Ancient DNA analysis of cyprinid remains from the Mesolithic-Neolithic Danube Gorges reveals an extirpated fish species Rutilus frisii (Nordmann, 1840)

The paper presents and discusses the results of the first ancient DNA (aDNA) analysis of cyprinid remains recovered from the Mesolithic-Neolithic sites of Vlasac, Lepenski Vir and Padina in the Danube Gorges (North-Central Balkans). Cyprinids constitute a significant portion of the identified fish remains recovered from these sites, which is indicative of their dietary role, and their large pharyngeal teeth have been worn as garment appliqués and associated with a great number of buried individuals. aDNA analysis (involving mitochondrial and nuclear markers) of pharyngeal bones with teeth corresponding to those used as appliqués has determined that they originate from anadromous Rutilus frisii (vyrezub), previously unrecorded in the Middle and Lower Danube. At present, the species inhabits the Black, Azov and Caspian Sea basins, but the only known populations in the Danube inhabit solely its upper reaches in Austria. The results of our study and the occurrence of R. frisii in the Danube Gorges further corroborate that its Upper Danube and Black Sea habitat had been connected in the past, i.e. that the species was entering the whole stretch of the river during its spawning migrations. Furthermore, precise taxonomic identification has important implications for a better understanding of fishing practices and their seasonal schedule in the Danube Gorges, and the distribution of cyprinid pharyngeal teeth ornaments in Europe during the Mesolithic.