First record of a Late Miocene hominid from North Macedonia

Known for over a century, the Late Miocene mammalian faunas of Veles, North Macedonia, have long been recognized for their scientific importance. However, hominid remains had not been previously reported from this fossil-rich area. Here, we report the discovery of an isolated upper molar from the vicinity of Veles-most likely from the Belushka locality-which constitutes the first known record of a Late Miocene hominid from the Republic of North Macedonia, and provide a review of the associated mammalian assemblages. The molar-probably an M³-exhibits a relatively large crown, broad and low cusps, inferred thick enamel, low dentine horns, and short, thick roots that are not markedly divergent. These characteristics, along with its inferred middle Turolian (MN12) age and Balkan provenance, suggest that the Veles molar belongs to an indeterminate member of the hominid tribe Graecopithecini. Together with the Graecopithecus freybergi mandible from Pyrgos Vassilissis, Greece, and an isolated graecopithecin upper premolar from Azmaka, Bulgaria, the Veles specimen provides rare additional evidence for the survival of apes well into the Turolian of the Balkan Peninsula. Furthermore, the Veles fossil faunas now document the co-occurrence of apes (Graecopithecini indet.) and monkeys (Mesopithecus), a pattern rarely observed in the Eurasian Miocene fossil record.