Non-lethal effects of predators on prey growth rates depend on prey density and nutrient additions

A number of studies show that predators can depress prey growth rates by inducing reductions in foraging activity, but the size of this non-lethal effect is quite variable. Here I investigate how prey density and resource productivity may alter the extent to which predators depress the growth rates of their prey. Theory predicts that when resources are overgrazed, an increase in predation risk will have little net effect on individual food intake because the decline in foraging effort will be offset by an increase in resource level. Thus, the non-lethal effects of predators on prey growth rates should depend upon prey density and resource productivity in a predictable manner, with the growth penalty imposed by predators being strongest when resources are undergrazed and weakest when resources are overgrazed. I tested this hypothesis by manipulating predation risk, prey density, and nutrient additions in a mesocosm experiment with the pulmonate snail Helisoma trivolvis. Refuge use by snails was 45% higher in the presence of caged crayfish than in their absence. Snail growth rates were reduced, on average, by 24% in the presence of caged crayfish. However, the magnitude of the growth penalty exacted by crayfish depended on snail density and nutrient additions. When snails were stocked at high density and nutrient additions were low, growth suppression was just 2.6%. At the other extreme, when snails were at low density and nutrient additions were high, growth suppression was 44.6%. Thus, the non-lethal effects of predators on prey growth depend on environmental context, illustrating an important link between individual traits and system-level properties.