Accounting for Age Structure and Spatial Structure in Eco-Evolutionary Analyses of a Large, Mobile Vertebrate.

The idealized concept of a population is integral to ecology, evolutionary biology, and natural resource management. To make analyses tractable, most models adopt simplifying assumptions, which almost inevitably are violated by real species in nature. Here, we focus on both demographic and genetic estimates of effective population size per generation (Ne), the effective number of breeders per year (Nb), and Wright's neighborhood size (NS) for black bears (Ursus americanus) that are continuously distributed in the northern lower peninsula of Michigan, United States. We illustrate practical application of recently developed methods to account for violations of 2 common, simplifying assumptions about populations: 1) reproduction occurs in discrete generations and 2) mating occurs randomly among all individuals. We use a 9-year harvest dataset of >3300 individuals, together with genetic determination of 221 parent-offspring pairs, to estimate male and female vital rates, including age-specific survival, age-specific fecundity, and age-specific variance in fecundity (for which empirical data are rare). We find strong evidence for overdispersed variance in reproductive success of same-age individuals in both sexes, and we show that constraints on litter size have a strong influence on results. We also estimate that another life-history trait that is often ignored (skip breeding by females) has a relatively modest influence, reducing Nb by 9% and increasing Ne by 3%. We conclude that isolation by distance depresses genetic estimates of Nb, which implicitly assume a randomly mating population. Estimated demographic NS (100, based on parent-offspring dispersal) was similar to genetic NS (85, based on regression of genetic distance and geographic distance), indicating that the >36000 km2 study area includes about 4-5 black-bear neighborhoods. Results from this expansive data set provide important insight into effects of violating assumptions when estimating evolutionary parameters for long-lived, free-ranging species. In conjunction with recently developed analytical methodology, the ready availability of nonlethal DNA sampling methods and the ability to rapidly and cheaply survey many thousands of molecular markers should facilitate eco-evolutionary studies like this for many more species in nature.