ALBERT

Description: Woodpeckers and other primary cavity excavators (PCEs) are important worldwide for excavating cavities in trees, and a large number of studies have examined their nesting preferences. However, quantitative measures of wood hardness have been omitted from most studies, and ecologists have focused on the effects of external tree- and habitat-level features on nesting. Moreover, information is lacking on the role of wood hardness in limiting nesting opportunities for this important guild. Here, we used an information theoretic approach to examine the role of wood hardness in multi-scale nest-site selection, and in limiting nesting opportunities for six species of North American PCEs. We found that interior wood hardness at nests (n = 259) differed from that at random sites, and all six species of PCE had nests with significantly softer interior wood than random trees (F(1,517) = 106.15, P 〈 0.0001). Accordingly, interior wood hardness was the most influential factor in our models of nest-site selection at both spatial scales that we examined - in the selection of trees within territories and in the selection of nest locations on trees. Moreover, regardless of hypothesized excavation abilities, all the species in our study appeared constrained by interior wood hardness, and only 4-14% of random sites were actually suitable for nesting. Our findings suggest that past studies that did not measure wood hardness counted many sites as available to PCEs when they were actually unsuitable, potentially biasing results. Moreover, by not accounting for nest-site limitations in PCEs, managers may overestimate the amount of suitable habitat. We therefore urge ecologists to incorporate quantitative measures of wood hardness into PCE nest-site selection studies, and to consider the limitations faced by avian cavity excavators in forest management decisions. # doi:10.1890/14-1042.1