Managing aquatic habitat structure for resilient trophic interactions.

Structural habitat (the three-dimensional arrangement of physical matter, abiotic and biotic, at a location) is a foundational element for the resilience and maintenance of biodiversity, yet anthropogenic development is driving the global simplification of aquatic environments. Resource managers regularly seek to conserve aquatic food webs by increasing structural habitat complexity with expected benefits to fisheries; however, the global effectiveness of such actions is unclear. Our synthesis and theoretical analyses found that the response of a consumer-resource interaction (predatory sportfish and forage fish prey) to the addition of prey refuge habitat differed among systems with low and high rates of biomass transfer from resource to consumer (i.e., biomass potential); stabilization was not the rule. Greater prey refuge habitat availability tended to stabilize systems characterized by high biomass potential while simultaneously increasing consumer densities. In contrast, increasing prey refuge habitat availability in systems characterized by low biomass potential tended to mute energy transfer and moved consumer densities toward local extinction. Importantly, biomass potential and prey refuge can have antagonistic effects on stability and relative consumer densities, and it is therefore important to consider the local conditions of a system when using habitat manipulation as a management measure. Further development of our context-dependent perspective to whole food webs, and across different environments, may help to guide structural habitat management to better restore and protect aquatic ecosystems.

Climate and landscape conditions indirectly affect fish mercury levels by altering lake water chemistry and fish size.

Mercury pollution is a global environmental problem that threatens ecosystems, and negatively impacts human health and well-being. Mercury accumulation in fish within freshwater lakes is a complex process that appears to be driven by factors such as individual fish biology and water chemistry at the lake-scale, whereas, climate, and land-use/land-cover conditions within lake catchments can be influential at relatively larger scales. Nevertheless, unravelling the intricate network of pathways that govern how lake-scale and large-scale factors interact to affect mercury levels in fish remains an important scientific challenge. Using structural equation models (SEMs) and multiple long-term databases we identified direct and indirect effects of lake-scale and larger-scale factors on mercury levels in Walleye and Northern Pike - two species that are valued in inland fisheries. At the lake-level, the most parsimonious path models contained direct effects of fish weight, DOC, and pH, as well as an indirect effect of DOC on fish mercury levels via fish weight. Interestingly, lakeshed-, climate-, and full-path models that combine the effects of both lakeshed and climate revealed indirect effects of surrounding landscape conditions and latitude via DOC, pH, and fish weight but no direct effects on fish mercury levels. These results are generally consistent across species and lakes, except for some differences between stratified and non-stratified lakes. Our findings imply that understanding climate and land-use driven alterations of water chemistry and fish biology will be critical to predicting and mitigating fish mercury bioaccumulation in the future.

Food web rewiring in a changing world.

Climate change is asymmetrically altering environmental conditions in space, from local to global scales, creating novel heterogeneity. Here, we argue that this novel heterogeneity will drive mobile generalist consumer species to rapidly respond through their behaviour in ways that broadly and predictably reorganize - or rewire - food webs. We use existing theory and data from diverse ecosystems to show that the rapid behavioural responses of generalists to climate change rewire food webs in two distinct and critical ways. First, mobile generalist species are redistributing into systems where they were previously absent and foraging on new prey, resulting in topological rewiring - a change in the patterning of food webs due to the addition or loss of connections. Second, mobile generalist species, which navigate between habitats and ecosystems to forage, will shift their relative use of differentially altered habitats and ecosystems, causing interaction strength rewiring - changes that reroute energy and carbon flows through existing food web connections and alter the food web's interaction strengths. We then show that many species with shared traits can exhibit unified aggregate behavioural responses to climate change, which may allow us to understand the rewiring of whole food webs. We end by arguing that generalists' responses present a powerful and underutilized approach to understanding and predicting the consequences of climate change and may serve as much-needed early warning signals for monitoring the looming impacts of global climate change on entire ecosystems.

Context-dependent interactions and the regulation of species richness in freshwater fish.

Species richness is regulated by a complex network of scale-dependent processes. This complexity can obscure the influence of limiting species interactions, making it difficult to determine if abiotic or biotic drivers are more predominant regulators of richness. Using integrative modeling of freshwater fish richness from 721 lakes along an 11o latitudinal gradient, we find negative interactions to be a relatively minor independent predictor of species richness in lakes despite the widespread presence of predators. Instead, interaction effects, when detectable among major functional groups and 231 species pairs, were strong, often positive, but contextually dependent on environment. These results are consistent with the idea that negative interactions internally structure lake communities but do not consistently 'scale-up' to regulate richness independently of the environment. The importance of environment for interaction outcomes and its role in the regulation of species richness highlights the potential sensitivity of fish communities to the environmental changes affecting lakes globally.

Blinded by the light? Nearshore energy pathway coupling and relative predator biomass increase with reduced water transparency across lakes.

Habitat coupling is a concept that refers to consumer integration of resources derived from different habitats. This coupling unites fundamental food web pathways (e.g., cross-habitat trophic linkages) that mediate key ecological processes such as biomass flows, nutrient cycling, and stability. We consider the influence of water transparency, an important environmental driver in aquatic ecosystems, on habitat coupling by a light-sensitive predator, walleye (Sander vitreus), and its prey in 33 Canadian lakes. Our large-scale, across-lake study shows that the contribution of nearshore carbon (δ13C) relative to offshore carbon (δ13C) to walleye is higher in less transparent lakes. To a lesser degree, the contribution of nearshore carbon increased with a greater proportion of prey in nearshore compared to offshore habitats. Interestingly, water transparency and habitat coupling predict among-lake variation in walleye relative biomass. These findings support the idea that predator responses to changing conditions (e.g., water transparency) can fundamentally alter carbon pathways, and predator biomass, in aquatic ecosystems. Identifying environmental factors that influence habitat coupling is an important step toward understanding spatial food web structure in a changing world.

The consistency of a species' response to press perturbations with high food web uncertainty.

Predicting species responses to perturbations is a fundamental challenge in ecology. Decision makers must often identify management perturbations that are the most likely to deliver a desirable management outcome despite incomplete information on the pattern and strength of food web links. Motivated by a current fishery decline in inland lakes of the Midwestern United States, we evaluate consistency of the responses of a target species (walleye [Sander vitreus]) to press perturbations. We represented food web uncertainty with 193 plausible topological models and applied four perturbations to each one. Frequently the direction of the focal predator response to the same perturbation is not consistent across food web topologies. Simultaneous application of management perturbations led to less consistent outcomes compared to the best single perturbation. However, direct manipulation of the adult focal predator produced a desirable outcome in 77% of 193 plausible topologies. Identifying perturbations that produce consistent outcomes in the face of food web uncertainty can have important implications for natural resource conservation and management efforts.

The body size dependence of trophic cascades.

Trophic cascades are indirect positive effects of predators on resources via control of intermediate consumers. Larger-bodied predators appear to induce stronger trophic cascades (a greater rebound of resource density toward carrying capacity), but how this happens is unknown because we lack a clear depiction of how the strength of trophic cascades is determined. Using consumer resource models, we first show that the strength of a trophic cascade has an upper limit set by the interaction strength between the basal trophic group and its consumer and that this limit is approached as the interaction strength between the consumer and its predator increases. We then express the strength of a trophic cascade explicitly in terms of predator body size and use two independent parameter sets to calculate how the strength of a trophic cascade depends on predator size. Both parameter sets predict a positive effect of predator size on the strength of a trophic cascade, driven mostly by the body size dependence of the interaction strength between the first two trophic levels. Our results support previous empirical findings and suggest that the loss of larger predators will have greater consequences on trophic control and biomass structure in food webs than the loss of smaller predators.

A bioenergetic framework for the temperature dependence of trophic interactions.

Changing temperature can substantially shift ecological communities by altering the strength and stability of trophic interactions. Because many ecological rates are constrained by temperature, new approaches are required to understand how simultaneous changes in multiple rates alter the relative performance of species and their trophic interactions. We develop an energetic approach to identify the relationship between biomass fluxes and standing biomass across trophic levels. Our approach links ecological rates and trophic dynamics to measure temperature-dependent changes to the strength of trophic interactions and determine how these changes alter food web stability. It accomplishes this by using biomass as a common energetic currency and isolating three temperature-dependent processes that are common to all consumer-resource interactions: biomass accumulation of the resource, resource consumption and consumer mortality. Using this framework, we clarify when and how temperature alters consumer to resource biomass ratios, equilibrium resilience, consumer variability, extinction risk and transient vs. equilibrium dynamics. Finally, we characterise key asymmetries in species responses to temperature that produce these distinct dynamic behaviours and identify when they are likely to emerge. Overall, our framework provides a mechanistic and more unified understanding of the temperature dependence of trophic dynamics in terms of ecological rates, biomass ratios and stability.

Effects of differential habitat warming on complex communities.

Food webs unfold across a mosaic of micro and macro habitats, with each habitat coupled by mobile consumers that behave in response to local environmental conditions. Despite this fundamental characteristic of nature, research on how climate change will affect whole ecosystems has overlooked (i) that climate warming will generally affect habitats differently and (ii) that mobile consumers may respond to this differential change in a manner that may fundamentally alter the energy pathways that sustain ecosystems. This reasoning suggests a powerful, but largely unexplored, avenue for studying the impacts of climate change on ecosystem functioning. Here, we use lake ecosystems to show that predictable behavioral adjustments to local temperature differentials govern a fundamental structural shift across 54 food webs. Data show that the trophic pathways from basal resources to a cold-adapted predator shift toward greater reliance on a cold-water refuge habitat, and food chain length increases, as air temperatures rise. Notably, cold-adapted predator behavior may substantially drive this decoupling effect across the climatic range in our study independent of warmer-adapted species responses (for example, changes in near-shore species abundance and predator absence). Such modifications reflect a flexible food web architecture that requires more attention from climate change research. The trophic pathway restructuring documented here is expected to alter biomass accumulation, through the regulation of energy fluxes to predators, and thus potentially threatens ecosystem sustainability in times of rapid environmental change.

Increased temperature variation poses a greater risk to species than climate warming.

Increases in the frequency, severity and duration of temperature extremes are anticipated in the near future. Although recent work suggests that changes in temperature variation will have disproportionately greater effects on species than changes to the mean, much of climate change research in ecology has focused on the impacts of mean temperature change. Here, we couple fine-grained climate projections (2050-2059) to thermal performance data from 38 ectothermic invertebrate species and contrast projections with those of a simple model. We show that projections based on mean temperature change alone differ substantially from those incorporating changes to the variation, and to the mean and variation in concert. Although most species show increases in performance at greater mean temperatures, the effect of mean and variance change together yields a range of responses, with temperate species at greatest risk of performance declines. Our work highlights the importance of using fine-grained temporal data to incorporate the full extent of temperature variation when assessing and projecting performance.

Food web expansion and contraction in response to changing environmental conditions.

Macroscopic ecosystem properties, such as major material pathways and community biomass structure, underlie the ecosystem services on which humans rely. While ecologists have long sought to identify the determinants of the trophic height of food webs (food chain length), it is somewhat surprising how little research effort is invested in understanding changes among other food web properties across environmental conditions. Here we theoretically and empirically show how a suite of fundamental macroscopic food web structures respond, in concert, to changes in habitat accessibility using post-glacial lakes as model ecosystems. We argue that as resource accessibility increases in coupled food webs, food chain length contracts (that is, reduced predator trophic position), habitat coupling expands (that is, increasingly coupled macrohabitats) and biomass pyramid structure becomes more top heavy. Our results further support an emerging theoretical view of flexible food webs that provides a foundation for generally understanding ecosystem responses to changing environmental conditions.