Predator functional responses: Discriminating between handling and digesting prey

We present a handy mechanistic functional response model that realistically incorporates handling (i.e., attacking and eating) and digesting prey. We briefly review current functional response theory and thereby demonstrate that such a model has been lacking so far. In our model, we treat digestion as a background process that does not prevent further foraging activities (i.e., searching and handling). Instead, we let the hunger level determine the probability that the predator searches for new prey. Additionally, our model takes into account time wasted through unsuccessful attacks. Since a main assumption of our model is that the predator''s hunger is in a steady state, we term it the steady-state satiation (SSS) equation. The SSS equation yields a new formula for the asymptotic maximum predation rate (i.e., asymptotic maximum number of prey eaten per unit time, for prey density approaching infinity). According to this formula, maximum predation rate is determined not by the sum of the time spent for handling and digesting prey, but solely by the larger of these two terms. As a consequence, predators can be categorized into two types: handling-limited predators (where maximum predation rate is limited by handling time) and digestion- limited predators (where maximum predation rate is limited by digestion time). We give examples of both predator types. Based on available data, we suggest that most predators are digestion limited. The SSS equation is a conceptual mechanistic model. Two possible applications of this model are that (1) it can be used to calculate the effects of changing predator or prey characteristics (e.g., defenses) on predation rate and (2) optimal foraging models based on the SSS equation are testable alternatives to other approaches. This may improve optimal foraging theory, since one of its major problems has been the lack of alternative models