Biomass distribution of fishes and mussels mediates spatial and temporal heterogeneity in nutrient cycling in streams

Animals can play important roles in cycling nutrients [hereafter consumer-driven nutrient dynamics (CND)], but researchers typically simplify animal communities inhabiting dynamic environments into single groups that are tested under relatively static conditions. We propose a conceptual framework and present empirical evidence for CND that considers the potential effects of spatially overlapping animal groups within dynamic ecosystems. Because streams can maintain high biomass of mussels and fish, we were able to evaluate this framework by testing if biogeochemical hotspots generated by stable aggregations of mussels attract fishes. We predicted that spatial overlap between these groups may increase the flux of mineralized nutrients. We quantified how different fish assemblage biomass was between mussel bed reaches and reaches without mussels. We compared fish and mussel biomass at mussel beds to test whether differences in animal biomass mediate their contributions to nutrient cycling through nitrogen and phosphorous excretion. We estimated areal excretion rates for each group by combining biomass estimates with measured excretion rates. Fish biomass was homogeneously distributed, except following a period of low flow when fish were more concentrated at mussel beds. Mussel biomass was consistently an order of magnitude greater than fish biomass and mussel areal excretion rates exceeded fish excretion rates. However, the magnitude of those differences varied spatially and temporally. Mussel excretion stoichiometry varied with changes in assemblage composition, while fish excretion stoichiometry varied little. Biogeochemical hotspots associated with mussels did not generally overlap with fish aggregations, thus, under these conditions, animal processes appear to exert additive ecosystem effects.