Null Models for the Opportunity for Selection

Crow’s “opportunity for selection” (I p variance in relative fitness) is an important albeit controversial eco-evolutionary concept, particularly regarding the most appropriate null model(s). Here, we treat this topic in a comprehensive way by considering op-portunities for both fertility selection (I_f) and viability selection (I_m) for discrete generations, both seasonal and lifetime reproductive success in age-structured species, and experimental designs that include either a full or partial life cycle, with complete enumeration or random subsampling. For each scenario, a null model that includes random demographic stochasticity can be constructed that follows Crow’s initial formulation that I p I_f 1 I_m. The two components of I are qualita-tively different. Whereas an adjusted I_f (D_If) can be computed that accounts for random demographic stochasticity in offspring number, I_m cannot be similarly adjusted in the absence of data on phenotypic traits under viability selection. Including as potential parents some individuals that die before reproductive age produces an overall zero-inflated Poisson null model. It is always important to remember that (1) Crow’s I represents only the opportunity for selection and not selection itself and (2) the species’ biology can lead to random sto-chasticity in offspring number that is either overdispersed or under-dispersed compared with the Poisson (Wright-Fisher) expectation.