The shape of density dependence in fragmented landscapes explains an inverse buffer effect in a migratory songbird

It is well known that forest fragmentation reduces fecundity in several avian species, including wood thrush, Hylocichla mustelina, a migratory songbird that has been declining for several decades. However, I found that landscape-scale density in wood thrush was lower and population declines steeper in higher quality, less-fragmented landscapes (an inverse buffer effect) than in poor quality landscapes. These patterns suggest that wood thrush was not limited by availability of breeding habitat but that declines were primarily driven by non-breeding season events. A two-season model predicts that if this hypothesis is correct, breeding population trends will be negatively related to the strength of density dependence (b') in the breeding season. To test this, a site-dependence model was used to construct fecundity-density curves and showed that landscape fragmentation affected the shape of density dependence. In good quality landscapes, the onset of strong density dependence was much more abrupt than in poorer quality landscapes and the realized strength of density dependence, b', was lower in good quality landscapes. Population trends were negatively associated with b', providing support for the non-breeding limitation hypothesis. The combination of the negative associations of trends with b' and b' with landscape quality explain the inverse buffer effect.