全部 标题 作者
关键词 摘要

OALib Journal期刊
ISSN: 2333-9721
费用:99美元

查看量下载量

相关文章

更多...
ISRN Zoology  2013 

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

DOI: 10.1155/2013/138319

Full-Text   Cite this paper   Add to My Lib

Abstract:

Predation onto cave bears (especially cubs) took place mainly by lion Panthera leo spelaea (Goldfuss), as nocturnal hunters deep in the dark caves in hibernation areas. Several cave bear vertebral columns in Sophie’s Cave have large carnivore bite damages. Different cave bear bones are chewed or punctured. Those lets reconstruct carcass decomposition and feeding technique caused only/mainly by Ice Age spotted hyenas Crocuta crocuta spelaea, which are the only of all three predators that crushed finally the long bones. Both large top predators left large tooth puncture marks on the inner side of cave bear vertebral columns, presumably a result of feeding first on their intestines/inner organs. Cave bear hibernation areas, also demonstrated in the Sophie’s Cave, were far from the cave entrances, carefully chosen for protection against the large predators. The predation stress must have increased on the last and larger cave bear populations of U. ingressus (extinct around 25.500?BP) in the mountains as result of disappearing other seasonally in valleys migrating mammoth steppe fauna due to climate change and maximum glacier extensions around 22.000?BP. 1. Introduction 1.1. Cave Bear Research History in Germany Many incomplete cave bear bones from the Late Pleistocene Ice Age (113.000–22.000?BP) have been found in Europe and focussed herein in Germany mainly from 42 famous cave sites but also a smaller number from open air sites (Figure 1(a)). The cave bear bones show always similar patterns of damage [1–3] including bones from the new described Sophie’s Cave site in Upper Franconia, Bavaria (Figure 1(b)) and the not far situated famous Zoolithen Cave with its Late Pleistocene herbivorous [4] cave bear “Ursus spelaeus Rosenmüller 1794” [5], whose holotype skull was recently rediscovered [6]. This most famous bone-rich Zoolithen Cave of central Europe with its long history of “cave bear bone hunt” [7] has also yielded the holotype skulls of predators such as the Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss 1823) [2, 8, 9], the steppe lion Panthera leo spelaea (Goldfuss 1810) [8, 10, 11], the “cave wolverine” Gulo gulo spelaeus (Goldfuss 1818) [12], and the “cave wolf” Canis lupus spelaeus Goldfuss 1823 [9]. Figure 1: (a) Geographical locations of German cave bear sites, both cave den and open air sites, including Sophie’s Cave in Bavaria, southern Germany. (b) Cave map showing cave bear, hyena, and wolf den areas during the early/middle Late Pleistocene in Upper Franconia (Bavaria, Germany). (c) The present-day entrance of Sophie’s Cave. (d)

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.
[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.
[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.
[4]  D. E. Nelson, A. Angerbj?rn 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.
[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 He?berga-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.
[7]  F. Heller, “Die berühmten Knochenh?hlen des fr?nkischen Jura und das Schicksal ihres Fundinhaltes. Nach zeitgen?sischen Berichten und Quellen,” Berichte der Naturwissenschaftlichen Gesellschaft Bayreuth, vol. 12, pp. 5–20, 1966.
[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.
[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.
[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.
[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 Vielfrasssch?dels aus der Geilenreuther H?hle,” Nova Acta Physico-Medica Academiea Caesarae Leopoldino-Carolinae Naturae Curiosorum, vol. 11, pp. 313–322, 1818.
[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 H?hlen um Muggendorf, Naturhistorische Gesellschaft Nürnberg, Nürnberg, Germany, 1993.
[16]  J. C. Rosenmüller, Merkwürdigkeiten der Gegend um Muggendorf, Unger, 1804.
[17]  H. Schabdach, Die Sophien-H?hle 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.
[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.
[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.
[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.
[22]  J. W. Holle, “Die neu entdeckte Kochsh?hle oder die H?hlenk?nigin im k?niglichen Landgerichte Hollfeld-Waischenfeld,” Bayerische Annalen, vol. 26, pp. 197–198, 1833.
[23]  R. Wagner, “Ueber die neu entdeckte Zoolithenh?hle bey Rabenstein,” Bayerische Annalen, vol. 47, pp. 313–315, 1833.
[24]  K. Sternberg, “Vortrag des Pr?sidenten Grafen Kaspar Sternberg in der allgemeinen Versammlung des b?hmischen Museums in Prag,” Verhandlungen der Gesellschaft des vaterl?ndischen Museums in B?hmen Prag, pp. 12–30, 1835.
[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 H?hlenb?rgebisses,” Mitteilung der Kommission für Quart?rforschung der ?sterreichischen Akademie der Wissenschaften, vol. 11, pp. 1–12, 1999.
[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.
[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.
[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.
[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.
[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.
[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.
[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.
[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.
[38]  A. Neischl, Die H?hlen der fr?nkischen Schweiz und ihre Bedeutung für die Entstehung der dortigen T?ler, Schrag, Nürnberg, Germany, 1904.
[39]  R. G. Sp?ker, 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 Fr?nkischen Alb—Formen, Prozesse, Datierungsprobleme. Die Fr?nkische Alb,” Schriften des Zentralen Instituted für fr?nkische Landeskunde Universit?t Erlangen, vol. 28, pp. 35–69, 1989.
[41]  J. H. Bretz, “Vadose and phreatic features of limestone caverns,” The Journal of Geology, vol. 50, no. 6, pp. 675–811, 1942.
[42]  D. C. Ford and P. W. Williams, Karst Geomorphology and Hydrology, Unwin-Hyman, London, UK, 1989.
[43]  F. Brand, “Was k?nnen wir aus lehmigen Ablagerungen der Bingh?hle ablesen?” in Die Bingh?hle bei Streitberg—Auf den Spuren eines unterirdischen Flusses, F, R. Brand, F. Illmann, D. Leja, Preu, and H. Schabdach, Eds., pp. 28–34, 2006.
[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.
[45]  D. Burger, “Dolomite weathering and micromorphology of paleosoils in the Frankonian Jura,” Catena Supplementaria, vol. 15, pp. 261–267, 1989.
[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.
[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.
[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.
[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.
[50]  J. Quilès, C. Petrea, O. Moldovan et al., “Cave bears (Ursus spelaeus) from the Pe?tera cu Oase (Banat, Romania): Paleobiology and taphonomy,” Comptes Rendus—Palevol, vol. 5, no. 8, pp. 927–934, 2006.
[51]  G. Rabeder, D. Nagel, and M. Pacher, Der H?hlenb?r, 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.
[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.
[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.
[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, ?kologie 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.
[60]  C. Diedrich and K. ?ák, “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.
[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.
[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.
[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.
[64]  G. Rabeder and M. Hofreiter, “Der neue Stammbaum der H?hlenb?ren,” Die H?hle, vol. 55, pp. 1–19, 2004.
[65]  G. Rabeder, M. Hofreiter, D. Nagel, S. Paabo, and G. Withalm, “Die neue Taxonomie der H?hlenbaren,” Abhandlungen zur Karst-und H?hlenkunde, vol. 34, pp. 68–69, 2002.
[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.
[67]  M. Hofreiter, “Genetic stability and replacement in Late Pleistocene cave bear populations,” Abhandlungen zur Karst-und H?hlenkunde, vol. 34, pp. 64–67, 2002.
[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.
[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.
[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.
[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.
[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.
[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.
[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 Ur?ilor Cave (Carpathians, Romania),” Ichnos, vol. 18, no. 1, pp. 9–26, 2010.
[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.
[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.
[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.
[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.
[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.
[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.
[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.
[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.
[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.
[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.
[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.
[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.
[93]  R. Musil, “Die H?hle, “Sveduv stul”, ein typischer H?hlenhy?nenhorst,” Anthropos N.S., vol. 5, no. 13, pp. 97–260, 1962.
[94]  J. F. Tournepiche and C. Couture, “The hyena den of Rochelot Cave (Charente, France),” Monographien des R?misch-Germanischen Zentralmuseums, vol. 42, pp. 89–101, 1999.
[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.
[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.
[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.
[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.
[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.
[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.
[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.
[102]  A. J. Sutcliffe, “Spotted hyaena: crusher, gnawer, digester and collector of bones,” Nature, vol. 227, no. 5263, pp. 1110–1113, 1970.
[103]  W. B. Ballard, “Brown bear kills gray wolf,” Canadian Field-Naturalist, vol. 94, atricle 91, 1980.
[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.
[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.
[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.
[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.
[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.
[109]  J. T. Groiss, “über pathologische Bildungen an Skelettresten jungquart?rer S?ugetiere aus der Zoolithenh?hle bei Burggeilenreuth,” Geologische Bl?tter NO-Bayern, vol. 28, no. 1, pp. 1–21, 1978.
[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.
[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.

Full-Text

Contact Us

service@oalib.com

QQ:3279437679

WhatsApp +8615387084133