New and revised middle Villafranchian canid material from Dafnero-3 (Greece): Implications to Nyctereutes systematics and zoogeography

New cranial, mandibular and dental canid material from Dafnero-3 site, in northern Greece confirms previous reports of Nyctereutes megamastoides and Vulpes sp. from either Dafnero-3 or/and Dafnero-1 sites but allows a reassignment of specimens previously referred to as Nyctereutes tingi from Dafnero-3 to Nyctereutes vulpinus . The results further support the previous hypothesis of a close phylogenetic link between N . tingi and N . vulpinus , revise the intraspecific diversity of the latter, and allow the establishment of a working phylogeographic hypothesis. Nyctereutes vulpinus is noted for the first time in southeast Europe, expanding its geographical range, confirming the simultaneous presence of two distinct Nyctereutes species in the Balkan area as in China.