Combined carbon flows through detritus, microbes, and animals in reference and experimentally enriched stream ecosystems.

Tracking carbon (C) flow through ecosystems requires quantification of myriad biophysical processes, including C routing through microbial and metazoan food webs. Yet detailed organic matter budgets are rarely combined with simultaneous measurement of C flows supporting microbial and animal production. Here, we synthesize concurrent data sets on organic matter, microbes, and macroinvertebrates from two detritus-based stream ecosystems, one of which was subject to experimental nitrogen (N) and phosphorus (P) enrichment. Our synthesis provides new insights into C flow through forest stream ecosystems. Over 3 yr, the reference stream showed a striking balance of inputs and outputs, with a mean surplus of only 7 g C·m-2 ·yr-1 (~1% of annual inputs), presumably stored in sediments as fine particulate organic matter (FPOM). In contrast, N and P enrichment over 2 yr resulted in severe deficits of C (-576 g C·m-2 ·yr-1 or ~170% of annual inputs), a shortfall presumably met by stored C. Our data set provides an ecosystem-based estimate of the fate of forest litter C at ambient nutrient concentrations: 6.2% was leached as dissolved organic C, 40.6% and 8.5% flowed to litter-associated fungi and bacteria, respectively, 7.5% was consumed by macroinvertebrates, 1.8% was exported as coarse particles, and the remainder (35.4%) was presumably fragmented by biophysical processes. Our calculations also allowed an estimate of inputs into the heterogeneous FPOM pool, which is otherwise difficult to obtain. At naturally low nutrient concentrations, 50.7% was derived from fragmented litter, 39.1% from microbial biomass (mostly fungal), and 10.2% from macroinvertebrate egesta. Nutrient addition drove large changes in C fluxes in the experimental stream, especially in flows of leaf litter to fungi (×1.7 pretreatment) and macroinvertebrates (×2.7), and of FPOM to hydrologic export (×2.6). Our results underscore the key roles of both microbes and metazoans in controlling C flow through detritus-based ecosystems, as well as how release from persistent nutrient limitation may perturb steady-state conditions of C inputs vs. outputs. Our analysis also suggests areas for future research, including assessing the relative importance of stored vs. recycled C in fueling detrital food webs subject to altered nutrient regimes and other global-change drivers.