Holocene extinction dynamics of Equus hydruntinus, a late-surviving European megafaunal mammal

The European wild ass (Equus hydruntinus) is a globally extinct Eurasian equid. This species was widespread in Europe and southwest Asia during the Late Pleistocene, but its distribution became restricted to southern Europe and adjacent geographic regions in the Holocene. Previous research on E. hydruntinus has focused predominantly on its taxonomy and Late Pleistocene distribution. However, its Holocene distribution and extinction remain poorly understood, despite the fact that the European wild ass represents one of Europe's very few globally extinct Holocene megafaunal mammal species. We summarise all available Holocene zooarchaeological spatio-temporal occurrence data for the species, and analyse patterns of its distribution and extinction using point pattern analysis (kernel density estimation and Clark Evans index) and optimal linear estimation. We demonstrate that the geographic range of E. hydruntinus became highly fragmented into discrete subpopulations during the Holocene, which were associated with separate regions of open habitat and which became progressively extinct between the Neolithic and Iron Age. These data challenge previous suggestions of the late survival of E. hydruntinus into the medieval period in Spain, and instead suggest that postglacial climate-driven vegetational changes were a primary factor responsible for extinction of the species, driving isolation of small remnant subpopulations that may have been increasingly vulnerable to human exploitation. This study contributes to a more nuanced understanding of Late Quaternary species extinctions in Eurasia, suggesting that they were temporally staggered and distinct in their respective extinction trajectories.