Habitat coupling in a large lake system: Delivery of an energy subsidy by an offshore planktivore to the nearshore zone of Lake Superior

We hypothesised that the autumn spawning migration of Lake Superior cisco (Coregonus artedi) provides a resource subsidy, in the form of energy-rich cisco eggs, from the offshore pelagic to the nearshore benthic community over winter, when alternate prey production is likely to be low. We tested this hypothesis using fish and macroinvertebrate surveys, fish population demographics, diet and stable isotope analyses, and bioenergetics modelling. The benthic, congeneric lake whitefish (C. clupeaformis) was a clear beneficiary of cisco spawning. Cisco eggs represented 16% of lake whitefish annual consumption in terms of biomass, but 34% of energy (because of their high energy density: >10 kJ g wet mass^-1). Stable isotope analyses were consistent with these results and suggest that other nearshore fish species may also rely on cisco eggs. The lipid content of lake whitefish liver almost doubled from 26 to 49% between November and March, while that of muscle increased from 14 to 26% over the same period, suggesting lake whitefish were building, rather than depleting, lipid reserves during winter. In the other Laurentian Great Lakes, where cisco populations remain very low and rehabilitation efforts are underway, the offshore-to-nearshore ecological link apparent in Lake Superior has been replaced by non-native planktivorous species. These non-native species spawn in spring have smaller eggs and shorter incubation periods. The rehabilitation of cisco in these systems should reinstate the onshore subsidy as it has in Lake Superior.