Inclusive Fitness

Although the concept of metaindividual is mathematically elegant, arguments expressed in more conventional biological terminology are usually more cogent. The concept of inclusive fitness is based on a decomposition of the (exponential of) invasion fitness into an individual component and a component that derives from the change in the reproductive output of other individuals of the same type caused by interactions with the focal individual. At the invasion stage, infinite dilution guarantees that other individuals in a metaindividual only have the focal type when those individuals are related by descent. The idea then is that the change in reproductive output of nonfocal individuals is weighed with the probability that these individuals are identical by descent to the focal one.

For simple ecological scenarios, with, for example, non-overlapping generations, fixed-size aggregates and mutants that closely resemble the residents, it is possible to separately calculate the various contributions before combining them. (The close resemblance is needed to guarantee that selection does not affect the probabilities of identity by descent and that fitness effects are approximately additive.)

When possible, calculations through the inclusive fitness route are preferable, as the resulting decomposition helps in interpreting trait evolution. When higher cognitive abilities do not come into play, metaindividual-based techniques usually have a larger reach than inclusive-fitness-based ones. When metaindividual-based techniques work, it is possible in principle to reconstruct the ingredients occurring in an inclusive fitness calculation, at least numerically. However, doing so often is far more complicated than directly calculating invasion fitness and hence may contribute less insight than one has come to expect from the simple cases.

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