We have previously reported the effect of predator learning on coexistence of prey populations in three-species experimental systems with two bruchid beetles, Callosobruchus chinensis and C. maculatus as the hosts, and the pteromalid wasp, Anisopteromalus calandrae. Introducing the parasitoid greatly enhanced the coexistence time, to a maximum of 118 weeks, where C. maculates and C. chinensis showed periodic anti-phase oscillations. Behavioral experiments confirmed the frequency-dependent predation of A. calandrae which was caused by learning. A. calandrae females learned the host-related olfactory cues during oviposition and increased their preference for the common host species.

We formulated the learning process of Individual-Based Model including operant conditioning of the parasitic wasp that has a success-experience-reward neural system. The preference for either host species was dynamically calculated using Rescorla-Wagner model in each step. Using the model, we could predict quantitative results on associative learning with olfactory cue of each host, frequency-dependent predation of the wasp, and the equivalent population dynamics to the experimental ones with anti-phase, two-week-lag oscillations of the two hosts.