Individual quality and age but not environmental or social conditions modulate costs of reproduction in a capital breeder

Costs associated with reproduction are widely known to play a role in the evolution of reproductive tactics with consequences to population and eco-evolutionary dynamics. Evaluating these costs as they pertain to species in the wild remains an important goal of evolutionary ecology. Individual heterogeneity, including differences in individual quality (i.e., among-individual differences in traits associated with survival and reproduction) or state, and variation in environmental and social conditions can modulate the costs of reproduction; however, few studies have considered effects of these factors simultaneously. Taking advantage of a detailed, long-term dataset for a population of feral horses (Sable Island, Nova-Scotia, Canada), we address the question of how intrinsic (quality, age), environmental (winter severity, location), and social conditions (group size, composition, sex ratio, density) influence the costs of reproduction on subsequent reproduction. Individual quality was me...