Natural selection by pulsed predation: Survival of the thickest

Selective predation can lead to natural selection in prey populations and may alleviate competition among surviving individuals. The processes of selection and competition can have substantial effects on prey population dynamics, but are rarely studied simultaneously. Moreover, field studies of predator-induced short-term selection pressures on prey populations are scarce. Here we report measurements of density dependence in body composition in a bivalve prey (edible cockle, Cerastoderma edule) during bouts of intense predation by an avian predator (red knot, Calidris canutus). We measured densities, patchiness, morphology, and body composition (shell and flesh mass) of cockles in a quasi-experimental setting, i.e. before and after predation in three similar plots of 1 ha each, two of which experienced predation, and one of which remained unvisited in the course of the short study period and served as a reference. An individual's shell and flesh mass declined with cockle density (negative density dependence). Before predation, cockles were patchily distributed. After predation, during which densities were reduced by 78% (from 232 m-2 to 50 m-2), the patchiness was substantially reduced, i.e. the spatial distribution was homogenized. Red knots selected juvenile cockles with an average length of 6.9 mm (SD 1.0). Cockles surviving predation had heavier shells than before predation (an increase of 21.5 percentage points), but similar flesh masses. By contrast, in the reference plot shell mass did not differ statistically between initial and final sampling occasions, while flesh mass was larger (an increase of 13.2 percentage points). In this field-study, we show that red knots imposed a strong selection pressure on cockles to grow fast with thick shells and little flesh mass, with selection gradients among the highest reported in the literature