The mechanisms causing extinction debts.

Extinction debts can result from many types of habitat changes involving mechanisms other than metapopulation processes. This is a fact that most recent literature on extinction debts pays little attention to. We argue that extinction debts can arise because (i) individuals survive in resistant life-cycle stages long after habitat quality change, (ii) stochastic extinctions of populations that have become small are not immediate, and (iii) metapopulations survive long after that connectivity has decreased if colonization-extinction dynamics is slow. A failure to distinguish between these different mechanisms and to simultaneously consider both the size of the extinction debt and the relaxation time hampers our understanding of how extinction debts arise and our ability to prevent ultimate extinctions.