Table of Contents
ISRN Zoology
Volume 2013 (2013), Article ID 138319, 25 pages
http://dx.doi.org/10.1155/2013/138319
Research Article

Extinctions of Late Ice Age Cave Bears as a Result of Climate/Habitat Change and Large Carnivore Lion/Hyena/Wolf Predation Stress in Europe

Paleologic, Private Research Institute, Petra Bezruce 96, CZ-26751 Zdice, Czech Republic

Received 16 September 2012; Accepted 5 October 2012

Academic Editors: L. Kaczmarek and C.-F. Weng

Copyright © 2013 Cajus G. Diedrich. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Linked References

  1. C. Diedrich, “Cave bear killers, scavengers between the Scandinavian and Alpine ice shields—the last hyenas and cave bears in antagonism—and the reason why cave bears hibernated deeply in caves,” Stalactite, vol. 58, no. 2, pp. 53–63, 2009. View at Google Scholar
  2. C. Diedrich, “The Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss 1823) population from the Zoolithen Cave at Gailenreuth (Bavaria, South Germany)—a hyena cub raising den of specialized cave bear scavengers in Boreal Forest environments of Central Europe,” Historical Biology, vol. 23, no. 4, pp. 335–367, 2011. View at Google Scholar
  3. C. Diedrich, “Cave bear killers and scavengers in the Final Ice Age of Europe—feeding specialization as reaction on the mammoth steppe fauna absence in mountainous regions,” Quaternary International, vol. 255, pp. 59–78, 2012. View at Publisher · View at Google Scholar
  4. D. E. Nelson, A. Angerbjörn K, Liden, and I. Turk, “Stable isotopes and the metabolism of the European cave bear,” Oecologia, vol. 116, no. 1-2, pp. 177–181, 1998. View at Publisher · View at Google Scholar
  5. J. C. Rosenmüller, “Quedam 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, Leipzig, Germany, 1794.
  6. C. Diedrich, “The rediscovered cave bear “Ursus spelaeus Rosenmüller 1794” holotype of the Zoolithen Cave (Germany) from the historic Rosenmüller collection,” Acta Carsologica Slovacia, vol. 47, no. 1, pp. 25–32, 2009. View at Google Scholar
  7. F. Heller, “Die berühmten Knochenhöhlen des fränkischen Jura und das Schicksal ihres Fundinhaltes. Nach zeitgenösischen Berichten und Quellen,” Berichte der Naturwissenschaftlichen Gesellschaft Bayreuth, vol. 12, pp. 5–20, 1966. View at Google Scholar
  8. C. Diedrich, “The rediscovered holotypes of the Upper Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss 1823) and the steppe lion Panthera leo spelaea (Goldfuss, 1810) and taphonomic discussion to the Zoolithen Cave hyena den at Geilenreuth (Bavaria, South-Germany),” Zoological Journal of the Linnean Society, vol. 154, no. 4, pp. 822–831, 2008. View at Publisher · View at Google Scholar
  9. G. A. Goldfuss, “Osteologische Beitraege zur Kenntnis verschiedener Saeugethiere der Vorwelt. VI. Ueber die Hoelen-Hyaene (Hyaena spelaea),” Nova Acta Physico-Medica Academiea Caesarae Leopoldino-Carolinae Naturae Curiosorum, vol. 3, no. 2, pp. 456–490, 1823. View at Google Scholar
  10. C. Diedrich, “The largest european lion Panthera leo spelaea (Goldfuss 1810) population from the zoolithen cave, germany: specialised cave bear predators of Europe,” Historical Biology, vol. 23, no. 2-3, pp. 271–311, 2011. View at Publisher · View at Google Scholar · View at Scopus
  11. G. A. Goldfuss, Die Umgebungen von Muggendorf. Ein Taschenbuch für Freunde der Natur und Alterthumskunde, Palm, Erlangen, Germany, 1810.
  12. G. A. Goldfuss, “Beschreibung eines fossilen Vielfrassschädels aus der Geilenreuther Höhle,” Nova Acta Physico-Medica Academiea Caesarae Leopoldino-Carolinae Naturae Curiosorum, vol. 11, pp. 313–322, 1818. View at Google Scholar
  13. W. Buckland, Reliquiae Diluvianae, or Observations on the Organic Remains Contained in Caves, Fissures, and Diluvial Gravel, and Other Geological Phenomena, Attesting the Action of an Universal Deluge, J. Murray, London, UK, 1823.
  14. J. F. Esper, Ausführliche Nachricht von neuentdeckten Zoolithen unbekannter vierfüssiger Thiere und denen sie enthaltenden, so wie verschiedenen andern, denkwürdigen Grüften der Obergebürgischen Lande des Marggrafthums Bayreuth, Knorr, Nürnberg, Germany, 1774.
  15. B. Kaulich and H. Schaaf, Kleiner Führer zu den Höhlen um Muggendorf, Naturhistorische Gesellschaft Nürnberg, Nürnberg, Germany, 1993.
  16. J. C. Rosenmüller, Merkwürdigkeiten der Gegend um Muggendorf, Unger, 1804.
  17. H. Schabdach, Die Sophien-Höhle im Ailsbachtal—Wunderwelt unter Tage, Reinhold Lippert, Ebermannstadt, Germany, 1998.
  18. M. Knapp, N. Rohland, J. Weinstock et al., “First DNA sequences from Asian cave bear fossils reveal deep divergences and complex phylogeographic patterns,” Molecular Ecology, vol. 18, no. 6, pp. 1225–1238, 2009. View at Publisher · View at Google Scholar · View at Scopus
  19. S. C. Münzel, M. Stiller, M. Hofreiter, A. Mittnik, N. J. Conard, and H. Bocherens, “Pleistocene bears in the Swabian Jura (Germany): genetic replacement, ecological displacement, extinctions and survival,” Quaternary International, vol. 245, no. 2, pp. 225–237, 2011. View at Publisher · View at Google Scholar
  20. M. Pacher and A. J. Stuart, “Extinction chronology and palaeobiology of the cave bear (Ursus spelaeus),” Boreas, vol. 38, no. 2, pp. 189–206, 2009. View at Publisher · View at Google Scholar · View at Scopus
  21. M. Stiller, G. Baryshnikov, H. Bocherens et al., “Withering away—25,000 years of genetic decline preceded cave bear extinction,” Molecular Biology and Evolution, vol. 27, no. 5, pp. 975–978, 2010. View at Publisher · View at Google Scholar · View at Scopus
  22. J. W. Holle, “Die neu entdeckte Kochshöhle oder die Höhlenkönigin im königlichen Landgerichte Hollfeld-Waischenfeld,” Bayerische Annalen, vol. 26, pp. 197–198, 1833. View at Google Scholar
  23. R. Wagner, “Ueber die neu entdeckte Zoolithenhöhle bey Rabenstein,” Bayerische Annalen, vol. 47, pp. 313–315, 1833. View at Google Scholar
  24. K. Sternberg, “Vortrag des Präsidenten Grafen Kaspar Sternberg in der allgemeinen Versammlung des böhmischen Museums in Prag,” Verhandlungen der Gesellschaft des vaterländischen Museums in Böhmen Prag, pp. 12–30, 1835. View at Google Scholar
  25. C. Diedrich, “Ice age geomorphological Ahorn Valley and Ailsbach River terrace evolution—and its importance for the cave use possibilities by cave bears, top predators (hyenas, wolves and lions) and humans (Neanderthals, Late Magdalénians) in the Frankonian Karst—case studies in the Sophie’s Cave near Kirchahorn, Bavaria,” In press, 2013.
  26. G. Rabeder, “Die Evolution des Höhlenbärgebisses,” Mitteilung der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften, vol. 11, pp. 1–12, 1999. View at Google Scholar
  27. M. Domínguez-Rodrigo and A. Piqueras, “The use of tooth pits to identify carnivore taxa in tooth-marked archaeofaunas and their relevance to reconstruct hominid carcass processing behaviours,” Journal of Archaeological Science, vol. 30, no. 11, pp. 1385–1391, 2003. View at Publisher · View at Google Scholar · View at Scopus
  28. J. T. Faith, “Sources of variation in carnivore tooth-mark frequencies in a modern spotted hyena (Crocuta crocuta) den assemblage, Amboseli Park, Kenya,” Journal of Archaeological Science, vol. 34, no. 10, pp. 1601–1609, 2007. View at Publisher · View at Google Scholar · View at Scopus
  29. M. M. Selvaggio and J. Wilder, “Identifying the involvement of multiple carnivore taxa with archaeological bone assemblages,” Journal of Archaeological Science, vol. 28, no. 5, pp. 465–470, 2001. View at Publisher · View at Google Scholar · View at Scopus
  30. A. Hill, “Bone modification by modern spotted hyenas,” in Bone Modification, R. Bonnichsen and M. H. Sorg, Eds., pp. 169–178, Center for the Study of the First Americans, Orono, Me, USA, 1989. View at Google Scholar
  31. C. Diedrich, “Cracking and nibbling marks as indicators for the Upper Pleistocene spotted hyena as a scavenger of cave bear (Ursus spelaeus Rosenmüller 1794) carcasses in the Perick Caves den of Northwest Germany,” Abhandlungen der Naturhistorischen Gesellschaft Nürnberg, vol. 45, pp. 73–90, 2005. View at Google Scholar
  32. C. Diedrich, “Late Pleistocene Crocuta crocuta spelaea (Goldfuss 1823) clans as prezewalski horse hunters and woolly rhinoceros scavengers at the open air commuting den and contemporary Neanderthal camp site Westeregeln (central Germany),” Journal of Archaeological Science, vol. 39, no. 6, pp. 1749–1767.
  33. C. Diedrich, “The largest Late Pleistocene hyena population from the Srbsko Chlum-Komín Cave (Czech Republic) and its prey in a commuting and prey depot cave den of Central Europe,” Historical Biology. In press.
  34. C. Diedrich, “Late Pleistocene Crocuta crocuta spelaea (Goldfuss 1823) populations from the Emscher River terrace open air hyena den near Bottrop and other sites in NW Germany: their bone accumulations along rivers in lowland mammoth steppe environments and scavenging activities on woolly rhinoceros,” Quaternary International, vol. 276-277, pp. 93–119, 2012. View at Publisher · View at Google Scholar
  35. C. Diedrich, “Late Pleistocene Eemian Ice Age spotted hyena feeding strategies and steppe lions on their largest prey—palaeoloxodon antiquus Falconer and Cautley, 1845 at the straight-tusked elephant graveyard and Neandertalian site Neumark-Nord Lake 1, Central Germany,” Archaeological and Anthropological Sciences. In press.
  36. B. van Valkenburgh, “Trophic diversity in past and present guilds of large predatory mammals,” Paleobiology, vol. 14, no. 2, pp. 155–173, 1988. View at Google Scholar · View at Scopus
  37. B. van Valkenburgh, “Tracking ecology over geological time: evolution within guilds of vertebrates,” Trends in Ecology and Evolution, vol. 10, no. 2, pp. 71–76, 1995. View at Publisher · View at Google Scholar · View at Scopus
  38. A. Neischl, Die Höhlen der fränkischen Schweiz und ihre Bedeutung für die Entstehung der dortigen Täler, Schrag, Nürnberg, Germany, 1904.
  39. R. G. Spöker, Zur Landschafts-Entwicklung im Karst des oberen und mittleren Pegnitz-Gebietes, Verlag des Amtes für Landeskunde, Remagenm, Germany, 1952.
  40. K. A. Habbe, “Der Karst der Fränkischen Alb—Formen, Prozesse, Datierungsprobleme. Die Fränkische Alb,” Schriften des Zentralen Instituted für fränkische Landeskunde Universität Erlangen, vol. 28, pp. 35–69, 1989. View at Google Scholar
  41. J. H. Bretz, “Vadose and phreatic features of limestone caverns,” The Journal of Geology, vol. 50, no. 6, pp. 675–811, 1942. View at Publisher · View at Google Scholar
  42. D. C. Ford and P. W. Williams, Karst Geomorphology and Hydrology, Unwin-Hyman, London, UK, 1989.
  43. F. Brand, “Was können wir aus lehmigen Ablagerungen der Binghöhle ablesen?” in Die Binghöhle bei Streitberg—Auf den Spuren eines unterirdischen Flusses, F, R. Brand, F. Illmann, D. Leja, Preu, and H. Schabdach, Eds., pp. 28–34, 2006. View at Google Scholar
  44. T. Dogwiler and C. M. Wicks, “Sediment entrainment and transport in fluviokarst systems,” Journal of Hydrology, vol. 295, no. 1–4, pp. 163–172, 2004. View at Publisher · View at Google Scholar · View at Scopus
  45. D. Burger, “Dolomite weathering and micromorphology of paleosoils in the Frankonian Jura,” Catena Supplementaria, vol. 15, pp. 261–267, 1989. View at Google Scholar
  46. C. Diedrich, “The Late Pleistocene Panthera leo spelaea (Goldfuss 1810) skeletons from the Sloup and Srbsko Caves in Czech Republic (Central Europe) and contribution to steppe lion cranial pathologies and postmortally damages as results of interspecies fights, hyena antagonism and cave bear attacks,” Bulletin of Geosciences, vol. 86, no. 4, pp. 817–840, 2011. View at Google Scholar
  47. C. Diedrich, “Pleistocene Panthera leo spelaea (Goldfuss 1810) remains from the Balve Cave (NW Germany)—a cave bear, hyena den and Middle Palaeolithic human cave, and review of the Sauerland Karst lion sites,” Quaternaire, vol. 22, no. 2, pp. 105–127, 2011. View at Google Scholar
  48. G. Harnes, “A guide for differentiating mammalian carnivore taxa responsible for gnaw damage to herbivore limb bones,” Paleobiology, vol. 9, no. 2, pp. 164–172, 1983. View at Google Scholar · View at Scopus
  49. A. C. Pinto Llona and P. J. Andrew, “Scavenging behaviour patterns in cave bears Ursus spelaeus,” Revue de Paléobiologie, vol. 23, pp. 845–885, 2004. View at Google Scholar
  50. J. Quilès, C. Petrea, O. Moldovan et al., “Cave bears (Ursus spelaeus) from the Peştera cu Oase (Banat, Romania): Paleobiology and taphonomy,” Comptes Rendus—Palevol, vol. 5, no. 8, pp. 927–934, 2006. View at Publisher · View at Google Scholar · View at Scopus
  51. G. Rabeder, D. Nagel, and M. Pacher, Der Höhlenbär, vol. 4 of Thorbecke Species, Thorbecke Jan, Stuttgart, Germany, 2000.
  52. C. Diedrich, “The Crocuta crocuta spelaea (goldfuss 1823) population and its prey from the Late Pleistocene Teufelskammer Cave hyena den besides the famous Paleolithic Neanderthal Cave (NRW, NW Germany),” Historical Biology, vol. 23, no. 2-3, pp. 237–270, 2011. View at Publisher · View at Google Scholar · View at Scopus
  53. M. C. Stiner, “Comparative ecology and taphonomy of spotted hyenas, humans, and wolves in Pleistocene Italy,” Revue de Paleobiologie, vol. 23, no. 2, pp. 771–785, 2004. View at Google Scholar · View at Scopus
  54. L. D. Mech, The Arctic Wolf: Living with the Pack, Voyageur Press, Stillwater, Minn, USA, 1988.
  55. J. A. Leonard, C. Vilà, K. Fox-Dobbs, P. L. Koch, R. K. Wayne, and B. van Valkenburgh, “Megafaunal extinctions and the disappearance of a specialized wolf ecomorph,” Current Biology, vol. 17, no. 13, pp. 1146–1150, 2007. View at Publisher · View at Google Scholar · View at Scopus
  56. D. I. Bibikow, Der Wolf—Canis Lupus, Neue Brehm-Bücherei, Wittenberg, Germany, 2003.
  57. R. M. Nowak, Walker’s Mammals of the World, Johns Hopkins University Press, Baltimore, Md, USA, 1999.
  58. E. Ziemen, Der Wolf, Verhalten, Ökologie und Mythos, Kosmos, Stuttgart, Germany, 2003.
  59. C. Diedrich, “Typology of Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) coprolite aggregate pellets from the European Late Pleistocene and their significance at dens and scavenging sites,” New Mexico Museum of Natural History and Science, Bulletin, vol. 57, pp. 369–377, 2012. View at Google Scholar
  60. C. Diedrich and K. Žák, “Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823) in horizontal and vertical caves of the Bohemian Karst (Czech Republic),” Bulletin of Geosciences, vol. 81, no. 4, pp. 237–276, 2006. View at Publisher · View at Google Scholar · View at Scopus
  61. I. Barja, F. J. de Miguel, and F. Bárcena, “The importance of crossroads in faecal marking behaviour of the wolves (Canis lupus),” Naturwissenschaften, vol. 91, no. 10, pp. 489–492, 2004. View at Publisher · View at Google Scholar · View at Scopus
  62. T. J. Floyd, L. D. Mech, and P. A. Jordan, “Relating wolf scat content to prey consumed,” Journal of Wildlife Management, vol. 42, no. 3, pp. 528–532, 1978. View at Publisher · View at Google Scholar
  63. L. D. Mech and J. M. Packard, “Possible use of (Canis lupus) den over several centuries,” Canadian Field Naturalist, vol. 104, pp. 484–485, 1990. View at Google Scholar
  64. G. Rabeder and M. Hofreiter, “Der neue Stammbaum der Höhlenbären,” Die Höhle, vol. 55, pp. 1–19, 2004. View at Google Scholar
  65. G. Rabeder, M. Hofreiter, D. Nagel, S. Paabo, and G. Withalm, “Die neue Taxonomie der Höhlenbaren,” Abhandlungen zur Karst-und Höhlenkunde, vol. 34, pp. 68–69, 2002. View at Google Scholar
  66. P. Gibbard and T. van Kolfschoten, “The Pleistocene and holocene epochs—chapter 22,” in A Geologic Time Scale 2004, F. M. Gradstein, G. Ogg, J. Smith, and A. Gilbert, Eds., pp. 441–452, Cambridge University Press, Cambridge, UK, 2004. View at Google Scholar
  67. M. Hofreiter, “Genetic stability and replacement in Late Pleistocene cave bear populations,” Abhandlungen zur Karst-und Höhlenkunde, vol. 34, pp. 64–67, 2002. View at Google Scholar
  68. W. von Koenigswald, Lebendige Eiszeit—Klima und Tierwelt im WandelTheiss-Verlag, Wissenschaftliche Buchgesellschaft, Darmstadt, Germany, 2002.
  69. J. Weinstock, “Late Pleistocene reindeer populations in Middle and Western Europe,” BioArchaeologica, vol. 3, pp. 1–307, 2000. View at Google Scholar
  70. I. Debeljak, “Fossil population structure and mortality of the cave bear from the Mokrica cave (North Slovenia),” Acta Carsologica, vol. 36, no. 3, pp. 475–484, 2007. View at Google Scholar · View at Scopus
  71. A. Grandal-D'Anglade and J. R. V. Romaní, “A population study on the cave bear (Ursus spelaeus Ros.-Hein.) from Cova Eirós (Triacastela, Galicia, Spain),” Geobios, vol. 30, no. 5, pp. 723–731, 1997. View at Google Scholar · View at Scopus
  72. P. Argenti and P. P. A. Mazza, “Mortality analysis of the Late Pleistocene bears from Grotta Lattaia, Central Italy,” Journal of Archaeological Science, vol. 33, no. 11, pp. 1552–1558, 2006. View at Publisher · View at Google Scholar · View at Scopus
  73. H. Bocherens, D. G. Drucker, D. Bonjean et al., “Isotopic evidence for dietary ecology of cave lion (Panthera spelaea) in North-Western Europe: prey choice, competition and implications for extinction,” Quaternary International, vol. 245, no. 2, pp. 249–261, 2011. View at Publisher · View at Google Scholar
  74. C. Diedrich, “Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone records from open air sites in Northern Germany—evidence of hyena-lion antagonism and scavenging in Europe,” Quaternary Science Reviews, vol. 30, no. 15-16, pp. 1883–1906, 2011. View at Publisher · View at Google Scholar · View at Scopus
  75. R. Estes, The Safari Companion: A Guide to Watching African Mammals, Chelsea Green Publishing, White River Junction Hartford, Vt, USA, 1999.
  76. G. Schaller, The Serengeti Lion. A Study of Predator-Prey Relations, The University of Chicago Press, Chicago, Ill, USA, 1972.
  77. C. Diedrich, “Ichnological and ethological studies in one of Europe's famous bear den in the Urşilor Cave (Carpathians, Romania),” Ichnos, vol. 18, no. 1, pp. 9–26, 2010. View at Google Scholar
  78. H. Kruuk, “Interactions between populations of spotted hyaenas (Crocuta crocuta Erxleben) and their prey species,” in Animal Populations in Relation to Their Food Resources, A. Watson, Ed., pp. 359–374, Blackwell, Oxford, UK, 1970. View at Google Scholar
  79. H. Kruuk, The Spotted Hyena—A Story of Predation and Social Behavior, The University of Chicago Press, Chicago, Ill, USA, 1972.
  80. N. Rohland, J. L. Pollack, D. Nagel et al., “The population history of extant and extinct hyenas,” Molecular Biology and Evolution, vol. 22, no. 12, pp. 2435–2443, 2005. View at Publisher · View at Google Scholar · View at Scopus
  81. C. Diedrich, “Freeland remains of the cave bear Ursus spelaeus Rosenmüller 1794 from the Upper Pleistocene of Northwest Germany,” Bulletin de la Société Histoire Naturelle Toulouse, vol. 141, no. 1, pp. 19–23, 2005. View at Google Scholar
  82. C. Diedrich, “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, vol. 98, pp. 193–199, 2006. View at Google Scholar
  83. C. Diedrich, “Periodical use of the Balve Cave (NW Germany) as a Late Pleistocene Crocuta crocuta spelaea (Goldfuss 1823) den: hyena occupations and bone accumulations versus human Middle Palaeolithic activity,” Quaternary International, vol. 233, no. 2, pp. 171–184, 2011. View at Publisher · View at Google Scholar · View at Scopus
  84. C. Diedrich, “Late Pleistocene Cave bear remains from the open air hyena den Emscher River terrace site Bottrop (NW Germany),” Stalactite, vol. 58, no. 2, pp. 42–47, 2009. View at Google Scholar
  85. J. Burger, W. Rosendahl, O. Loreille et al., “Molecular phylogeny of the extinct cave lion Panthera leo spelaea,” Molecular Phylogenetics and Evolution, vol. 30, no. 3, pp. 841–849, 2004. View at Publisher · View at Google Scholar · View at Scopus
  86. C. Diedrich, “Specialized horse killers in Europe: foetal horse remains in the Late Pleistocene Srbsko Chlum-Komín Cave hyena den in the Bohemian Karst (Czech Republic) and actualistic comparisons to modern African spotted hyenas as zebra hunters,” Quaternary International, vol. 220, no. 1-2, pp. 174–187, 2010. View at Publisher · View at Google Scholar · View at Scopus
  87. J. Ford, Predators at War, National Geographic, 2005.
  88. D. Joubert and B. Joubert, Eternal Enemies: Lions and Hyena, Wildlife Films Botswana for National Geographic, 2003.
  89. F. Palomares and T. M. Caro, “Interspecific killing among mammalian carnivores,” American Naturalist, vol. 153, no. 5, pp. 492–508, 1999. View at Publisher · View at Google Scholar · View at Scopus
  90. B. M. Rothschild and C. Diedrich, “Comparison of pathologies in the extinct Pleistocene Eurasian steppe lion Pantherea leo spelaea (Goldfuss, 1810) to those in the modern lion, Panthera leo—results of fights with hyenas, bears and lions and other ecological stress,” International Journal of Paleopathology, vol. 2, no. 4, pp. 187–198, 2012. View at Publisher · View at Google Scholar
  91. M. G. L. Mills and M. Mills, “An analysis of bones collected at hyaena breeding dens in the Gemsbok National Parks,” Annales of the Transvaal Museum, vol. 30, pp. 145–155, 1977. View at Google Scholar
  92. P. Fosse, J. P. Brugal, J. L. Guadelli, P. Michel, and J. F. Tournepiche, “Les repaires d'hyenes des cavernes en Europe occidentale: presentation et comparisons de quelques assemblages osseux,” in Economie Prehistorique: Les comportements de substance au Paleolithique. XVIII Rencontres internationales d'Archeologie et d'Historie d'Antibes, pp. 44–61, Editions APDCA, Sophia Antipolis Biot, France, 1998. View at Google Scholar
  93. R. Musil, “Die Höhle, “Sveduv stul”, ein typischer Höhlenhyänenhorst,” Anthropos N.S., vol. 5, no. 13, pp. 97–260, 1962. View at Google Scholar
  94. J. F. Tournepiche and C. Couture, “The hyena den of Rochelot Cave (Charente, France),” Monographien des Römisch-Germanischen Zentralmuseums, vol. 42, pp. 89–101, 1999. View at Google Scholar
  95. S. M. Cooper, “Denning behavior of spotted hyaenas (Crocuta crocuta) in Botswana,” African Journal of Ekology, vol. 31, no. 2, pp. 178–180, 1993. View at Publisher · View at Google Scholar
  96. S. W. Lansing, S. M. Cooper, E. E. Boydston, and K. E. Holekamp, “Taphonomic and zooarchaeological implications of spotted hyena (Crocuta crocuta) bone accumulations in kenya: a modern behavioral ecological approach,” Paleobiology, vol. 35, no. 2, pp. 289–309, 2009. View at Publisher · View at Google Scholar · View at Scopus
  97. Y. M. Lam, “Variability in the behaviour of spotted hyaenas as taphonomic agents,” Journal of Archaeological Science, vol. 19, no. 4, pp. 389–406, 1992. View at Google Scholar · View at Scopus
  98. J. T. Pokines and J. C. K. Peterhans, “Spotted hyena (Crocuta crocuta) den use and taphonomy in the Masai Mara National Reserve, Kenya,” Journal of Archaeological Science, vol. 34, no. 11, pp. 1914–1931, 2007. View at Publisher · View at Google Scholar · View at Scopus
  99. L, Scott, and R. G. Klein, “A hyena accumulated bone assemblage from late holocene deposits at Deelpan, Orange Free State,” Annals of the South African Museum, vol. 86, pp. 217–227, 1981. View at Google Scholar
  100. S. M. Cooper, “The hunting behaviour of spotted hyaenas (Crocuta crocuta) in a region containing both sedentary and migratory populations of herbivores,” African Journal of Ecology, vol. 28, no. 2, pp. 131–141, 1990. View at Google Scholar · View at Scopus
  101. I. Di Silvestre, O. Novell, and G. Bogliani, “Feeding habits of the spotted hyaena in the Niokolo Koba National Park, Senegal,” African Journal of Ecology, vol. 38, no. 2, pp. 102–107, 2000. View at Publisher · View at Google Scholar · View at Scopus
  102. A. J. Sutcliffe, “Spotted hyaena: crusher, gnawer, digester and collector of bones,” Nature, vol. 227, no. 5263, pp. 1110–1113, 1970. View at Publisher · View at Google Scholar · View at Scopus
  103. W. B. Ballard, “Brown bear kills gray wolf,” Canadian Field-Naturalist, vol. 94, atricle 91, 1980. View at Google Scholar
  104. K. A. Gunther and D. W. Smith, “Interactions between wolves and female grizzly bears with cubs in Yellowstone National Park,” Ursus, vol. 15, no. 2, pp. 232–238, 2004. View at Google Scholar · View at Scopus
  105. R. D. Hayes and D. H. Mossop, “Interactions of wolves, Canis lupus, and brown bears, Ursus arctos, at a wolf den in the Northern Yukon (Canada),” Canadian Field-Naturalist, vol. 101, pp. 603–604, 1987. View at Google Scholar
  106. R. D. Hayes and A. Baer, “Brown bear, Ursus arctos, preying upon gray wolf, Canis lupus, pack,” Canadian Field-Naturalist, vol. 107, pp. 373–374, 1992. View at Google Scholar
  107. E. S. Richardson and D. Andriashek, “Wolf (Canis lupus) predation of a polar bear (Ursus maritimus) cub on the sea ice off northwestern Banks Island, Northwest Territories, Canada,” Arctic, vol. 59, no. 3, pp. 322–324, 2006. View at Google Scholar · View at Scopus
  108. L. L. Rogers and D. Mech, “Interactions of wolves and black bears in Northeastern Minnesota,” Journal of Mammalogy, vol. 62, no. 2, pp. 434–436, 1981. View at Publisher · View at Google Scholar
  109. J. T. Groiss, “Über pathologische Bildungen an Skelettresten jungquartärer Säugetiere aus der Zoolithenhöhle bei Burggeilenreuth,” Geologische Blätter NO-Bayern, vol. 28, no. 1, pp. 1–21, 1978. View at Google Scholar
  110. M. Janik, “Biogeography, demography and management of Urus arctos in the Western Carpathians,” International Conference of Bear Research and Management, vol. 9, no. 2, pp. 125–128, 1997. View at Google Scholar
  111. C. Diedrich, “Freeland remains and den caves of the cave bear Ursus spelaeus Rosenmüller 1794 from the Upper Pleistocene of Bohemia (Czech Republic),” Scientific Annals, vol. 98, pp. 187–192, 2006. View at Google Scholar