Deconstructing responses of dragonfly species richness to area, nutrients, water plant diversity and forestry.

Understanding large-scale variation in species richness in relation to area, energy, habitat heterogeneity and anthropogenic disturbance has been a major task in ecology. Ultimately, variation in species richness results from variation in individual species occupancies. We studied whether the individual species occupancy patterns are determined by the same candidate factors as total species richness. We sampled 26 boreal forest ponds for dragonflies (Odonata) and studied the effects of shoreline length, water vascular plant species density (WVPSD), availability of nutrients, intensity of forestry, amount of Sphagnum peat cover and pH on dragonfly species richness and individual dragonfly species. WVPSD and pH had a strong positive effect on species richness. Removal of six dragonfly species experiencing strongest responses to WVPSD cancelled the relationship between species richness and WVPSD. By contrast, removal of nine least observed species did not affect the relationship between WVPSD and species richness. Thus, our results showed that relatively common species responding strongly to WVPSD shaped the observed species richness pattern whereas the effect of least observed, often rare, species was negligible. Also, our results support the view that, despite of the great impact of energy on species richness at large spatial scales, habitat heterogeneity can still have an effect on species richness in smaller scales, even overriding the effects of area.