At the beginning of the Middle Pleistocene (MIS 21, 800,000 BP), three distinct hyenas remained to appear in Europe with the spotted hyena (Crocuta), brown hyena (Parahyaena) and striped hyena ...(Hyaena). Each developed in monophyletic lineages, whereas rare brown hyena Parahyaena brunnea mosbachensis (Geib, 1915) and striped hyena Hyaena hyaena prisca De Serres, Dubreuil and Jeanjean, (1828) did not change in their dentition morphology much to their modern forms. Those warm period extinct hyenas did not migrate from Africa over the Rhine Graben migratory channel to Central Europe after the late Mid‐Pleistocene Holsteinian Interglacial (MIS 9). The spotted hyenas took over the niche of European hyenas with their appearance in warm and cold periods. Those used more and more cave entrances as dens between Spain and Siberia. Their more rapid dental change coevolved from the largest “giant hyena” Crocuta brevirostris Boule, (1893) (Early Pleistocene, MIS 40‐20), over Crocuta intermedia De Serres, Dubreuil and Jeanjean, 1828 (Early Mid‐Pleistocene, MIS 19‐12), C. praespelaea Schütt, 1971 (Early Mid‐Pleistocene, MIS 11‐6), to Crocuta crocuta spelaea (Goldfuss, 1823) (Early Mid‐Pleistocene, MIS 5‐3), which is genetically a subspecies to Modern African extant Crocuta crocuta crocuta Erxleben, 1777 (MIS 2‐1). Spotted hyenas developed a perfect thick‐skin cutter M1 by elongation and bone crusher conical P4 teeth. This adaptation to their main thick‐skin big game guilt: elephants, rhinos and hippos.
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Late Pleistocene Ice Age spotted hyena remains are described from the “Unicorn holotype skeleton” gypsum karst site Quedlinburg‐Sewecken‐Berge, Germany (Central Europe). The hyena population consists ...of adolescent to late adult individuals (96% of hyena NISP; 15% of megafauna NISP) indicating a commuting den site type. The comparisons to other European bone assemblages support hunting specialization on woolly rhinoceros (19% of NISP) and horses (27% of NISP). Specialization on bovids (Bison/Bos) can be added for this site. The megafauna contain few Eemian warm period remains of a large horse Equus ferus fossilis. Most (95%) of the megafauna is attributed to the Late Pleistocene glacial (Weichselian/Wuermian). Horse bones are dominated by distal leg elements from the smaller Przewalski horses Equus ferus przewalskii (26% of NISP). The Unicorn “holotype” skeleton originates from a composed horse skull, vertebrae and front legs, whereas the elephant remains added to this biologically not valid species must have been a straight‐tusked elephant tusk.
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In the fossil record, 3.097 studied bones including 40 skulls of grown‐up individuals from 81 German, Czech and English and Spanish extant spotted hyena Crocuta crocuta spelaea den sites let to ...distinguish two main types of pathologies. Ageing pathologies in the form of arthritis are typical for elderly hyenas. Their strong tooth use or losses and canine damages due to intensive bone crushing made them finally defenceless. Bite trauma on the sagittal crests of the crania is the secure proof for battles between top predators: lions and hyenas. A full crest healing resulted in a strong convex‐deformed skull shape. Other complete upper/lower jaw bites made by lions are found at a hyena skeleton shoulder, in which scapulae are partly healed. This hyena died in its den cave finally; as such, it is known for another Ice Age spotted hyena that was found recently as a skeleton in a Spanish cave, but with a non‐healed strong sagittal crest bite trauma. The postcranial bite traumas in the Ice Age spotted hyenas are predominantly on the distal hind legs (tibia/fibula—68% of all traumas). This correlates to African relatives because spotted hyenas attack its guilt and enemies from “behind.”
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New crania and cranial material from along the Vilyuy River of the Tynda region, North‐East Siberia (Russia), support the identification of open air hyaena den sites.
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Siberian extinct top predators—Ice Age spotted hyenas—are discovered in the permafrost of Yakutia in‐between frozen mammoth, rhinoceros, bison or horse carcasses, such as in Mongolia at new open‐air ...sites. Historic described European holotypes and new crania from Siberia, Mongolia, and Europe allow presenting the monophyly of Crocuta reaching back 2.53 Mio years (Late Pliocene). Spotted hyenas coevolved in dental change and body size to their largest guilt, thick‐skinned elephants and rhinoceroses. The Late Pleistocene (MIS 5d‐3, 113.00–25.000 BP) Crocuta crocuta spelaea (Goldfuss, 1823) is proven to have scavenged Siberian woolly rhinoceros Coeleodonta antiquitatis (Blumenbach, 1799) carcasses. Rhino mummy skins have scavenging signs attributable only to hyenas. Chew cuts of breaking‐cutting scissor dentition are found at mummy skins. The last Eurasian hyenas left always similar chew‐damaged woolly rhinoceroses’ crania with braincase openings and damaged mandibles, whereas their deep bite scratches are often found on the distal joints of long bones. Strongest rhino bone damage is found at natal den sites (cave or open air), to which hyenas imported guilt to their cubs. The recently known Ice Age spotted hyena palaeobiogeography overlaps exactly with those of woolly rhinoceros and mammoths. All reached Bering Straits.
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Extinct European Late Pleistocene (MIS 3‐5d) Crocuta crocuta spelaea and late Middle Pleistocene (MIS 11‐6) C. c. praespelaea and extant African spotted hyenas C. c. crocuta were/are alive and ...post‐mortem cannibals. 38 fossil crania and some braincases, as well as 3.097, studied Pleistocene Ice Age spotted bones from 81 European Pleistocene den sites expose many bite damages. Cranial damages correlate with the newest films about modern African spotted hyenas which cut and decompose the head first of their antagonistic female same clan leader or other clans. The skulls and jaws have repeatedly similar bite damages, whereas, at birth/natal dens, their cranial remain abundance is highest. At communal dens, Pleistocene spotted hyena carcasses were similarly decomposed—first head off antagonistic clan leading females, and further similar strategy decomposing of the legs. Large postcranial C. c. spelaea bones have three identical bone destruction stages. These damage stages are similar to those found at hyena guilt including competing carnivores such as cave bears, steppe lions, but also herbivores like horses, and even a few smaller‐sized Neanderthal long bones.
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Bone remains and a trackway of
Pantheraichnus bottropensis nov. ichg. ichnsp. of the Late Pleistocene lion
Panthera leo spelaea (
Goldfuss, 1810) have been recovered from Bottrop and other open air ...sites in northern Germany. Some of these bones are from open air hyena den sites. A relative high proportion of lion bones (20%) exhibit bite, chew or nibble marks, or bone crushing and nibbling caused by a large carnivore. Repeated patterns of similar bone damage have been compared to bone remains found at hyena dens in gypsum karst areas and cave sites in northern Germany. Ice Age spotted hyenas have been the main antagonists and the main scavengers on lion carcasses. The remains appear to have been imported often by hyenas into their communal dens, supporting the theory of strong hyena-lion antagonism, similar to the well documented antagonism between modern African lions and spotted hyenas. Most of the lion bones from the open air hyena den at Bottrop are probably a result of such antagonism, as are the rare remains of these carnivores found within large hyena prey bone accumulations along the Pleistocene rivers. The Emscher River terrace also has the largest quantity of hyena remains from open air river terrace sites in northern Germany. Their cub remains, and incomplete chewed prey bones from mammoths and woolly rhinoceroses, typical of hyena activity, underline the character of these sites as cub-raising and communal dens, where their prey was accumulated along the riverbanks in a similar manner to modern African hyenas.
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Punctured extinct cave bear femora were misidentified in southeastern Europe (Hungary/Slovenia) as ‘Palaeolithic bone flutes’ and the ‘oldest Neanderthal instruments’. These are not instruments, nor ...human made, but products of the most important cave bear scavengers of Europe, hyenas. Late Middle to Late Pleistocene (Mousterian to Gravettian) Ice Age spotted hyenas of Europe occupied mainly cave entrances as dens (communal/cub raising den types), but went deeper for scavenging into cave bear dens, or used in a few cases branches/diagonal shafts (i.e. prey storage den type). In most of those dens, about 20% of adult to 80% of bear cub remains have large carnivore damage. Hyenas left bones in repeating similar tooth mark and crush damage stages, demonstrating a butchering/bone cracking strategy. The femora of subadult cave bears are intermediate in damage patterns, compared to the adult ones, which were fully crushed to pieces. Hyenas produced round–oval puncture marks in cub femora only by the bone-crushing premolar teeth of both upper and lower jaw. The punctures/tooth impact marks are often present on both sides of the shaft of cave bear cub femora and are simply a result of non-breakage of the slightly calcified shaft compacta. All stages of femur puncturing to crushing are demonstrated herein, especially on a large cave bear population from a German cave bear den.
During the last Ice Age of central Europe, cave bears hibernated deep inside their caves and Ice Age spotted hyenas used the cave entrance areas as dens. Steppe lions were also sporadic cave dwellers ...in order to hunt the herbivorous cave bears or steal prey from hyena dens, or for antagonistic and territorial reasons. Steppe lion skeletons recently discovered in the midst of cave bear skeletons, deep inside the Urşilor Cave (Romania), and steppe lion remains from other European caves, in particular a large population from the Zoolithen Cave in Germany, provide evidence of active predation by lions on cave bears. However, it was not mainly lions that fed on the cave bears: the large quantities of damaged cave bear bones, including incomplete long bones that repeatedly show similar types of damage, crushed long bones, and damaged skulls, reveal that hyenas were the primary scavengers on cave bear carcasses and were largely responsible for the destruction of their carcasses and bones. Predation and scavenging on cave bears (mainly by these two Ice Age predators but also by wolves and leopards) particularly in mountainous boreal forest environments that were devoid of the mammoth steppe fauna, explains the large quantities of fragmented cave bear bones that have been found all over Eurasia. The stress caused by these carnivores, deprived of their specialized ecological niche, may have led the herbivorous cave bears to hibernate as deeply as possible within their hibernation caves, in an attempt to protect themselves against predation. The felid predators were sometimes killed inside the caves, probably in conflicts with adult cave bears or even with hyenas, and remained as complete or only partly disarticulated carcasses.
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