Foraging habitats and niche partitioning of European large herbivores during the Holocene - Insights from 3D dental microwear texture analysis

Environmental changes related to forest expansion and later to agricultural development and deforestation during the Holocene in Europe have strongly shaped herbivore distribution and habitat selection, leading to species extinction. In this paper, we examine, through dental microwear textural analysis, the foraging habitats, dietary flexibility and niche partitioning of large herbivores (100 to 1000 kg) including two bovids (European bison Bison bonasus and aurochs Bos primigenius) and two cervids (moose Alces alces and red deer Cervus elaphus). The study focuses on northeast (NE) and south central (SC) Europe during three periods of the Holocene: Neolithic (6500-3500 cal yrs BP), Roman Period and Middle Ages (~2000-450 cal yrs BP) and Recent times (1963-2014). Our study highlights differences in diet between species only in the Neolithic SC Europe. This may have resulted from diversified habitats and dietary niche separation in SC Europe. The absence of differences in NE Europe may suggest more homogeneous habitat conditions in combination with lower densities of large herbivores in forested habitats. Analysis shows that during the Neolithic, in the bison diet there was more herbaceous monocotyledons in SC Europe than in NE Europe. We also found temporal changes in microwear texture variables for the bison in SC Europe suggesting a more herbaceous diet in the Neolithic than during the Roman Period and Middle Ages. However, neither spatial nor temporal differences in microwear texture variables were found in the aurochs. Thus, the diet of the European bison has likely changed, which may indicate a greater plasticity that promoted survival in refuge habitats.