Effects of multiple, predator-induced behaviors on short-term producer-grazer dynamics in open systems

We investigated the population consequences of multiple behavioral responses of grazers to a foraging return-predation risk trade-off in an open system consisting of primary producers, grazers, and predators. Using a dynamical model where grazers adjust their foraging activity and emigration rate to the densities of predators and producers, we explored how changes in control variables (predator density, grazer immigration, and producer immigration and carrying capacity) affect the dynamics of producers and grazers at temporal scales shorter than consumer and predator reproduction. The model predicts that producer biomass increases and that both the density of foraging grazers and the feeding rate of predators decrease with predator density. These predictions hold although total (foraging + nonforaging) grazer density may actually increase with predator density. The latter will occur whenever the benefit of higher resource density outweighs the increased risk of predation. In this case. per capita grazer emigration decreases with predator density, which might be misinterpreted as a direct "freezing" response to predators. Increased grazer immigration is predicted to result in decreased producer density and increased densities of both foraging and total grazers, as well as increased grazer emigration and predator feeding rates. Increased producer immigration or carrying capacity should increase producer and grazer densities and predator feeding rate but decrease per capita grazer emigration. Manipulation of predator (trout) densities in a set of nine large (50 m(2)) stream channels produced results in broad agreement with model predictions. Most notably, a positive effect of trout on benthic algal biomass was mainly mediated through grazer behavior (changes in the use of epibenthic surfaces and in emigration rare) rather than through consumptive reductions of grazer numbers by trout.