We asked: (i) Which environmental factors determine the level of a-diversity at several scales and b-diversity in steppic grasslands? (ii) How do the effects of environmental factors on a- and b-diversity vary between the different taxonomic groups (vascular plants, bryophytes, lichens)? We sampled nested-plot series ranging from 0.0001 to 100 m2 and additional 10-m2 plots, covering different vegetation types and management regimes in steppes and semi-natural dry grasslands of Central Podolia (Ukraine). We recorded all terricolous taxa and used topographic, soil, land-use and climatic variables as predictors. Richness-environment relationships at different scales and across taxonomic groups were assessed with multimodel inference. We also fitted power-law species-area relationships, using the exponent (z value) as a measure of b-diversity. In general, the richness values in the study region were intermediate compared to those known from similar grasslands throughout the Palaearctic, but for 1 cm2 we found seven species of vascular plants, a new world record. Heat index was the most important factor for vascular plants and bryophytes (negative relation), while lichen diversity depended mainly on stone and rock cover (positive). The explanatory power of climate-related variables increased with increasing grain size, while anthropogenic burning was the most important factor for richness patterns at the finest grain sizes (positive effect). The z values showed more variation at the finest grain sizes, but no significant differences in their mean between scales. The results highlight the importance of integrating scale into ecological analyses and nature conservation assessments in order to understand and manage biological diversity in steppe ecosystems.
