How to measure inclusive fitness, revisited

  title={How to measure inclusive fitness, revisited},
  author={Jeffrey R. Lucas and Scott R. Creel and Peter M. Waser},
  journal={Animal Behaviour},
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… 


  • C. Goodnight
  • Biology, Psychology
    Evolution; international journal of organic evolution
  • 2013
The direct fitness approach and contextual analysis come from very different traditions, but both methods rely on the same underlying equation, with the primary difference between the two approaches being that thedirect fitness approach uses fitness optimization modeling, whereas with contextual analysis, the same equation is used to solve for the change in fitness associated with a change in phenotype when the population is away from the optimal phenotype.

Hamilton goes empirical: estimation of inclusive fitness from life-history data

  • M. Oli
  • Biology
    Proceedings of the Royal Society of London. Series B: Biological Sciences
  • 2003
The method presented here allows empirical estimation of inclusive fitness from life–history data, simultaneously considers all components of fitness, including timing and magnitude of reproduction, and adequately addresses shortcomings of existing methods of estimating inclusive fitness.

Comparative statics of games between relatives.

Duality Of Stochasticity And Natural Selection Shape The Ecology-driven Pattern Of Social Interactions: The Fall Of Hamilton's Rule

From microbes to mammals, cooperation is selected-for in harsh, uncertain and unpredictable environments, and the evolution of cooperation is a bet-hedging (risk spreading) strategy of risk-averse individuals in stochastic environments.

Kin selection in Columbian ground squirrels: direct and indirect fitness benefits

The presence of closely related and philopatric female kin appeared to result in considerable fitness benefits for female ground squirrels, perhaps through the behavioural mechanisms of lowered aggression and other forms of behavioural cooperation.


A quantitative genetic model of multivariate trait evolution that integrates the effects of both IGEs and social selection and provides a general predictive equation for the evolution of social phenotypes that encompasses specific cases such as kin selection and reciprocity is presented.

The measurement and meaning of inclusive fitness

Evolutionaryadaptationisageneticprocess,butfortunatelywecanoftenunderstandit,toafirstapproximation, without knowing the geneticdetails. We use what Grafen (1982) called

Reproductive skew in cooperative breeding: Environmental variability, antagonistic selection, choice, and control

The results suggest the primary drivers for the initial evolution of cooperative breeding may be a combination of limited suitable sites, choice across those sites, and parental manipulation of offspring into helping roles.

The Ecology of Social Life: A Synthesis

A comparative summary of the ecological and demographic factors favoring social life reveals some patterns of correlated traits (sociality syndromes) that accentuate that variation in genetic relatedness alone is not sufficient to explain the occurrence or details in the organization of social life.

Kin selection and cooperative courtship in wild turkeys

It is shown, using genetic measures of relatedness and reproductive success, that kin selection can explain the evolution of cooperative courtship in wild turkeys and extends recent findings of male relatedness on avian leks by quantifying the kin-selected benefits gained by non-reproducing males.



How to measure inclusive fitness

  • S. Creel
  • Biology
    Proceedings of the Royal Society of London. Series B: Biological Sciences
  • 1990
By re-examining Hamilton’s original model, a modified verbal definition of inclusive fitness can be justified and no one should ever attempt to breed among obligately cooperative breeders.

The genetical evolution of social behaviour. I.

Twelve Misunderstandings of Kin Selection

Abstract Hamilton's theory of kin selection is much misunderstood. This paper lists and refutes 12 of the commonest misunderstandings, for example: “Kin selection is a special, complex kind of

Death and disappearance: estimating mortality risks associated with philopatry and dispersal

A graphical framework within which census data and behavioral observations can be used to place bounds on e, s, and sp is described, which reveals unexpected differences in dispersal risk among sex and age classes of dwarf mongooses.

Social Factors Affecting Reproduction Amongst Red Foxes

In some habitats foxes live in social groups comprised of one adult male and several adult vixens. These groups occupy territories from which neighbouring groups are excluded. Elsewhere, foxes live

A molecular genetic analysis of kinship and cooperation in African lions

A new application of DNA fingerprinting is described that unequivocally demonstrates the kinship structure of lion 'prides': female companions are always closely related, male companions are either closely related or unrelated, and mating partners are usually unrelated.

The Florida scrub jay : demography of a cooperative-breeding bird

It is found that habitat restraints, rather than kin selection, are the main source of the behavior of Florida Scrub Jays: the goal of increasing the number of close relatives other than descendants in future generations is of relatively minor importance in their cooperative-breeding behavior.

Delayed Dispersal and Reproduction as a Life-History Tactic in Cooperative Breeders: Fitness Calculations from Red-Cockaded Woodpeckers

It is shown, in a population in which the fate of dispersers could be documented, that the fitness of individuals delaying dispersal and reproduction can equal or exceed that of individuals attempting early reproduction, even without indirect fitness benefits due to helping behavior.