New insights on Early Pleistocene Nyctereutes from the Balkans based on material from Dafnero-3 (Greece) and Varshets (Bulgaria)

The study examines the taxonomy of the newly excavated Nyctereutes material from Dafnero-3 (Greece) and the unpublished specimens from Varshets (Bulgaria). One cranium and a hemimandible from Dafnero-3 site are ascribed to the species Nyctereutes megamastoides, already known from Dafnero-1 and other Early Pleistocene sites of Greece. Two additional crania and a hemimandible from Dafnero-3, along with the Varshets dental material are attributed to N. tingi. This species was previously reported in the Early Pliocene Megalo Emvolon fauna of northern Greece but the record has been challenged and a review is provided. The new findings of Dafnero-3, and Varshets, extend the presence of N. tingi in southeastern Europe until the beginning of the middle Villafranchian and confirm the geographic expansion of the taxon towards West. Moreover, the co-existence of N. tingi and N. megamastoides in Dafnero-3 is reported for the first time in Europe. A paleoecological analysis based on the lower molar method agrees with the existing hypothesis of a more omnivorous N. megamastoides and a more carnivorous N. tingi ecomorph, which further justifies their local co-existence. Additionally, the quantitative analyses of N. tingi from Dafnero-3, and Varshets reveal a possible succession from N. tingi to N. vulpinus, as already supported by previous studies.