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Adaptation is often described in behavioral ecology as individuals maximizing their inclusive fitness. Under what conditions does this hold, and how does this relate to the gene-centered perspective of adaptation? We unify and extend the literature on these questions to class-structured populations. We demonstrate that the maximization (in the best-response sense) of class-specific inclusive fitness obtains in uninvadable population states (meaning that all deviating mutants become extinct). This defines a genuine actor-centered perspec
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