RESUMO
Understanding how nutrients flow through food webs is central in ecosystem ecology. Tracer addition experiments are powerful tools to reconstruct nutrient flows by adding an isotopically enriched element into an ecosystem and tracking its fate through time. Historically, the design and analysis of tracer studies have varied widely, ranging from descriptive studies to modeling approaches of varying complexity. Increasingly, isotope tracer data are being used to compare ecosystems and analyze experimental manipulations. Currently, a formal statistical framework for analyzing such experiments is lacking, making it impossible to calculate the estimation errors associated with the model fit, the interdependence of compartments, and the uncertainty in the diet of consumers. In this article we develop a method based on Bayesian hidden Markov models and apply it to the analysis of N15-NH4+ tracer additions in two Trinidadian streams in which light was experimentally manipulated. Through this case study, we illustrate how to estimate N fluxes between ecosystem compartments, turnover rates of N within those compartments, and the associated uncertainty. We also show how the method can be used to compare alternative models of food web structure, calculate the error around derived parameters, and make statistical comparisons between sites or treatments.
Assuntos
Ecossistema , Cadeia Alimentar , Modelos Estatísticos , Nitrogênio/metabolismo , Compostos de Amônio/química , Animais , Luz , Cadeias de Markov , Isótopos de Nitrogênio , Plantas/metabolismo , Rios , Trinidad e Tobago , Água/químicaRESUMO
Decades of ecological study have demonstrated the importance of top-down and bottom-up controls on food webs, yet few studies within this context have quantified the magnitude of energy and material fluxes at the whole-ecosystem scale. We examined top-down and bottom-up effects on food web fluxes using a field experiment that manipulated the presence of a consumer, the Trinidadian guppy Poecilia reticulata, and the production of basal resources by thinning the riparian forest canopy to increase incident light. To gauge the effects of these reach-scale manipulations on food web fluxes, we used a nitrogen (15 N) stable isotope tracer to compare basal resource treatments (thinned canopy vs. control) and consumer treatments (guppy introduction vs. control). The thinned canopy stream had higher primary production than the natural canopy control, leading to increased N fluxes to invertebrates that feed on benthic biofilms (grazers), fine benthic organic matter (collector-gatherers), and organic particles suspended in the water column (filter feeders). Stream reaches with guppies also had higher primary productivity and higher N fluxes to grazers and filter feeders. In contrast, N fluxes to collector-gatherers were reduced in guppy introduction reaches relative to upstream controls. N fluxes to leaf-shredding invertebrates, predatory invertebrates, and the other fish species present (Hart's killifish, Anablepsoides hartii) did not differ across light or guppy treatments, suggesting that effects on detritus-based linkages and upper trophic levels were not as strong. Effect sizes of guppy and canopy treatments on N flux rates were similar for most taxa, though guppy effects were the strongest for filter feeding invertebrates while canopy effects were the strongest for collector-gatherer invertebrates. Combined, these results extend previous knowledge about top-down and bottom-up controls on ecosystems by providing experimental, reach-scale evidence that both pathways can act simultaneously and have equally strong influence on nutrient fluxes from inorganic pools through primary consumers.