Density- and size-dependent mechanisms modulate the outcome of stocking in a naturally recruiting freshwater piscivore (northern pike, Esox lucius): A replicated whole-lake experiment

Despite the widespread use of stock enhancements to improve fisheries across the world, there is a lack of robust knowledge under which conditions fish stocking provides additive effects rather than merely replacing a fraction of natural recruitment. Fully controlled and replicated studies at the level of entire ecosystems are needed to provide answers. Properly monitored experimental releases also allow testing mechanisms of density- and sizedependent population regulation. In this study, the population-level outcome of stocking juvenile pike (Esox lucius L.) in naturally reproducing lentic stocks was investigated. We used a replicated before-after-control-impact design in 15 experimental lakes involving two stocking densities and unstocked controls. Releasing age-0 pike failed to generate additive effects at the age-2 cohort. As expected from theory, we observed density-dependent mortality and differential survival of wild and stocked pike. Stocked and wild fishes showed contrasting responses in terms of growth to variation in predator density, competitor density and the forage base, suggesting both subpopulations differed in their response to stocking-induced changes in population traits (e.g., density). Despite the lack of additive effects caused by stocking, a fraction of the stocked individuals established in the stock-enhanced cohort, indicating that replacement of wild recruits by stocked conspecifics had occurred. Depending on the origin of the stocking material, pike stocking thus has the potential for genetic hybridization, while not necessarily benefiting fisheries catch. We conclude that whenever a natural pike population exists, stocking juvenile pike will not produce additive effects in lakes and that enhanced pike populations will be strongly regulated by size- and density-dependent juvenile mortality and less by density-dependent growth.