Foraging for intermittently refuged prey: Theory and field observations of a parasitoid

1. Many insect herbivores feed in concealed locations but become accessible intermittently, creating windows of greater vulnerability to attack, and generating a proportion of the prey population that is readily accessible to foraging natural enemies. We incorporated accessible prey into an extant optimal foraging model, and found that this addition allowed opportunistic exploitation of prey that have already emerged from refugia (the leaving strategy) as a viable strategy, in addition to waiting at refugia for prey to emerge (the waiting strategy). 2. We parameterized the model empirically for the parasitoid Macrocentrus grandii and its host, Ostrinia nubilalis, under field conditions. The model predicted that M. grandii should adopt a leaving strategy when host patch density is high (travel time between patches is short), but a waiting strategy when host patch density is low (travel time between patches is long). 3. Field observations of M. grandii patch tenure were consistent with model predictions, indicating that M. grandii exhibited flexible behaviour based on experience within a foraging bout, and that these behavioural shifts improved foraging efficiency. 4. Behaviour of M. grandii was responsive to heterogeneity in host emergence rates, and appeared to be driven by the relatively small proportion of the host population that became accessible at a fast rate. Therefore understanding forager responses to intermittently refuged prey may require characterization of the behaviour of a subset of the prey population, rather than the average prey individual. 5. The model can potentially be used as a framework for comparative studies across forager taxa, to understand when foragers on intermittently accessible prey should adopt fixed waiting or leaving strategies vs. a flexible strategy that is responsive to the current environment.