State-dependent mortality can enhance behavioral unpredictability

BACKGROUND: Although behavioral unpredictability is widely described within-individual variability in behavior, its adaptive significance is little understood. Using a dynamic state variable model, this study investigated the conditions under which behavioral unpredictability (a component of within-individual variability) in foraging behavior is advantageous. The model considers a situation in which a forager forages for a fixed period, represented by discrete time steps. The outcome of foraging may change the level of a state (e.g., size and fat storage) of the forager at each time step, and variability in the foraging outcome is assumed to be positively correlated with behavioral unpredictability. The probability of death at each time step is influenced by the state at the same time step. Reproduction occurs after all the foraging steps and is influenced by the state level of a forager at the time of reproduction. According to the expected utility hypothesis, the relationship (e.g., curvature) between the state and fitness will determine the role of behavioral unpredictability. In the model, the relationship was obtained by using the backward iteration method for each foraging time step. RESULTS: State-dependent mortality adds curvature to the relationship between the state and fitness, which makes the effect of behavioral unpredictability on fitness either positive or negative. This conclusion holds for any state-dependent mortality (i.e., as long as mortality is not independent of the state factor). Given that state-dependent mortality is commonly described, conditions that benefit behavioral unpredictability are likely also common. CONCLUSIONS: When mortality depends on a state that is influenced by behavior, conditions that favor behavioral unpredictability may become common. How behavioral unpredictability influences the variability of behavioral outcomes is as important as how it influences the expectation of behavioral outcomes when studying the adaptive significance of behavioral unpredictability.