Abstract
Organisms exert multiple, often-contrasting, influences on ecosystems. Migrating Pacific salmon (Oncorhynchus spp.) deliver nutrients to freshwater ecosystems, but also disturb benthic sediments during upstream migration and nest building. The relative importance of these contrasting roles is not well understood, especially not in relation to the temporal dynamics of other environmental drivers. To assess the influence of salmon-mediated enrichment and disturbance, we measured stream biofilm metrics (production, respiration, chlorophyll a [chla], ash-free dry mass, stable isotope signatures) and environmental variables (spawner and carcass abundance, dissolved nutrients, temperature, discharge, light) from July through September in multiple habitats of a southeast Alaska stream. Biofilm production and biomass increased before and early in the salmon run, but declined later in the run. Biofilm stable isotope composition indicated incorporation of salmon-derived carbon and nitrogen (N) during the latter part of the run. Biofilm biomass differed among benthic habitat types (i.e., riffles, pools stream edges) but temporal patterns were generally similar, suggesting salmon and environmental influences were not habitat-specific. We used this high-frequency field data to parameterize an ordinary differential equation model for dissolved inorganic N, chla, and cellular N, and estimated model parameters using Marcov Chain Monte Carlo. Posterior distributions indicated that 1) habitats and locations were generally similar in model parameters, 2) removing the effect of salmon resulted in no change in biofilm chla early in the run (mid-August), but did show a decrease for some habitats later in the run (September), and 3) the overall integrated salmon effect over the run was one of biofilm loss. Only the combination of high frequency biofilm and environmental data collections with a process-based model, allowed us to determine how environmental context dynamics interact with salmon run dynamics to modulate the biofilm response in natal spawning streams. High-frequency field data combined with modeling provides a critical tool to gain a mechanistic understanding of resource subsidies, especially those from organisms that have other influences and roles, in relation to the environmental context and will allow us to evaluate the small and large scale importance of subsidies.
Highlights
The physical and chemical template of ecosystems combined with biological processes provide the environmental context that influences ecosystem structure and dynamics, including the response to resource subsidies
We examined the influence of salmon density on biofilm accrual using a process-based model to disentangle the salmon influence from changes in the other environmental characteristics, such as discharge which is generally higher during salmon runs in our study system
The first live salmon in Twelve Mile Creek were observed on 5 August, and abundance increased to >1000 salmon in the 300 m reach within 2 weeks, before declining near the end of September (Figure 1A)
Summary
The physical and chemical template of ecosystems combined with biological processes provide the environmental context (after Janetski et al, 2009) that influences ecosystem structure and dynamics, including the response to resource subsidies (sensu Polis et al, 2004). The influence of subsidies on a recipient ecosystem varies in relation to the available endogenous resources and the environmental context (Marczak et al, 2007; Subalusky and Post, 2019). Both the subsidy and environmental context can be spatially and temporally dynamic, creating complex patterns in responses, and currently limiting our understanding of the overall importance of resources subsidies. Understanding the effects of salmon is challenging because it requires explicit quantification of the enrichment and disturbance effects in the context of subsidy and environmental dynamics
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