Recycling of Badger/Fox Burrows in Late Pleistocene Loess by Hyenas at the Den Site Bad Wildungen-Biedensteg (NW, Germany): Woolly Rhinoceros Killers and Scavengers in a Mammoth Steppe Environment of Europe
The Late Pleistocene (MIS 5c-d) Ice Age spotted hyena open air den and bone accumulation site Bad Wildungen-Biedensteg (Hesse, NW, Germany) represents the first open air loess fox/badger den site in Europe, which must have been recycled by Crocuta crocuta spelaea (Goldfuss, 1823) as a birthing den. Badger and fox remains, plus remains of their prey (mainly hare), have been found within the loess. Hyena remains from that site include parts of cub skeletons which represent 10% of the megafauna bones. Also a commuting den area existed, which was well marked by hyena faecal pellets. Most of the hyena prey bones expose crack, bite, and nibbling marks, especially the most common bones, the woolly rhinoceros Coelodonta antiquitatis (NISP？？=？？32%). The large amount of woolly rhinoceros bones indicate hunting/scavenging specializing on this large prey by hyenas. Other important mammoth steppe hyena prey remains are from Mammuthus primigenius, Equus caballus przewalskii, Bison/Bos, Megaloceros giganteus, Cervus elaphus, and Rangifer tarandus. The few damaged bone remains of a scavenged cave bear Ursus spelaeus subsp. are unique for an open air situation. Abundant micromammal, frog, and some fish remains were concentrated in “pellets” that contain mainly mammoth steppe micromammals and also frog and fish remains that seem to originate from the nearby river/lake. 1. Introduction Late Pleistocene European bone assemblages have been produced mainly by late Ice Age spotted hyena Crocuta crocuta spelaea [1] and were first recognized by Buckland [2] in the “Kuhloch Cave” (K？nig-Ludwigs Cave, Bavaria, Germany) and the Kirkdale Cave (Kent, England). More recent studies provide information on the hyena prey bone assemblages (e.g., [3–10]) as well as on the new subdivided fossil hyena den types (e.g., [11]). These identifications of three classified Ice Age den forms are particularly important also to distinguish bone accumulations made by hyenas from those accumulated by Middle Palaeolithic humans (e.g., [9, 12–15]). Few contemporary used hyena and Neanderthal sites have been described from hyena dens in mammoth steppe lowlands and adjacent cave-rich region environments of north-central Europe, in England and Germany [9, 16]. The degree of prey bone damage and presence/absence of “nibbling sticks” and faecal pellets or hyena population structure and their individual amount allow the reconstruction, much better, of the ethology of the last hyenas of Europe. The discussions for nonarchaeological sites no longer focus only on the human/carnivore origin discussion. Although
References
[1]
G. A. Goldfuss, “Osteologische Beitr？ge zur Kenntnis verschiedener S？ugethiere der Vorwelt. VI. Ueber die H？len-Hy？ne (Hy？na spelaea),” Nova Acta Physico-Medica Academiea Caesarae Leopoldino-Carolinae Naturae Curiosorum, vol. 3, no. 2, pp. 456–490, 1823.
[2]
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.
[3]
K. T. Liebe, “Die Lindentaler Hy？nenh？hle und andere diluviale Knochenfunde in Ostthüringen,” Archiv des Anthropologischen Organs der Deutschen Gesellschaft für Anthropologie, Ethnographie und Urgeschichte, vol. 9, pp. 1–55, 1876.
[4]
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.
[5]
R. Musil, “Die H？hle “Sveduv stul”, ein typischer H？hlenhy？nenhorst,” Anthropos NS, vol. 5, no. 13, pp. 97–260, 1962.
[6]
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.
[7]
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.
[8]
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.
[9]
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, 0212.
[10]
C. 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.
[11]
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, vol. 24, no. 2, pp. 161–185, 2012.
[12]
J. P. Brugal, P. Fosse, and J. P. Guadeli, “Comparative study of bone assemblages made by recent and Plio-Pleistocene Hyaenids (Hyena, Crocuta),” in Proceedings of the 1983 Bone Modification conference, Hot Springs, A. Hannus, L. Rossum, and R. P. Winham, Eds., pp. 158–187, Arachaeological Laboratory, Augustana Collegue, Sioux Falls, SD, USA, 1997.
[13]
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.
[14]
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.
[15]
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.
[16]
S. Aldhouse-Green, Scott, K. Schwarcz H et al., “Coygan Cave, Laugharne, 1995. South Wales, a Mousterian site and hyaena den: a report on the University of Cambridge excavations,” Proceedings of the Prehistoric Society London, vol. 61, pp. 37–80, 1995.
[17]
C. Diedrich, “A clan of Late Pleistocene hyenas, Crocuta crocuta spelaea (Goldfuss 1823), from the R？senbeck Cave (Germany) and a contribution to cranial shape variability,” Biological Journal of the Linnean Society, vol. 103, no. 1, pp. 191–220, 2011.
[18]
C. Diedrich, “Late Pleistocene Crocuta crocuta spelaea (Goldfuss 18) 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.
[19]
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.
[20]
C. 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, no. 1, pp. 33–41, 2008.
[21]
C. Diedrich, “Seltene Freilandfunde der sp？teiszeitlichen Fleckenhy？ne Crocuta crocuta spelaea (Goldfuss 1823) in Sachsen-Anhalt und Beitrag zu Horsttypen der letzten Hy？nen im Jung-Pleistoz？n von Mitteldeutschland,” Zeitschrift für Mitteldeutsche Vorgeschichte, vol. 94, pp. 1–19, 2013.
[22]
C. Diedrich, “Crocuta crocuta spelaea (Goldfuss 1823) remains from the Upper Pleistocene hyaena Teufelskammer Cave den site near Hochdahl in the Neander valley (NRW, NW Germany),” Cranium, vol. 24, no. 2, pp. 39–44, 2007.
[23]
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 vs. human Middle Palaeolithic activity,” Quaternary International, vol. 233, no. 2, pp. 171–184, 2011.
[24]
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.
[25]
C. Diedrich, “Crocuta crocuta spelaea (Goldfuss 1823) remains from the Upper Pleistocene hyaena Teufelskammer Cave den site near Hochdahl in the Neander valley (NRW, NW Germany),” Cranium, vol. 24, no. 2, pp. 39–44, 2007.
[26]
C. 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.
[27]
C. 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.
[28]
D. Walther, “Eine Siedlungsstelle des Neandertalers bei Buhlen, Kr. Waldeck-Frankenberg,” Geschichtsbl？tter für Waldeck, vol. 93, pp. 6–25, 2005.
[29]
C. Diedrich, “By ice age spotted hyenas protracted, cracked, nibbled and chewed skeleton remains of Coelodonta antiquitatis (Blumenbach, 1807) from the Lower Weichselian (Upper Pleistocene) open air prey deposit site Bad Wildungen-Biedensteg (Hesse, NW Germany),” Journal of Taphonomy, vol. 4, no. 4, pp. 173–205, 2006.
[30]
R. Huckriede and V. Jacobshagen, “Die Fundschichten,” in Eine Faunenfolge aus dem jungpleistoz？nen L？？ bei Bad Wildungen, E. Jacobshagen, R. Huckriede, and V. Jacobshagen, Eds., vol. 44 of Abhandlungen des Hessischen Landesamtes für Bodenforschung, pp. 93–105, 1963.
[31]
A. Semmel, “Studien über den Verlauf jungpleistoz？ner Formung in Hessen,” Frankfurter Geographische Hefte, vol. 45, pp. 1–133, 1968.
[32]
J. Kulick and Quart？r, in Erl？uterungen zur Geologische Karte von Hessen, M. Horn, J. Kulick, and D. Meischner, Eds., pp. 184–228, Wiesbaden, Germany, 1973.
[33]
G. Storch, “über Kleins？uger der Tundra und Steppe in jungpleistoz？nen Eulengew？llen aus dem nordhessischen L？？,” Natur und Museum, vol. 99, pp. 541–551, 1969.
[34]
E. Jacobshagen, “Die Faunen und ihre Bindung an Klima und Umwelt,” in Eine Faunenfolge aus dem jungpleistoz？nen L？？ bei Bad Wildungen, E. Jacobshagen, R. Huckriede, and V. Jacobshagen, Eds., vol. 44 of Abhandlungen des Hessischen Landesamtes für Bodenforschung, pp. 5–92, 1963.
[35]
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 (Hesse, Germany),” Scientific Annals, School of Geology Aristotle University of Thessaloniki (AUTH), vol. 98, pp. 193–199, 2006.
[36]
W. Sp？ttel, “Equus przewalskii Poljakov 1881 mit besonderer Berücksichtigung der im Tierzuchtinstitut der Universit？t Halle gehaltenen Tiere,” Kühn Archiv, vol. 11, pp. 89–137, 1926.
[37]
C. Diedrich, “Tracking Late Pleistocene steppe lion predation in a mammoth steppe near a spotted hyena open air den of Northwestern Germany—comparisons to modern African lion trackways—new picture of top predator prey specializations, and global Pleistocene megafauna track site review,” Ichnos, 2013.
[38]
A. Forsten, “The small caballoid horse of the Upper Pleistocene and Holocene,” Journal of Animal Breeding Genetics, vol. 105, pp. 161–176, 1987.
[39]
B. Cramer, Morphometrische untersuchungen an quart？ren Pferden in Mitteleuropa [Ph.D. dissertation], Universit？t Tübingen, Tübingen, Germany, 2002.
[40]
C. 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)—Late Pleistocene woolly rhinoceros scavengers,” Historical Biology, vol. 24, no. 1, pp. 63–89, 2012.
[41]
C. Diedrich, “The Ice Age spotted Crocuta crocuta spelaea (Goldfuss 1823) population, their excrements and prey from the Late Pleistocene hyena den Sloup Cave in the Moravian Karst; Czech Republic,” Historical Biology, vol. 24, no. 2, pp. 161–185, 2012.
[42]
C. 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.
[43]
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.
[44]
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.
[45]
W.-D. Heinrich, G. Peters, K.-D. J？ger, and E. G. B？hme, “Erdbaue von S？ugetieren—zusammenfassende Kennzeichnung eines neuen Fundst？ttentyps im baltischen Vereisungsgebiet,” Wissenschaftliche Zeitschrift der Humboldt-Universit？t zu Berlin, Mathemathisch-Naturwissenschaftliche Reihe, vol. 6, pp. 777–781, 1983.
[46]
T. Rathgeber, “Pleistoz？ne und holoz？ne Tierreste aus H？hlen im Kartenblatt 7521 Reutlingen (Schw？bische Alb),” Laichinger H？hlenfreund, vol. 43, pp. 27–34, 2008.
[47]
P. Lüps and T. J. Roper, “Cannibalism in a female badger (Meles meles): infanticide orpredation?” Journal of Zoology London, vol. 221, pp. 314–315, 1990.
[48]
G. Clau？en, Der J？ger und Sein Wild, Paul Parey, Berlin, Germany, 1986.
[49]
M. Stubbe, “Biometrie und Morphologie des mitteleurop？ischen Dachses Meles meles (L., 1758),” S？ugetierkundliche Informationen, vol. 1, pp. 3–26, 1980.
[50]
A. V. Abramov and A. Y. Puzachenko, “Sexual dimorphism of craniological characters in Eurasian badgers, Meles spp. (Carnivora, Mustelidae),” Zoologischer Anzeiger, vol. 244, no. 1, pp. 11–29, 2005.
[51]
P. Lüps, “Geschlechtsdimorphismus beim Dachs Meles meles L. Auspr？gungsgrad und Versuch einer Interpretation. Populations？kologie marderartiger S？ugetiere 2,” Wissenschaftliche Beitr？ge der Universit？t Halle, vol. 37, no. P39, pp. 531–542, 1989.
[52]
R. Boesi and C. Biancardi, “Diet of the Eurasian badger Meles meles (Linnaeus, 1758) in the natural reserve of Lago di Piano, Northern Italy,” Mammalian Biology, vol. 67, no. 2, pp. 120–125, 2002.
[53]
G. Peters, W.-D. Heinrich, P. Beurton, and K.-D. J？ger, “Fossile und rezente Dachsbauten mit Massenanreicherungen von Wirbeltierknochen,” Mitteilungen des Zoologischen Museums Berlin, vol. 48, pp. 415–435, 1972.
[54]
F. Winterfeld, “über quart？re Mustelidenreste Deutschlands. Zeitschrift der deutschen geologischen Gesellschaft,” Geologischen Gesellschaft, vol. 37, pp. 826–864, 1885.
[55]
A. Nehring, “übersicht über 24 mitteleurop？ische Quart？r-Faunen,” Zeitschrift der Deutschen Geologische Gesellschaft, vol. 32, pp. 468–509, 1890.
[56]
C. Diedrich, “Eine oberpleistoz？ne Population von Crocuta crocuta spelaea (Goldfuss 1823) aus dem eiszeitlichen Fleckenhy？nenhorst Perick-H？hlen von Hemer (Sauerland, NW Deutschland) und ihr Kannibalismus,” Philippia, vol. 12, no. 2, pp. 93–115, 2005.
[57]
L. G. Frank, “When hyenas kill their own,” New Scientist, vol. 141, pp. 38–41, 1994.
[58]
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.
[59]
P. P. White, “Maternal response to neonatal sibling conflict in the spotted hyena, Crocuta crocuta,” Behavioral Ecology and Sociobiology, vol. 62, no. 3, pp. 353–361, 2008.
[60]
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.
[61]
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.
[62]
P. Fosse, “La grotte no. 1 de Lunel-Viel (Hérault, France): repaire d’hyènes du pleistocène moyen,” Paléo, vol. 8, pp. 47–81, 1996.
[63]
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, supplement 3, pp. 3–47, 1998.
[64]
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 Fran？aise pour l’étude du Quaternaire, vol. 2, pp. 91–100, 1989.
[65]
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. Rencontres Internationales d'Archéologie et d'Histoire d'Antibes, pp. 44–61, APDCA, Sophia Antipolis, France, 1998.
[66]
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.
[67]
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.
[68]
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.
[69]
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.
[70]
C. Diedrich, “Von eiszeitlichen Fleckenhy？nen benagte Mammuthus primigenius (Blumenbach 1799)-Knochen und -Knabbersticks aus dem oberpleistoz？nen Perick-H？hlenhorst (Sauerland) und Beitrag zur Taphonomie von Mammutkadavern,” Philippia, vol. 12, no. 1, pp. 63–84, 2005.
[71]
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.
[72]
H. Kruuk, The Spotted Hyena. A Story of Predation and Social Behavior, University of Chicago Press, Chicago, Ill, USA, 1972.
[73]
J. R. Henschel, R. Tilson, and F. von Blottnitz, “Implications of a spotted hyaena bone assemblage in the Namib Desert,” South African Archaeological Bulletin, vol. 34, pp. 127–131, 1979.
[74]
A. Hill, “A modern hyena den in Amboseli National Park, Kenya, Nairobi,” in Proceedings of the 8th Pan-African Congress on Prehistory and Quaternary Studies, pp. 137–138, 1980.
[75]
A. K. Behrensmeyer and D. Boaz, “The recent bones of Amboseli Park Kenya in relation to East African paleoecology,” in Fossils in the Making: Vertebrate Taphonomy and Paleoecology, A. K. Behrensmeyer and A. P. Hill, Eds., pp. 72–92, University of Chicago Press, Chicago, Ill, USA, 1980.
[76]
C. K. Brain, “Some criteria for the recognition of bone collecting agencies in African caves,” in Fossils in the Making: Vertebrate Taphonomy and Paleoecology, A. K. Behrensmeyer and A. P. Hill, Eds., pp. 107–130, University of Chicago Press, Chicago, Ill, USA, 1980.
[77]
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.
[78]
J. R. Henschel, J. D. Skinner, and A. S. van Jaarsveld, “The diet of the spotted hyaenas Crocuta crocuta in Kruger National Park,” South African Journal of Zoology, vol. 21, pp. 303–308, 1986.
[79]
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.
[80]
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.
[81]
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.
[82]
C. Diedrich, “Die sp？teiszeitlichen Fleckenhy？nen und deren Exkremente aus Neumark-Nord,” Arch？ologie in Sachsen-Anhalt Sonderband, vol. 62, pp. 440–448, 2010.
[83]
S. K. Bearder and R. M. Randall, “The use of fecal marking sites by spotted hyaenas and civets,” Carnivore, vol. 1, pp. 32–48, 1978.
[84]
T. Keller and G. F？rsterling, “Unterbrochene Mahlzeit? Eiszeitliche Knochenfunde aus dem Gipskarst von Morschen-Konnefeld,” in Arch？ologische und Pal？ontologische Bodendenkmalpflege des Landesamtes für Denkmalpflege Hessen, vol. 202, pp. 18–21, 2002.
[85]
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.
[86]
G. Bosinski, “Tierdarstellungen von G？nnersdorf. Nachtr？ge zu Mammut und Pferd sowie die übrigen Tierdarstellungen,” Monographien des R？misch-Germanischen Zentralmuseums, vol. 72, pp. 1–175, 2008.
[87]
R. Springhorn, “A wild horse (Equus przewalskii Poljakov 1881) of Mesolithic age from Kempen (Germany, Northrhine-Westphalia, Lippe County),” Eiszeitalter und Gegenwart, vol. 52, pp. 40–46, 2003.
[88]
M. Baales, “Auf der F？hrte sp？tglazialer Pferde bei Mertloch (Neuwieder Becken),” Bonner Zoologische Beitr？ge, vol. 50, no. 3, pp. 109–133, 2002.
[89]
A. Leroi-Gourhan, “Préhistoire de l’art occidental,” Ars Antiqua, vol. 1, pp. 1–601, 1971.
[90]
C. 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.
[91]
P. Wernert, “Beutestücke der H？hlenhy？nen im anatomischen Verband aus Achenheimer L？ssen,” Quart？r, vol. 19, pp. 55–64, 1968.
[92]
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.
[93]
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.
[94]
C. Diedrich, “Ein Sch？delfund und von Hy？nen angenagter Oberschenkelknochen des Wollnashorns Coelodonta antiquitatis Blumenbach 1799 aus den pleistoz？nen Weserkiesen bei Minden (Norddeutschland),” Philippia, vol. 11, no. 3, pp. 211–217, 2004.
[95]
C. Diedrich, “Eingeschleppte und benagte Knochenreste von Coelodonta antiquitatis (Blumenbach, 1807) aus dem oberpleistoz？nen Fleckenhy？nenhorst Perick-H？hlen im Nordsauerland (NW Deutschland) und Beitrag zur Taphonomie von Wollnashornknochen in Westfalen,” Mitteilungen der H？hlen und Karstforscher, vol. 4, pp. 100–117, 2008.
[96]
F. Heller, “Hy？nenfra？-Reststücke von Sch？deln des Wollhaarigen Nashorns Coelodonta antiquitatis blumenbach,” Quart？r, vol. 14, pp. 89–93, 1962.
[97]
C. Diedrich, “Die oberpleistoz？ne Population von Ursus spelaeus Rosenmüller 1794 aus dem eiszeitlichen Fleckenhy？nenhorst Perick-H？hlen von Hemer (Sauerland, NW Deutschland),” Philippia, vol. 12, no. 4, pp. 275–346, 2006.
[98]
C. Diedrich, “Evolution, Horste, Taphonomie und Pr？datoren der Rübel？nder H？hlenb？ren, Harz (Norddeutschland),” Mitteilungen des Verbandes der deutschen H？hlen- und Karstforscher, vol. 59, no. 1, pp. 4–29, 2013.
[99]
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.
[100]
W. von Koenigswald, Lebendige Eiszeit, Thorbecke, Darmstadt, Germany, 2002.
