Inclusive fitness

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In evolutionary biology, inclusive fitness is one of two metrics of evolutionary success as defined by W. D. Hamilton in 1964:

• Personal fitness is the number of offspring that an individual begets (regardless of who rescues/rears/supports them)
• Inclusive fitness is the number of offspring equivalents that an individual rears, rescues or otherwise supports through its behaviour (regardless of who begets them)

An individual's own child, who carries one half of the individual's genes, is defined as one offspring equivalent. A sibling's child, who will carry one-quarter of the individual's genes, is 1/2 offspring equivalent. Similarly, a cousin's child, who has 1/16 of the individual's genes, is 1/8 offspring equivalent.

From a gene's point of view, evolutionary success ultimately depends on leaving behind the maximum number of copies of itself in the population. Prior to Hamilton's work, it was generally assumed that genes only achieved this through the number of viable offspring produced by the individual organism they occupied. However, this overlooked a wider consideration of a gene's success, most clearly in the case of the social insects where the vast majority of individuals do not produce (their own) offspring.