Ursus spelaeus spelaeus Rosenmüller, 1794

Cave bear



Taxonomy & Nomenclature



Conservation Status
















NB: many of the references listed below may refer to other taxa.


Original scientific description:

Rosenmüller, Johann Christian. (1794). Quaedam de ossibus fossilibus animalis cuiusdam, historiam eius et cognitionem accuratiorem illustrantia, dissertatio, quam d. 22. Octob. 1794 ad disputandum proposuit Ioannes Christ. Rosenmüller Heßberga-Francus, LL.AA.M. in Theatro anatomico Lipsiensi Prosector assumto socio Io. Chr. Aug. Heinroth Lips. Med. Stud. Cum tabula aenea.- 34 S., Leipzig.


Other references:

Ábelová, M. – Sabol, M. 2009. Attempt on the reconstruction of palaeotemperature and palaeoenvironment in the territory of Slovakia during the Last Glacial based on oxygen and carbon isotopes from tooth enamel and bone collagen of cave bears. Slovenský kras, 47, Suppl. 1, 5-18. 

Abrams, G., Bello, S. M., Di Modica, K., Pirson, S. and Bonjean, D. (2014). When Neanderthals used cave bear (Ursus spelaeus) remains: bone retouchers from unit 5 of Scladina Cave (Belgium). Quatern. Int. 326-327: 274-287.

Athen, K. (2006). Biostatistical investigation of long bones and metapodial bones of Ursus spelaeus and Ursus deningeri. Scientific Annals, School of Geology Aristotle University of Thessaloniki 98: 159-162.

Augusta, J. – Hokr, Z. 1948a. O mláděti jeskynního medvěda (Ursus spelaeus Blmb.) z jaskyně „Domica“ ve Slovenskem krasu. Práce Štátneho geologického ústavu v Bratislave, 17, 29–34. 

Baca, M., Nadachowski, A., Lipecki, G., Mackiewicz, P., Marciszak, A., Popoviæ, D., Socha, P., Stefaniak, K., Wojtal, P. (2017). Impact of climatic changes in the Late Pleistocene on migrations and extinction of mammals in Europe: four case studies. Geological Quarterly 61(2): [pagination?]. doi: 10.7306/gq.1319

Baca, Mateusz, Popović, Danijela, Stefaniak, Krzysztof, Marciszak, Adrian, Urbanowski, Mikołaj, Nadachowski, Adam and Mackiewicz, Paweł. (2016). Retreat and extinction of the Late Pleistocene cave bear ([i]Ursus spelaeus[/i] sensu lato). The Science of Nature 103: 92.

Barlow, Axel et al. (2018). Partial genomic survival of cave bears in living brown bears. Nature Ecology & Evolution 2: 1563-1570. [Abstract]

Baryshnikov, G. F. (2006). Morphometrical variability of cheek teeth of cave bears. Scientific Annals, School of Geology. Aristotle University of Thessaloniki. Spec. 98: 81-102.

Baryshnikov, G. F., Germonpré, M. and Baryshnikova, S. V. (2010). Cave bear ([i]Ursus spelaeus[/i]) from Chamber B of the Goyet Cave in Belgium. Russian Journal of Theriology 9(2): 93-104.

Baryshnikov, Gennady F., Germonpré, M. and Sablin, M. (2003). Sexual dimorphism and morphometric variability of cheek teeth of the cave bear ([i]Ursus spelaeus[/i]). Belgian Journal of Zoology 133(2): 111-119.

Baryshnikov, Gennady F., Gimranov, Dmitry and Kosintsev, Pavel. (In Press, 2018). Variability of the upper incisors in the cave bears (Carnivora, Ursidae) from the Caucasus and Urals. Comptes Rendus Palevol. [Abstract]

Baryshnikov, G. and Puzachenko, A. (2011). Craniometrical variability in the cave bears (Carnivora, Ursidae): multivariate comparative analysis. Quaternary International 245: 350-368.

Baryshnikov, Gennady F. and Puzachenko, Andrey Yu. (2018). Morphometry of upper cheek teeth of cave bears (Carnivora, Ursidae). Boreas. doi: [Abstract]

Baryshnikov, Gennady F. and Puzachenko, Andrey Yu. (2020). Morphometry of lower cheek teeth of cave bears (Carnivora, Ursidae) and general remarks on the dentition variability. Boreas. [Abstract]

Baryshnikov, G. and Tsoukala, E. (2010). New analysis of the Pleistocene carnivores from Petralona Cave (Macedonia, Greece) based on the Collection of the Thessaloniki Aristotle University. Geobios 43: 389-402.

Bate, Dorothea M. A. (1937). Note on recent finds of Dama clactoniana (Cervus browni auctt.) in London and Swanscombe. Proceedings of the Prehistoric Society 3(1-2): 460-463. [First page]

Bendík, A. – Sabol, M. 2007. Cave Bears from the Cave of Izabela Textorisová (the Veľká Fatra Mts., Slovakia) – a state of the art. Scripta Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis, Geology, 35, 150–156.

Bendík, A. – Štuller, F. – Straka, Ľ. – Novomeský, F. – Nečas, L. – Strecha, J. – Sabol, M. 2009. Pathological modifications on bones of cave bears from the Veľká Fatra Mts. (Central Western Carpathians). Slovenský kras, 47, Suppl. 1, 19–24. 

Bocherens, Hervé. (2015). Isotopic tracking of large carnivore palaeoecology in the mammoth steppe. Quaternary Science Reviews 117: 42-71. [Abstract]

Bocherens, Hervé. (2019). Isotopic insights on cave bear palaeodiet. Historical Biology 31(4): 410-421. [Abstract]

Bocherens, Hervé, Bridault, A., Drucker, D.G., Hofreiter, M., Münzel, S.C., Stiller, M., van der Plicht, J. (2014). The last of its kind? Radiocarbon, ancient DNA and stable isotope evidence from a late cave bear (Ursus spelaeus Rosenmüller, 1794) from Rochedane (France). Quaternary International 339-340: 179-188.

Hervé Bocherens, Aurora Grandal-d'Anglade and Keith A. Hobson. (In Press, 2014). Pitfalls in comparing modern hair and fossil bone collagen C and N isotopic data to reconstruct ancient diets: a case study with cave bears (Ursus spelaeus). Isotopes in Environmental and Health Studies. [Abstract]

Bocherens, H., Stiller, M., Hobson, K. A., Pacher, M., Rabeder, G., Burns, J. A., et al. (2011). Niche partitioning between two sympatric genetically distinct cave bears (Ursus spelaeus and Ursus ingressus) and brown bear (Ursus arctos) from Austria: isotopic evidence from fossil bones. Quaternary International 245(2): 238-248.

Boeskorov G.G., S. E. Grigoriev, Baryshnikov G.F. (2012). New evidence for the existence of Pleistocene cave bears in Arctic Siberia. Doklady Biological Sciences 445(2): 226-230.

Bon C, Caudy N, de Dieuleveult M, Fosse P, Philippe M, Maksud F, Beraud-Colomb E, Bouzaide E, Kefie R, Laugier C, Rousseau B, Casane D, van der Plicht J, Elalouf J-M. (2008). Deciphering the complete mitochondrial genome and phylogeny of the extinct cave bear in the Paleolithic painted cave of Chauvet. Proceedings of the National Academy of Science of the United States of America 105(45): 17447-17452.

Bona, Fabio, Baioni, Marco and Cilli, Elisabetta. (2023). Paleoclimate and paleoenvironment in north-western Italy (Southern Alps) during the Last Glacial Stage: the small-mammal record from the Buco del Frate Cave (Prevalle – Brescia). Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy) 129(1): 79-90.

Boule, M. (1887). Ours et hyènes Quaternaires. Le Naturaliste revue illustrée des sciences naturelles (2) 5: 53-54.

Bray, Fabrice et al. (2022). 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. archaeometry.

Buccheri, Fabio et al. (2014). New evidence of interaction between Neandertal and cave bear during Middle Pleistocene in North Italy (Borgosesia, VC). Poster presented at the 7th International Meeting on Taphonomy and Fossilization Ferrara, Italy, September 10-13, 2014.

Buccheri, Fabio et al. (2016). Taphonomic analysis on fossil remains from the Ciota Ciara Cave (Piedmont, Italy) and new evidence of cave bear and wolf exploitation with simple quartz flakes by Neanderthal. Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy) 122(3): 41-54.

Čeklovský, T. – Sabol, M. 2012. Cave Bear Assemblage from the Šarkanova diera Cave (the Volovské vrchy Mts., Eastern Slovakia). Braunschweiger Naturkundliche Schriften 11: 25-39.

Channarayapatna, Sharada, Valensi, Patricia and Hohenstein, Ursula Thun. (2016). Archaeozoological analyses of large mammals from the prehistoric cave site of Lazaret, France: A case study of Archaeostratigraphic Unit 28, pp. 399-407. In: Pérez, Lucía Agudo et al. (eds.). IX Jornadas de Jóvenes en Investigación Arqueológica. Santander 8-11 de junio de 2016.

Charters, Daniel et al. (2019). Temporal variation in cave bear (Ursus spelaeus) dentition: The stratigraphic sequence of Scladina Cave, Belgium. Quaternary Science Reviews 205: 76-85. [Abstract]

Charters, Daniel et al. (2022). Morphological evolution of the cave bear (Ursus spelaeus) mandibular molars: coordinated size and shape changes through the Scladina Cave chronostratigraphy. Palaeogeography, Palaeoclimatology, Palaeoecology 587: 110787.

Christiansen, Per. (1999). What size were Arctodus simus and Ursus spelaeus (Carnivora: Ursidae)? Ann. Zool. Fennici 36: 93-102.

Daschek, Éva J. and Mester, Zsolt. (2020). A site with mixed occupation: Neanderthals and carnivores at Érd (Hungary). Journal of Archaeological Science: Reports 29: 102116.

Dawkins, William Boyd. (1868). On the Pleistocene Mammals of Yorkshire. Proceedings of the Geological and Polytechnic Society of the West Riding of Yorkshire 4: 502-510.

Debeljak, Irena. (2007). Fossil population structure and mortality of the cave bear from the Mokrica Cave (northern Slovenia). Acta Carsologica 36(3): 475-484.

Diedrich, Cajus G. (2005). Cracking and nibbling marks as indicators for the Upper Pleistocene spotted hyaena as a scavenger of cave bear (Ursus spelaeus Rosenmüller, 1794) carcasses in the Perick Caves den of Northwest Germany. Abhandlungen der Naturhistorischen Gesellschaft Nürnberg 45: 73-90.

Diedrich, Cajus G. (2006a). Ice age spotted hyenas ?hunting or only scavenging on a cave bear Ursus spelaeus Rosenmüller at the Ice Age spotted hyena freeland den and prey deposit site Bad Wildungen-Biedensteg (Hessia, Germany). Scientific Annals, School of Geology Aristotle University of Thessaloniki (AUTH), special volume 98: 193-199.

Diedrich, Cajus G. (2006b). Die oberpleistozäne Population von Ursus spelaeus Rosenmüller 1794 aus dem eiszeitlichen Fleckenhyänenhorst Perick-Höhlen von Hemer (Sauerland, NW Deutschland). Philippia 12(4): 275-346.

Diedrich, Cajus G. (2009). The rediscovered cave bear “Ursus spelaeus Rosenmüller 1794” holotype of the Zoolithen Cave (Germany) from the historic Rosenmüller collection. Acta Carsologica Slovacia 47(1): 25-32.

Diedrich, Cajus G. (2012c). Cave bear killers and scavengers from the last ice age of central Europe: Feeding specializations in response to the absence of mammoth steppe fauna from mountainous regions. Quaternary International, special issue Mammoths and Their Relatives 1: Biotopes, Evolution and Human Impact V International Conference, Le Puy-en-Velay, 2010 255: 59-78. [Abstract]

Diedrich, Cajus G. (2015). ‘Neanderthal bone flutes’: simply products of Ice Age spotted hyena scavenging activities on cave bear cubs in European cave bear dens. R. Soc. open sci. 2: 140022.

Dong, Wei and Zhang, Yingqi. (2022). Biochronologic sequences of the Pleistocene mammalian faunas in China and correlations with numeric dating. Quaternary Sciences 42(5): 1227-1246.

Döppes, Doris et al. (2019). Extinction pattern of Alpine cave bears - new data and climatological interpretation. Historical Biology 31(4): 422-428. [Abstract]

Döppes, Doris et al. (2021). The late pleistocene cave bear fauna of the Torrener Bärenhöhle in the northern alps (Salzburg, Austria). Historical Biology. doi: [Abstract]

Estraviz López, Darío. (2019). Quaternary fossil vertebrates from continental Portugal: Paleobiodiversity, revision of specimens and new localities. Dissertation. xxv + 170 pp.

Estraviz López, Darío et al. (2021). Quantitative classification of metapodial bones of Ursus spelaeus and Ursus arctos from Northwestern Iberia using multivariate analysis. Ciências da Terra Procedia 1: 92-95. doi:10.21695/cterraproc.v1i0.418

Farina, Simone. (2016). Catalogue of the late Pleistocene-Holocene fossil mammalian collection from “Canale delle Acque Alte (Canale Mussolini)” (Natural History Museum, University of Pisa). Atti Soc. Tosc. Sci. Nat., Mem., Serie A, 123: 47-53, 3 figs.

Figueirido, Borja and van Heteren, Anneke H. (2019). The story continues: recent advances on the life and death of the Pleistocene cave bear. Historical Biology 31(4): 405-409. [Abstract]

Fortes, Gloria G. et al. (2016). Ancient DNA reveals differences in behaviour and sociality between brown bears and extinct cave bears. Molecular Ecology 25(19): 4907-4918. [Abstract]

Fourvel, Jean-Baptiste, Fosse, Philippe, Fernandez, Philippe and Antoine, Pierre-Olivier. (2014). La grotte de Fouvent, dit l’Abri Cuvier (Fouvent-le-Bas, Haute-Saône, France): analyse taphonomique d’un repaire d’hyènes du Pléistocène supérieur (OIS 3). PALEO 25: 79-99.

Fuentes-Sánchez, Daniel et al. (2019). Evidence of congenital block vertebra in Pleistocene Cave Bear (Ursus spelaeus) from Cueva de Guantes (Palencia, Spain). International Journal of Paleopathology 24: 165-170.


Galdies, Johann. (2021). The bears of the European steppe: a review. Quaternaire 33(1): 47-62. [Abstract]

García-Vázquez, Ana, Pinto-Llona, Ana C. and Grandal-d’Anglade, Aurora. (In Press, 2017). Brown bear (Ursus arctos L.) palaeoecology and diet in the Late Pleistocene and Holocene of the NW of the Iberian Peninsula: A study on stable isotopes. Quaternary International. [Abstract]

García-Vázquez, Ana et al. (2023). Characterising the cave bear Ursus spelaeus Rosenmüller by ZooMS: a review of peptide mass fingerprinting markers. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. doi:10.1017/S1755691023000038

Ghezzo, Elena, Palchetti, Alessandro and Rook, Lorenzo. (2014). Recovering data from historical collections: stratigraphic and spatial reconstruction of the outstanding carnivoran record from the Late Pleistocene Equi cave (Apuane Alps, Italy). Quaternary Science Reviews 96: 168-179.

Gimranov, Dmitry, Bocherens, Hervé, Kavcik-Graumann, Nadja, Nagel, Doris and Rabeder, Gernot. (2022). The cave bears from Imanay Cave (Southern Urals, Russia). Historical Biology 35(4): 580-588.

Gimranov, D. O. and Kosintsev, P. A. (2022). Cave Bears (Ursus spelaeus sensu lato) of the Urals. Paleontological Journal 56: 97-105.

Aurora Grandal-d'Anglade and Fernando Lopez-Gonzalez. (2004). A study of the evolution of the Pleistocene cave bear by a morphometric analysis of the lower carnassial. Oryctos 5: 83-94.

Grandal-d’Anglade, Aurora et al. (2019). The cave bear’s hibernation: reconstructing the physiology and behaviour of an extinct animal. Historical Biology 31(4): 429-441. [Abstract]

Grandal-D'Anglade, A. and Vidal Romaní, J. R. (1997). A population study on the Cave Bear (Ursus spelaeus Ros.-Hein.) from Cova Eirós (Triacastela, Galicia, Spain). Geobios 30(5): 723-731.

Gretzinger, Joscha et al. (2019). Large-scale mitogenomic analysis of the phylogeography of the Late Pleistocene cave bear. Scientific Reports 9: 10700.

Gutiérrez, Eliécer E. and Pine, Ronald H. (2015). No need to replace an “anomalous” primate (Primates) with an “anomalous” bear (Carnivora, Ursidae). ZooKeys 487: 141-154.

Hajna, Nadja Zupan et al. (2021). Pliocene to Holocene chronostratigraphy and palaeoenvironmental records from cave sediments: Račiška pečina section (SW Slovenia). Quaternary International. doi:

Hänni C, Laudet V, Stehelin D, Taberlet P. (1994). Tracking the origins of the cave bear (Ursus spelaeus) by mitochondrial DNA sequencing. Proceedings of the National Academy of Sciences, USA  91(25): 12336-12340.

van Heteren, Anneke H. and Figueirido, Borja. (2019). Diet reconstruction in cave bears from craniodental morphology: past evidences, new results and future directions. Historical Biology 31(4): 500-509. [Abstract]

van Heteren, Anneke H., MacLarnon, Ann, Soligo, Christophe and Rae, Todd C. (2015). Functional morphology of the cave bear (Ursus spelaeus) mandible: a 3D geometric morphometric analysis. Organisms Diversity & Evolution. doi:10.​1007/​s13127-015-0238-2 [Abstract]

Hilderbrand, G. V., Farley, S. D., Robbins, C. T., Hanley, T. A., Titus, K. and Servheen, C. (1996). Use of stable isotopes to determine diets of living and extinct bears. Can. J. Zool. 74: 2080-2088. 

Hofreiter, Michael. (2007). Sudden replacement of cave bear mitochondrial DNA in the late Pleistocene. Current Biology 17(4): R122-123.

Indelicato, M., Rossi, M. and Santi, G. (2023). Cave bear skulls from Buco del Frate (Brescia, N. Italy). First morphometric analysis. e-Research Reports of Museum Burg Golling 10: 1-6.

Jones, D. Brent and DeSantis, Larisa R. G. (2016). Dietary ecology of the extinct cave bear: Evidence of omnivory as inferred from dental microwear textures. Acta Palaeontologica Polonica 61.

Knapp, Michael. (2019). From a molecules’ perspective – contributions of ancient DNA research to understanding cave bear biology. Historical Biology 31(4): 442-447. [Abstract]

Knapp M., Rohland N., Weinstock J., Baryshnikov G., Sher A., Nagel D., Rabeder G., Pinhasi R.O.N., Schmidt H.A., Hofreiter M. 2009 First DNA sequences from Asian cave bear fossils reveal deep divergences and complex phylogeographic patterns. Molecular Ecology 18(6), 1225-1238.

Kosintsev, P.A. and Vorob’ev, A.A., in Mamont i ego okruzhenie: 200 let izucheniya (The Mammoth and Its Surroundings: 200 Years of Study), Moscow: Geos, 2001, pp. 266-278.

Kowalski, K., 1961, [Traces of a cave bear]: Priroda, 1961, v. 2, p. 110-112.

Krajcarz, Magdalena, Pacher, M. – Krajcarz, M. T. – Laughlan, L. – Rabeder, G. – Sabol, M. – Wojtal, P. and Bocherens, H. (2016). Isotopic variability of cave bears (δ15N, δ13C) across Europe during MIS 3. Quaternary Science Reviews 131A: 51-72. [Abstract]

Krause J, Unger T, Noçon A, Malaspinas A-S, Kolokotronis S-O, Stiller M, Soibelzon L, Spriggs H, Dear PH, Briggs AW, Bray SCE, O’Brien SJ, Rabeder G, Matheus P, Cooper A, Slatkin M, Pääbo S, Hofreiter M (2008) Mitochondrial genomes reveal an explosive radiation of extinct and extant bears near the Miocene–Pliocene boundary. BMC Evolutionary Biology 8: 220.

Kurtén, Björn Olof Lennartson. (1955). Sex dimorphism and size trends in the cave bear Ursus spelaeus Rosenmüller and Heinroth. Acta Zool Fennica 90: 1-48.

Kurten, B., Acta Zool. Fen., 1958, vol. 90, p. 59.

Kurtén B. 1968 Pleistocene mammals of Europe. London, AldineTransaction.

Kurtén, Björn Olof Lennartson. (1976). The Cave Bear Story: Life and Death of a Vanished Animal. New York: Columbia University Press. 163 pp.

Loreille O, Orlando L, Patou-Mathis M, Philippe M, Taberlet P, Hänni C. (2001). Ancient DNA analysis reveals divergence of the cave bear, Ursus spelaeus, and brown bear, Ursus arctos, lineages. Current Biology 11: 200-203.

Majkić, Ana et al. (2017). Sequential Incisions on a Cave Bear Bone from the Middle Paleolithic of Pešturina Cave, Serbia. Journal of Archaeological Method and Theory. doi:10.1007/s10816-017-9331-5 [Abstract]

Markova A.K., Puzachenko A.Y., van Kolfschoten T. 2010 The North Eurasian mammal assemblages during the end of MIS 3 (Brianskian–Late Karginian–Denekamp Interstadial). Quaternary International 212(2), 149-158.

Ivan Martini, Mauro Coltorti, Paul P.A. Mazza, Marco Rustioni, Fabio Sandrelli. (In Press, 2013). The latest Ursus spelaeus in Italy, a new contribution to the extinction chronology of the cave bear. Quaternary Research. [Abstract]

Marciszak, Adrian and Matyaszczyk, Lena. (2021). Południowa Cave - first mid-Middle Pleistocene site from Studety Mts (Silesia, SW Poland) and their biostratigraphical significance, p. 64. In: Ratajczak-Skrzatek U., Kovalchuk O., Stefaniak K. (eds.). Proceedings of INQUA SEQS 2021 Conference, Wrocław, Poland. University of Wrocław & Polish Geological Society. 119 pp.

Marciszak, Adrian, Kropczyk, Aleksandra, Gornig, Wiktoria, Demidziuk, Krzysztof  and Nowakowski, Dariusz. (2023). Carnivores record and Palaeolithic bear cult in Europe: a review from Radochowska Cave (Sudety Mts, SW Poland). Historical Biology.

Marciszak, Adrian Przemysław and Lipecki, Grzegorz. (2020). The history of bears (Ursidae, Carnivora, Mammalia) from Silesia (southern Poland) and the neighbouring areas. Geological Quarterly 64(4): 876-897. doi: 10.7306/gq.1565

Mattson, David J. (1998). Diet and Morphology of Extant and Recently Extinct Northern Bears. A Selection of Papers from the Tenth International Conference on Bear Research and Management, Fairbanks, Alaska, July 1995, and Mora, Sweden, September 1995. Ursus 10: 479-496. [Abstract]

Micó, C., Arilla, M., Rosell, J., Villalba, M., Santos, E., Rivals, F., et al. (2020). Among goats and bears: A taphonomic study of the faunal accumulation from Tritons Cave (Lleida, Spain). Journal of Archaeological Science: Reports 30: 102194. [Abstract]

Mihevc, A. (2003). Sledovi jamskega medveda v Račiški pečini. Naše jame, 45 (2003), pp. 48-55

Moncel, M.-H. (2001). Microlithic Middle Palaeolithic assemblages in Central Europe and elephant remains, pp. 314-317. In: Cavarretta, C., Gioia, P., Mussi, M. and Palombo, M. R. (eds.). The World of Elephants – International Congress, Rome. Proceedings of the 1st international congress – Consiglio nazionale delle ricerche, Rome. [automatic download]

Mondanaro, Alessandro et al. (2019). Additive effects of climate change and human hunting explain population decline and extinction in cave bears. Boreas. doi: [Abstract]

Münzel, S. C. and Conard, N. J. (2004). Cave bear hunting in Hohle Fels Cave in the Ach Valley of Swabian Jura. Revue de Paléobiologie 23(2): 877-885.

Münzel, Susanne C. et al. (2011). Pleistocene bears in the Swabian Jura (Germany): Genetic replacement, ecological displacement, extinctions and survival. Quaternary International 245(2): 225-237.

Münzel, Susanne C. et al. (In Press, 2013). Behavioural ecology of Late Pleistocene bears (Ursus spelaeus, Ursus ingressus): Insight from stable isotopes (C, N, O) and tooth microwear. Quaternary International. [Abstract]

Nadachowskia., Lipeckig., Stefaniak K., Wojtal P. (2010). Radiocarbon dates on cave bear (Ursus spelaeus) and brown bear (Ursus arctos) from Late Pleistocene of Poland. Geophysical Research Abstracts. Vol. 12, EGU2010-5815, 2010. EGU General Assembly 2010.

Naito, Yuichi I. et al. (2016). Evidence for herbivorous cave bears (Ursus spelaeus) in Goyet Cave, Belgium: implications for palaeodietary reconstruction of fossil bears using amino acid δ15N approaches. Journal of Quaternary Science. DOI: 10.1002/jqs.2883 [Abstract]

Naito, Yuichi I. et al. (2020). Heavy reliance on plants for Romanian cave bears evidenced by amino acid nitrogen isotope analysis. Scientific Reports 10: 6612.

Nikolov, I. (1983). Nyakoi belezhki varhu peshternata fosilna bozayna fauna v Balgariya (Some notes on the cave fossil mammalian fauna in Bulgaria). In: IV Evropeyska regionalna konferentsiya po speleologiya (IVth European Regional Conference on Speleology). October 1981, Sofia, 215-218. (In Bulgarian)

Noonan, J.P., M. Hofreiter, D. Smith, J.R.Priest,N.Rohland,G.Rabeder,J.Krause, J.C.Detter, et al. 2005. Genomic sequencing of Pleistocene cave bears. Science 309: 597-600.

Orlando, L., Bonjean, D., Bocherens, H., Thenot, A., Argant, A., Otte, M., Hanni, C., (2002). Ancient DNA and the population genetics of Cave Bears (Ursus spelaeus) through space and time. Mol. Biol. and Evol. 19: 1920-1933.

Pacher, Martina. (2017). Anterior premolar variability in Pleistocene cave and brown bears and its significance in species determination. Fossil Imprint 73(3-4): 482-494.

Pacher, M. and Quilès, J. (2013). Cave Bear Paleontology and Paleobiology at the Pestera cu Oase: Fossil Population Structure and Size Variability, in Trinkhaus E., Constantin S. & Zilhao J. (eds), Life and Death at the Pestera cu Oase. A Setting for Modern Human Emergence in Europe. Oxford University Press: 127–146.

Pacher, M., Stuart, A., Baryshnikov, G., Stiller, M., Kosintsev, P., Vorobiev, A. (2009). Cave bears of the Ural Mountains – a survey based on direct radiocarbon dates, aDNA and morphometrical analysis. 15th International Cave Bear Symposium – Spisska Nova Ves, Slovakia. Abstract book, Bratislava, 14-17.

Pedraza, Iván Ramírez et al. (2022). Paleodiet and niche partitioning among the easternmost European cave bears based on tooth wear analysis. Historical Biology 34(6): 1063-1071.

Peigné S, Goillot C, Germonpré M, Blondel C, Bignon O, et al. (2009). Predormancy omnivory in European cave bears evidenced by a dental microwear analysis of Ursus spelaeus from Goyet, Belgium. Proc Natl Acad Sci USA 106: 15390-15393.

Peigné, Stéphane and Merceron, Gildas. (2019). Palaeoecology of cave bears as evidenced by dental wear analysis: a review of methods and recent findings. Historical Biology 31(4): 448-460. [Abstract]

Pérez-Ramos, Alejandro et al. (2019). A three-dimensional analysis of tooth-root morphology in living bears and implications for feeding behaviour in the extinct cave bear. Historical Biology 31(4): 461-473. [Abstract]

Pérez-Ramos, Alejandro et al. (2020a). Biomechanical simulations reveal a trade-off between adaptation to glacial climate and dietary niche versatility in European cave bears. Sci. Adv. 6: eaay9462.

Pérez-Ramos, Alejandro, Romero, Alejandro, Rodriguez, Ernesto and Figueirido, Borja. (2020b). Three-dimensional dental topography and feeding ecology in the extinct cave bear. Biology Letters. doi:

Petrikovich, J. 1897. Nález kostí jaskynného medveďa (Ursus spelaeus Rosenmüller) v jaskyni Lôm, v Gaderskej doline v Turci. Sborník Muzeálnej slovenskej spoločnosti, 2, 231–235. 

Petrikovich, J. 1907. Nález kostí jaskynného medveďa (Ursus spelaeus Rosenmülleri) v jaskyni Lôm v Gaderi a kosti z obecného medveďa (Ursus arctos L.) na Balovom grúni v Belianskej doline v Turci. Sborník Muzeálnej slovenskej spoločnosti, 12, 24–28. 

Pinto-Llona, A. C. (2013). Macrowear and occlusal microwear on teeth of cave bears Ursus spelaeus and brown bears Ursus arctos: Inferences concerning diet. Palaeogeography, Palaeoclimatology, Palaeoecology 370: 41-50.

Pinto Llona, A. C. and Andrew, P. J. (2004). Scavenging behaviour patterns in cave bears Ursus spelaeus, pp. 845-853. In: Brugal, J.-P. and Fosse, P. (eds.). 14ème congrès UISPP, Hommes et Carnivores au Paléolithique. Revue de Paléobiologie 23. Genève, Switzerland.

Pohar, V. (1997). Late Glacial mammal macrofauna in Slovenia. Quartär, 47-48: 149-158.

Popov, R. (1936). Fosilni I subfosilni zhivotinski ostanki v izsledvanite do sega peshteri na Balgariya (Fossil and subfossil animal remains in the explored until now caves of Bulgaria). Izvestiya na Balgarskoto peshterno druzhestvo (Bulletin of the Bulgarian Speleological Society). Sofia, Vol. I: 1-12. (In Bulgarian).

Prat-Vericat, M.; Rufí Casals, Isaac; Jovells-Vaqué, S.; Sorbelli, L.; Terradas-Batlle, Xavier ; Madurell-Malapeira, Joan. (2021). The Middle to Late Pleistocene vertebrate assemblage from Cova 120 (Alta Garrotxa, Eastern Pyrenees). 5th International Meeting of Early-stage Researchers in Palaeontology Online event, May 18-21 (2021). [Abstract]

Rabal-Garcés, Raquel, Cuenca-Bescós G., Ignacio Canudo J., De Torres T. 2012 Was the European cave bear an occasional scavenger? Lethaia 45(1), 96-108.

Rabal-Garcés, Raquel et al. (2021). A Cave Occupied by Cave Bears for Thousands of Years in the Sobrarbe-Pirineos UNESCO Global Geopark (Huesca, Aragon, Spain). Geoconservation Research 4(6): 710-719.

Rabeder, G. – Hofreiter, M. – Nagel, D. – Withalm, G. 2004. New taxa of Alpine cave bears (Ursidae, Carnivora). Cahiers scientifiques, 2, 49–67.

Ramírez-Pedraza, Iván et al. (2019a). Microwear and isotopic analyses on cave bear remains from Toll Cave reveal both short-term and long-term dietary habits. Scientific Reports 9: 5716.

Ramírez-Pedraza, Iván et al. (In Press, 2019b). Dietary habits of the cave bear from the Late Pleistocene in the northeast of the Iberian Peninsula. Quaternary International. doi: [Abstract]

Riehecky, Janet. (2008). Cave Bear.  Blazers Extinct Monsters series. Capstone Press.

Robu, Marius. (In Press, 2015). The assessment of the internal architecture of a MIS 3 cave bear bone assemblage. Case study: Urşilor cave, Western Carpathians, Romania. Palaeogeography, Palaeoclimatology, Palaeoecology. doi:10.1016/j.palaeo.2015.12.012 [Abstract]

Robu, Marius et al. (2016). Size of the cave bear population and skeletal distribution from Urşilor Cave, Romania. Trav. Inst. Spéol. 55: 63-80.

Marius Robu, Jonathan G. Wynn, Ionuţ C. Mirea, Alexandru Petculescu, Marius Kenesz, Cristina M. Puşcaş, Marius Vlaicu, Erik Trinkaus, Silviu Constantin. (2017). The diverse dietary profiles of MIS 3 cave bears from the Romanian Carpathians: insights from stable isotope (δ13C and δ15N) analysis. Palaeontology. DOI: 10.1111/pala.12338 [Abstract]

Robu, Marius et al. (In Press, 2018). Palaeoichnology of an MIS 3 cave bear settlement – Urşilor Cave (Western Carpathians, Romania). Palaeogeography, Palaeoclimatology, Palaeoecology. [Abstract]

Robu, Marius et al. (2019). Palaeoecology and palaeoclimatic context of Romanian Carpathian MIS 3 cave bears using stable isotopes (δ13C and δ18O). Palaeogeography, Palaeoclimatology, Palaeoecology 534: 109288. [Abstract]

Rodríguez-Almagro, Manuel et al. (In Press, 2024). Taphonomic study of the cave bears (Ursus cf. deningeri and U. spelaeus) from the Sima I of the El Polvorín cave (Northern Iberian Peninsula). Quaternary Science Advances.

Rosendahl, W. – Döppes, D. 2012. About the true nature of dragons in caves. In Silviu, C. (Ed.): Abstract Book, The 18th International Cave Bear Symposium (20–22 September, Baile Herculane, Romania), p. 66. [relevant reference?]

Rosendahl, Wilfried and Kempe, Stephan. (2005). Ursus spelaeus Rosenmüller 1794 and not Rosenmüller & Heinroth - Johann Christian Rosenmüller, his life and the Ursus spelaeus. Naturhistorische Gesellschaf 5: 191-198.

Rossi, Mario; Santi, Giuseppe; Zorzin, Roberto; Döppes, Doris; Friedrich, Ronny; Lindauer, Susanne; Rosendahl, Wilfried. (2018). The first directly dated cave bear from the Covoli di Velo Cave (Verona Province, Veneto, northern Italy) with some discussion of Italian Alps cave bears. Journal of Cave & Karst Studies 80(3): 145-152. [Abstract]

Sabol, M. 1998. Jaskynný medveď (Ursus spelaeus Rosenmüller et Heinroth) z jaskyne Tmavá skala. Mineralia Slovaca, 30(4): 285-308.

Sabol, M. 1999. The fossil bears from the Okno cave (Demänová valley, Slovakia). Mineralia Slovaca, 31, 2, 87–109.

Sabol, M. 2000. Fosílne a subfosílne medveďovité mäsožravce (Ursidae, Carnivora) z územia Slovenska. Manuskript, Comenius University, Bratislava, 149 p.

Sabol, M. 2001. Geographical distribution of Cave Bears (Ursus spelaeus Rosenmüller et Heinroth, 1794) in the territory of Slovakia. Beiträge zur Paläontologie, 26,


Sabol, M. 2002a. Fossil findings of cave bears from the Upper Pleistocene sediments of selected caves in Slovakia. Mineralia Slovaca, 34, 1, 35–52.

Sabol, M. 2002b. Morfometrická analýza osteologických zvyškov medveďov jaskynných (Ursus spelaeus Rosenmüller et Heinroth, 1794) z depozitu Považského múzea

v Žiline. Vlastivedný zborník Považia, 21, 227–236.

Sabol, M. 2005. Cave Bears (Carnivora, Mammalia) from the Trojuholník Cave (Slovakia). Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften, Band 14: 161-176.

Sabol, Martin. (2017). A short history of cave bear research in Slovakia. Aragonit 22(1): 3-6.

Sabol, M. – Bendík, A. – Grivalský, M. – Grivalský, M. – Lizák, J. – Michlík, I. 2014. Latest and highest fossil record of cave bears (Ursus ex gr. spelaeus) in Slovakian Western Carpathians. Quaternary International, 339–340, 189–196.

Sabol, M. – Bendik, A. – Stuller, F. – Novomeský, F. – Necas, L. 2008a. A record of a three-legged cave bear female from the Cave of Izabela Textorisova (the Velka Fatra Mountains, northern Slovakia). Stalactite, 58, 2, 31–34.

Sabol, M. – Döppes, D. – Pacher, M. – Rabeder, G. – Withalm, G. 2008b. Cave bears from the Medvedia jaskyna in the Slovensky raj Mountains (Slovakia): preliminary results. Stalactite 58(2): 74-77.

Sabol, M. – Kozáková, E. 2007. Cave Bears from the Medvedia Cave in the Jánska Valley (the Low Tatras Mts., Northern Slovakia). Scripta Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis, Geology, 35, 97–102.

Sabol, M. – Kozáková, E. – Sliva, Ľ. 2009. Late Pleistocene cave bears (Ursus ex gr. spelaeus) from the Medvedia Cave in the Jánska Valley (the Low Tatras Mts., Slovakia). Slovenský kras, 47, Suppl. 1, 85–104. 

Sabol, M. – Struhár, V. 2002. Fosílne a subfosílne nálezy medveďov (Carnivora, Ursidae) z územia Liptova (severné Slovensko). Slovenský kras, 40, 49–88. 

Sabol, M. – Višňovská, Z. 2007. Cave Bears from the Važecká (Northern Slovakia) science and exhibitions. Scripta Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis, Geology, 35, 145–150. 

Sanchis, Alfred et al. (In Press, 2018). Neanderthal and carnivore activities at Llonin Cave, Asturias, northern Iberian Peninsula: Faunal study of Mousterian levels (MIS 3)Activités de Néandertaliens et carnivores dans la grotte de Llonin, Asturies, Nord de la péninsule Ibérique : étude de la faune des niveaux moustériens (MIS 3). Comptes Rendus Palevol. [Abstract]

Santi, Giuseppe and Rossi, Mario. (2020). Ursus gr. spelaeus from Grotta del Bandito (Piedmont, Northern Italy). 1. Morphodynamics of teeth and indications of an evolutionary step. Geologia Croatica 73(1): 1-12.

Santos, Elena, Gómez-Olivencia, Asier, Arlegi, Mikel and Arsuaga, Juan Luis. (In Press, 2015). Cranial morphological differences within [i]U. deningeri[/i] – [i]U. spelaeus[/i] lineage: A double traditional and geometric morphometrics approach. Quaternary International. doi:10.1016/j.quaint.2015.11.096 [Abstract]

Scheer, A. (2001). The utilisation of mammoth remains as raw material and its importance for the Gravettian people of the German Danube, pp. 455-459. In: Cavarretta, C., Gioia, P., Mussi, M. and Palombo, M. R. (eds.). The World of Elephants – International Congress, Rome. Proceedings of the 1st international congress – Consiglio nazionale delle ricerche, Rome. [automatic download]

Schmidt, Z. 1970. Výskyt a geografické rozšírenie medveďov (Ursinae) na území slovenských Karpát. Slovenský kras, 8, 7–20.

Schmidt, Z. – Chrapan, J. 1970. Datovanie osteologického materiálu fosílnych Ursidae metódou 14C z Medvedej jaskyne v Slovenskom raji. Slovenský kras, 8, 69–82. 

Spahni, J.-G. (1954). Les gisements à Ursus spelaeus de l'Autriche et leurs problèmes. Bulletin de la Société préhistorique de France 51 (7): 346-367.

SPÖTL, C., REIMER, P. J., RABEDER, G. and SCHOLZ, D. (2014). Presence of cave bears in western Austria before the onset of the Last Glacial Maximum: new radiocarbon dates and palaeoclimatic considerations. J. Quaternary Sci. 29: 760-766. doi: 10.1002/jqs.2747 [Abstract]

Spötl, Christoph et al. (2019). Cave bear occupation in Schwabenreith Cave, Austria, during the early last glacial: constraints from 230Th/U‐dated speleothems. Journal of Quaternary Science.

Stiller, M., Baryshnikov, G., Bocherens, H., Grandal d'Anglade, A.; Hilpert, B., Munzel, S.C., Pinhasi, R., Rabeder, G., Rosendahl, W., Trinkaus, E., Hofreiter, M., Knapp, M. (2010). Withering away – 25,000 years of genetic decline preceded cave bear extinction. Molecular Biology and Evolution 27(5): 975-978.

Stiller M., Molak M., Prost S., Pacher M., Rabeder G., Baryshnikov G., Rosendahl W., Muenzel S., Bocherens H., Grandal-d'Anglade A., Hilpert B., Germonpré M., Stasyk O., Pinhasi R., Ho S., Hofreiter M., Knapp M. (2014). Mitochondrial DNA diversity and evolution of the Pleistocene cave bear complex. Quaternary International 339-340: 224-231.

Štuller, F. – Novomeský, F. – Krajčovič, J. – Straka, Ľ. – Bendík, A. – Sabol, M. – Nečas, L. – Strecha, J. 2011. The interdisciplinary cooperation of forensic medicine, clinical medicine and palaeozoology: A case of cave bears (Ursus spelaeus) bones. Patologie a soudní lékařství, 54, 1, 39–41.

Terlato, Gabriele. (2019?). Fate of cave bear in northeastern italy: Anthropic impact, extinction chronology, stable isotope and genetic evidence. [Abstract]

Terlato, G.; Bocherens, H.; Romandini, M.; Nannini, N.; Hobson, K. A.; Peresani, M. (2018). Extinction and paleoecology of the Late Pleistocene cave bear from northeastern Italy: radiocarbon and stable isotope evidence. Quaternary: Past, Present, Future - AIQUA Conference. [Abstract]

Terlato, Gabriele, Bocherens, Hervé, Romandini, Matteo, Nannini, Nicola, Hobson, Keith A. and Peresani, Marco. (2019). Chronological and Isotopic data support a revision for the timing of cave bear extinction in Mediterranean Europe. Historical Biology 31(4): 474-484. [Abstract]

Toškan, Borut, Dirjec, Janez and Bavdek, Alma. (2014). Lost in time? Repatriated animal remains from Anelli’s excavations at Betalov spodmol (SW Slovenia). RMZ – M&G 61: 141-155.

Veitschegger, Kristof. (2017). The effect of body size evolution and ecology on encephalization in cave bears and extant relatives. BMC Evolutionary Biology 17: 124.

Veitschegger, Kristof et al. (2019). Longevity and life history of cave bears – a review and novel data from tooth cementum and relative emergence of permanent dentition. Historical Biology 31(4): 510-516. [Abstract]

Vlček, L. – Sabol, M. 2006. Vyhodnotenie nálezu fosílií medveďa jaskynného (Ursus spelaeus Rosenmüller, 1794) z jaskyne Teplica na Muránskej planine. Aragonit, 11, 23–25.

Vlček, L. – Sabol, M. – Kučerová, J. 2005. Správa o náleze osteologických zvyškov medveďa jaskynného (Ursus spelaeus Rosenmüller, 1794) z jaskyne Teplica na Muránskej planine. Reussia, 2, 2, 52–54. 

Weinstock, Jaco. (1999). The Upper Pleistocene mammalian faunafrom the Große Grotte near Blaubeuren (southwestern Germany). Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 277: 49pp., 13 figs., 32 tabs.

Wojtal, Piotr et al. (In Press, 2014). Gravettian hunting and exploitation of bears in Central Europe. Quaternary International. doi:10.1016/j.quaint.2014.10.017

Wojtal, Piotr et al. (2020). Carnivores in the everyday life of Gravettian hunters-gatherers in Central Europe. Journal of Anthropological Archaeology 59: 101171.

Zilio, Leandro et al. (2021). Examining Neanderthal and carnivore occupations of Teixoneres Cave (Moià, Barcelona, Spain) using archaeostratigraphic and intra-site spatial analysis. Sci Rep. 11: 4339.

Zunino, Marta et al. (2022). New insights into taphonomic analysis of the Upper Pleistocene Ursus spelaeus bone deposit from Bàsura cave (Toirano, NW Italy). Journal of Quaternary Science.


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