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Hamilton’s theory of inclusive fitness is a widely used framework for studying the evolution of social behavior, but controversy surrounds its status. Hamilton originally derived his famous rb > c rule for the spread of a social gene by assuming additivity of costs and benefits. However, it has recently been argued that the additivity assumption can be dispensed with, so long as the −c and b terms are suitably defined, as partial regression coefficients. I argue that this way of generalizing Hamilton’s rule to the nonadditive case, while formally correct, faces conceptual problems.

On Hamilton’s Rule and Inclusive Fitness Theory with Nonadditive Payoffs