Woodpeckers and other excavators maintain the diversity of cavity-nesting vertebrates

Woodpeckers and other excavators create most of the holes used by secondary cavity nesters (SCNs) in North American temperate mixedwood forests, but the degree to which excavators release SCNs from nest-site limitation is debated. Our goal was to quantify how excavators maintain the diversity and abundance of secondary cavity nesters in a temperate forest through the creation of tree cavities. We examined the short- and long-term (legacy) effects of excavators (principally woodpeckers, but also red-breasted nuthatches and black-capped chickadees) on forest biodiversity using longitudinal monitoring data (1,732 nest cavities, 25 sites, 16 years) in British Columbia, Canada. Sites with higher densities of excavator nests had more cavities available, higher species richness of SCNs and higher nest density of SCNs, indicating the importance of a standing stock of cavities. Years with higher nesting densities of excavators were followed by years with higher SCN diversity, indicating that the creation of nesting opportunities through fresh excavation releases SCNs from community-wide nest-site limitation. We also show that excavators leave a 'legacy' of biodiversity (species richness and abundance) at a site by accumulating cavities at rates faster than they become unusable by decay or destruction. By quantifying site-level effects of cavity excavation on the SCN community, our study highlights the key role of excavators as ecosystem engineers that maintain forest wildlife biodiversity.