Editing Cave bear

{{Short description|Extinct species of carnivore}}
{{Speciesbox
| name = Cave bear
| fossil_range = Middle to Late [[Pleistocene]], {{fossil range|0.25|0.024}}
| image = Teufelshöhle-Höhlenbär-Dreiviertelprofil.jpg
| image_upright = 1.2
| image_caption = Mounted cave bear skeleton at [[Devil's Cave (near Pottenstein)|Devil's Cave]], Germany
| genus = Ursus
| species = spelaeus
| authority = [[Johann Christian Rosenmüller|Rosenmüller]], 1794
| extinct = yes
}}

The '''cave bear''' ('''''Ursus spelaeus''''') is a prehistoric [[species]] of [[bear]] that lived in [[Europe]] and [[Asia]] during the [[Pleistocene]] and became [[extinct]] about 24,000 years ago during the [[Last Glacial Maximum]].

Both the word ''cave'' and the scientific name ''spelaeus'' are used because fossils of this species were mostly found in [[cave]]s. This reflects the views of experts that cave bears spent more time in caves than the [[brown bear]], frequently using them to hibernate during the winter months. Unlike brown bears, cave bears are thought to have been almost entirely or exclusively herbivorous.

Cave bears exhibit a great degree of size, morphological and genetic variability, and Late Pleistocene cave bears are often (though not universally) considered to be [[species complex]] of up to 6 different species.<ref>{{Cite journal |last=Knapp |first=Michael |date=2019-04-21 |title=From a molecules' perspective – contributions of ancient DNA research to understanding cave bear biology |url=https://www.tandfonline.com/doi/full/10.1080/08912963.2018.1434168 |journal=Historical Biology |language=en |volume=31 |issue=4 |pages=442–447 |doi=10.1080/08912963.2018.1434168 |issn=0891-2963}}</ref><ref name=":1" />

==Taxonomy==
[[File:Cave bear AMNH.jpg|thumb|upright|Rearing ''Ursus spelaeus'' skeleton at the [[AMNH]]]]
Cave bear skeletons were first described in 1774 by Johann Friedrich Esper, in his book ''Newly Discovered Zoolites of Unknown Four Footed Animals''. While scientists at the time considered that the skeletons could belong to [[ape]]s, [[canid]]s, [[felid]]s, or even [[dragon]]s or [[unicorn]]s, Esper postulated that they actually belonged to [[polar bear]]s. Twenty years later, [[Johann Christian Rosenmüller]], an [[anatomist]] at [[Leipzig University]], gave the species its binomial name. The bones were so numerous that most researchers had little regard for them. During [[World War I]], with the scarcity of [[phosphate]] dung, earth from the caves where cave bear bones occurred was used as a source of phosphates. When the "dragon caves" in [[Austria]]’s [[Duchy of Styria|Styria]] region were exploited for this purpose, only the skulls and leg bones were kept.<ref name="Brunner2007">{{cite book|author=Bernd Brunner|title=Bears: A Brief History|url=https://books.google.com/books?id=1W98s7aVy8cC&pg=PA41|year=2007|publisher=Yale University Press|isbn=978-0-300-12299-2|page=41}}</ref>

Many caves in [[Central Europe]] and [[North America]]  have skeletons of cave bears inside, such as the ''Heinrichshöhle'' in [[Hemer, Germany|Hemer]] and the ''[[Dechen Cave|Dechenhöhle]]'' in [[Iserlohn]], [[Germany]]. A complete skeleton, five complete skulls, and 18&nbsp;other bones were found inside [[Kletno Bear Cave]], in 1966 in [[Poland]].<ref>{{cite book |author=Praca Zbiorowa |title=Jaskinia Niedźwiedzia w Kletnie. Badanie i udostępnianie |publisher=Polska Akademia Nauk, Ossolineum |location=Wrocław |year=1989 |isbn=8304030373 |language=pl}} with summary {{in lang|en}}</ref> In [[Romania]], in a cave called [[Bears' Cave]], 140&nbsp;cave bear skeletons were discovered in 1983.<ref>[https://web.archive.org/web/20070202235231/http://www.personal.psu.edu/users/w/x/wxk116/cavebears/ Cave Bears]. Jan Kowalski. psu.edu</ref>

Remains assigned to "cave bears" ''sensu lato'' from the Late Pleistocene exhibit a strong degree of morphological and size variability, and have often been assigned to their own species, including ''[[Ursus rossicus]]'' (Eastern Europe, Central Asia and Siberia)'', [[Ursus ingressus]]'' (Central Europe to the Urals), ''[[Ursus kanivetz]],'' (Urals) ''[[Ursus kudarensis]]'' (the Caucasus), ''[[Ursus eremus]]'' (Central Europe, possibly a subspecies of ''U. spelaeus'' s.s.) and ''Ursus spelaeus'' sensu stricto (Western, Central and Southeast Europe).<ref name=":0" /><ref name=":1">{{Cite journal |last=Barlow |first=Axel |last2=Paijmans |first2=Johanna L.A. |last3=Alberti |first3=Federica |last4=Gasparyan |first4=Boris |last5=Bar-Oz |first5=Guy |last6=Pinhasi |first6=Ron |last7=Foronova |first7=Irina |last8=Puzachenko |first8=Andrey Y. |last9=Pacher |first9=Martina |last10=Dalén |first10=Love |last11=Baryshnikov |first11=Gennady |last12=Hofreiter |first12=Michael |date=2021-04-26 |title=Middle Pleistocene genome calibrates a revised evolutionary history of extinct cave bears |url=https://linkinghub.elsevier.com/retrieve/pii/S096098222100138X |journal=Current Biology |language=en |volume=31 |issue=8 |pages=1771–1779.e7 |doi=10.1016/j.cub.2021.01.073}}</ref> These populations/species show considerable genetic divergence from each other, (with genetic divergences estimated at hundreds of thousands to a million years)<ref name=":1" /> though whether these species should be regarded as synonyms of ''U. speleus'' is debated.<ref name=":0">{{cite journal |last1=Galdies |first1=Johann |title=The bears of the European steppe: a review |journal=Quaternaire |date=2022-03-01 |volume=33 |issue=1 |pages=47–62 |doi=10.4000/quaternaire.16605 |url=https://journals.openedition.org/quaternaire/16605 |issn=1142-2904 |access-date=2025-02-07}}</ref>

===Evolution===
Both the cave bear and the brown bear are thought to be descended from the [[Plio-Pleistocene]] Etruscan bear (''[[Ursus etruscus]]'')<ref name=Kurten1976>{{cite book |author=Kurtén, B. |year=1976 |title=The Cave Bear Story: Life and death of a vanished animal |publisher=Columbia University Press |location=New York, NY}}</ref><ref name=Rabader2000>{{cite book |vauthors=Rabeder G, Nagel D, Pacher M |year=2000 |title=Der Höhlenbär. Species 4 |publisher=Thorbecke Verlag |location=Stuttgart, DE}}</ref><ref>{{cite book |vauthors=Argant A, Crégut-Bonnoure E |year=1996 |contribution=Famille des Ursidae |editor1=Guérin, C. |editor2=Patou-Mathis, M. |title=Les grands mammiferes Plio-Pleistocenes d'Europe |pages=167–177 |publisher=Masson |location=Paris, FR}}</ref> that lived about 5.3&nbsp;[[Mya (unit)|Mya]] to 100,000&nbsp;years ago. The last common ancestor of cave bears and brown bears lived between 1.2&ndash;1.4&nbsp;Mya.<ref name="Loreille2001">{{Cite journal |author=Loreille, O. |display-authors=etal |year=2001 |title=Ancient DNA analysis reveals divergence of the cave bear, ''Ursus spelaeus'', and brown bear, ''Ursus arctos'', lineages |journal=[[Current Biology]] |volume=11 |issue=3 |pages=200–203 |bibcode=2001CBio...11..200L |doi=10.1016/S0960-9822(01)00046-X |pmid=11231157 |doi-access=free}}</ref> Middle Pleistocene remains of the cave bear lineage are assigned to ''[[Ursus deningeri]]'' (Deninger's bear),<ref name=":0" /> which is known from remains spanning from Europe to China.<ref>{{Cite journal |last1=Jiangzuo |first1=Qigao |last2=Wagner |first2=Jan |last3=Chen |first3=Jin |last4=Dong |first4=Cuiping |last5=Wei |first5=Jianhua |last6=Ning |first6=Juan |last7=Liu |first7=Jinyi |date=2018-11-01 |title=Presence of the Middle Pleistocene cave bears in China confirmed – Evidence from Zhoukoudian area |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379118303354 |journal=Quaternary Science Reviews |language=en |volume=199 |pages=1–17 |doi=10.1016/j.quascirev.2018.09.012}}</ref> The transition between Deninger's bear and the cave bear is given as the last interglacial, although the boundary between these forms is arbitrary, and intermediate or transitional [[taxa]] have been proposed, e.g. ''Ursus spelaeus deningeroides'',<ref name=Argant1991>{{Cite journal|author=Argant, A. |title=Carnivores quaternaires de Bourgogne |year=1991 |journal=Documents des Laboratoires de Géologie de la Faculté des Sciences de Lyon |volume=115 |pages=1–301}}</ref> while other authorities consider both taxa to be chronological variants of the same species.<ref name=MazzaRustioni1994>{{Cite journal |author1=Mazza, P. |author2=Rustioni, M. |title=On the phylogeny of Eurasian bears |year=1994 |journal=Palaeontographica Abteilung A |volume=230 |issue=1–3 |pages=1–32|doi=10.1127/pala/230/1994/1 |bibcode=1994PalAA.230....1M |s2cid=247508689 }}</ref>

Cave bears found anywhere will vary in age, thus facilitating investigations into evolutionary trends. The three anterior [[premolar]]s were gradually reduced, then disappeared, possibly in response to a largely vegetarian diet. In a fourth of the skulls found in the ''Conturines'', the third premolar is still present, while more [[Synapomorphy|derived]] specimens elsewhere lack it. The last remaining premolar became conjugated with the true [[Molar (tooth)|molars]], enlarging the crown and granting it more cusps and cutting borders. This phenomenon, called ''molarization'', improved the [[mastication]] capacities of the molars, facilitating the processing of tough vegetation. This allowed the cave bear to gain more energy for hibernation, while eating less than its ancestors.<ref>{{cite web |url=http://www.altabadia.it/orsispelei/evoluzione_i.htm |title=Gli orsi spelèi delle Conturines / Ursus Spelaeus |website=Altabadia.it |access-date=2011-09-26 |df=dmy-all}}</ref>

In 2005, scientists recovered and [[DNA sequencing|sequenced]] the [[nuclear DNA]] of a cave bear that lived between 42,000 and 44,000 years ago. The procedure used genomic DNA extracted from one of the animal's [[teeth]]. Sequencing the DNA directly (rather than first replicating it with the [[polymerase chain reaction]]), the scientists recovered 21 cave bear [[gene]]s from remains that did not yield significant amounts of DNA with traditional techniques.<ref name="Noonan2005">{{Cite journal |author=Noonan, James P. |display-authors=etal |year=2005 |title=Genomic Sequencing of Pleistocene Cave Bears |url=https://digital.library.unt.edu/ark:/67531/metadc786565/ |journal=[[Science (journal)|Science]] |volume=309 |issue=5734 |pages=597–599 |bibcode=2005Sci...309..597N |doi=10.1126/science.1113485 |pmid=15933159 |s2cid=34704597 |doi-access=free}}</ref> This study confirmed and built on results from a previous study using [[mitochondrial DNA]] extracted from cave bear remains ranging from 20,000 to 130,000 years old.<ref name=Loreille2001/> Both show that the cave bear was more closely related to the [[brown bear]] and [[polar bear]] than it was to the [[American black bear]], but had split from the brown bear lineage before the distinct eastern and western brown bear lineages diversified, and before the split of brown bears and polar bears. The [[Genetic divergence|divergence]] date estimate of cave bears and brown bears is about 1.2–1.4&nbsp;Mya.<ref name=Loreille2001/> However, a recent study showed that both species had some hybridization between them.<ref>{{Cite journal|last1=Barlow|first1=Axel|last2=Cahill|first2=James A.|last3=Hartmann|first3=Stefanie|last4=Theunert|first4=Christoph|last5=Xenikoudakis|first5=Georgios|last6=Fortes|first6=Gloria G.|last7=Paijmans|first7=Johanna L. A.|last8=Rabeder|first8=Gernot|last9=Frischauf|first9=Christine|date=27 August 2018|title=Partial genomic survival of cave bears in living brown bears|journal=[[Nature Ecology & Evolution]]|language=En|volume=2|issue=10|pages=1563–1570|doi=10.1038/s41559-018-0654-8|issn=2397-334X|url=https://escholarship.org/content/qt6qr394rt/qt6qr394rt.pdf?t=poma92|pmid=30150744|pmc=6590514|bibcode=2018NatEE...2.1563B }}</ref>

==Description==
[[File:Ursus spelaeus Sergiodlarosa.jpg|thumb|Life restoration.]]
The cave bear had a very broad, domed skull with a steep forehead; its stout body had long thighs, massive shins and in-turning feet, making it similar in skeletal structure to the brown bear.<ref name="Bearalmanac">{{cite book | author = Brown, Gary | title = Great Bear Almanac | year = 1996 | page = [https://archive.org/details/greatbearalmanac00gary/page/340 340] | publisher = Lyons & Burford | isbn = 1-55821-474-7 | url = https://archive.org/details/greatbearalmanac00gary/page/340 }}</ref> Cave bears were comparable in size to, or larger than, the largest modern-day bears, measuring up to {{convert|2|m|ft|abbr=on}} in length.<ref>{{cite book |editor=Palmer, D.|year=1999 |title= The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|publisher= Marshall Editions|location=London|page= 217|isbn= 1-84028-152-9}}</ref> The average weight for males was {{convert|350|to|600|kg|lb|abbr=on}},<ref name="weight">{{cite journal|url=https://www.jstor.org/stable/23735739 |journal=Annales Zoologici Fennici |volume=36 |issue=2 |pages=93–102 |year=1999 |title=What size were Arctodus simus and Ursus spelaeus (Carnivora: Ursidae)? |author=Per Christiansen|jstor=23735739 }}</ref> while females weighed {{convert|225|to|250|kg|lb|abbr=on|round=5}}.<ref name="weight" /> Of cave bear skeletons in museums, 90% are classified as male due to a misconception that the female skeletons were merely "dwarfs". Cave bears grew larger during glaciations and smaller during interglacials, probably to adjust heat loss rate.<ref name="CLAW">{{cite book | author= Macdonald, David | title= The Velvet Claw | year= 1992 | page= [https://archive.org/details/velvetclawnatura00macd/page/256 256] | isbn= 0-563-20844-9 | publisher= Parkwest | location= New York | url= https://archive.org/details/velvetclawnatura00macd/page/256 }}</ref>

Cave bears of the [[Last Glacial Maximum|last Ice Age]] lacked the usual two or three premolars present in other bears; to compensate, the last molar is very elongated, with supplementary cusps.<ref>[http://www.altabadia.it/orsispelei/nozioni_i.htm Gli orsi spelèi delle Conturines/ Ursus Spelaeus]. Altabadia.it. Retrieved on 2011-09-26.</ref> The [[humerus]] of the cave bear was similar in size to that of the [[polar bear]], as were the [[femora]] of females. The femora of male cave bears, however, bore more similarities in size to those of [[Kodiak bear]]s.<ref name="weight"/>

==Behaviour==

===Dietary habits===
[[File:Ours des carvernes - Crâne.jpg|thumb|''Ursus spelaeus'' lacked the usual two or three premolars present in other bear species.]]
Cave bear teeth were very large and show greater wear than most modern bear species, suggesting a diet of tough materials. However, [[tubers]] and other gritty food, which cause distinctive tooth wear in modern brown bears, do not appear to have constituted a major part of cave bears' diets on the basis of [[dental microwear]] analysis.<ref>Pinto Llona, A. C., Andrews, P. & Etxeberrı´a, P. 2005: Taphonomy and Palaeoecology of Cave Bears from the Quaternary of Cantabrian Spain. Fondacio´n de Asturias/Du Pont Ibe´rica/The Natural History Museum, Grafinsa, Oviedo.</ref> Seed fruits are documented to have been consumed by cave bears.<ref>{{Cite journal |last=Duñó-Iglesias |first=Paulo |last2=Ramírez-Pedraza |first2=Iván |last3=Rivals |first3=Florent |last4=Mirea |first4=Ionuț-Cornel |last5=Faur |first5=Luchiana-Maria |last6=Constantin |first6=Silviu |last7=Robu |first7=Marius |date=15 February 2024 |title=Palaeodiet during the pre-dormancy period of MIS 3 Romanian cave bears as inferred from dental microwear analysis |url=https://linkinghub.elsevier.com/retrieve/pii/S0031018223006065 |journal=Palaeogeography, Palaeoclimatology, Palaeoecology |language=en |volume=636 |pages=111988 |doi=10.1016/j.palaeo.2023.111988|doi-access=free }}</ref>

The morphological features of the cave bear chewing apparatus, including loss of premolars, have long been suggested to indicate their diets displayed a higher degree of [[herbivory]] than the [[Eurasian brown bear]].<ref name=Kurten1976/> Indeed, a solely vegetarian diet has been inferred on the basis of tooth morphology.<ref name=Rabader2000/> Results obtained on the stable isotopes of cave bear bones also point to a largely vegetarian diet in having low levels of [[nitrogen-15]] and [[carbon-13]], which are accumulated at a faster rate by carnivores as opposed to herbivores.<ref name=Bocherens2006>{{Cite journal|author= Bocherens, H. |title=Bears and humans in Chauvet Cave (Vallon-Pont-d'Arc, Ardeche, France): Insights from stable isotopes and radiocarbon dating of bone collagen |year=2006 |journal=[[Journal of Human Evolution]] |volume=50 |pages=370–376|pmid= 16442587|issue= 3|doi= 10.1016/j.jhevol.2005.12.002|display-authors=etal}}</ref><ref name="Omnivore"/>

[[File:Osso e molari di Ursus spelaeus della mandibola inferiore.jpg|thumb|Detail of the molars of the lower jaw]]

However, some evidence points toward the occasional inclusion of animal protein in cave bear diets. For example, toothmarks on cave bear remains in areas where cave bears are the only recorded potential [[carnivore]]s suggests occasional cannibalistic scavenging,<ref name="Foxbear">{{cite news | url = http://www.foxnews.com/story/0,2933,320971,00.html | title = Prehistoric Cave Bears Weren't So Cuddly After All | work = FOXNews | access-date = 2008-01-11 | date = 2008-01-09 | archive-url = https://web.archive.org/web/20100101013103/http://www.foxnews.com/story/0,2933,320971,00.html | archive-date = 2010-01-01 | url-status = dead }}</ref><ref name=Pacher2000>{{Cite journal|author=Pacher, M. |title=Taphonomische Untersuchungen der Höhlenbären-Fundstellen in der Schwabenreith-Höhle bei Lunz am See (Niederösterreich) |year=2000 |journal=Beiträge zur Paläontologie |volume=25 |pages=11–85}}</ref> possibly on individuals that died during hibernation, and dental microwear analysis indicates the cave bear may have fed on a greater quantity of bone than its contemporary, the smaller Eurasian brown bear.<ref name="cavebear">{{Cite journal | url = http://www.geo.auth.gr/12icbs/proceedings/pinto.pdf| author=Pinto Llono, A.C. | series = Special |title = Comparative dental microwear analysis of cave bears Ursus spelaeus Rosenmüller, 1794 and brown bears Ursus arctos Linnaeus ,1758 |year=2006 |journal = Scientific Annals, School of Geology Aristotle University of Thessaloniki | volume= 98 |pages=103–108}}</ref> The dental microwear patterns of cave bear [[Molar (tooth)|molars]] from the northeastern [[Iberian Peninsula]] show that cave bears may have consumed more meat in the days and weeks leading up to hibernation.<ref>{{Cite journal |last1=Ramírez-Pedraza |first1=Iván |last2=Pappa |first2=Spyridoula |last3=Blasco |first3=Ruth |last4=Arilla |first4=Maite |last5=Rosell |first5=Jordi |last6=Millán |first6=Ferran |last7=Maroto |first7=Julià |last8=Soler |first8=Joaquim |last9=Soler |first9=Narcís |last10=Rivals |first10=Florent |date=20 August 2020 |title=Dietary habits of the cave bear from the Late Pleistocene in the northeast of the Iberian Peninsula |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618219307931 |journal=[[Quaternary International]] |language=en |volume=557 |pages=63–69 |doi=10.1016/j.quaint.2019.09.043 |bibcode=2020QuInt.557...63R |s2cid=204261124 |access-date=4 February 2024 |via=Elsevier Science Direct}}</ref> Additionally, cave bear remains from [[Peștera cu Oase]] in the southwestern tip of the Romanian part of the [[Carpathian Mountains]] had elevated levels of nitrogen-15 in their bones, indicative of [[omnivore|omnivorous]] diets,<ref name="Omnivore"/><ref name="Richards2008">{{Cite journal |author=Richards |first1=Michael P. |last2=Pacher |first2=Martina |last3=Stiller |first3=Mathias |last4=Quilès |first4=Jérôme |last5=Hofreiter |first5=Michael |last6=Constantin |first6=Silviu |last7=Zilhão |first7=João |last8=Trinkaus |first8=Erik |display-authors= |date=15 January 2008 |title=Isotopic evidence for omnivory among European cave bears: Late Pleistocene Ursus spelaeus from the Pestera cu Oase, Romania. |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=105 |issue=2 |pages=600–604 |bibcode=2008PNAS..105..600R |doi=10.1073/pnas.0711063105 |pmc=2206582 |pmid=18187577 |doi-access=free}}</ref><ref>{{Cite journal |last=Robu |first=Marius |last2=Wynn |first2=Jonathan G. |last3=Mirea |first3=Ionuţ C. |last4=Petculescu |first4=Alexandru |last5=Kenesz |first5=Marius |last6=Puşcaş |first6=Cristina M. |last7=Vlaicu |first7=Marius |last8=Trinkaus |first8=Erik |last9=Constantin |first9=Silviu |date=10 November 2017 |editor-last=O'Regan |editor-first=Hannah |title=The diverse dietary profiles of MIS 3 cave bears from the Romanian Carpathians: insights from stable isotope (δ 13 C and δ 15 N) analysis |url=https://onlinelibrary.wiley.com/doi/10.1111/pala.12338 |journal=[[Palaeontology (journal)|Palaeontology]] |language=en |volume=61 |issue=2 |pages=209–219 |doi=10.1111/pala.12338 |issn=0031-0239 |access-date=24 March 2024 |via=Wiley Online Library}}</ref> although the values are within the range of those found for the strictly herbivorous [[mammoth]].<ref>Bocherens, H. 2003: Isotopic biogeochemistry and the paleoecology of the mammoth steppe fauna. In Reumer, F., Braber, F., Mol, D. & de Vos, J. (eds.): Advances in Mammoth Research, 57–76. Deinsea 9.</ref> One isotopic study concluded that cave bears displayed omnivorous habits similar to those of modern brown bears.<ref>{{Cite journal |last1=Robu |first1=Marius |last2=Fortin |first2=Jennifer K. |last3=Richards |first3=Michael P. |last4=Schwartz |first4=Charles C. |last5=Wynn |first5=Jonathan G. |last6=Robbins |first6=Charles T. |last7=Trinkaus |first7=Erik |date=13 February 2013 |title=Isotopic evidence for dietary flexibility among European Late Pleistocene cave bears ( Ursus spelaeus ) |url=http://www.nrcresearchpress.com/doi/10.1139/cjz-2012-0222 |journal=[[Canadian Journal of Zoology]] |language=en |volume=91 |issue=4 |pages=227–234 |doi=10.1139/cjz-2012-0222 |issn=0008-4301 |access-date=13 January 2024}}</ref>

Although the current prevailing opinion concludes that cave bears were largely herbivorous, and more so than any modern species of the genus ''[[Ursus (genus)|Ursus]]'',<ref name="extinction"/> increasing evidence points to omnivorous diets, based both on regional variability of isotopic composition of bone remains indicative of dietary plasticity,<ref name="Omnivore"/><ref name=Richards2008/> and on a recent re-evaluation of craniodental morphology that places the cave bear squarely among omnivorous modern bear species with respect to its skull and tooth shapes.<ref name=Figueirido2009>{{Cite journal|doi=10.1111/j.1469-7998.2008.00511.x|author=Figueirido, B. |title=Ecomorphological correlates of craniodental variation in bears and paleobiological implications for extinct taxa: an approach based on geometric morphometrics |year=2009 |journal=Journal of Zoology |volume=277|issue=1 |pages=70–80|display-authors=etal}}</ref>

===Mortality===
[[File:Ursus spelaeus juvenile.jpg|thumb|upright|Standing skeleton of juvenile cave bear]]
Death during hibernation was a common end for cave bears, mainly befalling specimens that failed ecologically during the summer season through inexperience, sickness or old age.<ref name="Bjorn">{{cite book | author = Kurten, Bjorn | title = Pleistocene Mammals of Europe | year = 1968 | page = 325 | isbn = 0-202-30953-3 | publisher = AldineTransaction | location = New Brunswick, N.J.}}</ref> Some cave bear bones show signs of numerous ailments, including [[spinal fusion]], [[bone tumour]]s, [[Dental caries|cavities]], [[Root resorption|tooth resorption]], [[necrosis]] (particularly in younger specimens), [[osteomyelitis]], [[periostitis]], [[rickets]] and [[kidney stones]].<ref name="Bearalmanac"/> There is also evidence that cave bears suffered from tuberculosis.<ref>{{Cite journal |last=Marciszak |first=Adrian |last2=Mackiewicz |first2=Paweł |last3=Borówka |first3=Ryszard K. |last4=Capalbo |first4=Chiara |last5=Chibowski |first5=Piotr |last6=Gąsiorowski |first6=Michał |last7=Hercman |first7=Helena |last8=Cedro |first8=Bernard |last9=Kropczyk |first9=Aleksandra |last10=Gornig |first10=Wiktoria |last11=Moska |first11=Piotr |last12=Nowakowski |first12=Dariusz |last13=Ratajczak-Skrzatek |first13=Urszula |last14=Sobczyk |first14=Artur |last15=Sykut |first15=Maciej T. |last16=Zarzecka-Szubińska |first16=Katarzyna |last17=Kovalchuk |first17=Oleksandr |last18=Barkaszi |first18=Zoltán |last19=Stefaniak |first19=Krzysztof |last20=Mazza |first20=Paul P. A. |date=29 April 2024 |title=Fate and preservation of the Late Pleistocene cave bears from Niedźwiedzia Cave in Poland, through taphonomy, pathology, and geochemistry |url=https://www.nature.com/articles/s41598-024-60222-3 |journal=[[Scientific Reports]] |language=en |volume=14 |issue=1 |doi=10.1038/s41598-024-60222-3 |issn=2045-2322 |pmc=11059340 |pmid=38684693 |access-date=24 July 2024}}</ref> Male cave bear skeletons have been found with broken [[baculum|bacula]], probably due to fighting during the breeding season.<ref name="Bjorn"/> Cave bear [[longevity]] is unknown, though it has been estimated that they seldom exceeded twenty years of age.<ref name="bieder">{{cite book | author = Bieder, Robert | title = Bear | year = 2005 | page = 192 | isbn = 1-86189-204-7 | publisher = Reaktion Books | place = London }}</ref> Paleontologists doubt adult cave bears had any natural predators, save for pack-hunting [[Gray wolf|wolves]] and [[cave hyena]]s, which would probably have attacked sick or infirm individuals.<ref name="bieder"/> Cave hyenas are thought to be responsible for the disarticulation and destruction of some cave bear skeletons. Such large carcasses were an optimal food resource for the hyenas, especially at the end of the winter, when food was scarce.<ref name="hyena">{{cite web|url=https://nts1.cgu.cz/bulletin/contents/2006/vol81no4/237_diedrich.pdf |title=Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823)in horizontal and vertical caves of the Bohemian Karst |work=CAJUSG. DIEDRICH & KARELŽÁK |access-date=2008-01-20 }}{{dead link|date=November 2016 |bot=InternetArchiveBot |fix-attempted=yes }}</ref> The presence of fully articulated adult [[Panthera leo spelaea|cave lion]] skeletons, deep in cave bear dens, indicates the lions may have occasionally entered dens to prey on hibernating cave bears, with some dying in the attempt.<ref>[http://www.fns.uniba.sk/fileadmin/kalendar/2009/symposium/Abstract_book.pdf 15th International Cave Bear Symposium – Spišská Nová Ves, Slovakia] {{webarchive |url=https://web.archive.org/web/20100331031106/http://www.fns.uniba.sk/fileadmin/kalendar/2009/symposium/Abstract_book.pdf |date=March 31, 2010 }}. 17–20 September 2009. (PDF). Retrieved on 2011-09-26.</ref>

==Range and habitat==
The cave bear's range stretched across Europe; from [[Spain]]<ref>{{Cite journal |last=Rodríguez-Almagro |first=Manuel |last2=Gómez-Olivencia |first2=Asier |last3=Villalba de Alvarado |first3=Mónica |last4=Arsuaga |first4=Juan Luis |last5=Sala |first5=Nohemi |date=January 2024 |title=Taphonomic study of the cave bears (Ursus cf. deningeri and U. spelaeus) from the Sima I of the El Polvorín cave (Northern Iberian Peninsula) |url=https://linkinghub.elsevier.com/retrieve/pii/S2666033424000091 |journal=Quaternary Science Advances |language=en |volume=13 |pages=100171 |doi=10.1016/j.qsa.2024.100171 |access-date=3 May 2024 |via=Elsevier Science Direct|doi-access=free }}</ref> and the [[British Isles]] in the west,<ref name="extinction" /> Belgium,<ref>{{Cite journal |last=Bray |first=Fabrice |last2=Flament |first2=Stéphanie |last3=Abrams |first3=Grégory |last4=Bonjean |first4=Dominique |last5=Rolando |first5=Christian |last6=Tokarski |first6=Caroline |last7=Auguste |first7=Patrick |date=19 June 2022 |title=Extinct species identification from late middle Pleistocene and earlier Upper Pleistocene bone fragments and tools not recognizable from their osteomorphological study by an enhanced proteomics protocol |url=https://onlinelibrary.wiley.com/doi/10.1111/arcm.12800 |journal=[[Archaeometry]] |language=en |volume=65 |issue=1 |pages=196–212 |doi=10.1111/arcm.12800 |issn=0003-813X |access-date=13 May 2024 |via=Wiley Online Library|hdl=20.500.12210/77657 |hdl-access=free }}</ref> [[Italy]],<ref>{{Cite journal |last=Martini |first=Ivan |last2=Coltorti |first2=Mauro |last3=Mazza |first3=Paul P.A. |last4=Rustioni |first4=Marco |last5=Sandrelli |first5=Fabio |date=January 2014 |title=The latest Ursus spelaeus in Italy, a new contribution to the extinction chronology of the cave bear |url=https://www.cambridge.org/core/product/identifier/S0033589400002118/type/journal_article |journal=[[Quaternary Research]] |language=en |volume=81 |issue=1 |pages=117–124 |doi=10.1016/j.yqres.2013.10.003 |issn=0033-5894 |access-date=3 May 2024 |via=Cambridge Core}}</ref> parts of [[Germany]],<ref>{{Cite journal |last=Münzel |first=Susanne C. |last2=Stiller |first2=Mathias |last3=Hofreiter |first3=Michael |last4=Mittnik |first4=Alissa |last5=Conard |first5=Nicholas J. |last6=Bocherens |first6=Hervé |date=6 December 2011 |title=Pleistocene bears in the Swabian Jura (Germany): Genetic replacement, ecological displacement, extinctions and survival |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618211002345 |journal=[[Quaternary International]] |language=en |volume=245 |issue=2 |pages=225–237 |doi=10.1016/j.quaint.2011.03.060 |access-date=3 May 2024 |via=Elsevier Science Direct}}</ref> [[Poland]],<ref>{{Cite journal |last=Baca |first=Mateusz |last2=Popović |first2=Danijela |last3=Stefaniak |first3=Krzysztof |last4=Marciszak |first4=Adrian |last5=Urbanowski |first5=Mikołaj |last6=Nadachowski |first6=Adam |last7=Mackiewicz |first7=Paweł |date=11 October 2016 |title=Retreat and extinction of the Late Pleistocene cave bear (Ursus spelaeus sensu lato) |url=http://link.springer.com/10.1007/s00114-016-1414-8 |journal=[[The Science of Nature]] |language=en |volume=103 |issue=11-12 |doi=10.1007/s00114-016-1414-8 |issn=0028-1042 |pmc=5059403 |pmid=27730265 |access-date=3 May 2024 |via=Springer Link}}</ref> the [[Balkans]],<ref>{{Cite journal |last=Cvetković |first=Nevena J. |last2=Dimitrijević |first2=Vesna M. |date=7 August 2014 |title=Cave bears (Carnivora, Ursidae) from the Middle and Late Pleistocene of Serbia: A revision |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618213008252 |journal=[[Quaternary International]] |language=en |volume=339-340 |pages=197–208 |doi=10.1016/j.quaint.2013.10.045 |access-date=3 May 2024 |via=Elsevier Science Direct}}</ref><ref>{{Cite journal |last=Seetah |first=T. Krish |last2=Cardini |first2=Andrea |last3=Miracle |first3=Preston T. |date=February 2012 |title=Can morphospace shed light on cave bear spatial-temporal variation? Population dynamics of Ursus spelaeus from Romualdova pećina and Vindija, (Croatia) |url=https://linkinghub.elsevier.com/retrieve/pii/S0305440311003645 |journal=[[Journal of Archaeological Science]] |language=en |volume=39 |issue=2 |pages=500–510 |doi=10.1016/j.jas.2011.10.005 |access-date=3 May 2024 |via=Elsevier Science Direct}}</ref> [[Romania]],<ref>{{Cite journal |last=Quilès |first=Jérôme |last2=Petrea |first2=Catalin |last3=Moldovan |first3=Oana |last4=Zilhão |first4=João |last5=Rodrigo |first5=Ricardo |last6=Rougier |first6=Hélène |last7=Constantin |first7=Silviu |last8=Milota |first8=Ştefan |last9=Gherase |first9=Mircea |last10=Sarcină |first10=Laurenţiu |last11=Trinkaus |first11=Erik |date=December 2006 |title=Cave bears (Ursus spelaeus) from the Peştera cu Oase (Banat, Romania): Paleobiology and taphonomy |url=https://linkinghub.elsevier.com/retrieve/pii/S1631068306001205 |journal=[[Comptes Rendus Palevol]] |language=en |volume=5 |issue=8 |pages=927–934 |doi=10.1016/j.crpv.2006.09.005 |access-date=3 May 2024 |via=Elsevier Science Direct}}</ref><ref>{{Cite journal |last=Bojar |first=Ana-Voica |last2=Piotrowska |first2=Natalia |last3=Barbu |first3=Victor |last4=Bojar |first4=Hans-Peter |last5=Pawełczyk |first5=Fatima |last6=Smeu |first6=Andrei |last7=Guja |first7=Ovidiu |date=24 July 2024 |title=Ursus spelaeus (Rosenmüller, 1794) during the MIS 3: new evidence from the Cioclovina Uscată Cave and radiocarbon age overview for the Carpathians* |url=https://www.tandfonline.com/doi/abs/10.1080/10256016.2024.2376730%4010.1080/tfocoll.2024.0.issue-ESIR2023 |journal=Isotopes in Environmental and Health Studies |volume=0 |issue=0 |pages=1–13 |doi=10.1080/10256016.2024.2376730 |issn=1025-6016 |pmid=39049521 |access-date=14 March 2025 |via=Taylor and Francis Online|doi-access=free }}</ref> [[Georgia (country)|Georgia]], and parts of [[Russia]],<ref>{{Cite journal |last=Kosintsev |first=P.A. |last2=Gasilin |first2=V.V. |last3=Gimranov |first3=D.O. |last4=Bachura |first4=O.P. |date=28 October 2016 |title=Carnivores (Mammalia, Carnivora) of the Urals in the Late Pleistocene and Holocene |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618215010757 |journal=[[Quaternary International]] |language=en |volume=420 |pages=145–155 |doi=10.1016/j.quaint.2015.10.089 |access-date=3 May 2024 |via=Elsevier Science Direct}}</ref> including the [[Caucasus]]; and northern [[Iran]].<ref>{{Cite journal |last=Amiri |first=S. |last2=Biglari |first2=F. |last3=Argant |first3=A. |last4=Crégut-Bonnoure |first4=E. |last5=Mohaseb |first5=A. F. |last6=Noseda |first6=A. |last7=Moradi Bistouni |first7=A. |last8=Mashkour |first8=M. |date=November 2022 |title=The Pleistocene Biodiversity of the Zagros Mountains: Carnivores of the Chenar Cave (Kermanshah- Iran) |url=https://hal.science/hal-04036403 |journal=ASWA Archaeozoology of SouthWest Asia and Adjacent Areas. Tokyo Japan |location=Tokyo, Japan}}</ref> No traces of cave bears have been found in the northern British Isles, [[Scandinavia]] or the [[Baltic countries]], which were all covered in extensive [[glacier]]s at the time. The largest numbers of cave bear remains have been found in [[Austria]], [[Switzerland]], northern Italy, northern Spain, southern France, and Romania, roughly corresponding with the Pyrenees, Alps, and Carpathians. The huge number of bones found in southern, central and eastern Europe has led some scientists to think Europe may have once had herds of cave bears. Others, however, point out that, though some caves have thousands of bones, they were accumulated over a period of 100,000 years or more, thus requiring only two deaths in a cave per year to account for the large numbers.<ref name="bieder"/>

The cave bear inhabited low mountainous areas, especially in regions rich in [[limestone]] caves. They seem to have avoided open plains, preferring forested or forest-edged terrains.<ref name="bieder"/>

==Relationship with humans==
[[File:De nyeste Kunstretninger og smitsomme Sindslidelser-Vægtegning i Combarelles.png|thumb|Cave bear (upper right) along with other animals depicted in [[rock art]] from the [[Les Combarelles]] cave]] 
Between the years 1917 and 1923, the Drachenloch cave in Switzerland was excavated by Emil Bächler. The excavation uncovered more than 30,000 cave bear skeletons. It also uncovered a stone chest or [[cist]], consisting of a low wall built from limestone slabs near a cave wall with a number of bear skulls inside it. A cave bear skull was also found with a femur bone from another bear stuck inside it. Scholars speculated that it was proof of prehistoric human religious rites involving the cave bear, or that the Drachenloch cave bears were hunted as part of a hunting ritual, or that the skulls were kept as trophies.<ref>{{Cite web |url=http://www.showcaves.com/english/ch/caves/Drachenloch.html |title=Caves of Switzerland: Drachenloch |access-date=2013-04-10 |archive-url=https://web.archive.org/web/20130508045303/http://www.showcaves.com/english/ch/caves/Drachenloch.html |archive-date=2013-05-08 |url-status=dead }}</ref> In ''Archaeology, Religion, Ritual'' (2004), archaeologist [[Timothy Insoll]] strongly questions whether the Drachenloch finds in the stone cist were the result of human interaction. Insoll states that the evidence for religious practices involving cave bears in this time period is "far from convincing". Insoll also states that comparisons with the religious practices involving bears that are known from historic times are invalid.<ref>Insoll, Timothy, ''Archaeology, Religion, Ritual'' (2004), Routledge (London), {{ISBN|0415253136}}</ref>

A similar phenomenon was encountered in Regourdou, southern France. A rectangular pit contained the remains of at least twenty bears, covered by a massive stone slab. The remains of a [[Neanderthal]] lay nearby in another stone pit, with various objects, including a bear [[humerus]], a scraper, a core, and some flakes, which were interpreted as grave offerings.

An unusual discovery in a deep chamber of [[Toirano Caves|Basura Cave]] in [[Savona]], Italy, is thought to be related to cave bear worship, because there is a vaguely [[zoomorphism|zoomorphic]] [[stalagmite]] surrounded by clay pellets. It is thought to have been used by Neanderthals for a ceremony; bear bones scattered on the floor further suggests it was likely to have had some sort of ritual purpose.<ref>{{cite book|title=Humankind emerging|year=1996 |publisher=HarperCollins |location=New York|pages=440–441|edition=7th |isbn= 0-673-52364-0|author1=B.G. Campbell |author2=J.D. Loy }}</ref>

==Extinction==
[[File:Medvebarlang21.jpg|thumb|Skeleton of a cave bear in the '"[[Bears' Cave]]", [[Chișcău]], [[Romania]]]]
Reassessment of fossils in 2019 indicate that the cave bear probably died out 24,000 years ago.<ref>{{Cite journal|last1=Terlato|first1=Gabriele|last2=Bocherens|first2=Hervé|last3=Romandini|first3=Matteo|last4=Nannini|first4=Nicola|last5=Hobson|first5=Keith A.|last6=Peresani|first6=Marco|date=21 April 2019|title=Chronological and Isotopic data support a revision for the timing of cave bear extinction in Mediterranean Europe|url=https://www.tandfonline.com/doi/full/10.1080/08912963.2018.1448395|journal=[[Historical Biology]]|language=en|volume=31|issue=4|pages=474–484|doi=10.1080/08912963.2018.1448395|bibcode=2019HBio...31..474T |s2cid=90029163|issn=0891-2963}}</ref> A complex set of factors, rather than a single factor, are suggested to have led to the extinction.<ref name="Withering">{{cite journal |author=Stiller, Mathias |display-authors=etal |year=2010 |title=Withering Away—25,000 Years of Genetic Decline Preceded Cave Bear Extinction |journal=[[Molecular Biology and Evolution]] |volume=27 |issue=5 |pages=975–978 |doi=10.1093/molbev/msq083 |pmid=20335279 |doi-access=}}</ref>

Compared with other [[megafauna]]l species that also became extinct during the Last Glacial Maximum, the cave bear was believed to have had a more specialized diet of high-quality plants and a relatively restricted geographical range. This was suggested as an explanation as to why it died out so much earlier than the rest.<ref name="extinction">{{cite journal |author1=Pacher M. |author2=Stuart A.J. |year=2009 |title=Extinction chronology and palaeobiology of the cave bear (''Ursus spelaeus'') |journal=[[Boreas (journal)|Boreas]] |volume=38 |issue=2 |pages=189–206 |bibcode=2009Borea..38..189P |doi=10.1111/j.1502-3885.2008.00071.x |s2cid=128603825 |via=Wiley Online Library}}</ref> Some experts have disputed this claim, as the cave bear had survived multiple [[climate change]]s prior to extinction. Additionally, [[mitochondrial DNA]] research indicated that the genetic decline of the cave bear began long before it became extinct, demonstrating habitat loss due to climate change was not responsible.<ref name="Withering"/> Finally, high [[Nitrogen-15|δ<sup>15</sup>N]] levels were found in cave bear bones from [[Romania]], indicating wider dietary possibilities than previously believed.<ref name="Omnivore">{{cite journal |author1=Trinkaus, Erik  |author2=Richards, Michael P. |title=Reply to Grandal and Fernández: Hibernation can also cause high δ15N values in cave bears |doi=10.1073/pnas.0801137105 |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=105 |issue=11 |pages=E15 |year=2008 |pmc=2393794 |bibcode=2008PNAS..105E..15T |doi-access=free}}</ref>

Some evidence indicates that the cave bear used only caves for hibernation and was not inclined to use other locations, such as [[thicket]]s, for this purpose, in contrast to the more versatile [[brown bear]]. This specialized hibernation behavior would have caused a high winter mortality rate for cave bears that failed to find available caves. Therefore, as human populations slowly increased, the cave bear faced a shrinking pool of suitable caves, and slowly faded away to extinction, as both Neanderthals and anatomically modern humans sought out caves as living quarters, depriving the cave bear of vital habitat. This hypothesis is being researched {{as of|lc=y|2010}}. According to the research study, published in the journal ''[[Molecular Biology and Evolution]]'', radiocarbon dating of the fossil remains shows that the cave bear ceased to be abundant in Central Europe around 35,000 years ago.<ref>{{cite web|title=True Causes for Extinction of Cave Bear Revealed: More Human Expansion Than Climate Change|url=https://www.sciencedaily.com/releases/2010/08/100824082230.htm|website=ScienceDaily|publisher=Plataforma SINC|date=25 August 2010}}</ref>

In addition to environmental change, human hunting has also been implicated in the ultimate extinction of the cave bear.<ref>{{Cite journal |last=Mondanaro |first=Alessandro |last2=Di Febbraro |first2=Mirko |last3=Melchionna |first3=Marina |last4=Carotenuto |first4=Francesco |last5=Castiglione |first5=Silvia |last6=Serio |first6=Carmela |last7=Danisi |first7=Simone |last8=Rook |first8=Lorenzo |last9=Diniz‐Filho |first9=Jose Alexandre F. |last10=Raia |first10=Pasquale |date=9 February 2019 |title=Additive effects of climate change and human hunting explain population decline and extinction in cave bears |url=https://onlinelibrary.wiley.com/doi/10.1111/bor.12380 |journal=[[Boreas (journal)|Boreas]] |language=en |volume=48 |issue=3 |pages=605–615 |doi=10.1111/bor.12380 |issn=0300-9483 |access-date=15 March 2024 |via=Wiley Online Library}}</ref> In 2019, the results of a large scale study of 81 bone specimens (resulting in 59 new sequences) and&nbsp;64 previously published complete mitochondrial genomes of cave bear mitochondrial DNA remains found in Switzerland, Poland, France, Spain, Germany, Italy and Serbia, indicated that the cave bear population drastically declined starting around 40,000 years ago at the onset of the [[Aurignacian]], coinciding with the arrival of anatomically modern humans.<ref>{{Cite web|url=https://www.bbc.co.uk/news/science-environment-49345392|title=Extinction: Humans played big role in demise of the cave bear|last=Briggs|first=Helen|date=16 August 2019|website=BBC News|access-date=17 August 2019}}</ref><ref name="Gretzinger2019">{{cite journal |last1=Gretzinger |first1=J. |last2=Molak |first2=M. |last3=Reiter |first3=E. |last4=Pfrengle |first4=S. |last5=Urban |first5=C. |last6=Neukamm |first6=J. |last7=Blant |first7=M. |last8=Conard |first8=N. J. |last9=Cupillard |first9=C. |last10=Dimitrijević |first10=V. |last11=Drucker |first11=D. G. |last12=Hofman-Kamińska |first12=E. |last13=Kowalczyk |first13=R. |last14=Krajcarz |first14=M. T. |last15=Krajcarz |first15=M. |date=15 August 2019 |title=Large-scale mitogenomic analysis of the phylogeography of the Late Pleistocene cave bear |journal=[[Scientific Reports]] |volume=9 |issue=1 |page=10700 |bibcode=2019NatSR...910700G |doi=10.1038/s41598-019-47073-z |pmc=6695494 |pmid=31417104 |last16=Münzel |first16=S. C. |last17=Peresani |first17=M. |last18=Romandini |first18=M. |last19=Rufí |first19=I. |last20=Soler |first20=J. |last21=Terlato |first21=G. |last22=Krause |first22=J. |last23=Bocherens |first23=H. |last24=Schuenemann |first24=V. J.}}</ref> It was concluded that human hunting and/or competition played a major role in their decline and ultimate disappearance, and that climate change was not likely to have been the dominant factor.<ref name="Gretzinger2019" /> In a study of Spanish cave bear mtDNA, each cave used by cave bears was found to contain almost exclusively a unique lineage of closely related haplotypes, indicating a homing behaviour for birthing and hibernation. The conclusion of this study is cave bears could not easily colonize new sites when in competition with humans for these resources.<ref>{{Cite journal|last1=Fortes|first1=Gloria G.|last2=Grandal-d'Anglade|first2=Aurora|last3=Kolbe|first3=Ben|last4=Fernandes|first4=Daniel|last5=Meleg|first5=Ioana N.|last6=García-Vázquez|first6=Ana|last7=Pinto-Llona|first7=Ana C.|last8=Constantin|first8=Silviu|last9=Torres|first9=Trino J. de|date=10 August 2016|title=Ancient DNA reveals differences in behaviour and sociality between brown bears and extinct cave bears|journal=Molecular Ecology|language=en|volume=25|issue=19|pages=4907–4918|doi=10.1111/mec.13800|issn=1365-294X|url=http://irep.ntu.ac.uk/id/eprint/38241/1/1233179_Barlow.pdf|pmid=27506329|bibcode=2016MolEc..25.4907F |s2cid=18353913}}</ref>

Overhunting by [[human]]s has been dismissed by some as human populations at the time were too small to pose a serious threat to the cave bear's survival. However, the two species may have competed for living space in caves.<ref name="bieder" /><ref name="Withering" /> The [[Chauvet Cave]] contains around 300 "bear hollows" created by cave bear hibernation.<ref>{{Cite web |title=Bear hollows {{!}} The Chauvet-Pont d'Arc cave |url=https://archeologie.culture.gouv.fr/chauvet/en/bear-hollows |access-date=6 September 2023 |website=archeologie.culture.gouv.fr}}</ref> Unlike brown bears, cave bears are seldom represented in cave paintings, leading some experts to believe the cave bear may have been avoided by human hunters<ref>''The Walking Larder: Patterns of Domestication, Pastoralism, and Predation'' by Juliet Clutton-Brock, published by Routledge, 1990, {{ISBN|0-04-445900-9}}</ref> or their habitat preferences may not have overlapped. Paleontologist [[Björn Kurtén]] [[hypothesis|hypothesized]] cave bear populations were fragmented and under stress even before the advent of the glaciers.<ref name="bieder" /> Populations living south of the [[Alps]] possibly survived significantly longer.<ref name="extinction" />

==See also==
* [[Azykh Cave]]
* [[Kletno Bear Cave]]
* [[Darband Cave]]
* [[Divje Babe Flute]]

==References==
{{Reflist|colwidth=30em}}

==External links==
{{Commons category|Ursus spelaeus}}
* [http://www.showcaves.com/english/ro/showcaves/Ursilor.html Show Caves of Romania]
* [https://web.archive.org/web/20120402115230/http://worldmuseumofman.org/cavebear.php Cave Bear Fossils]

{{Wikispecies|Ursus spelaeus}}

{{Ursidae extinct nav}}
{{Taxonbar|from=Q188851}}
{{Authority control}}

[[Category:Cave bear| ]]
[[Category:Pleistocene bears]]
[[Category:Pleistocene mammals of Europe]]
[[Category:Pleistocene mammals of Asia]]
[[Category:Pleistocene carnivorans]]
[[Category:Fossil taxa described in 1794]]
[[Category:Fossils of Serbia]]
[[Category:Ursus (mammal)]]