Red squirrel middens influence abundance but not diversity of other vertebrates.

Some animals modify the environment in ways that can influence the resources available to other species. Because red squirrels (Tamiasciurus hudsonicus) create large piles of conifer-cone debris (middens) in which they store cones, squirrels concentrate resources that might affect biodiversity locally. To determine whether other animals are attracted to midden sites beyond their affinity for the same resources that attract red squirrels, we assessed associations between middens, mammals, and birds at population and community levels. We surveyed 75 middens where residency rates of red squirrels varied during the previous five years; sampling along this residency gradient permitted us to evaluate the influence of resources at middens beyond the influence of a resident squirrel. At each location, we quantified vegetation, landscape structure, abundance of conifer cones, and midden structure, and used capture-recapture, distance sampling, and remote cameras to quantify presence, abundance, and species richness of mammals and birds. Red squirrels and the resources they concentrated at middens influenced mammals and birds at the population scale and to a lesser extent at the community scale. At middens with higher residency rates of red squirrels, richness of medium and large mammals increased markedly and species richness of birds increased slightly. After accounting for local forest characteristics, however, only species richness of medium-to-large mammals was associated with a red squirrel being resident during surveys. In areas where red squirrels were resident during surveys or in areas with greater amounts of resources concentrated by red squirrels, abundances of two of four small mammal species and two of four bird species increased. We conclude that the presence of this ecosystem modifier and the resources it concentrates influence abundance of some mammals and birds, which may have implications for maintaining biodiversity across the wide geographic range inhabited by red squirrels and other larderhoarding animals.