Whole-system nutrient enrichment increases secondary production in a detritus-based ecosystem

Although the effects of nutrient enrichment on consumer-resource dynamics are relatively well studied in ecosystems based on living plants, little is known about the manner in which enrichment influences the dynamics and productivity of consumers and resources in detritus-based ecosystems. Because nutrients can stimulate loss of carbon at the base of detrital food webs, effects on higher consumers may be fundamentally different than what is expected for living-plant-based food webs in which nutrients typically increase basal carbon. We experimentally enriched a detritus-based headwater stream for two years to examine the effects of nutrient-induced changes at the base of the food web on higher metazoan (predominantly invertebrate) consumers. Our paired-catchment design was aimed at quantifying organic matter and invertebrate dynamics in the enriched stream and an adjacent reference stream for two years prior to enrichment and two years during enrichment. Enrichment had a strong negative effect on standing crop of leaf litter, but no apparent effect on that of fine benthic organic matter. Despite large nutrient-induced reductions in the quantity of leaf litter, invertebrate secondary production during the enrichment was the highest ever reported for headwater streams at this Long Term Ecological Research site and was 1.2-3.3 times higher than predicted based on 15 years of data from these streams. Abundance, biomass, and secondary production of invertebrate consumers increased significantly in response to enrichment, and the response was greater among taxa with larval life spans < or = 1 yr than among those with larval life spans >1 yr. Production of invertebrate predators closely tracked the increased production of their prey. The response of invertebrates was largely habitat-specific with little effect of enrichment on food webs inhabiting bedrock outcrops. Our results demonstrate that positive nutrient-induced changes to food quality likely override negative changes to food quantity for consumers during the initial years of enrichment of detritus-based stream ecosystems. Longer-term enrichment may impact consumers through eventual reductions in the quantity of detritus.