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How to measure inclusive fitness, revisited

An individual’s inclusive fitness is derived by augmenting its traditional Darwinian fitness by certain components, and stripping it of others (Hamilton 1964). The component to be added is the sum of all eVects of the individual on his neighbours, weighted by the coeYcient of genetic relationship (r) between them. In the original derivation of inclusive fitness, the component to be subtracted was verbally defined as ‘all components which can be considered as due to the individual’s social environment’ (Hamilton 1964). It is not widely appreciated that this verbal definition of the component to be subtracted does not agree with its algebraic definition. Creel (1990a) used Hamilton’s algebraic definition of inclusive fitness to show that the component to be subtracted, e 0 , is actually equal to the average eVect