan (Pliocene) short-faced bear from El Salvador and its implications for Tremarctines in South America |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |volume=250 |issue=1 |pages=1–8 |doi=10.1127/0077-7749/2008/0250-0001 |url=http://sedici.unlp.edu.ar/handle/10915/5361 }}</ref><ref name="Schubert-2010a">{{Cite journal |last1=Schubert |first1=Blaine |last2=Hulbert |first2=Richard |last3=MacFadden |first3=Bruce |last4=Searle |first4=Michael |last5=Searle |first5=Seina |date=2010-01-01 |title=Giant Short-faced Bears (Arctodus simus) in Pleistocene Florida USA, a Substantial Range Extension |url=https://www.researchgate.net/publication/250071137 |journal=Journal of Paleontology |volume=84 |issue=1 |pages=79–87 |doi=10.1666/09-113.1 |bibcode=2010JPal...84...79S |s2cid=131532424}}</ref> The genetic divergence date for ''Arctodus'' is ~5.5 million years ago,<ref name="Pedersen 2728–2736.e8" /> around the [[Late Miocene|Miocene]]-[[Zanclean|Pliocene]] boundary (~5 Mya), when tremarctine bears, along with other ursids, experienced an explosive radiation in diversity, as [[C4 carbon fixation|C<sub>4</sub> vegetation]] ([[Poaceae|grasses]]) and open habitats dominated. The world experienced a major temperature drop and increased seasonality, and a faunal turnover which extinguished 70–80% of North American genera.<ref>{{Cite journal |last1=Krause |first1=Johannes |last2=Unger |first2=Tina |last3=Noçon |first3=Aline |last4=Malaspinas |first4=Anna-Sapfo |last5=Kolokotronis |first5=Sergios-Orestis |last6=Stiller |first6=Mathias |last7=Soibelzon |first7=Leopoldo |last8=Spriggs |first8=Helen |last9=Dear |first9=Paul H |last10=Briggs |first10=Adrian W |last11=Bray |first11=Sarah CE |last12=O'Brien |first12=Stephen J |last13=Rabeder |first13=Gernot |last14=Matheus |first14=Paul |last15=Cooper |first15=Alan |date=December 2008 |title=Mitochondrial genomes reveal an explosive radiation of extinct and extant bears near the Miocene-Pliocene boundary |journal=BMC Evolutionary Biology |language=en |volume=8 |issue=1 |page=220 |doi=10.1186/1471-2148-8-220 |issn=1471-2148 |pmc=2518930 |pmid=18662376 |bibcode=2008BMCEE...8..220K |doi-access=free }}</ref><ref name="Donohue2" />
''Arctodus pristinus'' was mostly restricted to the more densely forested thermal enclave in [[Southeastern United States#Climate|eastern North America]].<ref name="Bearalmanac">{{Cite book |last=Brown, Gary |url=https://archive.org/details/greatbearalmanac00gary/page/340 |title=Great Bear Almanac |year=1996 |isbn=978-1-55821-474-3 |page=[https://archive.org/details/greatbearalmanac00gary/page/340 340] |publisher=Lyons & Burford |url-access=registration}}</ref><ref name="Russell-2009">{{Cite journal |last1=Russell |first1=Dale A. |last2=Rich |first2=Fredrick J. |last3=Schneider |first3=Vincent |last4=Lynch-Stieglitz |first4=Jean |date=May 2009 |title=A warm thermal enclave in the Late Pleistocene of the South-eastern United States |journal=Biological Reviews |volume=84 |issue=2 |pages=173–202 |doi=10.1111/j.1469-185X.2008.00069.x |pmid=19391200 |s2cid=9609391 }}</ref> ''A. pristinus'' has the greatest concentration of fossils in Florida,<ref name="Emslie-1995" /> with the earliest finds being from the Late Blancan [[Kissimmee River]] [[Okeechobee County, Florida|6]] (2.7 - 2.2 Mya) and [[Santa Fe River (Florida)|Santa Fe River]] [[Columbia County, Florida|1]] sites.<ref name="Florida Museum-2017" /><ref name="Emslie-1995" /><ref>{{Cite journal |last=Morgan |first=Gary S. |date=2005 |title=The Great American Biotic Interchange in Florida |url=https://www.floridamuseum.ufl.edu/wp-content/uploads/sites/35/2017/03/bulletin-Morganlowres.pdf |journal=Bulletin of the Florida Museum of National History |volume=45 |issue=4 |pages=271–311|doi=10.58782/flmnh.pkqn7297 }}</ref> During the early [[Irvingtonian|Irvingtonian age]] (~1.36 million years ago), a western population of ''A. pristinus'' evolved into the enormous ''A. simus,'' being first recorded from the [[Fremont, California#Irvington District|Irvington type locality]] in California.<ref name="Richards-1996" /><ref name="Bell-2004">{{Cite book |last1=Bell |first1=Christopher |url=https://www.researchgate.net/publication/263425514 |title=Late Cretaceous and Cenozoic Mammals of North America |last2=Lundelius |first2=Ernest L. |last3=Barnosky |first3=Anthony D. |last4=Zarzewski |first4=Richard J. |last5=Graham |first5=Russell |last6=Lindsay |first6=Everett H. |last7=Ruez |first7=Dennis R. |last8=Semken |first8=Holmes A. |last9=Webb |first9=S. David |editor-first1=Michael O. |editor-last1=Woodburne |date=2004-04-21 |publisher=Columbia University Press |isbn=978-0-231-50378-5 |language=en |chapter=Chapter 7: The Blancan, Irvingtonian, and Rancholabrean Mammal Ages |doi=10.7312/wood13040}}</ref> Correspondingly, ''A. simus'' is most plentiful from western North America,<ref name="Emslie-1985" /><ref name="Martin-1978">{{Cite journal |last1=Martin |first1=Larry |last2=Neuner |first2=A. |date=1978-01-01 |title=The End of the Pleistocene in North America |url=https://digitalcommons.unl.edu/tnas/337 |journal=Transactions of the Nebraska Academy of Sciences and Affiliated Societies}}</ref> albeit preferring mixed habitat such as temperate open woodlands.<ref name="Trayler-2015">{{Cite journal |last1=Trayler |first1=Robin B. |last2=Dundas |first2=Robert G. |last3=Fox-Dobbs |first3=Kena |last4=Van De Water |first4=Peter K. |date=2015-11-01 |title=Inland California during the Pleistocene—Megafaunal stable isotope records reveal new paleoecological and paleoenvironmental insights |url=https://www.sciencedirect.com/science/article/pii/S0031018215004010 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=437 |pages=132–140 |bibcode=2015PPP...437..132T |doi=10.1016/j.palaeo.2015.07.034 |issn=0031-0182}}</ref><ref name="Pérez-Crespo-2018">{{Cite journal |last1=Pérez-Crespo |first1=Víctor Adrián |last2=Arroyo-Cabrales |first2=Joaquín |last3=Morales-Puente |first3=Pedro |last4=Cienfuegos-Alvarado |first4=Edith |last5=Otero |first5=Francisco J. |date=March 2018 |title=Diet and habitat of mesomammals and megamammals from Cedral, San Luis Potosí, México |journal=Geological Magazine |volume=155 |issue=3 |pages=674–684 |bibcode=2018GeoM..155..674P |doi=10.1017/S0016756816000935 |s2cid=132502543}}</ref><ref name="Harris-1985" /><ref name="Esker-2010" /><ref>{{Citation |last=Akersten |first=William A. |title=Diversity bottlenecks, oddball survivors, and negative keys |date=1996-12-31 |work=Palaeoecology and Palaeoenvironments of Late Cenozoic Mammals |pages=1–15 |url=https://www.degruyter.com/document/doi/10.3138/9781487574154-004/html |access-date=2024-01-23 |publisher=University of Toronto Press |doi=10.3138/9781487574154-004 |isbn=978-1-4875-7415-4}}</ref> Their ranges may have met in the Middle Pleistocene of Kansas,<ref name="Richards-1996" /> with ''A. simus'' migrating east in the Late Pleistocene (around the extinction of ''A. pristinus'').<ref name="Schubert-2010a" /><ref>{{Cite journal |last=Phillips |first=George Edward |date=6 August 2006 |title=Paleofaunistics of Nonmammalian Vertebrates from the Late Pleistocene of the Mississippi-Alabama Black Prairie |url=https://repository.lib.ncsu.edu/items/185efe46-230f-4ff1-87be-12da682be6e8 |journal=North Carolina State University (Masters) |via=North Carolina State University Library}}</ref> Although both ''Arctodus'' species co-inhabited North America for at least a million years during the Middle Pleistocene (''A. pristinus'' went extinct ~300,000 [[Before Present|BP]]), there is no direct evidence of [[sympatry|overlap]] or [[Competition (biology)|competition]] in the fossil record as of yet, as both species established largely [[Allopatric speciation|separate ranges]].<ref name="Schubert-2010a" />	''Arctodus pristinus'' was mostly restricted to the more densely forested thermal enclave in [[Southeastern United States#Climate|eastern North America]].<ref name="Bearalmanac">{{Cite book |last=Brown, Gary |url=https://archive.org/details/greatbearalmanac00gary/page/340 |title=Great Bear Almanac |year=1996 |isbn=978-1-55821-474-3 |page=[https://archive.org/details/greatbearalmanac00gary/page/340 340] |publisher=Lyons & Burford |url-access=registration}}</ref><ref name="Russell-2009">{{Cite journal |last1=Russell |first1=Dale A. |last2=Rich |first2=Fredrick J. |last3=Schneider |first3=Vincent |last4=Lynch-Stieglitz |first4=Jean |date=May 2009 |title=A warm thermal enclave in the Late Pleistocene of the South-eastern United States |journal=Biological Reviews |volume=84 |issue=2 |pages=173–202 |doi=10.1111/j.1469-185X.2008.00069.x |pmid=19391200 |s2cid=9609391 }}</ref> ''A. pristinus'' has the greatest concentration of fossils in Florida,<ref name="Emslie-1995" /> with the earliest finds being from the Late Blancan [[Kissimmee River]] [[Okeechobee County, Florida|6]] (2.7 - 2.2 Mya) and [[Santa Fe River (Florida)|Santa Fe River]] [[Columbia County, Florida|1]] sites.<ref name="Florida Museum-2017" /><ref name="Emslie-1995" /><ref>{{Cite journal |last=Morgan |first=Gary S. |date=2005 |title=The Great American Biotic Interchange in Florida |url=https://www.floridamuseum.ufl.edu/wp-content/uploads/sites/35/2017/03/bulletin-Morganlowres.pdf |journal=Bulletin of the Florida Museum of Natural History |volume=45 |issue=4 |pages=271–311|doi=10.58782/flmnh.pkqn7297 }}</ref> During the early [[Irvingtonian|Irvingtonian age]] (~1.36 million years ago), a western population of ''A. pristinus'' evolved into the enormous ''A. simus,'' being first recorded from the [[Fremont, California#Irvington District|Irvington type locality]] in California.<ref name="Richards-1996" /><ref name="Bell-2004">{{Cite book |last1=Bell |first1=Christopher |url=https://www.researchgate.net/publication/263425514 |title=Late Cretaceous and Cenozoic Mammals of North America |last2=Lundelius |first2=Ernest L. |last3=Barnosky |first3=Anthony D. |last4=Zarzewski |first4=Richard J. |last5=Graham |first5=Russell |last6=Lindsay |first6=Everett H. |last7=Ruez |first7=Dennis R. |last8=Semken |first8=Holmes A. |last9=Webb |first9=S. David |editor-first1=Michael O. |editor-last1=Woodburne |date=2004-04-21 |publisher=Columbia University Press |isbn=978-0-231-50378-5 |language=en |chapter=Chapter 7: The Blancan, Irvingtonian, and Rancholabrean Mammal Ages |doi=10.7312/wood13040}}</ref> Correspondingly, ''A. simus'' is most plentiful from western North America,<ref name="Emslie-1985" /><ref name="Martin-1978">{{Cite journal |last1=Martin |first1=Larry |last2=Neuner |first2=A. |date=1978-01-01 |title=The End of the Pleistocene in North America |url=https://digitalcommons.unl.edu/tnas/337 |journal=Transactions of the Nebraska Academy of Sciences and Affiliated Societies}}</ref> albeit preferring mixed habitat such as temperate open woodlands.<ref name="Trayler-2015">{{Cite journal |last1=Trayler |first1=Robin B. |last2=Dundas |first2=Robert G. |last3=Fox-Dobbs |first3=Kena |last4=Van De Water |first4=Peter K. |date=2015-11-01 |title=Inland California during the Pleistocene—Megafaunal stable isotope records reveal new paleoecological and paleoenvironmental insights |url=https://www.sciencedirect.com/science/article/pii/S0031018215004010 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=437 |pages=132–140 |bibcode=2015PPP...437..132T |doi=10.1016/j.palaeo.2015.07.034 |issn=0031-0182}}</ref><ref name="Pérez-Crespo-2018">{{Cite journal |last1=Pérez-Crespo |first1=Víctor Adrián |last2=Arroyo-Cabrales |first2=Joaquín |last3=Morales-Puente |first3=Pedro |last4=Cienfuegos-Alvarado |first4=Edith |last5=Otero |first5=Francisco J. |date=March 2018 |title=Diet and habitat of mesomammals and megamammals from Cedral, San Luis Potosí, México |journal=Geological Magazine |volume=155 |issue=3 |pages=674–684 |bibcode=2018GeoM..155..674P |doi=10.1017/S0016756816000935 |s2cid=132502543}}</ref><ref name="Harris-1985" /><ref name="Esker-2010" /><ref>{{Citation |last=Akersten |first=William A. |title=Diversity bottlenecks, oddball survivors, and negative keys |date=1996-12-31 |work=Palaeoecology and Palaeoenvironments of Late Cenozoic Mammals |pages=1–15 |url=https://www.degruyter.com/document/doi/10.3138/9781487574154-004/html |access-date=2024-01-23 |publisher=University of Toronto Press |doi=10.3138/9781487574154-004 |isbn=978-1-4875-7415-4}}</ref> Their ranges may have met in the Middle Pleistocene of Kansas,<ref name="Richards-1996" /> with ''A. simus'' migrating east in the Late Pleistocene (around the extinction of ''A. pristinus'').<ref name="Schubert-2010a" /><ref>{{Cite journal |last=Phillips |first=George Edward |date=6 August 2006 |title=Paleofaunistics of Nonmammalian Vertebrates from the Late Pleistocene of the Mississippi-Alabama Black Prairie |url=https://repository.lib.ncsu.edu/items/185efe46-230f-4ff1-87be-12da682be6e8 |journal=North Carolina State University (Masters) |via=North Carolina State University Library}}</ref> Although both ''Arctodus'' species co-inhabited North America for at least a million years during the Middle Pleistocene (''A. pristinus'' went extinct ~300,000 [[Before Present|BP]]), there is no direct evidence of [[sympatry|overlap]] or [[Competition (biology)|competition]] in the fossil record as of yet, as both species established largely [[Allopatric speciation|separate ranges]].<ref name="Schubert-2010a" />
Irvingtonian age (1,900,000 BP - 250,000 BP) specimens of ''Arctodus simus'' are particularly sparse. Finds are mostly from California, with additional remains from Texas, Kansas, Nebraska, and Montana.<ref name="Scott-1993" /><ref name="Hill-2000" /><ref name="Kurtén-1967">{{Cite book |last=Kurtén |first=Björn |url=https://books.google.com/books?id=GkCFtAEACAAJ |title=Pleistocene Bears of North America: Genus Arctodus, short-faced bears |date=1967 |publisher=Societas pro Fauna et Flora Fennica |language=en}}</ref> However, ''A. simus'' became a pan-continental species in the [[Rancholabrean]] (Late Pleistocene), sharing that distinction with the [[American black bear|black bear]].<ref name="Bell-2004" /><ref name="Esker-2010">{{Cite web |last1=Esker |first1=Donald |last2=Wilkins |first2=William |last3=Agenbroad |first3=Larry |date=2010-08-13 |title=Esker, Wilkins, and Agenbroad—Multivariate Analysis Of Ursids: A multivariate analysis of the ecology of North American Pleistocene bears, with a focus on ''Arctodus simus'' |url=https://www.researchgate.net/publication/314037201 |website=ResearchGate}}</ref> Despite ''Arctodus simus''{{'}} large temporal and geographic range, fossil remains are comparatively rare (109 finds as of 2010, in otherwise well-sampled localities).<ref name="Schubert-2010a" /><ref name="Pedersen 2728–2736.e8">{{Cite journal|last1=Pedersen|first1=Mikkel Winther|last2=De Sanctis|first2=Bianca|last3=Saremi|first3=Nedda F.|last4=Sikora|first4=Martin|last5=Puckett|first5=Emily E.|last6=Gu|first6=Zhenquan|last7=Moon|first7=Katherine L.|last8=Kapp|first8=Joshua D.|last9=Vinner|first9=Lasse|last10=Vardanyan|first10=Zaruhi|last11=Ardelean|first11=Ciprian F.|date=2021-06-21|title=Environmental genomics of Late Pleistocene black bears and giant short-faced bears |journal=Current Biology |volume=31|issue=12|pages=2728–2736.e8|doi=10.1016/j.cub.2021.04.027|pmid=33878301|bibcode=2021CBio...31E2728P |s2cid=233303447 |hdl=10037/22808|hdl-access=free}}</ref>	Irvingtonian age (1,900,000 BP - 250,000 BP) specimens of ''Arctodus simus'' are particularly sparse. Finds are mostly from California, with additional remains from Texas, Kansas, Nebraska, and Montana.<ref name="Scott-1993" /><ref name="Hill-2000" /><ref name="Kurtén-1967">{{Cite book |last=Kurtén |first=Björn |url=https://books.google.com/books?id=GkCFtAEACAAJ |title=Pleistocene Bears of North America: Genus Arctodus, short-faced bears |date=1967 |publisher=Societas pro Fauna et Flora Fennica |language=en}}</ref> However, ''A. simus'' became a pan-continental species in the [[Rancholabrean]] (Late Pleistocene), sharing that distinction with the [[American black bear|black bear]].<ref name="Bell-2004" /><ref name="Esker-2010">{{Cite web |last1=Esker |first1=Donald |last2=Wilkins |first2=William |last3=Agenbroad |first3=Larry |date=2010-08-13 |title=Esker, Wilkins, and Agenbroad—Multivariate Analysis Of Ursids: A multivariate analysis of the ecology of North American Pleistocene bears, with a focus on ''Arctodus simus'' |url=https://www.researchgate.net/publication/314037201 |website=ResearchGate}}</ref> Despite ''Arctodus simus''{{'}} large temporal and geographic range, fossil remains are comparatively rare (109 finds as of 2010, in otherwise well-sampled localities).<ref name="Schubert-2010a" /><ref name="Pedersen 2728–2736.e8">{{Cite journal|last1=Pedersen|first1=Mikkel Winther|last2=De Sanctis|first2=Bianca|last3=Saremi|first3=Nedda F.|last4=Sikora|first4=Martin|last5=Puckett|first5=Emily E.|last6=Gu|first6=Zhenquan|last7=Moon|first7=Katherine L.|last8=Kapp|first8=Joshua D.|last9=Vinner|first9=Lasse|last10=Vardanyan|first10=Zaruhi|last11=Ardelean|first11=Ciprian F.|date=2021-06-21|title=Environmental genomics of Late Pleistocene black bears and giant short-faced bears |journal=Current Biology |volume=31|issue=12|pages=2728–2736.e8|doi=10.1016/j.cub.2021.04.027|pmid=33878301|bibcode=2021CBio...31E2728P |s2cid=233303447 |hdl=10037/22808|hdl-access=free}}</ref>

Okumaya devam et...