Litter P content drives consumer production in detritus-based streams spanning an experimental N:P gradient

Abstract

Ecological stoichiometry theory (EST) is a key framework for predicting how variation in N:P supply ratios influences biological processes, at molecular to ecosystem scales, by altering the availability of C, N, and P relative to organismal requirements. We tested EST predictions by fertilizing five forest streams at different dissolved molar N:P ratios (2, 8, 16, 32, 128) for two years and tracking responses of macroinvertebrate consumers to the resulting steep experimental gradient in basal resource stoichiometry (leaf litter %N, %P, and N:P). Nitrogen and P content of leaf litter, the dominant basal resource, increased in all five streams following enrichment, with steepest responses in litter %P and N:P ratio. Additionally, increases in primary consumer biomass and production occurred in all five streams following N and P enrichment (averages across all streams: biomass by 1.2×, production by 1.6×). Patterns of both biomass and production were best predicted by leaf litter N:P and %P and were unrelated to leaf litter %N. Primary consumer production increased most in streams where decreases in leaf litter N:P were largest. Macroinvertebrate predator biomass and production were also strongly positively related to litter %P, providing robust experimental evidence for the primacy of P limitation at multiple trophic levels in these ecosystems. However, production of predatory macroinvertebrates was not related directly to primary consumer production, suggesting the importance of additional controls for macroinvertebrates at upper trophic positions. Our results reveal potential drivers of animal production in detritus-based ecosystems, including the relative importance of resource quality vs. quantity. Our study also sheds light on the more general impacts of variation in N:P supply ratio on nutrient-poor ecosystems, providing strong empirical support for predictions that nutrient enrichment increases food web productivity whenever large elemental imbalances between basal resources and consumer demand are reduced.

Keywords: carbon; detritus; ecological stoichiometry; nitrogen; nutrient enrichment; phosphorus; rivers; secondary production; streams.