Slow response of plant species richness to habitat loss and fragmentation.

We examined the response of vascular plant species richness to long-term habitat loss and fragmentation of Estonian calcareous grasslands (alvars). The current number of habitat specialist species in 35 alvars was not explained by their current areas and connectivities but it was explained by their areas and connectivities 70 years ago (R(2) = 0.27). We estimated the magnitude of extinction debt in local communities by assuming an equilibrium species richness in 14 alvars that had lost only a small amount of area and by applying this model to the remaining alvars, in which the average area has declined from 3.64 km(2) in the 1930s to 0.21 km(2) at present. The extinction debt estimated for individual alvars was around 40% of their current species number. Our conclusions are applicable to temperate grasslands in general, which have lost much area because of agricultural intensification and cessation of traditional management.