Forage fish, their fisheries, and their predators: Who drives whom?

The North Sea has a diverse forage fish assemblage, including herring, targeted for human consumption; sandeel, sprat, and Norway pout, exploited by industrial fisheries; and some sardine and anchovy, supporting small-scale fisheries. All show large abundance fluctuations, impacting on fisheries and predators. We review field, laboratory, and modelling studies to investigate the drivers of this complex system of forage fish. Climate clearly influences forage fish productivity; however, any single-species considerations of the influence of climate might fail if strong interactions between forage fish exist, as in the North Sea. Sandeel appears to be the most important prey forage fish. Seabirds are most dependent on forage fish, due to specialized diet and distributional constraints (breeding colonies). Other than fisheries, key predators of forage fish are a few piscivorous fish species including saithe, whiting, mackerel, and horse-mackerel, exploited in turn by fisheries; seabirds and seals have a more modest impact. Size-based foodweb modelling suggests that reducing fishing mortality may not necessarily lead to larger stocks of piscivorous fish, especially if their early life stages compete with forage fish for zooplankton resources. In complex systems, changes in the impact of fisheries on forage fish may have potentially complex (and perhaps unanticipated) consequences on other commercially and/or ecologically important species.