Wetenschappelijk artikelBalancing food and density-dependence in the spatial distribution of an interference-prone forager.

Foraging distributions are thought to be density-dependent, because animals not only select for a high availability and quality of resources, but also avoid conspecific interference. Since these processes are confounded, their relative importance in shaping foraging distributions remains poorly understood. Here we aimed to rank the contribution of density-dependent and density-independent effects on the spatio-temporal foraging patterns of eurasian oystercatchers. In our intertidal study area, tides caused continuous variation in oystercatcher density, providing an opportunity to disentangle conspecific interference and density-independent interactions with the food landscape. Spatial distributions were quantified using high-resolution individual tracking of foraging activity and location. In a model environment that included a realistic reconstruction of both the tides and the benthic food, we tested a family of behaviour-based optimality models against these tracking data. Density-independent interactions affected spatial distributions much more strongly than conspecific interference, even in an interference-prone species like oystercatchers. Spatial distributions were governed by avoidance of bill injury costs, selection for high interference-free intake rates and a decreasing availability of benthic bivalve prey after their exposure. These density-independent interactions outweighed interference competition in terms of effect size. We suggest that the bottleneck in our mechanistic understanding of foraging distributions may be primarily the role of density-independent prey attributes unrelated to intake rates, like damage costs in the case of oystercatchers foraging on perilous prey. At a landscape scale, above the finest inter-individual distances, effects of conspecific interaction on spatial distributions may have been overemphasised