Mid-Holocene bottleneck for central European dry grasslands: Did steppe survive the forest optimum in northern Bohemia, Czech Republic?

Revisiting the classical Gradmann's steppe theory' for central Europe, we examine whether the early Holocene steppe habitats survived the critical period of maximum Holocene afforestation: the mid-Holocene bottleneck. Despite the undisputable fact that afforestation was a dominant ecological factor in this period, our parallel analyses of pollen and molluscs from sedimentary sequences discovered in the dry lowland area of northern Bohemia, Czech Republic (Zahaji and Suchy potok sites, lower Ohe area) provide strong evidence for uninterrupted local occurrence of steppe grasslands throughout the Holocene. At the onset of the Neolithic agriculture, this area was covered by forest-steppe. Analogously to the present forest-steppe landscapes of eastern Europe and south-western Siberia, dry areas of northern Bohemia were dominated by open-canopy pine-birch forests that enabled continuous survival of many light-demanding plant species from the late Glacial and early Holocene to the Neolithic. Later on, anthropogenic deforestation and livestock grazing created a semi-natural steppe. Our data suggest that this secondary steppe can be viewed as a direct continuation of the late Pleistocene and early Holocene natural steppe rather than a purely cultural steppe developed only after deforestation of a continuously forested mid-Holocene landscape by humans. At the same time, we provide evidence supporting Gradmann's steppe theory', assuming that in central Europe, Neolithic farming started in those areas that were not completely forested but contained remnants of natural steppes. This finding has important implications for the interpretation of present biodiversity patterns in central Europe.