Species accumulation over space and time in European Plio-Holocene mammals

The rate of increase in species number with sampled area is one issue of major interest in ecology. Species number increases with sampled time as well, though this kind of analysis is much rarer in literature. Species-area and species-time relationships have been recently integrated in a single model, which allows studying how time and area interact with each other in determining the cumulative increase in species richness. Here we studied species-area, species-time, and species-time-area relationships in Plio-Holocene large mammals of Western Eurasia, by using an extensive database including 184 species distributed in 685 fossil sites. We found that the increase of species number with time is much higher than with area. When sampling inequality of fossil localities in time and space is accounted for, time and area interact with each other in a negative, though non-linear fashion. The intense climatic changes that characterized the Plio-Holocene period apparently affected both species-area and species-time relationships in large mammals, by increasing the slope of the former during the Pliocene and middle Pleistocene, and of the latter during younger, climatically harsher, late Pleistocene times. This study emphasizes the importance of accounting for time and space in tracing paleodiversity curves.