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Alexander K. Fremier

Publications and source records attributed to Alexander K. Fremier.

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Clarifying the trophic state concept to advance macroscale freshwater science and management

For over a century, ecologists have used the concept of trophic state (TS) to characterize an aquatic ecosystem's biological productivity. However, multiple TS classification schemes, each relying on a variety of measurable parameters as proxies for productivity, have emerged to meet use-specific needs. Frequently, chlorophyll a, phosphorus, and Secchi depth are used to classify TS based on autotrophic production, whereas phosphorus, dissolved organic carbon, and true color are used to classify TS based on both autotrophic and heterotrophic production. Both classification approaches aim to characterize an ecosystem's function broadly, but with varying degrees of autotrophic and heterotrophic processes considered in those characterizations. Moreover, differing classification schemes can create inconsistent interpretations of ecosystem integrity. For example, the US Clean Water Act focuses exclusively on algal threats to water quality, framed in terms of eutrophication in response to nutrient loading. This usage lacks information about non-algal threats to water quality, such as dystrophication in response to dissolved organic carbon loading. Consequently, the TS classification schemes used to identify eutrophication and dystrophication may refer to ecosystems similarly (e.g., oligotrophic and eutrophic), yet these categories are derived from different proxies. These inconsistencies in TS classification schemes may be compounded when interdisciplinary projects employ varied TS frameworks. Even with these shortcomings, TS can still be used to distill information on complex aquatic ecosystem function into a set of generalizable expectations. The usefulness of distilling complex information into a TS index is substantial such that usage inconsistencies should be explicitly addressed and resolved. To emphasize the consequences of diverging TS classification schemes, we present three case studies for which an improved understanding of the TS concept advances freshwater research, management efforts, and interdisciplinary collaboration. To increase clarity in TS, the aquatic sciences could benefit from including information about the proxy variables, ecosystem type, as well as the spatiotemporal domains used to classify TS. As the field of aquatic sciences expands and climatic irregularity increases, we highlight the importance of re-evaluating fundamental concepts, such as TS, to ensure their compatibility with evolving science.

Ecosphere

Stream metabolism increases with drainage area and peaks asynchronously across a stream network

Quantifying the spatial and temporal dynamics of stream metabolism across stream networks is key to understanding carbon cycling and stream food web ecology. To better understand intra-annual temporal patterns of gross primary production (GPP) and ecosystem respiration (ER) and their variability across space, we continuously measured dissolved oxygen and modeled stream metabolism for an entire year at ten sites across a temperate river network in Washington State, USA. We expected GPP and ER to increase with stream size and peak during summer and autumn months due to warmer temperatures and higher light availability. We found that GPP and ER increased with drainage area and that only four sites adhered to our expectations of summer peaks in GPP and autumn peaks in ER while the rest either peaked in winter, spring or remained relatively constant. Our results suggest the spatial arrangement and temporal patterns of discharge, temperature, light and nutrients within watersheds may result in asynchronies in GPP and ER, despite similar regional climatic conditions. These findings shed light on how temporal dynamics of stream metabolism can shift across a river network, which likely influence the dynamics of carbon cycling and stream food webs at larger scales.

Washington

Regime shifts and panarchies in regional scale social-ecological water systems

In this article we summarize histories of nonlinear, complex interactions among societal, legal, and ecosystem dynamics in six North American water basins, as they respond to changing climate. These case studies were chosen to explore the conditions for emergence of adaptive governance in heavily regulated and developed social-ecological systems nested within a hierarchical governmental system. We summarize resilience assessments conducted in each system to provide a synthesis and reference by the other articles in this special feature. We also present a general framework used to evaluate the interactions between society and ecosystem regimes and the governance regimes chosen to mediate those interactions. The case studies show different ways that adaptive governance may be triggered, facilitated, or constrained by ecological and/or legal processes. The resilience assessments indicate that complex interactions among the governance and ecosystem components of these systems can produce different trajectories, which include patterns of (a) development and stabilization, (b) cycles of crisis and recovery, which includes lurches in adaptation and learning, and (3) periods of innovation, novelty, and transformation. Exploration of cross scale (Panarchy) interactions among levels and sectors of government and society illustrate that they may constrain development trajectories, but may also provide stability during crisis or innovation at smaller scales; create crises, but may also facilitate recovery; and constrain system transformation, but may also provide windows of opportunity in which transformation, and the resources to accomplish it, may occur. The framework is the starting point for our exploration of how law might play a role in enhancing the capacity of social-ecological systems to adapt to climate change.

Ecology and Society

Balancing stability and flexibility in adaptive governance: An analysis of tools available in U.S. environmental law

Adaptive governance must work “on the ground,” that is, it must operate through structures and procedures that the people it governs perceive to be legitimate and fair, as well as incorporating processes and substantive goals that are effective in allowing social-ecological systems (SESs) to adapt to climate change and other impacts. To address the continuing and accelerating alterations that climate change is bringing to SESs, adaptive governance generally will require more flexibility than prior governance institutions have often allowed. However, to function as good governance, adaptive governance must pay real attention to the problem of how to balance this increased need for flexibility with continuing governance stability so that it can foster adaptation to change without being perceived or experienced as perpetually destabilizing, disruptive, and unfair. Flexibility and stability serve different purposes in governance, and a variety of tools exist to strike different balances between them while still preserving the governance institution’s legitimacy among the people governed. After reviewing those purposes and the implications of climate change for environmental governance, we examine psychological insights into the structuring of adaptive governance and the variety of legal tools available to incorporate those insights into adaptive governance regimes. Because the substantive goals of governance systems will differ among specific systems, we do not purport to comment on what the normative or substantive goals of law should be. Instead, we conclude that attention to process and procedure (including participation), as well as increased use of substantive standards (instead of rules), may allow an increased level of substantive flexibility to operate with legitimacy and fairness, providing the requisite levels of psychological, social, and economic stability needed for communities to adapt successfully to the Anthropocene.

Ecology and Society

Critical considerations for the application of environmental DNA methods to detect aquatic species

Species detection using environmental DNA (eDNA) has tremendous potential for contributing to the understanding of the ecology and conservation of aquatic species. Detecting species using eDNA methods, rather than directly sampling the organisms, can reduce impacts on sensitive species and increase the power of field surveys for rare and elusive species. The sensitivity of eDNA methods, however, requires a heightened awareness and attention to quality assurance and quality control protocols. Additionally, the interpretation of eDNA data demands careful consideration of multiple factors. As eDNA methods have grown in application, diverse approaches have been implemented to address these issues. With interest in eDNA continuing to expand, supportive guidelines for undertaking eDNA studies are greatly needed. Environmental DNA researchers from around the world have collaborated to produce this set of guidelines and considerations for implementing eDNA methods to detect aquatic macroorganisms. Critical considerations for study design include preventing contamination in the field and the laboratory, choosing appropriate sample analysis methods, validating assays, testing for sample inhibition and following minimum reporting guidelines. Critical considerations for inference include temporal and spatial processes, limits of correlation of eDNA with abundance, uncertainty of positive and negative results, and potential sources of allochthonous DNA. We present a synthesis of knowledge at this stage for application of this new and powerful detection method.

Methods in Ecology and Evolution

The response of stream periphyton to Pacific salmon: using a model to understand the role of environmental context

1. In stream ecosystems, Pacific salmon deliver subsidies of marine-derived nutrients and disturb the stream bed during spawning. The net effect of this nutrient subsidy and physical disturbance on biological communities can be hard to predict and is likely to be mediated by environmental conditions. For periphyton, empirical studies have revealed that the magnitude and direction of the response to salmon varies from one location to the next. Salmon appear to increase periphyton biomass and/or production in some contexts (a positive response), but decrease them in others (a negative response). 2. To reconcile these seemingly conflicting results, we constructed a system dynamics model that links periphyton biomass and production to salmon spawning. We used this model to explore how environmental conditions influence the periphyton response to salmon. 3. Our simulations suggest that the periphyton response to salmon is strongly mediated by both background nutrient concentrations and the proportion of the stream bed suitable for spawning. Positive periphyton responses occurred when both background nutrient concentrations were low (nutrient limiting conditions) and when little of the stream bed was suitable for spawning (because the substratum is too coarse). In contrast, negative responses occurred when nutrient concentrations were higher or a larger proportion of the bed was suitable for spawning. 4. Although periphyton biomass generally remained above or below background conditions for several months following spawning, periphyton production returned quickly to background values shortly afterwards. As a result, based upon our simulations, salmon did not greatly increase or decrease overall annual periphyton production. This suggests that any increase in production by fish or invertebrates in response to returning salmon is more likely to occur via direct consumption of salmon carcasses and/or eggs, rather than the indirect effects of greater periphyton production. 5. Overall, our simulations suggest that environmental factors need to be taken into account when considering the effects of spawning salmon on aquatic ecosystems. Our model offers researchers a framework for testing periphyton response to salmon across a range of conditions, which can be used to generate hypotheses, plan field experiments and guide data collection.

Freshwater Biology