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The final European Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss 1823) was the most important megamammal at all, proved by the study of 50 caves and 15 open air localities in Germany and Czech Republic, between the Skandinavian and Alpine ice shields. The carnivore impact is astonishing; nearly all non-anthropogenous or cave bear bone sites are the result of hyena prey accumulations around their dens, whereas up to several thousands of bones resulted from prey body pieces, and only few collected bones, deposited along river mud pits, in muddy loess or in caves. Therefore most of the bone accumulations in middle Europe can be referred more and more to hyena activity.
Hyenas raised up their cubs in the entrance areas of caves and imported prey to them. Not only at caves a specialization onto cave bear scavenging by hyenas can be proved, in which hyenas left bones always similar incomplete. Enrichments of bone fragments in caves resulted primary from hyena activities, too. Hyenas crushed bones even of their own species, whereas many cannibalistic traces can be observed. Normally at dens 10-45% bones are from hyenas including isolated milk teeth. For the den marking they used excrements, coprolites.
Nibbling sticks, on one or two sides intensively for the tooth change support chewed bone fragments are very typical for cub activities. Different hyena den types or prey depots explain most of the bone accumulations in Central Europe. In horizontal caves hyenas left strongly fragmented bones more at the entrance area, because juveniles and adult ones could use those easily. In more vertical caves the penetration was more difficult; here articulated prey remains were found. Even complete skeletons of killed and imported lions and hyenas survived in such caves. Such vertical caves were in a few cases only used as food storage sites.
Some very small cave dens were accessible only for cubs. Additionally gypsum karst cavities were used in several sites, which huge bone accumulations around are of hyena origin. The last type of dens are reused badger/red fox den burrows in the open mammoth steppe. Here a few localities in loess and in river sand bar deposits in northern Germany and northern Czech Republic prove this for the first time. Generally at hyena dens the bones have typical chewing marks or are bone shafts or are crushed, but much better is the identification by similar preserved bone shafts. The most perfect den markers are woolly rhinoceros bones, which are indestructible.
In their spongiosa the tooth scratch marks are generally well visible. Hyenas hunted actively in some regions as the Bohemian Karst mainly Przewalski horses (um to 50% prey animal bones), whereas in the Sauerland Karst of northern Germany the prey was more mixed of all mega mammals including up to 10% mammoth. Even wolves, lions and wolverines were eaten around their dens. Finally those second large predators besides the lions became extinct around 24.000 BP.
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