Revision of the early Middle Pleistocene bears (Ursidae, Mammalia) of Central Europe, with special respect to possible co-occurrence of spelaeoid and arctoid lineages

The bears from the Late Biharian localities Koneprusy Caves, C 718 and Kozi Grzbiet differ significantly from U. arctos, particularly in morphology of metaconid and entoconid complex of m1. In contrast, they exhibit a number of dental morphometric characters typical for spelaeoid lineage for which we interpret them as the early members of U. deningeri clade. Also the nominal taxa U. suessenbornensis, U. etruscus gombaszogensis and U. savini are assigned to spelaeoid lineage. U. sackdillingensis is tentatively assigned to arctoid lineage, similarly as few single specimens from Late Biharian sites Chlum 4, Kovesvarad, Prezletice and Voigtstedt. All other material from these sites was identified as U. deningeri. A smaller part of the studied material, including the type series of U. eberbachensis, is of ambiguous taxonomical status and do not allow an exact species determination.