Spatial analysis of North Sea cod recruitment: concurrent effects of changes in spawning stock biomass, temperature and herring abundance

The decline of the North Sea cod Gadus morhua has been attributed to both overfish- ing and ocean warming. However, another hypothesis is that overfishing of piscivorous cod has caused an increased abundance of formerly predatory-controlled pelagic fish including herring Clupea harengus, which in turn has suppressed the recruitment of cod through predation on the early life stages of cod. Here we analyze 40 yr of trawl survey data in order to explore how the abundance of young herring affects cod recruitment, and how cod biomass affects the abundance of herring. In both cases we also take into account the effects of spawner biomass (of cod and her- ring, respectively) and sea temperature. We take a novel, explicitly spatial, approach by analyzing these effects on a local (185 × 167 km) scale. Our results indicate large spatial variability in eco- logical mechanisms. In the German Bight, high cod recruitment is associated with low herring abundance, low temperatures and high overall cod spawner biomass. This area used to contain a large portion of the cod recruits, and there is a strong correlation between the fraction of cod recruits found in this area and overall recruitment. In this area, herring recruitment is also nega- tively associated with the abundance of large cod. Thus, for this part of the North Sea, our findings are consistent with a reversal of dominance between cod and herring; however, herring may affect the cod by competition rather than by predation.