Stream invertebrate productivity linked to forest subsidies: 37 stream-years of reference and experimental data.

Riparian habitats provide detrital subsidies of varying quantities and qualities to recipient ecosystems. We used long-term data from three reference streams (covering 24 stream-years) and 13-year whole-stream organic matter manipulations to investigate the influence of terrestrial detrital quantity and quality on benthic invertebrate community structure, abundance, biomass, and secondary production in rockface (RF) and mixed substrates (MS) of forested headwater streams. Using a mesh canopy covering the entire treatment stream, we examined effects of litter ex'clusion, small- and large-wood removal, and addition of artificial wood (PVC) and leaves of varying quality on organic matter standing crops and invertebrate community structure and function. We assessed differences in functional feeding group distribution between substrate types as influenced by organic matter manipulations and long-term patterns of predator and prey production in manipulated vs. reference years. Particulate organic matter standing crops in MS of the treatment stream declined drastically with each successive year of litter exclusion, approaching zero after three years. Monthly invertebrate biomass and annual secondary production was positively related to benthic organic matter in the MS habitats. Rockface habitats exhibited fewer changes than MS habitats across all organic matter manipulations. With leaf addition, the patterns of functional group distribution among MS and RF habitats returned to patterns seen in reference streams. Secondary production per unit organic matter standing crop was greatest for the leaf addition period, followed by the reference streams, and significantly less for the litter exclusion and wood removal periods. These data indicate that the limited organic matter remaining in the stream following litter exclusion and wood removal was more refractory than that in the reference streams, whereas the added leaf material was more labile and readily converted into invertebrate production. Predator production and total production were tightly coupled in reference and treatment streams, indicating strong relationships between predators and their prey. Results from the artificial wood addition demonstrate that physical structure alone will not restore invertebrate productivity without detrital resources from the riparian forest. Our long-term studies conducted over three decades at the ecosystem scale unequivocally show the necessity of maintaining and restoring aquatic-terrestrial linkages in forested headwater streams.