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Fruitbat's Pdf Library - Superfamily Ursoidea - 'bear-Dogs', Bears And Their Relatives

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These are a few of the pdf files (and a few Microsoft Word documents) that I've accumulated in my web browsing.

MOST of these are hyperlinked to their source.

If you want one that is not hyperlinked or if the link isn't working, e-mail me at joegallo1954@gmail.com and I'll be happy to send it to you.


Please note that this list will be updated continuously as I find more available resources. All of these files are freely available on the Internet so there should be no copyright issues.



Articles with author names in RED are new additions since May 12, 2018.



Order Carnivora


Superfamily Ursoidea


Family Amphicyonidae (†) - 'Bear Dogs'


Amphicyonidae - Africa/Middle East


Gurbuz, M. (1974). Amphicyon major Blainville Discovered in the Middle Miocene Beds of Candir. Mineral Research and Exploration Institute of Turkey.

Morales, J., M. Pickford and A. Valenciano (2016). Systematics of African Amphicyonidae, with descriptions of new material from Napak (Uganda) and Grillental (Namibia). Journal of Iberian Geology, 42(2).

Morales, J., P. Brewer and M. Pickford (2010). Carnivores (Creodonta and Carnivora) from the Basal Middle Miocene of Gebel Zelten, Libya, With a Note on a Large Amphicyonid from the Middle Miocene of Ngorora, Kenya. Bulletin of the Tethys Geological Society, Cairo, Vol.5.

Werdelin, L. and S.W. Simpson (2009). The last amphicyonid (Mammalia, Carnivora) in Africa. Geodiversitas, 31(4).


Amphicyonidae - Asia/Malaysia/Pacific Islands


Peigne, S., et al. (2006). A new amphicyonid (Mammalia, Carnivora, Amphicyonidae) from the late middle Miocene of northern Thailand and a review of the amphicyonine record in Asia. Journal of Asian Earth Sciences, 26.

Kohno, N. (1997). The first record of an amphicyonid (Mammalia: Carnivora) from Japan, and its implications for amphicyonid paleobiogeography. Paleontological Research, Vol.1, Number 4.

Viranta, S., S. Taseer Hussain and R.L Bernor (2004). The Anatomical Characteristics of Giant Miocene Amphicyonid (Carnivora) Humerus from Pakistan. Pakistan J.Zool., Vol. 36(1).

Wang, X.-M., J.-H. Wang and Q.-G. Jiangzuo (2016). New record of a haplocyonine amphicyonid in Early Miocene of Nei Mongol fills a long-suspected geographic hiatus. Vertebrata PalAsiatica, 54(1).

Zhai, R., et al. (2003). An aberrant amphicyonid mammal from the latest Eocene of the Bose Basin, Guangxi, China. Acta Palaeontologica Polonica, 48(2).


Amphicyonidae - Europe (including Greenland and Siberia)


De Bonis, L. (2015). Revival of a Species of the Rare European Oligocene Amphicyonid Goupilictis Ginsburg, 1969 (Mammalia, Carnivora, Amphicyonidae). Journal of Vertebrate Paleontology, e969401.

Pairó, M.C. and B. Kurtén (1976). Bears and Bear-Dogs from the Vallesian of of the Vallés-Penedés Basin, Spain. Acta Zoologica Fennica, 144.

Peigne, S. and E.P.J. Heizmann (2003). The Amphicyonidae (Mammalia: Carnivora) from Ulm-Westtangente (MN2, Early Miocene), Baden-Wurttemberg, Germany - Systematics and ecomorphology. Stuttgarter Beitr.Naturk., Ser.B, Number 343.

Peigne, S., et al. (2008). A New Amphicyonine (Carnivora: Amphicyonidae) from the Upper Miocene of Batallones-1, Madrid, Spain. Palaeontology, Vol.54, Part 4.

Siliceo, G., et al. (2014). Comparative Anatomy of the Shoulder Region in the Late Miocene Amphicyonid Magericyon anceps (Carnivora): Functional and Paleoecological Inferences. J.Mammal.Evol.

Viranta, S. (1996). European Miocene Amphicyonidae - taxonomy, systematics and ecology. Acta Zoologica Fennica, Number 204.


Amphicyonidae - North America


Berta, A. and H. Galiano (1984). A Miocene Amphicyonid (Mammalia: Carnivora) from the Bone Valley Formation of Florida. Journal of Vertebrate Paleongology, 41(1).

Boardman, G.S. and R.M. Hunt (2015). New material and evaluation of the chronostratigraphic position of Daphoenictis tedfordi (Mammalia, Carnivora, Amphicyonidae), a cat-like carnivoran from the latest Eocene of northwestern Nebraska, USA. Palaeontologia Electronica, 18.2.25A.

Boyd, C.A. and E. Welsh (2018). Biochronology and biogeography of Paradaphoenus (Carnivora: Amphicyonidae) within the Great Plains Region of North America. PeerJ Preprints. (Not peer reviewed)

Cook, H.J. A New Gigantic Fossil Dog from Colorado.

Hunt, R.M. (2011). Evolution of Large Carnivores During the Mid-Cenozoic of North America: the Temnocyonine Radiation (Mammalia, Amphicyonidae). Bulletin of the American Museum of Natural History, Number 358. (54 MB download).
Hunt, R.M. (2009). Long-legged pursuit carnivorans (Amphicyonidae, Daphoeninae) from the early Miocene of North America. Bulletin of the American Museum of Natural History, Number 318. (Note: This is a 20MB download)

Hunt, R.M. (2003). Intercontinental Migration of Large Mammalian Carnivores: Earliest Occurrence of the Old World Beardog Amphicyon (Carnivora, Amphicyonidae) in North America. Bulletin American Museum of Natural History, Number 279, Chapter 4.
Hunt, R.M. (2002). New Amphicyonid Carnivorans (Mammalia, Daphoeninae) from the Early Miocene of Southeastern Wyoming. American Museum Novitates, Number 3385.

Hunt, R.M. (2002). Intercontinental Migration of Neogene Amphicyonids (Mammalia, Carnivora): Appearance of the Eurasian Beardog Ysengrinia in North America. American Museum Novitates, Number 3384.
Hunt, R.M. (2001). Small Oligocene Amphicyonids from North America (Paradaphoenus, Mammalia, Carnivora). American Museum Novitates, Number 3331.

Hunt, R.M. (1972). Miocene Amphicyonids (Mammalia, Carnivora) from the Agate Spring Quarries, Sioux County, Nebraska. American Museum Novitates, Number 2506.

McGrew, P.O. (1939). A New Amphicyon from the Deep River Miocene. Geological Series of Field Museum of Natural History, Vol.VI, Number 23.

Tomiya, S. and Z.J. Tseng (2016). Whence the beardogs? Reappraisal of the Middle to Late Eocene 'Miacis' from Texas, USA, and the origin of Amphicyonidae (Mammalia, Carnivora). R.Soc. open sci., 3.


General Amphicyonidae


Figueirido, B., et al. (2011). Body mass estimation in amphicyonid carnivoran mammals: A multiple regression approach from the skull and skeleton. Acta Palaeontologica Polonica, 56(2).

Radinsky, L. (1980) Endocasts of Amphicyonid Carnivorans.American Museum Novitates, Number 2694.

Sorkin, B. (2006). Ecomorphology of the giant bear-dogs Amphicyon and Ischyrocyon. Historical Geology, 18(4).

Family Ursidae - The Bears and Their Allies.


Subfamily Agriotheriinae (†)


Hendey, Q.B. (1977). Fossil Bear from South Africa. South African Journal of Science, Vol.73.

Martin, J.E. (2013). A Late Occurrence of the Bear Agriotherium from the Blancan Ringold Formation of Southeastern Washington. Proceedings of the South Dakota Academy of Science, Vol.92.

Miller, W.E. and O. Carranza-Castaneda (1996). Agriotherium schneideri from the Hemphillian of Central Mexico. Journal of Mammalogy, 77(2).

Ogino, S., et al. (2011). New species of Agriotherium (Mammalia, Carnivora) from the late Miocene to early Pliocene of central Myanmar. Journal of Asian Earth Sciences, 42.

Oldfield, C.C., et al. (2012). Finite element analysis of ursid cranial mechanics and the prediction of feeding behaviour in the extinct giant Agriotherium africanum. Journal of Zoology, 286.

Samuels, J.X., J.A. Meachen-Samuels and P.A. Gensler (2009). The First Mid-Blancan Occurrence of Agriotherium (Ursidae) in North America: A Record from Hagerman Fossil Beds National Monument, Idaho. J.Paleont., 83(4).

Sorkin, B. (2006). Ecomorphology of the giant short-faced bears Agriotherium and Arctodus. Historical Biology, 8(1).

Stach, J. (1957). Agriotherium intermedium N.Sp. from the Pliocene Bone Breccia of Weze. Acta Palaeontologica Polonica, Vol.II, Number 1.


Subfamily Amphicynodontinae (†)


Clark, J. and T.E. Guensburg (1972). Arctoid Genetic Characters as Related to the Genus Parictis. Fieldiana Geology, Vol.26, Number 1.


Subfamily Ailuropodinae - Giant Pandas and their relatives


Abella, J., P. Montoya and J. Morales (2011). A New species of Agriarctos (Ailuropodinae, Ursidae, Carnivora) in the locality of Nombrevilla 2 (Zaragoza, Spain). Estudios Geológicos, 67(2).

Abella, J., et al. (2012). Kretzoiarctos gen.nov., the Oldest Member of the Giant Panda Clade. PLoS ONE, 7(11).

Baryshnikov, G.F. and P.A. Tleuberdina (2017). Late Miocene Indarctos (Carnivora: Ursidae) from the Karabulak Formation of the Kalmakpai River (Zaisan Depression, Eastern Kazakhstan). Proceedings of the Zoological Institute RAS, Vol.321, Number 1.
Beninda-Emonds, O.R.P. (2004). Phylogenetic Position of the Giant Panda. In: Giant Pandas: Biology and Conservation. Lindburg, D.G. and K. Baragona (eds.), University of California Press, Berkeley, CA.

de Bonis, L., et al. (2017). A new late Miocene ailuripodine (Giant Panda) from Rudabanya (North-central Hungary). Geobios, 2017 (accepted manuscript)

D*ng, W. (2008). Virtual cranial endocast of the oldest giant panda (Ailuropoda microta) reveals great similarity to that of its extant relative. Naturwissenschaften, 95.

D*ng, W. and J.-F. Zhang (2011). Evolution of Cranial Cavities in Giant Pandas (Ailuropoda, Carnivora, Mammalia). Vertebrata PalAsiatica, 49(2).

Figueirido, B., et al. (2011). Cranial shape transformation in the evolution of the giant panda (Ailuropoda melanoleuca). Naturwissenschaften, 98.

Gregory, W.K. (1936). On the Phylogenetic Relationships of the Giant Panda (Ailuropoda) to Other Arctoid Carnivora. American Museum Novitates, Number 878.

Jin, C, et al. (2007). The first skull of the earliest giant panda. PNAS, Vol.104, Number 26.

Qiu, Z.-X. and R.H. Tedford (2003). A New Species of Indarctos from Baode, China. Vertebrata PalAsiatica, 41(4).

Qiu, Z.-X. and G. Qi (1989). Ailuropod Found from the Late Miocene Deposits in Lufeng, Yunnan. Vertebrata PalAsiatica, 27(3).

Roussiakis, S.J. (2001). Postcranial remains of Indarctos atticus (Ursidae, Mammalia) from the classical locality of Pikermi (Attica, Greece), with a description of the front limb. Senckenbergiana lethaea, 81(2).

Tougard, C., et al. (1996). Extension of the geographic distribution of the giant panda (Ailuropoda) and search for the reasons for its progressive disappearance in Southeast Asia during the Latest Middle Pleistocene. C.R.Acad.Sci. Paris, Vol.323.

Wang, L. and M. Wu (1976). A Dental Anomaly of Ailuropoda melanoleuca baconi. Vertebrata PalAsiatica, 14(4).

Zhou, Z. and Y.-h. Li (1987). The Ultrastructure of the Enamel in the Giant Panda of Pleistocene. Vertebrata PalAsiatica, 25(4).


Subfamily Tremarctinae - Short-Faced Bears and Spectacled Bears


Tremarctinae - North America


Emslie, S.D. (1995). The Fossil Record of Arctodus pristinus (Ursidae: Tremarctinae) in Florida. Bulletin of the Florida Museum of Natural History, Vol.37, Part II, Number 15.

Emslie, S.D. and N.J. Czaplewski (1985). A New Record of Giant Short-faced Bear, Arctodus simus, from Western North America With a Re-evaluation of its Paleobiology. Contributions in Science, Natural History Museum of Los Angeles County, Number 371.

Gillette, D.D. and D.B. Madsen (1992). The Short-Faced Bear Arctodus simus from the Late Quaternary in the Wasatch Mountains of Central Utah. Journal of Vertebrate Paleontology, 12(1).

Kurtén, B. (1963). Fossil Bears from Texas. The Pearce-Sellards Series - Texas Memorial Museum, Number 1.

Kurtén, B. and E. Anderson (1974). Association of Ursus arctos and Arctodus simus (Mammalia: Ursidae) in the Late Pleistocene of Wyoming. Breviora, Number 426.

Puckette, W.L. (1976). Notes on the Occurrence of the Short-Faced Bear (Arctodus) in Oklahoma.Proc. Okla. Acad. Sci., 56.

Richards, R.L. and W.D. Turnbull (1995). Giant Short-faced Bear (Arctodus simus yukonensis) Remains from Fulton County, Northern Indiana. Fieldiana Geology, New Series Number 30.

Rinker, G.C. (1949). Tremarctotherium from the Pleistocene of Meade County, Kansas. Contributions from the Museum of Paleontology - University of Michigan, Vol. VII, Number 6.

Schubert, B.W. (2004) A Full-Glacial Short-Faced Bear (Arctodus simus) from Perkins Cave, Missouri. CRP, 21.

Schubert, B.W. and S.C. Wallace (2009). Late Pleistocene short-faced bears, mammoths, and large carcass scavenging in the Saltville Valley of Virginia, USA. Boreas, Vol.38.
Schubert, B.W. and J.E. Kaufmann (2003). A Partial Short-faced Bear Skeleton from an Ozark Cave with Comments on the Paleobiology of the Species.Journal of Cave and Karst Studies, 65(2).

Schubert, B.W., et al. (2010). Giant Short-Faced Bears (Arctodus simus) In Pleistocene Florida, USA, A Substantial Range Extension. J. Paleont., 84(1).

Scott, E. and S.M. Cox (1993). Arctodus simus (Cope, 1879) from Riverside County, California. PaleoBios, 15(2).


Tremarctinae - South America/Central America/Caribbean


Arnuado, M.E., et al. (2013). First Description of the Auditory Region of a Tremarctine (Ursidae, Mammalia) Bear: The Case of Arctotherium angustidens. J.Mammal.Evol.

Garcia Lopez, D.A., et al. (2008). First Record of Arctotherium (Ursidae, Tremarctinae) in Northwestern Argentina and Its Paleobiogeographic Significance. Journal of Vertebrate Paleontology, 28(4).

Mendoza, P.L., F.M. Larrain and E. Bostelmann (2015). Presence of Arctotherium (Carnivora, Ursidae, Tremarctinae) in a pre-cultural level of Bano Nuevo-1 cave (Central Patagonia, Chile). Estudios Geologicos, 71(2).

Perez-Crespo, V.A., et al. (2016). Carbon and Oxygen Isotopic Values for a Short Faced Bear Individual (Arctodus simus) from Cedral, San Luis Potosi, Mexico. ICBS Proceedings.

Prevosti, F.J., et al. (2003). The Southernmost Bear: Pararctotherium (Carnivora, Ursidae, Tremarctinae) in the Latest Pleistocene of Southern Patagonia, Chile. Journal of Vertebrate Paleontology, 23(3).

Soibelzon, L.H. and B.W. Schubert (2011). The Largest Known Bear, Arctotherium angustidens, from the Early Pleistocene Pampean Region of Argentina: With a Discussion of Size and Diet Trends in Bears. Journal of Paleontology, 85(1).

Soibelzon, L.H. and A.D. Rincon (2007). The fossil record of the short-faced bears (Ursidae, Tremarctinae) from Venezuela. Systematic, biogeographic and paleoecological implications. N.Jb.Geol.Palaont.Abh., Vol. 244/3.

Soibelzon, L.H. and A. Carlini (2004). Deciduous teeth morphology of some tremarctines (Ursidae, Tremarctinae). Descriptions, comparisons and possible phylogenetic implications. Ameghiniana, 41(2).

Soibelzon, L.H., E.P. Tonni and M. Bond (2005). The fossil record of South American short-faced bears (Ursidae, Tremarctinae). Journal of South American Earth Sciences, 20.

Soibelzon, L.H., et al. (2014). South American Giant Short-Faced Bear (Arctotherium angustidens) Diet: Evidence from Pathology, Morphology, Stable Isotopes, and Biomechanics. Journal of Paleontology, 88(6).

Soibelzon, L.H., et al. (2009). First report of a South American short-faced bears' den (Arctotherium angustidens): palaeobiological and palaeoecological implications. Alcheringa, 33.

Soibelzon, L.H., et al. (2008). A Blancan (Pliocene) short-faced bear from El Salvador and its implications for Tremarctines in South America. N.Jb.Geol.Paläont.Abh., Vol.250/1.

Stucci, M., et al. (2009). A 6000+ year-old specimen of a spectacled bear from an Andean cave in Peru. Ursus, 20(1).

Trajano, E. and H. Ferrarezzi (1994). A Fossil Bear from Northeastern Brazil, With a Phylogenetic Analysis of the South American Extinct Tremarctinae (Ursidae).Journal of Vertebrate Paleontology, 14(4).


General Tremarctinae


Christiansen, P. (1999). What size were Arctodus simus and Ursus spelaeus (Carnivora, Ursidae)? Ann.Zool. Fennici, 36.

Donohue, S.L., et al. (2013). Was the Giant Short-Faced Bear a Hyper-Scavenger? A New Approach to the Dietary Study of Ursids Using Dental Microwear Textures. PLoS ONE, 8(10).

Figueirido, B. and L.H. Soibelzon (2009). Inferring palaeoecology in extinct tremarctine bears (Carinvora, Ursidae) using geometric morphometrics. Lethaia.

Figueirido, B., et al. (2010). Demythologizing Arctodus simus, the 'Short Faced' Long Legged and Predaceous Bear that Never Was. Journal of Vertebrate Paleontology, 30(1).

Lynch, E.R. (2012). Cursorial Adaptations in the Forelimb of the Giant Short-Faced Bear, Arctodus simus, Revealed by Traditional and 3D Landmark Morphometrics. Masters Thesis - East Tennessee State University.

Matheus, P.E. (2003). Locomotor Adaptations and Ecomorphology of Short-Faced Bears (Arctodus simus) in Eastern Beringia. Palaeontology Program, Government of Yukon, Occasional Papers in Earth Sciences, Number 7.

Mitchell, K.J., et al. (2016). Ancient mitochondrial DNA reveals convergent evolution of giant short-faced bears (Tremarctinae) in North and South America. Biol.Lett., 12.

Salesa, M.J., et al. (2006). Anatomy of the "false thumb" of Tremarctos ornatus (Carnivora, Ursidae, Tremarctinae): phylogenetic and funtional implications. Estudios Geológicos, 62(1).

Sorkin, B. (2006). Ecomorphology of the giant short-faced bears Agriotherium and Arctodus. Historical Biology, 8(1).


Subfamily Ursavinae (†)


Qiu, Z.-X., T. Deng and B.-Y. Wang (2014). A Late Miocene Ursavus skull from Guanghe, Gansu, China. Vertebrata PalAsiatica, 52(3).

Subfamily Ursinae - Bears


Ursinae - Africa/Middle East


Stiner, M.C., G. Arsebuk and F.C. Howell (1996). Cave Bears and Paleolithic Artifacts in Yarimburgaz Cave, Turkey: Dissecting a Palimpsest. Geoarchaeology: An International Journal, Vol.11, Number 4.

Stiner, M.C., et al. (1998). Reconstructing cave bear paleoecology from skeletons: a cross-disciplinary study of middle Pleistocene bears from Yarimburgaz Cave, Turkey. Paleobiology, 74(1).


Ursinae - Asia/Malaysia/Pacific Islands


Baryshnikov, G.E. and A.F. Lavrov (2015). Early Miocene Bear Ballusia (Carnivora, Ursidae) from the Locality Khirgis-Nur-I in Mongolia. Proceedings of the Zoological Institute RAS, Vol.319, Number 3.

Baryshnikov, G.F. and A.V. Lavrov (2013). Pliocene bear Ursus minimus Deveze de Chabriol et Bouillet, 1827 (Carnivora, Ursidae) in Russia and Kazakhstan. Russian J.Theriol., 12(2).

Knapp, M., et al. (2009). First DNA sequences from Asian cave bear fossils reveal deep divergences and complex phylogeographic patterns. Molecular Ecology, 18.

Lan, T., et al. (2017). Evolutionary history of enigmatic bears in the Tibetan Plateau - Himalaya region and the identity of the yeti. Proc.R.Soc. B, 284. (Thanks to Oxytropidoceras for locating this one!)

Qiu, Z.-X., T.Deng and B.-Y. Wang (2009). First Ursine Bear Material from Dongxiang, Gansu - Addition to the Longdan Mammalian Fauna (2). Vertebrata PalAsiatica, 47(4).
Sykes, B.C., et al. (2014). Genetic analysis of hair samples attributed to yeti, bigfoot and other anomalous primates. Proc.R.Soc. B, 281. (Thanks to Oxytropidoceras for pointing this one out!)


Ursinae - Europe (including Greenland and Siberia)


Abella, J., et al. (2013). On the Socio-Sexual Behaviour of the Extinct Ursid Indarctos arctoides: An Approach Based on Its Baculum Size and Morphology. PLoS ONE, 8(9).

Ábelová, M. (2007). Paleodiet Inferred from Ursus spelaeus (Rosenmüller et Heinroth) Tooth from Tmavá Cave (Slovak Republic) Using Carbon Isotope Analyses. Acta Mus. Moraviae, Sci.geol., LXXXXII.

Andrews, P. and A. Turner (1992). Life and death of the Westbury bears. Acta. Zool. Fennici, 28.

Argenti, P. and P.P.A. Mazza (2006). Mortality analysis of the Late Pleistocene bears from Grotta Lattaia, central Italy. Journal of Archaeological Science, 33.

Baca, M., et al. (2013). Ancient DNA and dating of cave bear remains from Niedwiedzia Cave suggest early appearance of Ursus ingressus in Sudetes. Quaternary International, xxx. (Article in press)

Baryshnikov, G.F. (2010). Late Pleistocene brown bear (Ursus arctos) from the Caucasus. Russ.J.Theriol., 9(1).

Baryshnikov, G.F. (2010). Middle Pleistocene Ursus thibetanus (Mammalia, Carnivora) from Kudaro Caves in the Caucasus. Proceedings of the Zoological Institute RAS, Vol.314, Number 1.

Baryshnikov, G.F. (2002). Late Miocene Indarctos punjabicus atticus (Carnivora, Ursidae) in Ukraine, with survey of Indarctos records from the former USSR. Russian J. Theriol., 1(2).

Baryshnikov, G.F. (2000). Late Pleistocene cave bear (Ursus deningeri kudarensis) from the Akhstyrskaya Cave in the Caucasus (Russia). Beitr.Palaont., 25.

Baryshnikov, G.F. (1998). Cave Bears from the Paleolithic of the Greater Caucasus. Zoological Institute RAS.

Baryshnikov, G.F. and A.V. Lavrov (2013). Pliocene bear Ursus minimus Deveze de Chabriol et Bouillet, 1827 (Carnivora, Ursidae) in Russia and Kazakhstan. Russian J.Theriol., 12(2).

Baryshnikov, G.F. and D.S. Zakharov (2013). Early Pliocene Bear Ursus thibetanus (Mammalia, Carnivora) from Priozernoe Locality in the Dniester Basin (Moldova Republic). Proceedings of the Zoological Institute RAS, Vol.317, Number 1.

Baryshnikov, G.F. and G.G. Boeskorov (2004). Skull of the Pleistocene brown bear (Ursus arctos) from Yakutia, Russia. Russian J.Theriol., 3(2).

Baryshnikov, G.F. and I. Foronova (2001). Pleistocene small cave bear (Ursus rossicus) from the South Siberia, Russia. Cadernos Lab. Xeoloxico de Laxe, Vol.26.

Baryshnikov, G.F., M. Germonpré and S.V. Baryshnikova (2010). Cave bear (Ursus spelaeus) from Chamber B of the Goyet Cave in Belgium. Russian Journal of Theriology, 9(2).

Bocherens, H., et al. (2013). The last of its kind? Radiocarbon, ancient DNA and stable isotope evidence from a late cave bear (Ursus spelaeus Rosenmuller, 1794) from Rochedane, France. Quaternary International, xxx. (Article in press)

Bocherens, H., 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.

Bon, C., et al. (2008). Deciphering the complete mitochondrial genome and phylogeny of the extinct cave bear in the Paleolithic Painted Cave of Chauvet. PNAS, Vol.105, Number 45.

de Carlis, A., et al. (2005). Morphometry of Ursus spelaeus Remains from Valstrona (Northern Italy). Geo. Alp, Vol.2, S.

De Torres Perezhidalgo, T. (1992). The European descendants of Ursus etruscus C.Cuvier (Mammalia, Carnivora, Ursidae). Boletino Geologico y Minero, Vol. 103-4.
Debeljak, I. (2007). Fossil Population Structure and Mortality of the Cave Bear from the Mokrica Cave (North Slovenia). Acta Carsologica, 36/3.

Diedrich, C.G. (2013). Extinction of Late Ice Age Cave Bears as a Result of Climate/Habitat Change and Large Carnivore Lion/Hyena/Wolf Predation Stress in Europe. Hindawi Publishing Corporation ISRN Zoology, Vol.2013, Article ID 138319.

Diedrich, C.G. (2009). The Rediscovered Cave Bear "Ursus spelaeus Rosenmüller 1794" Holotype of the Zoolithin Cave (Germany) from the Historic Rosenmüller Collection. Slovenský Kras Acta Carsologica Slovaca, 47, Suppl. 1.

Diedrich, C.G. (2006). Cave Bear Open Air Site Remains and Den Caves from the Upper Pleistocene of Central Bohemia (Czech Republic). Scientific Annals, School of Geology, Aristotle University of Thessaloniki, Special Volume 98.

Edwards, C.J., et al. (2011). Ancient Hybridization and an Irish Origin for the Modern Polar Bear Matriline. Current Biology, 21.

Garcia, N. (2004). New results on the remains of Ursidae from Untermassfeld: comparisons with Ursus dolinensis from Atapuerca and other Early and Middle Pleistocene sites. 18th International Senckenberg Conference in Weimar.

Garcia, N., et al. (2007). Endocranial Morphology of the Ursus deningeri von Reichenau 1904 from the Sima de los Huesos (Sierra de Atapuerca) Middle Pleistocene Site.  Journal of Vertebrate Paleontology, 27(4).

Grandal-d'Anglade, A. (2010). Bite force of the extinct Pleistocene Cave bear Ursus spelaeus  Rosenmüller from Europe. C.R. Palevol, 9.

Grandal-d'Anglade, A. and F. López-González (2005). Sexual dimorphism and ontogenetic variation in the skull of the cave bear (Ursus spelaeus Rosenmüller) of the European Upper Pleistocene. Geobios, 38.

Grandal-d'Anglade, A. and F. López-González (2004). A Study of the Evolution of the Pleistocene Cave Bear by a Morphometric Analysis of the Lower Carnassial. Oryctos, Vol.5.

Ingólfsson, Ó and Ø. Wiig (2008). Late Pleistocene fossil find in Svalbard: the oldest remains of a polar bear (Ursus maritimus Phipps, 1744) ever discovered. Polar Research, 2008.

Jambresic, G. and M. Paunovic (2002). Osteometry, Variability, Biomechanics, and Locomotion Pattern of the Cave Bear Limb Bones from Croatian Localities. Geologia Croatica, 55(1).

Jones, D.B. and L.R.G. DeSantis (2016). Dietary ecology of the extinct cave bear: Evidence of omnivory as inferred from dental microwear textures. Acta Palaeontologica Polonica, 61(4). 

Kavcik-Graumann, N., et al. (2016). The Bears of Illinka Cave Near Odessa (Ukraine). ICBS Proceedings.

Kosintsev, P.A. and O.P. Bachura (2015). A Mass Burial of Brown Bears (Ursus arctos L., 1758) from the Upper Pleistocene of the Northern Urals. Doklady Biological Sciences, Vol.462.

Kosintsev, P.A., et al. (2016). The First Finding of Asian Black Bear (Carnivora, Ursidae, Ursus (Euarctos) thibetanus G. Cuvier, 1823) in the Late Pleistocene of Northern Eurasia. Doklady Biological Sciences, Vol.471.

Kostopoulos, D.S. and K. Vasileiadou (2006). The Greek Late Neogene-Quaternary Ursids in Relation to Palaeogeography and Palaeoenvironment.Scientific Annals School of Geology, Aristotle University of Thessaloniki, Special Volume 98.

Marciszak, A., et al. (2015). Ursus arctos L.,1758 from Bukovynka Cave (W Ukraine) in an overview on the fossil brown bears size variability based on cranial material. Quaternary International, 357.

Martini, I., et al. (2013). The latest Ursus spelaeus in Italy, a new contribution to the extinction chronology of the cave bear. Quaternary Research, xxx. (Article in press)

McFarlane, D.A., M. Sabol and J. Lundberg (2010). A Unique Population of Cave Bears (Carnivora: Ursidae) from the Middle Pleistocene of Kents Cavern, England, Based on Dental Morphometrics. W.M. Keck Science Faculty Papers, Paper 77.

Medin, T., et al. (2015). Late Villafranchian Ursus etruscus and other large carnivorans from the Orce sites (Gaudix-Baza basin, Andalusia, southern Spain): Taxonomy, biochronology, paleobiology and ecogeographical context. Quaternary International, xxx. (Article in press)

Montoya, P., et al. (2001). Indarctos (Ursidae, Mammalia) from the Spanish Turolian (Upper Miocene). Scripta Geol., 122.

Munzel, S.C. and N.J. Conard (2004). Cave Bear Hunting in the Hohle Fels, a Cave Site in the Ach Valley, Swabian Jura. Revue de Paleobiologie, Geneve, 23(2).

Munzel, S.C., et al. (2014). Behavioural ecology of Late Pleistocene bears (Ursus spelaeus, Ursus ingressus): Insight from stable isotopes (C,N,O) and tooth microwear. Quaternary International, 339-340.

Nowakowski, D. (2018). Frequency of appearance of transverse (Harris) lines reflects living conditions of the Pleistocene bear - Ursus ingressus - (Sudety Mts., Poland). PLoS ONE, 13(4).

Pairó, M.C. and B. Kurtén (1976). Bears and Bear-Dogs from the Vallesian of of the Vallés-Penedés Basin, Spain. Acta Zoologica Fennica, 144.

Peigné, S., et al. (2009). Predormancy omnivory in European cave bears evidenced by a dental microwear analysis of Ursus spelaeus from Goyet, Belgium. PNAS, Vol.106, Number 36.

Perez-Hidalgo, T. De Torres (1992). The European Descendants of Ursus etruscus C. Cuvier (Mammalia, Carnivora, Ursidae). Boletin Geologico y Minero, Vol.103-4.

Quiles, J., et al. (2006). Cave bears (Ursus spelaeus) from the Pestera cu Oase (Banat, Romania): Paleobiology and taphonomy. C.R. Palevol, 5.
Rabeder, G., et al. (2008). Morphological responses of cave bears (Ursus spelaeus group) to high-alpine habitats. Die Hohle, 59.
Rabeder, G., et al. (2004). New Taxa of Alpine Cave Bears (Ursidae, Carnivora). Cahiers scientifiques/Hors serie number 2.

Richards, M.P., et al. (2008). Isotopic evidence for omnivory among European cave bears: Late Pleistocene Ursus spelaeus from the Pestera cu Oase, Romania. PNAS, Vol.105, Number 2.

Robu, M. (2016). The assessment of the internal architecture of an MIS 3 cave bear bone assemblage. Case study: Ursilor Cave, Western Carpathians, Romania. Palaeogeography, Palaeoclimatology, Palaeoecology, 444.

Robu, M. (2016). Fossil population structure and mortality analysis of the cave bears from Ursilor Cave, north-western Romania. Acta Palaeontologica Polonica, 61(2).
Rosendahl, W. and D. Doppes (2006). Trace Fossils from Bears in Caves of Germany and Austria. Scientific Annals, School of Geology, Aristotle University of Thessaloniki, Special Volume 98.
Rosendahl, W., et al. (2005). New radiometric datings of different cave bear sites in Germany - results and interpretations. Bull.Soc.Hist.Nat., Toulouse, 141-1.

Rossi, M. and G. Santi (2015). Observations on the Ursus gr. spelaeus remains from the Pocala cave (Trieste, Friuli Venezia Giulia, N. Italy). Revue de Paleobiologie, Geneve, 34(1).

Saarma, U., et al. (2007). Mitogenomic structure of brown bears (Ursus arctos L.) in northeastern Europe and a new time frame for the formation of European brown bear lineages. Molecular Ecology, 16.

Sabol, M. (2001). Geographical Distribution of Cave Bears (Ursus spelaeus Rosenmüller et Heinroth, 1794) in the Territory of Slovakia. Beitr.Palaont., 26.

Sabol, M. (2001). Fossil Brown Bears of Slovakia. Cedernos Lab. Xeoloxico de Laxe, Vol.26.

Santi, G. and M. Rossi (2005). Ursus spelaeus from the Buco dell'Orso Cave (Laglio, Lombardy, North Italy): an evolutionary hypothesis. PalArch, 3(3).

Spassov, N., et al. (2017). First Record of the "Small Cave Bear" in Bulgaria and the Taxonomic Status of Bears of the Ursus savini Andrews - Ursus rossicus Borissiak Group. Fossil Imprint, Vol.73, Numbers 3-4.

Spotl, C., et al. (2014). Presence of cave bears in western Austria before the onset of the Last Glacial Maximum: new radiocarbon dates and palaeoclimatic considerations. Journal of Quaternary Science, 29(8).

Torres, T., et al. (2007). Hominid exploitation of the environment and cave bear populations. The case of Ursus spelaeus Rosenmüller-Heinroth in Amutxate cave (Aralar, Navarra-Spain). Journal of Human Evolution, 52(1).

Toskan, B. (2006). Cave Bear Metapodials from Divje Babe I (Western Slovenia). Scientific Annals, School of Geology Aristotle University of Thessaloniki, Special Volume 98.

Tsoukala, E., G. Rabeder and S. Verginis (2001). Ursus spelaeus and Late Pleistocene associated faunal remains from Loutraki (Pella, Macedonia, Greece) - Excavations of 1999. Cadernos Lab. Xeoloxico de Laxe, Vol.26.

Vercoutere, C., C. San Juan-Foucher and P. Foucher (2006). Human Modifications on Cave Bear Bones from the Gargas Cave (Haute-Pyrenees, France). Scientific Annals, School of Geology, Aristotle University of Thessaloniki, Special Volume 98.

Villaluenga, A., et al. (2012). Cave Bear (Ursus spelaeus Rosenmüller Heinroth, 1794) and Humans During the Early Upper Pleistocene (Lower and Middle Palaeolithic) in Lezetxici , Lezetxici II and Astigarragako Kobea (Basque Country, Spain). Preliminary Approach. Journal of Taphonomy, Vol.10, Issue 3-4.

Viranta, S. (2004). Habitat preferences of European Middle Miocene omnivorous ursids. Acta Paleontologica Polonica, 49(2).

Viranta, S. and A. Grandal-d'Anglade (2012). Late Pleistocene Large Mammal Paleocommunities: A Comparative Study Between Localities with Brown Bear (Ursus arctos), Cave Bear (U. spelaeus) and Mousterian Lithic Assemblage. Journal of Taphonomy, Vol.10 (Issue 3-4).

Wagner, J. (2010). Pliocene to early Middle Pleistocene ursine bears in Europe: a taxonomic overview. Journal of the National Museum (Prague), Natural History Series, Vol.179(20).

Wagner, J. (2006). A List of Craniodental Material of Pliocene Ursids (Genus Ursus) in the Collection of Naturhistorisches Museum Basel. Scientific Annals, School of Geology, Aristotle University of Thessaloniki, Special Volume 98.

Wagner, J. and M. Gasparik (2014). Research history of Pleistocene faunas in Gombasek quarry (Slovakia), with comments to the type specimen and the type locality of Ursus deningeri gombaszogenis Kretzoi, 1938. Fragmenta Palaeontologica Hungarica, Vol.31.

Wagner, J. and S. Ҫermák (2012). Revision of the early Middle Pleistocene bears (Ursidae, Mammalia) of Central Europe, with special respect to possible co-occurrence of spelaeoid and arctoid lineages. Bulletin of Geosciences, 87(3).

Wagner, J., S. Cermák and I. Horacek (2011). The Presence of Ursus ex gr. minimus-thibetanus in the Late Villányian and Its Position Among the Pliocene and Pleistocene Black Bears in Europe. Quaternaire, Hors-serie, 4.

Weinstock, J. (2000). Cave Bears from Southern Germany: sex Ratios and Age Structure. A Contribution Toward a Better Understanding of the Palaeobiology of Ursus spelaeus. Archaeofauna, 9.


Ursinae - North America


Czaplewski, N.J. and W.L. Puckette (2014). Late Pleistocene Remains of an American Black Bear (Ursus americanus) and Two Small Vertebrates from an Oklahoma Ozark Cave. Bull.Okla.Acad.Sci., 94.

Czaplewski, N.J. and S. Willsey (2013). Late Quaternary Brown Bear (Ursidae: Ursus cf. arctos) From a Cave in the Huachuca Mountains, Arizona. USDA Forest Service Proceedings RMRS-P-67.

Harington, C.R., et al. (2014). Brown bear (Ursus arctos) (9880 +/- 35 BP) from late-glacial Champlain Sea deposits at Saint-Nicholas, Quebec, Canada, and the dispersal history of brown bears. Can.J. Earth Sci., 51.

Herrero, S. Aspects of Evolution and Adaptation in American Black Bears (Ursus americanus Pallas) and Brown and Grizzly Bears (U. arctos Linné.) of North America. Panel 4. Bear Behavior. Ursus, Vol.2.

Kurtén, B. (1963). Fossil Bears from Texas. The Pearce-Sellards Series - Texas Memorial Museum, Number 1.

Kurtén, B. (1960). A Skull of the Grizzly Bear (Ursus arctos L.) from Pit 10, Rancho la Brea. Los Angeles County Museum - Contributions in Science, Number 39.

Kurtén, B. and E. Anderson (1974). Association of Ursus arctos and Arctodus simus (Mammalia: Ursidae) in the Late Pleistocene of Wyoming. Breviora, Number 426.

Matheus, P., et al. (2004). Pleistocene Brown Bears in the Mid-Continent of North America. Science (Breviora), Vol.306.

McClaren, D., et al. (2005). Bear Hunting at the Pleistocene/Holocene Transition of the Northern Northwest Coast of North America. Canadian Zooarchaeology, Number 22. (MS Word document)

Mustoe, G.E. and C.A. Carlstad (1995). A Late Pleistocene Brown Bear (Ursus arctos) from Northwest Washington. Northwest Science, Vol.69, Number 2.

Samuels, J.X., J.A. Meachen-Samuels and P.A. Gensler (2009). The First Mid-Blancan Occurrence of Agriotherium (Ursidae) in North America: A Record from Hagerman Fossil Beds National Monument, Idaho. J.Paleont.,83(4).

Stangl, F.B., A.C. Evans and R.D. Bradley (2014). Comments on Late Quaternary Ursids from the Texas/Oklahoma Southern Plain, with Documentation of the Last Known Native Black Bear (Ursus americanus) from the Texas Hill Country. Museum of Texas Tech University, Occasional Papers Number 321.

Wang, X., et al. (2017). A basal ursine bear (Protarctos abstrusus) from the Pliocene High Arctic reveals Eurasian affinities and a diet rich in fermentable sugars. Scientific Reports, 7: 17722.

Wolverton, S. and R.L. Lyman (1998). Measuring Late Quaternary Ursid Diminution in the Midwest. Quaternary Research, 49.

Wooding, S. and R. Ward (1997). Phylogeography and Pleistocene Evolution in the North American Black Bear. Mol.Biol.Evol., 14(11).


General Ursinae


Baca, M., et al. (2016). Retreat and extinction of the Late Pleistocene cave bear (Ursus spelaeus sensu lato). Sci.Nat., 103(92).

Barnes, I., et al. (2002). Dynamics of Pleistocene Population Extinctions in Beringian Brown Bears. Science, Vol.295.

Christiansen, P. (1999). What size were Arctodus simus and Ursus spelaeus (Carnivora, Ursidae)?Ann.Zool. Fennici, 36.

Debeljak, I. (1996). Ontogenetic development of dentition in the cave bear. Geologija, 39.

Fernández-Mosquera, D., M. Vila-Taboada and A. Grandal-d'Anglade (2001). Stable isotopes data (δ13C and δ15N) from the cave bear (Ursus spelaeus): a new approach to its palaeoenvironment and dormancy. Proc.R.Soc.Lond. B, 268.

Georgiev, D., et al. (2010). New localities of Quaternary fossil bears (Ursus sp. L.) (Mammalia: Carnivora: Ursidae). ZooNotes, 8: 1-4.

Hailer, F., et al. (2012). Nuclear Genomic Sequences Reveal that Polar Bears Are an Old and Distinct Bear Lineage. Science, Vol.336.

Hofreiter, M., et al. (2004). Evidence for Reproductive Isolation between Cave Bear Populations. Current Biology, Vol.14.

Kurtén, B. (1964). The evolution of the Polar Bear, Ursus maritimus Phipps. Acta Zoologica Fennica, 108.

Kurtén, B. (1958). Life and Death of the Pleistocene Cave Bear. A Study in Paleoecology. Acta Zoologica Fennica, 95.

Leiss-Holzinger, E., et al. (2015). Imaging of the inner structure of cave bear teeth by novel non-destructive techniques. Palaeontologia Electronica, PE Article Number 18.1.1T.

Lindqvist, C., et al. (2010). Complete mitochondrial genome of a Pleistocene jawbone unveils the origin of polar bear. PNAS, Vol.107, Number 11.

Mackiewicz, P., et al. (2017). Estimating the extinction time of two cave bears, Ursus spleaeus and U. ingressus. Acta zoologica cracoviensia, 60(2).

Pinto Llona, A.C. (2006). Comparative Dental Microwear Analysis of Cave Bears Ursus spelaeus Rosenmüller, 1794 and Brown Bears Ursus arctos Linnaeus,1758. Scientific Annals, School of Geology, Aristotle University of Thessaloniki, Special Volume 98.

Stiller, M., et al. (2013). Mitochondrial DNA diversity and evolution of the Pleistocene cave bear complex. Quaternary International, xxx. (Article in press)

Stiner, M.C. (1999). Cave Bear Ecology and Interactions with Pleistocene Humans. Ursus, 11.
Stiner, M.C. (1998). Mortality analysis of Pleistocene bears and its paleoanthropological relevance. Journal of Human Evolution, 34.

van Heteren, A.H., et al. (2015). Functional morphology of the cave bear (Ursus spelaeus) mandible: a 3D geometric morphometric analysis. Organisms Diversity & Evolution. (Author's personal copy)

van Heteren, A.H., et al. (2014). Functional morphology of the cave bear (Ursus spelaeus) cranium: a three-dimensional geometric morphometric analysis. Quaternary International, 339-340.

van Heteren, A.H., et al. (2009). Cave Bears and Their Closest Living Relatives: A 3D Geometric Morphometrical Approach to the Functional Morphology of the Cave Bear Ursus spelaeus. Acta Carsologica Slovaca, 47 Suppl. 1.

Veitschegger, K., et al. (2018). Longevity and life history of cave bears - a review and novel data from tooth cementum and relative emergence of permanent dentition. Historical Biology, 2018.

von Koenigswald, W. (1992). Tooth enamel of the cave bear (Ursus spelaeus) and the relationship between diet and enamel structures. Acta. Zool. Fennici, 28.

Wolverton, S. (2001). Caves, Ursids and Artifacts: A Natural-Trap Hypothesis. Journal of Ethnobiology, 21(2).


General Ursoidea


Abella, J., P. Montoya and J. Morales (2014). Paleodiversity of the Superfamily Ursoidea (Carnivora, Mammalia) in the Spanish Neogene, related to environmental changes. Journal of Iberian Geology, 40(1).

Arroyo-Cabrales, J., et al. (2016). North American Ursid (Mammalia: Ursidae) Defaunation from Pleistocene to Recent. ICBS Proceedings, Cranium, 2016.

Donohue, S.L. (2013). Using Dental Microwear Textures to Assess Feeding Ecology of Extinct and Extant Bears. Masters Thesis - Vanderbilt University.

Figueirido, B., P. Palmqvist and J.A. Perez-Claros (2009). Ecomorphological correlates of craniodental variation in bears and paleobiological implications for extinct taxa: an approach based on geometric morphometrics. Journal of Zoology, 277.

Krause, J., et al. (2008). Mitochondrial genomes reveal an explosive radiation of extinct and extant bears near the Miocene-Pliocene boundary. BMC Evolutionary Biology, 8:220.

Mclellan, B. and D.C. Reiner (1994). A Review of Bear Evolution. Int.Conf. Bear Res. and Manage., 9(1).

van Heteren, A.H. (2011). Masticatory adaptations of extant and extinct Ursidae: an assessment using three-dimensional geometric morphometrics. Ph.D.Thesis - University of Roehampton.

Waits, L.P., et al. (1999). Rapid Radiation Events in the Family Ursidae Indicated by Likelihood Phylogenetic Estimation from Multiple Fragments of mtDNA. Molecular Phylogenetics and Evolution, Vol.13, Number 1.

Yu, L, et al. (2004). Phylogeny of the bears (Ursidae) based on nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution, 32.

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Updated October 6, 2010.


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Updated February 6, 2011.


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Updated March 20, 2011.


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Updated April 8, 2012.


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Updated December 23, 2016.



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