Relationships between direct predation and risk effects.

Risk effects arise when prey alter their behavior in response to predators, and these responses carry costs. Empirical studies have found that risk effects can be large. Nonetheless, studies of predation in vertebrate conservation and management usually consider only direct predation. Given the ubiquity and strength of behavioral responses to predators by vertebrate prey, it is not safe to assume that risk effects on dynamics can be ignored. Risk effects can be larger than direct effects. Risk effects can exist even when the direct rate of predation is zero. Risk effects and direct effects do not necessarily change in parallel. When risk effects reduce reproduction rather than survival, they are easily mistaken for limitation by food supply.