Extinction debt at extinction threshold

To allow for long-term metapopulation persistence, a network of habitat fragments must satisfy a certain condition in terms of number, size, and spatial configuration of the fragments. The influence of landscape structure on the threshold condition can be measured by a quantity called metapopulation capacity, which can be calculated for real fragmented landscapes. Habitat loss and fragmentation reduce the metapopulation capacity of a landscape and make it less likely that the threshold condition can be met. If the condition is not met, the metapopulation is expected to go extinct, but it takes some time following habitat loss before the extinction will occur, which generates an extinction debt in a community of species. We show that extinction debt is especially great in a community in which many species are close to their extinction threshold following habitat loss because the metapopulation-dynamic time delay is especially long in such species. A corollary is that landscapes that have recently experienced substantial habitat loss and fragmentation are expected to show a transient excess of rare species, which represents a previously overlooked signature of extinction debt. We consider a putative example of extinction debt on forest-inhabiting beetles in Finland. At present, the few remaining natural-like forests are distributed evenly throughout southern Finland, but the number of regionally extinct old-growth forest beetles is much greater in the southwestern coastal areas, where human impact on forests has been lengthy, than in the northeastern inland areas, where intensive forestry started only after World War II. Ignoring time delays in population and metapopulation dynamics will lead to an underestimate of the number of effectively endangered species.