When in life does density dependence occur in fish populations?

Fisheries advice is based on demographic calculations, which assume that density-dependent processes regulating recruitment occur only in early life. This assumption is challenged by laboratory and lake studies and some recent indications from marine systems that demonstrate density-dependent regulation late in life. By accounting for spatial dynamics of a population, something that has previously been ignored in models of fish, we show that density-dependent regulation is determined by the size of the habitat: in small habitats, for example small lakes, regulation occurs late in life, while it can occur early in large habitats. When regulation happens late in life, fisheries yield is maximized by exploitation of mainly juvenile fish, while exploiting mature fish maximizes yield if regulation happens early. We review and interpret observations of density dependence in the light of the theory. Our results challenge the current assumption that density dependence always occurs early in life and highlights the need for an increased understanding of density-dependent processes. This can only come about by a change of focus from determining stock-recruitment relationships towards understanding when and how density-dependent regulation occurs in nature.