Late Pleistocene hyenas Crocuta crocuta spelaea of Upper Rhine valley open air sites and a contribution to skull shape variability

Six skulls, two mandibles and a few postcranial bones of the Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss 1823) are described from the open air Rhine river gravel pit sites Brühl, Ketsch and Altlussheim of southwestern Germany (Baden-Württemberg). Additionally, an overview of one open air loess den site Bad Cannstadt, and six hyena den caves in Franconia are briefly presented to get a first impression of the taphonomy and palaeoecology of those last hyenas in southwestern Germany. The megafauna, besides the hyenas, found in the Rhine River gravel pits is a typical Weichselian/Wuermian fauna consisting of Mammuthus primigenius, Coelodonta antiquitatis, Ovibos moschatus, Bison priscus, Equus ferus przewalskii, Equus hydruntinus, Megaloceros giganteus, Cervus elaphus, Rangifer tarandus, Capra ibex and even remains of some other carnivore species, Panthera leo spelaea and either Ursus spelaeus or Canis lupus, are recognized. The selected large hyena material consists mainly of brain cases and one complete skull, and shows three different types of brain case shape. The saggital crest is highly convex, hardly convex or even flat. The three skull morphs were compared to 40 skulls from European hyena den sites such as northern German caves and open air den sites. Especially the high amount of skulls from the hyena den site in the gypsum karst of Westeregeln near Magdeburg shows the same large variability in skull morphology. Therefore, the phenomenon does not seem to be typical for a certain region, subspecies or related to sexual dimorphism.