On Procobus Khomenko, 1913 (Mammalia: Artiodactyla: Bovidae), with new evidence from the Late Miocene of Greece

Procobus is a rare and controversial Late Miocene bovid genus (Artiodactyla, Bovidae), originally known by a few cranial remains from the northern shores of the Black Sea. New material from Dytiko-1 fossil site in Axios Valley, northern Greece expands its geographic and chronological distribution and provides additional craniodental characters for its definition. A thorough description and comparison of all available specimens from Greece, Moldova and Ukraine highlight differences from the similar-sized and sympatric Palaeoryx and permit re-discussing its phylogenetic relationships. Our data reject previously proposed relations with Reduncini and suggests Procobus as a pan-Caprini, unique among Late Miocene bovid genera by the combination of large size and long, paedomorphic horns curving forward. A preliminary phylogenetic analysis indicates Procobus as the sister taxon of crown Caprini+Pantholops (Tibetan antelope) lineage.