Productivity and recovery of forage fish under climate change and fishing: North Sea sandeel as a case study

Forage fish occupy a central position in marine food-webs worldwide by mediating the transfer of energy and organic matter from lower to higher trophic levels. The lesser sandeel (Ammodytes marinus) is one of the ecologically and economically most important forage fish species in the North-east Atlantic, acting as a key prey for predatory fish and sea birds, as well as supporting a large commercial fishery. In this case study, we investigate the underlying factors affecting recruitment and how these in turn affect productivity of the North Sea sandeel using long-term data and modelling. Our results demonstrate how sandeel productivity in the central North Sea (Dogger Bank) depends on a combination of external and internal regulatory factors, including fishing and climate effects, as well as density dependence and food availability of the preferred zooplankton prey (Calanus finmarchicus and Temora longicornis). Furthermore, our model scenarios suggest that while fishing largely contributed to the abrupt stock decline during the late 1990s and the following period of low biomass, a complete recovery of the stock to the highly productive levels of the early 1980s would only be possible through changes in the surrounding ecosystem, involving lower temperatures and improved feeding conditions. To that end, we stress the need for ecosystem-based management accounting for multiple internal and external factors occurring within the broader context of the ecosystem in which forage fish species, such as sandeel, play an important and integral part