Understanding spatial distributions: negative density-dependence in prey causes predators to trade-off prey quantity with quality.

Negative density-dependence is generally studied within a single trophic level, thereby neglecting its effect on higher trophic levels. The 'functional response' couples a predator's intake rate to prey density. Most widespread is a type II functional response, where intake rate increases asymptotically with prey density; this predicts the highest predator densities at the highest prey densities. In one of the most stringent tests of this generality to date, we measured density and quality of bivalve prey (edible cockles Cerastoderma edule) across 50 km² of mudflat, and simultaneously, with a novel time-of-arrival methodology, tracked their avian predators (red knots Calidris canutus). Because of negative density-dependence in the individual quality of cockles, the predicted energy intake rates of red knots declined at high prey densities (a type IV, rather than a type II functional response). Resource-selection modelling revealed that red knots indeed selected areas of intermediate cockle densities where energy intake rates were maximized given their phenotype-specific digestive constraints (as indicated by gizzard mass). Because negative density-dependence is common, we question the current consensus and suggest that predators commonly maximize their energy intake rates at intermediate prey densities. Prey density alone may thus poorly predict intake rates, carrying capacity and spatial distributions of predators.

Birds influence vegetation coverage and structure on sandy biogeomorphic islands in the Dutch Wadden Sea.

Small uninhabited islands form important roosting and breeding habitats for many coastal birds. Previous studies have demonstrated that guano can promote ecosystem productivity and functionality on island ecosystems. Here, we assess the role of external nutrient input by coastal birds on the vegetation structure and coverage on sandy biogeomorphic islands, where island-forming processes depend on vegetation-sedimentation feedbacks. As a first step, we investigated whether breeding birds affect vegetation productivity on sandy back-barrier islands in the Wadden Sea. Using a combination of bird observations and plant stable isotope (δ15N) analyses, we demonstrate that (i) breeding birds transport large quantities of nutrients via their faecal outputs to these islands annually and that (ii) this external nitrogen source influences vegetation development on these sandy, nutrient-limited, islands. Based on these results we discuss how this avian nutrient pump could impact island development and habitat suitability for coastal birds and discuss future directions for research. In general, we conclude that avian subsidies have the potential to affect both the ecological and biogeomorphic functioning of coastal soft-sediment systems. However, the strength and scale of especially these biogeomorphic interactions are not fully understood. For the conservation of both threatened coastal birds and sandy back-barrier islands and the design of appropriate management strategies, we argue that three-way interactions between birds, vegetation and sandy island morphodynamics need to be further elucidated.