| Summary: | 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. spatial raster with cockle densities corresponding to Fig3AThe coordinate reference system is EPSG:32631 - WGS 84 / UTM zone 31N, and cockle density is presented in numbers per square meter.resource_landscape-Fig3A_cockle_density.tifSpatial raster with relative AFDMflesh corresponding to Fig3BThe coordinate reference system is EPSG:32631 - WGS 84 / UTM zone 31N. Relative AFDMflesh is presented as the ratio of AFDMflesh to average AFDMflesh for cockles of identical length (see main article).resource_landscape-Fig3B_relative_AFDMflesh.tifspatial raster with predicted intake rates corresponding to ...
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