Table of Contents
Paleontology Journal
Volume 2014 (2014), Article ID 106203, 34 pages
http://dx.doi.org/10.1155/2014/106203
Research Article

Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition about Prey

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

Received 21 March 2013; Accepted 11 June 2013; Published 20 February 2014

Academic Editor: Vlad Codrea

Copyright © 2014 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. 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,” Knorrs, Nürnberg, Germany, 1774.
  2. C. G. 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
  3. 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, pp. 1–34, Cum tabula aenea, Leipzig, Germany, 1794. View at Google Scholar
  4. G. L. C. F.D. Cuvier, “Sur les ossements fossiles des Hyènes,” Annales du Musée Histoire Naturelle Paris, vol. 6, p. 127, 1805. View at Google Scholar
  5. G. A. Goldfuss, “Die Umgebungen von Muggendorf. Ein Taschenbuch für Freunde der Natur und Altertumskunde,” Erlangen, Germany, 1810.
  6. G. A. Goldfuss, “Osteologische Beitraege zur Kenntnis verschiedener Saeugethiere der Vorwelt—IV. Ueber den Schaedel des Hoehlenloewen,” Verhandlungen der Kaiserlichen Leopoldinischen Carolinaeischen Akademie der Naturfreunde, vol. 10, no. 2, pp. 489–494, 1821. View at Google Scholar
  7. G. A. Goldfuss, “Osteologische Beitraege zur Kenntnis verschiedener Saeugethiere der Vorwelt—VI. Ueber die Hoehlen-Hyäne (Hyaena spelaea),” Nova Acta Physico-Medica Academiea Caesarae Leopoldino-Carolinae Naturae Curiosorum, vol. 3, no. 2, pp. 456–490, 1823. View at Google Scholar
  8. C. G. Diedrich, “The holotypes of the upper Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823: Hyaenidae) and Panthera leo spelaea (Goldfuss, 1810: Felidae) of the Zoolithen Cave hyena den (South Germany) and their palaeo-ecological interpretation,” Zoological Journal of the Linnean Society, vol. 154, no. 4, pp. 822–831, 2008. View at Publisher · View at Google Scholar · View at Scopus
  9. 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.
  10. C. G. Diedrich, “Eine oberpleistozäne Population von Crocuta crocuta spelaea (Goldfuss, 1823) aus dem eiszeitlichen Fleckenhyänenhorst Perick-Höhlen von Hemer (Sauerland, NW Deutschland) und ihr Kannibalismus,” Philippia, vol. 12, no. 2, pp. 93–115, 2005. View at Google Scholar
  11. C. G. Giebel, “Die antediluvialianische Saeugethierfauna Deutschlands,” Jahresberichte der Naturwissenschaftlichen Vereinigung Halle, vol. 3, pp. 219–236, 1852. View at Google Scholar
  12. W. Boyd Dawkins and B. A. Oxen, “On a hyæna-den at Wookey hole, near wells. no. II,” Quarterly Journal of the Geological Society of London, vol. 19, no. 1-2, pp. 260–274, 1863. View at Publisher · View at Google Scholar · View at Scopus
  13. E.K. Tratman, D. T. Donovan, and J. B. Campbell, “The hyena den (Wookey Hole) Mendip Hills, Somerset,” Proceedings of the University Bristol Spelaeological Society, vol. 12, no. 3, pp. 245–279, 1971. View at Google Scholar
  14. K. T. Liebe, “Die Lindentaler Hyänenhöhle und andere diluviale Knochenfunde in Ostthüringen,” Archäologisches und Anthropologisches Organ der Deutschen Gesellschaft für Anthropologie Ethnographie und Urgeschichte, vol. 9, p. 155, 1876. View at Google Scholar
  15. E. Thenius, “Hyänenfrassspuren aus dem Pleistozän von Kärnten. Ein Beitrag zur Frage der sog. “osteodontokeratischen Kultur” der Australopithecinen (Hominidae),” Carinthia II, vol. 72, pp. 88–101, 1961. View at Google Scholar
  16. A. Nehring, “Ausgrabungen bei thiede und westeregeln,” Verhandlungen der Berliner Gesellschaft für Anthropologie, Ethnographie und Urgeschichte, vol. 1876, pp. 206–209, 1876. View at Google Scholar
  17. C. G. Diedrich, “A clan of Late Pleistocene hyenas, Crocuta crocuta spelaea (Goldfuss 1823), from the Rösenbeck Cave (Germany) and a contribution to cranial shape variability,” Biological Journal of the Linnean Society, vol. 103, no. 1, pp. 191–220, 2011. View at Publisher · View at Google Scholar · View at Scopus
  18. E. Haarlé, “Présentation dos de repas d`hyène tachetées,” Bulletin de la Socièté d’Histoire Naturelle de Toulouse, vol. 26, pp. 22–25, 1892. View at Google Scholar
  19. S. H. Reynolds, “A monograph of the British Pleistocene Mammalia—Volume II—Part II. The Cave Hyena,” Palaeontological Society Monographs, vol. 1902, pp. 1–25, 1902. View at Google Scholar
  20. A. Currant, “The Quaternary mammal collections at the Sommerset Country Museum Taunton,” in The Quaternary Mammals of Southern and Eastern England, Field Guide, D. C. Schreve, Ed., pp. 101–109, 2004. View at Google Scholar
  21. K. Ehrenberg, O. Sickenberg, and A. Stifft-Gottlieb, “Die Fuchs- oder Teufelslucken bei Eggenburg, Niederdonau. 1 Teil,” Abhandlungen der Zoologisch-Botanischen Gesellschaft, vol. 17, no. 1, pp. 1–130, 1938. View at Google Scholar
  22. H. Zapfe, “Die Lebensspuren der eiszeitlichen Höhlenhyäne. Die urgeschichtliche Bedeutung der Lebensspuren knochenfressender Raubtiere,” Palaeobiologica, vol. 7, pp. 111–146, 1939. View at Google Scholar
  23. A.J. Sutcliffe and F. E. Zeuner, “Excavations in the Torbryan Cave, Devonshire I Tonewton Cave,” Proceedings of the Devon Arachaeological Explorations Society, vol. 5, pp. 127–145, 1958. View at Google Scholar
  24. R. Musil, “Die Höhle “Svedův stul”, ein typischer Höhlenhyänenhorst,” Anthropos New Series, vol. 5, no. 13, pp. 97–260, 1962. View at Google Scholar
  25. P. Wernert, “Beutetierstücke der Höhlenhyänen im anatomischen Verband aus Achenheimer Lössen,” Quartär, vol. 19, pp. 55–64, 1968. View at Google Scholar
  26. T. Rathgeber, “Neue jungpleistozäne Säugetier-Reste aus der Aufhauser Höhle (Kat.-Nr. 7424/13) bei Geislingen an der Steige (Schwäbische Alb),” Mitteilungen des Verbandes der deutschen Höhlen- und Karstforscher, vol. 28, no. 1, pp. 9–12, 1982. View at Google Scholar
  27. D. P. B. Erdbrink, “Eleven bones: more fossil remains of cave lions and cave hyenas from the North Sea area,” Bijdragen tot de Dierkunde, vol. 53, no. 1, pp. 1–12, 1983. View at Google Scholar
  28. J. L. Guadelli, “Étude taphonomique du repairesd’hyènes de Camiac (Gironde, France), élements de comparisation entrée un site naturel et un gisement préhistorique,” Bulletin de l’Association Française pour l’Étude du Quaternaire, vol. 2, pp. 91–100, 1989. View at Google Scholar
  29. E. S. Tsoukala, “The Pleistocene large mammals from the Agios Georgios Cave, Kilkis (Macedonia, N. Greece),” Geobios, vol. 25, no. 3, pp. 415–433, 1992. View at Google Scholar · View at Scopus
  30. J. F. Tournepiche, “Les grand mammiferes Pleistocenes de Poitou-charentes,” Paléo, vol. 8, pp. 109–141, 1996. View at Google Scholar
  31. P. Fosse, “Variabilité des assamblages osseux créés par l’hyena des cavernes,” Paléo, vol. 9, pp. 15–54, 1997. View at Google Scholar
  32. 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, 1998. View at Google Scholar
  33. P. Fosse, “Cave occupation during Palaeolithic times: man and/or Hyena?” Verlag des Römisch-Germanischen Zentralmuseums Bonn Monographien, vol. 42, pp. 73–88, 1999. View at Google Scholar
  34. 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
  35. J. G. Enloe, F. David, and G. Baryshnikov, “Hyenas and hunters: zooarchaeological investigations at Prolom II Cave, Crimea,” International Journal of Osteoarchaeology, vol. 10, no. 5, pp. 310–324, 2000. View at Google Scholar · View at Scopus
  36. 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
  37. G. Mangano, “An exclusively hyena-collected bone assemblage in the Late Pleistocene of Sicily: taphonomy and stratigraphic context of the large mammal remains from San Teodoro Cave (North-Eastern Sicily, Italy),” Journal of Archaeological Science, vol. 38, no. 12, pp. 3584–3595, 2011. View at Publisher · View at Google Scholar · View at Scopus
  38. E. Barycka, “Middle and late Pleistocene Felidae and Hyaenidae of Poland. Fauna Poloniae. Fauna Polski 2ns,” Museum and Institute of Zoology Polish Academy of Science, Warszawa, Poland, 2008.
  39. C. G. 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
  40. C. G. Diedrich, “Europe's first Upper Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823) skeleton from the Koněprusy Caves: a hyena cave prey depot site in the Bohemian Karst (Czech Republic),” Historical Biology, vol. 24, no. 1, pp. 63–89, 2012. View at Publisher · View at Google Scholar · View at Scopus
  41. C. G. 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, 2014. View at Google Scholar
  42. C. G. Diedrich, “An Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) population, their excrements and prey from the Late Pleistocene hyena den of the Sloup Cave in the Moravian Karst, Czech Republic,” Historical Biology, vol. 24, no. 2, pp. 161–185, 2012. View at Publisher · View at Google Scholar · View at Scopus
  43. C. G. Diedrich, “The Crocuta crocuta spelaea (Goldfuss, 1823) population and its prey from the Late Pleistocene teufelskammer Cave hyena den besides the famous Paleolithic Neandertal 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
  44. C. G. Diedrich, “Periodical use of the Balve Cave (NW Germany) as a Late Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823) den: Hyena occupations and bone accumulations vs. human Middle Palaeolithic activity,” Quaternary International, vol. 233, no. 2, pp. 171–184, 2011. View at Publisher · View at Google Scholar · View at Scopus
  45. C. G. Diedrich, “The Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) population with its type specimens from the Zoolithen Cave at Gailenreuth (Bavaria, South Germany): a hyena cub raising den of specialised cave bear scavengers in boreal forest environments of Central Europe,” Historical Biology, vol. 23, no. 4, pp. 335–367, 2011. View at Publisher · View at Google Scholar · View at Scopus
  46. C. G. 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
  47. C. G. Diedrich, “Late Ice Age wolves as cave bear scavengers in the Sophie’s Cave of Germany—extinctions of cave bears as result of climate/habitat change and large carnivore predation stress in Europe,” ISRN Zoology, 2013. View at Google Scholar
  48. C. Diedrich, “Late Pleistocene hyenas Crocuta crocuta spelaea (Goldfuss, 1823) from Upper Rhine valley open air sites and the contribution to skull shape variability,” Cranium, vol. 25, no. 2, pp. 31–42, 2008. View at Google Scholar
  49. C. G. Diedrich, “Oberpleistozäne fleckenhyänenreste (Crocuta crocuta spelaea (Goldfuss, 1823)) aus flussterrassenablagerungen in der münsterländer bucht (NW Deutschland),” Philippia, vol. 11, no. 3, pp. 227–234, 2004. View at Google Scholar
  50. C. G. Diedrich, “The Late Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823) population from the Emscher River terrace hyena open air den Bottrop and other sites in NW-Germany—woolly rhinoceros scavengers and their bone accumulations along rivers in lowland mammoth steppe environments,” Quaternary International, vol. 276-277, pp. 93–119, 2012. View at Google Scholar
  51. C. G. Diedrich, “The Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) open air birth and communal den site Bad Wildungen-Biedensteg (Hesse, NW Germany)—woolly rhinoceros killers and scavengers in a mammoth steppe environment of Europe—recycling of badger/fox burrows in loess by hyenas in the Ice Age,” Journal of Geological Research, vol. 276-277, pp. 93–119, 2013. View at Publisher · View at Google Scholar · View at Scopus
  52. C. G. Diedrich, “Seltene Freilandfunde der späteiszeitlichen Fleckenhyäne Crocuta crocuta spelaea (Goldfuss, 1823) in Sachsen-Anhalt und Beitrag zu Horsttypen der letzten Hyänen im Jung-Pleistozän von Mitteldeutschland,” Zeitschrift für Mitteldeutsche Vorgeschichte, vol. 93, 2011. View at Google Scholar
  53. C. G. 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, 2012. View at Publisher · View at Google Scholar · View at Scopus
  54. C. G. Diedrich, “The Unicornu fossile Valentini, 1714 “holotype” skeleton from the Ice Age spotted hyena commuting den at the open air gypsum karst Quedlinburg Sewecken-Berge (Central Germany),” Earth Science Reviews, 2012. View at Google Scholar
  55. C. G. Diedrich, “Late Pleistocene Hystrix (Acanthion) brachyura Linnaeus, 1758 from the Fuchsluken cave at the Rote Berg near Saalfeld (Thuringia, Germany)—a porcupine and hyena den and contribution to their palaeobiogeography,” The Open Palaeontological Journal, vol. 1, pp. 33–41, 2008. View at Google Scholar
  56. C. G. Diedrich, “The Late Pleistocene spotted hyena populations and their den sites in the Thuringian Mountains—rhino scavengers during Neanderthal settlement times in the Zechstein karst near Pößneck, Thuringia (Central Germany),” Eiszeitalter und Gegenwart. Quaternary Science Research, 2014. View at Google Scholar
  57. C. G. Diedrich, “Mammoth scavengers in Europe—the last Ice Age spotted hyenas and steppe lions and their feeding strategies on their largest prey,” Journal of Archaeological Science, 2014. View at Google Scholar
  58. C. G. 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 Neanderthalian site Neumark-Nord Lake 1, Central Germany,” Archaeological and Anthropologcal Sciences, 2013. View at Google Scholar
  59. C. G. Diedrich, “Cannibalism and inter-/intraspecies fight pathologies within Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) populations of Europe,” Pal Arch, 2014. View at Google Scholar
  60. C. G. Diedrich, “An Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) population, their excrements and prey from the Late Pleistocene hyena den of the Sloup Cave in the Moravian Karst, Czech Republic,” Historical Biology, vol. 24, no. 2, pp. 161–185, 2012. View at Publisher · View at Google Scholar · View at Scopus
  61. C. G. Diedrich, “A diseased Panthera leo spelaea (Goldfuss, 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord, central Germany,” Historical Biology, vol. 23, no. 2-3, pp. 195–217, 2011. View at Publisher · View at Google Scholar · View at Scopus
  62. C. G. Diedrich, “Late Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeletons from the Czech Republic (centeral Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears,” Bulletin of Geosciences, vol. 86, no. 4, pp. 817–840, 2011. View at Publisher · View at Google Scholar · View at Scopus
  63. C. G. Diedrich, “Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) skeleton remains from Praha-Podbaba and other lion finds from loess and river terrace sites in Central Bohemia (Czech Republic),” Bulletin of Geosciences, vol. 82, no. 2, pp. 99–117, 2007. View at Publisher · View at Google Scholar · View at Scopus
  64. C. G. Diedrich, “Steppe lion remains imported by Ice Age spotted hyenas into the Late Pleistocene Perick Caves hyena den in northern Germany,” Quaternary Research, vol. 71, no. 3, pp. 361–374, 2009. View at Publisher · View at Google Scholar · View at Scopus
  65. C. G. Diedrich, “Cave bear killers and scavengers from the last ice age of central Europe: feeding specializations in response to the absence of mammoth steppe fauna from mountainous regions,” Quaternary International, vol. 255, pp. 59–78, 2012. View at Publisher · View at Google Scholar · View at Scopus
  66. C. G. Diedrich and T. Rathgeber, “Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss 1810) skeleton remains of the Upper Rhine Valley (SW Germany) and contributions to their sexual dimorphism, taphonomy and habitus,” Historical Biology, vol. 24, no. 1, pp. 1–28, 2012. View at Publisher · View at Google Scholar · View at Scopus
  67. C. G. 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
  68. 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, 2012. View at Google Scholar
  69. W. D. Dawkins and A. Sandfort, “A monograph of the British pleistocene mammalia,” in The British Pleistocene Felidae. Palaeontographical Society Monographs, vol. 1(1866–1872), pp. 1–194, 1900. View at Google Scholar
  70. M. Boule, “Les grands chats des caverns,” Annales de Paléontologie, vol. 1, pp. 69–95, 1906. View at Google Scholar
  71. F. Heller, “Ein Schädel von Felis spelaea Goldf. aus der Frankenalb,” Erlanger Geologische Abhandlungen, vol. 7, pp. 1–24, 1953. View at Google Scholar
  72. W. O. Dietrich, “Fossile Löwen im Europäischen und Afrikanischen Pleistozän,” Paläontologische Abhandlungen A, vol. 3, no. 2, pp. 323–366, 1968. View at Google Scholar
  73. R. Ballesio, “Étude de Panthera (leo) spelaea (Goldfuss) nov. subsp. (Mammalia, Carnivora, Felidae) du gisement du Pleistocène moyen des Abimes de la Fage à Noailles (Corrèze),” Nouveaux Archaeologie du Musée Histoire Naturelle Lyon, vol. 13, pp. 47–55, 1975. View at Google Scholar
  74. J. Altuna, “Fund eines Skelettes des Höhlenlöwen (Panthera leo spelaea Goldfuss) in Arrikrutz, Baskenland,” Bonner Zoologische Beiträge, vol. 32, no. 1-2, pp. 31–46, 1981. View at Google Scholar
  75. G. Tichy, “Über der Fund eines Höhlenlöwen (Panthera felis spelaea Goldfuss) aus dem Tennengebirge bei Salzburg,” Mitteilungen der Gesellschaft für Salzburger Landeskunde, vol. 125, pp. 845–864, 1985. View at Google Scholar
  76. A. Argant, “Étude de ‘exemplaire de Panthera spelaea (Goldfuss, 1810) (Mammalia, Carnivora, Felidae) du gisement Pleistocène moyen recent de la grotte d” Aze (Saone et Loire),” Revue de Paléobiologie, vol. 7, no. 2, pp. 449–466, 1988. View at Google Scholar
  77. C. Gross, Das Skelett des Höhlenlöwen (Panthera leo spelaea, Goldfuss, 1810) aus Siegsdorf/Ldkr. Traunstein im Vergleich mit anderen Funden aus Deutschland und den Niederlanden [dissertation], Tierärztliche Fakultät der Ludwig-Maximilians-Universität München, 1992.
  78. K. Fischer, “Neufunde von jungpleistozänen Höhlenlöwen Panthera leo spelaea (Goldfuss, 1810) in Rübeland, Harz,” Braunschweiger Naturkundliche Schriften, vol. 4, no. 3, pp. 455–471, 1994. View at Google Scholar · View at Scopus
  79. G. Gužvica, “Panihera spelaea (Goldfuss, 1810) from North-Western Croatia,” Geologia Croatica, vol. 51, no. 1, pp. 59–73, 1998. View at Google Scholar · View at Scopus
  80. G. F. Baryshnikov and G. Boeskorov, “The Pleistocene cave lion, Panthera spelaea (Carnivora, Felidae) from Yakutia, Russia,” Cranium, vol. 18, pp. 7–24, 2001. View at Google Scholar
  81. F. Bona, “Systematic position of a complete lion-like cat skull from the Eemian ossiferous rubble near Zandobbio (Bergamo, North Italy),” Rivista Italiana di Paleontologia e Stratigrafia, vol. 112, no. 1, pp. 157–166, 2006. View at Google Scholar · View at Scopus
  82. C. G. Diedrich, “Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from an open air loess bone accumulation and human campsite in Freyburg a. d. U. (Saxony-Anhalt),” Jahresschrift für Mitteldeutsche Vorgeschichte, vol. 92, pp. 9–24, 2008. View at Google Scholar
  83. C. G. Diedrich, “Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy, palaeobiology and sexual dimorphism,” Annales de Paleontologie, vol. 95, no. 3, pp. 117–138, 2009. View at Publisher · View at Google Scholar · View at Scopus
  84. C. G. Diedrich, M. Robu, O. Moldovan et al., “New Upper Pleistocene steppe lion skeleton finds between the cave bear hibernation plateaus of the Urşilor Cave bear den, Romania,” in Proceedings of the 15th International Cave Bear Symposium, p. 10, Bratislava, Slovakia, 2009.
  85. 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
  86. M. Sabol, “A record of pleistocene lion-like felids in the territory of Slovakia,” Quaternaire, no. 4, pp. 215–228, 2011. View at Google Scholar · View at Scopus
  87. 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
  88. S. K. Bearder, “Feeding habits of spotted hyaenas in a woodland habitat,” East African Wildlife Journal, vol. 15, pp. 263–280, 1977. View at Google Scholar
  89. L. Werdelin and N. Solounias, “The Hyaenidae: taxonomy, systematics and evolution,” Fossils & Strata, vol. 30, pp. 1–104, 1991. View at Google Scholar · View at Scopus
  90. 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
  91. R. G. Klein and K. Scott, “Glacial/interglacial size variation in fossil spotted hyenas (Crocuta crocuta) from Britain,” Quaternary Research, vol. 32, no. 1, pp. 88–95, 1989. View at Google Scholar · View at Scopus
  92. G. F. Baryshnikov, “Chronological and geographical variability of Crocuta spelaea (Carnivora, Hyaenidae) from the Pleistocene of Russia,” Deinsea, vol. 6, pp. 155–174, 1999. View at Google Scholar
  93. C. G. Diedrich, “Disappearance of the last hyenas and lions of Europe in the Late Quaternary—a chain reaction of large mammal mammoth and woolly rhino prey extinction,” in Proceedings of the European Geoscience Union General Assembly, Vienna, Austria, 2010.
  94. T. Lynn, Morphological variation in a durophagous carnivore, the spotted hyena Crocuta crocuta [dissertation], Michigan University, Michigan, Ill, USA, 2009.
  95. H. Kruuk, “The spotted hyena,” in A Story of Predation and Social Behavior, p. 335, The University of Chicago Press, Chicago, Ill, USA, 1972. View at Google Scholar
  96. 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
  97. H. Hofer and M. East, “Population dynamics, population size, and the commuting system of Serengeti spotted hyenas,” in Serengeti II: Dynamics, MAnagement, and Conservation of an Ecosystem, A. R. E. Sinclair and P. Arcese, Eds., pp. 332–363, University of Chicago Press, Chicago, Ill, USA, 1995. View at Google Scholar
  98. S. T. Von Soemmerring, “Ueber die geheilte Verletzung eines fossilen Hyaenenschädels,” Nova Acta Physico-Medica Academiae Caesar Leopoldiana, vol. 14, pp. 1–44, 1828. View at Google Scholar
  99. L. G. Frank, “When hyenas kill their own,” New Scientist, vol. 141, pp. 38–41, 1994. View at Google Scholar
  100. P. A. White, “Maternal rank is not correlated with cub survival in the spotted hyena, Crocuta crocuta,” Behavioral Ecology, vol. 16, no. 3, pp. 606–613, 2005. View at Publisher · View at Google Scholar · View at Scopus
  101. S. M. Cooper, “Denning behavior of spotted hyaenas (Crocuta crocuta) in Botswana,” African Journal of Ecology, vol. 31, pp. 178–180, 1993. View at Google Scholar
  102. C. Stuart and T. Stuart, Field Guide to the Larger Mammals of Africa, Struik, Cape Town, South Africa, 1997.
  103. M. L. East, H. Hofer, and A. Türk, “Functions of birth dens in spotted hyaenas (Crocuta crocuta),” Journal of Zoology London, vol. 219, pp. 690–697, 1989. View at Google Scholar
  104. J. T. Pokines and J. C. Kerbis 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
  105. J. Brunner, “Beobachtungen zu den Lebensspuren der Hyänen an den Knochen der Huftiere aus dem Unter-Pliozän von Pikermi,” Palaeobiologica, vol. 8, pp. 120–126, 1944. View at Google Scholar
  106. J. P. Brugal, P. Fosse, and J. P. Guadeli, “Comparative study of bone assemblages made by recent and Plio-Pleistocene Hyaenids (Hyena, Crocuta),” Hot Springs South Dakota Occasional Publications, vol. 1, no. 1, pp. 158–187, 1997. View at Google Scholar
  107. A. Arribas and P. Palmqvist, “Taphonomy and palaeoecology of an assemblage of large mammals: Hyaenid activity in the lower Pleistocene site at Venta Micena (Orce, Guadix-Baza basin, Granada, Spain),” Geobios, vol. 31, no. 3, pp. 3–47, 1998. View at Google Scholar · View at Scopus
  108. R.-D. Kahlke, “Untermassfeld—a late Early Pleistocene (Epivillafranchian) fossil site near Meiningen (Thuringia, Germany) and its position in the development of the European mammal fauna,” BAR International Series, vol. 1578, pp. 1–114, 2006. View at Google Scholar
  109. P. Fosse, “La grotte no. 1 de Lunel-Viel (Hérault, France): repaire d’hyènes du Pleistocène Moyen,” Paléo, vol. 8, pp. 47–81, 1996. View at Google Scholar
  110. J. Lindly, “Hominid and carnivore activity at Middle and Upper Palaeolithic cave sites in eastern Spain,” Munibe, vol. 40, pp. 45–70, 1988. View at Google Scholar
  111. K. Cruz-Uribe, “Distinguishing hyena from hominid bone accumulations,” Journal of Field Archaeology, vol. 18, pp. 467–486, 1991. View at Google Scholar
  112. H. T. Bunn, “Comparative analyses of modern bone assemblages from a San hunter-gatherer camp in the Kalahari Desert, Botswana, and from a spotted hyena den near Nairobi, Keynia,” British Archaeological Reports, vol. 283, pp. 143–148, 1983. View at Google Scholar
  113. T. R. Pickering, “Reconsideration of criteria for differentiating faunal assemblages accumulated by hyenas and hominids,” International Journal of Osteoarchaeology, vol. 12, no. 2, pp. 127–141, 2002. View at Publisher · View at Google Scholar · View at Scopus
  114. P. Villa, J.-C. Castel, C. Beauval, V. Bourdillat, and P. Goldberg, “Human and carnivore sites in the European Middle and Upper Paleolithic: similarities and differences in bone modification and fragmentation,” Revue de Paleobiologie, vol. 23, no. 2, pp. 705–730, 2004. View at Google Scholar · View at Scopus
  115. 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
  116. B. F. Kuhn, L. R. Berger, and J. D. Skinner, “Examining criteria for identifying and differentiating fossil faunal assemblages accumulated by hyenas and hominins using extant hyenid accumulations,” International Journal of Osteoarchaeology, vol. 20, no. 1, pp. 15–35, 2010. View at Publisher · View at Google Scholar · View at Scopus
  117. 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
  118. J. R. Henschel, R. Tildin, and F. von Blottnitz, “Implications of a spotted hyaena bone assemblage in the Namib Desert,” South African Archaeoligical Bulletin, vol. 34, pp. 127–131, 1979. View at Google Scholar
  119. A. Hill, “A modern hyena den in Amboseli National Park, Kenya,” in Proceedings of the 8th Panafrican Congress of Prehistory and Quaternary Studies, vol. 1980, pp. 137–138, 1980.
  120. L. Scott and R. G. Klein, “A hyena accumulated bone assemblage from late Holocene deposits at Deelpan, Orange Free State, South Africa,” Annals of the South African Museum, vol. 86, pp. 217–227, 1981. View at Google Scholar
  121. G. Avery, D. M. Avery, S. Braine, and R. Loutit, “Bone accumulation by hyenas and jackals a taphonomic study,” South African Journal of Science, vol. 80, pp. 186–187, 1984. View at Google Scholar
  122. E. E. Boydston, K. M. Kapheim, and K. E. Holekamp, “Patterns of den occupation by the spotted hyaena (Crocuta crocuta),” African Journal of Ecology, vol. 44, no. 1, pp. 77–86, 2006. View at Publisher · View at Google Scholar · View at Scopus
  123. 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
  124. 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
  125. P. R. K. Richardson, “Carnivore damage to antelope bones and its archaeological implications,” Palaeontologica Africana, vol. 23, pp. 109–125, 1980. View at Google Scholar
  126. C.W. Marean and L. M. Spencer, “Impact of carnivore revenging on zooarchaeological measures of element abundance,” American Antiquity, vol. 56, pp. 645–658, 1991. View at Google Scholar
  127. R. J. Blumenschine, “An experimental model of the timing of hominid and carnivore influence on archaeological bone assemblages,” Journal of Archaeological Science, vol. 15, no. 5, pp. 483–502, 1988. View at Google Scholar · View at Scopus
  128. 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, Maine, Canada, 1989. View at Google Scholar
  129. C. W. Marean, L. M. Spencer, R. J. Blumenschine, and S. D. Capaldo, “Captive hyaena bone choice and destruction, the Schlepp effect and olduvai archaeofaunas,” Journal of Archaeological Science, vol. 19, no. 1, pp. 101–121, 1992. View at Google Scholar · View at Scopus
  130. 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
  131. R. F. Ewer, “Some adaptive features in the dentition of hyenas,” Annals and Magazine of Natural History, vol. 12, no. 7, pp. 188–194, 1954. View at Google Scholar
  132. A. S. Van Jaarsveld, J. D. Skinner, and M. Lindeque, “Growth, development and parental investment in the spotted hyaena, Crocuta crocuta,” Journal of Zoology, vol. 216, no. 1, pp. 45–53, 1988. View at Google Scholar · View at Scopus
  133. C. G. 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
  134. C. G. Diedrich, “Benagte und zerknackte Knochen des eiszeitlichen Pferdes Equus ferus przewalskii Poljakoff, 1881 aus einem oberpleistozänen Fleckenhyänenhorst des Nordsauerlandes und westfälischen Freilandfundstellen,” Philippia, vol. 12, no. 1, pp. 47–62, 2005. View at Google Scholar
  135. C. G. Diedrich, “Von eiszeitlichen Fleckenhyänen eingeschleppte Reste des Steppenwisents Bison priscus Bojanus 1827 aus dem oberpleistozänen Fleckenhyänenhorst des Perick-Höhlensystems (NW Deutschland),” Philippia, vol. 12, no. 1, pp. 21–30, 2005. View at Google Scholar
  136. C. G. Diedrich, “Von oberpleistozänen Fleckenhyänen gesammelte, versteckte, verbissene, zerknackte Knochen und Geweihe des Riesenhirsches Megaloceros giganteus (Blumenbach, 1799) aus den Perick-Höhlen im Nordsauerland (NW Deutschland),” Philippia, vol. 12, no. 1, pp. 31–46, 2005. View at Google Scholar
  137. C. Diedrich, “The Late Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823) population and its prey from the Fuchsluken Cavities near Saalfeld (Thuringia, Central Germany)—a specialized bovid and equuid hunters commuting and cub raising hyena and porcupine overlapping den site,” Pal Arch, 2014. View at Google Scholar
  138. S. K. Bearder and R. M. Randal, “The use of fecal marking sites by spotted hyaenas and civets,” Carnivore, vol. 1, pp. 32–48, 1978. View at Google Scholar
  139. L. K. Horwitz and P. Goldberg, “A study of Pleistocene and Holocene hyaena coprolites,” Journal of Archaeological Science, vol. 16, no. 1, pp. 71–94, 1989. View at Google Scholar · View at Scopus
  140. C. F. Rodríguez, P. R. Rego, and A. M. Cortizas, “Characterization and depositional evolution of Hyaena (Crocuta crocuta) coprolites from La Valina Cave (Northwest Spain),” Journal of Archaeological Science, vol. 22, no. 5, pp. 597–607, 1995. View at Google Scholar · View at Scopus
  141. N. R. Larkin, J. Alexander, and M. D. Lewis, “Using experimental studies of recent faecal material to examine hyaena coprolites from the West Runton freshwater bed, Norfolk, U.K,” Journal of Archaeological Science, vol. 27, no. 1, pp. 19–31, 2000. View at Publisher · View at Google Scholar · View at Scopus
  142. J. Reumer, D. Mol, and W. Borst, “The first Late Pleistocene coprolite of Crocuta crocuta spelaea from the North Sea,” Deinsea, vol. 14, pp. 15–18, 2010. View at Google Scholar
  143. J. S. Carrión, J. A. Riquelme, C. Navarro, and M. Munuera, “Pollen in hyaena coprolites reflects late glacial landscape in southern Spain,” Palaeogeography, Palaeoclimatology, Palaeoecology, vol. 176, no. 1–4, pp. 193–205, 2001. View at Publisher · View at Google Scholar · View at Scopus
  144. C. Sillero-Zubiri and D. Gottelli, “Feeding ecology of spotted hyaena (Mammalia: Crocuta crocuta) in a mountain forest habitat,” Journal of African Zoology, vol. 106, no. 2, pp. 169–176, 1992. View at Google Scholar · View at Scopus
  145. L. D. Mech, The Arctic Wolf: Living With the Pack, Voyageur Press, Stillwater, Minn, USA, 1988.
  146. C. G. Diedrich, “A skeleton of an injured Coelodonta antiquitatis (Blumenbach 1807) from the Upper Pleistocene of north-western Germany,” Cranium, vol. 25, no. 1, pp. 1–16, 2008. View at Google Scholar
  147. P. White and C. G. Diedrich, “Taphonomy story of a modern African elephant Loxodonta africana carcass on a lakeshore in Zambia (Africa),” Quaternary International, vol. 276-277, pp. 287–296, 2012. View at Google Scholar
  148. D. Joubert and B. Joubert, Eternal Enemies: Lions & Hyenas, Wildlife Films Botswana for National Geographic, 2003.
  149. W. von Koenigswald, H. Müller-Beck, and E. Pressmar, “Archäologie und paläontologie in den weinberghöhlen von Mauern (Bayern), Grabungen 1937–1967,” Archaeologica Venatoria, vol. 3, pp. 1–152, 1999. View at Google Scholar
  150. G. Rabeder, D. Nagel, and M. Pacher, Der Höhlenbär, Thorbecke Species 4, Stuttgart, Germany, 2000.
  151. D. E. Nelson, A. Angerbjörn, K. Lidén, 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 · View at Scopus
  152. C. G. Diedrich, “Cracking and nibbling marks as indicators for the Upper Pleistocene spotted hyaena as a scavenger of cave bear (Ursus spelaeus Rosenmüller, 1794) carcasses in the Perick Caves den of Northwest Germany,” Abhandlungen der Naturhistorischen Gesellschaft Nürnberg, vol. 45, pp. 73–90, 2005. View at Google Scholar
  153. J. Quilès, C. Petrea, O. Moldovan et al., “Cave bears from P. cu Oase (Banat, Romania): taphonomy and paleobiology,” Comptes Rendus-Palevol, vol. 5, no. 8, pp. 927–934, 2006. View at Publisher · View at Google Scholar · View at Scopus
  154. J. F. Tournepiche, “Un Neandertalien devore par des hyenas?” Paleo, vol. 6, pp. 319–321, 1994. View at Google Scholar
  155. I. Turk, J. Dirjec, and G. Bastiani, “Nove analize piscali iz Divji bab I (Slovenija),” Archeološki Vestnik, vol. 52, pp. 25–79, 2001. View at Google Scholar
  156. 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
  157. R. Barnett, B. Shapiro, I. Barnes et al., “Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity,” Molecular Ecology, vol. 18, no. 8, pp. 1668–1677, 2009. View at Publisher · View at Google Scholar · View at Scopus
  158. G. Schaller, The Serengeti Lion. A Study of Predator-Prey Relations, University of Chicago Press, Chicago, Ill, USA, 1972.
  159. R. Estes, The Safari Companion: A Guide to Watching African Mammals, Chelsea Green Publishing Company, Vermont, USA, 1999.
  160. A. Turner, “Dental sex dimorphism in European lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications,” Annales Zoologici Fennici, vol. 21, pp. 1–8, 1984. View at Google Scholar
  161. 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 · View at Scopus
  162. D. Joubert, “Hunting behaviour of lions (Panthera leo) on elephants (Loxodonta africana) in the Chobe National Park, Botswana,” African Journal of Ecology, vol. 44, no. 2, pp. 279–281, 2006. View at Publisher · View at Google Scholar · View at Scopus
  163. A. Grandal-D'Anglade and J. R. Vidal 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
  164. 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
  165. 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
  166. A. C. Pinto Llona and P. J. Andrew, “Scavenging behaviour patterns in cave bears Ursus spelaeus,” in 14ème congrès UISPP, Hommes et Carnivores au Paléolithique, J.-P. Brugal and P. Fosse, Eds., Revue de Paléobiologie 23, pp. 845–853, Genève, Switzerland, 2004.
  167. 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
  168. C. G. Diedrich, “Pleistocene Stratigraphy and bone taphonomy in the Zoolithen Cave—and new theory about Esper’s “biblic flood” with a first geomorphological Wiesent Valley and river terrace evolution model—small glaciers and high/postglacial flood events in Upper Franconia. Eiszeitalter und Gegenwart,” Quaternary Science Journal, 2013. View at Google Scholar
  169. 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 · View at Scopus
  170. C. G. Diedrich, “Oldest and most Northern late palaeolithic cave bear hunters in Europe,” Archaeology, Ethnology and Anthropology of Eurasia, 2014. View at Google Scholar
  171. 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