Modelling effects of nonbreeders on population growth estimates.

Adult individuals that do not breed in a given year occur in a wide range of natural populations. However, such nonbreeders are often ignored in theoretical and empirical population studies, limiting our knowledge of how nonbreeders affect realized and estimated population dynamics and potentially impeding projection of deterministic and stochastic population growth rates. We present and analyse a general modelling framework for systems where breeders and nonbreeders differ in key demographic rates, incorporating different forms of nonbreeding, different life histories and frequency-dependent effects of nonbreeders on demographic rates of breeders. Comparisons of estimates of deterministic population growth rate, λ, and demographic variance, σd2, from models with and without distinct nonbreeder classes show that models that do not explicitly incorporate nonbreeders give upwardly biased estimates of σd2, particularly when the equilibrium ratio of nonbreeders to breeders, Nnb∗/Nb∗, is high. Estimates of λ from empirical observations of breeders only are substantially inflated when individuals frequently re-enter the breeding population after periods of nonbreeding. Sensitivity analyses of diverse parameterizations of our model framework, with and without negative frequency-dependent effects of nonbreeders on breeder demographic rates, show how changes in demographic rates of breeders vs. nonbreeders differentially affect λ. In particular, λ is most sensitive to nonbreeder parameters in long-lived species, when Nnb∗/Nb∗>0, and when individuals are unlikely to breed at several consecutive time steps. Our results demonstrate that failing to account for nonbreeders in population studies can obscure low population growth rates that should cause management concern. Quantifying the size and demography of the nonbreeding section of populations and modelling appropriate demographic structuring is therefore essential to evaluate nonbreeders' influence on deterministic and stochastic population dynamics.