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The application of metacommunity theory to the management of riverine ecosystems

River managers strive to use the best available science to sustain biodiversity and ecosystem function. To achieve this goal requires consideration of processes at different scales. Metacommunity theory describes how multiple species from different communities potentially interact with local-scale environmental drivers to influence population dynamics and community structure. However, this body of knowledge has only rarely been used to inform management practices for river ecosystems. In this article, we present a conceptual model outlining how the metacommunity processes of local niche sorting and dispersal can influence the outcomes of management interventions and provide a series of specific recommendations for applying these ideas as well as research needs. In all cases, we identify situations where traditional approaches to riverine management could be enhanced by incorporating an understanding of metacommunity dynamics. A common theme is developing guidelines for assessing the metacommunity context of a site or region, evaluating how that context may affect the desired outcome, and incorporating that understanding into the planning process and methods used. To maximize the effectiveness of management activities, scientists, and resource managers should update the toolbox of approaches to riverine management to reflect theoretical advances in metacommunity ecology.

WIREs Water

Nonlinear relationships can lead to bias in biomass calculations and drift-foraging models when using summaries of invertebrate drift data

Drift-foraging models offer a mechanistic description of how fish feed in flowing water and the application of drift-foraging bioenergetics models to answer both applied and theoretical questions in aquatic ecology is growing. These models typically include nonlinear descriptions of ecological processes and as a result may be sensitive to how model inputs are summarized because of a mathematical property of nonlinear equations known as Jensen’s inequality. In particular, we show that the way in which continuous size distributions of invertebrate prey are represented within foraging models can lead to biases within the modeling process. We begin by illustrating how different equations common to drift-foraging models are sensitive to invertebrate inputs. We then use two case studies to show how different representations of invertebrate prey can influence predictions of energy intake and lifetime growth. Greater emphasis should be placed on accurate characterizations of invertebrate drift, acknowledging that inferences from drift-foraging models may be influenced by how invertebrate prey are represented.

Environmental Biology of Fishes

Linking biodiversity to ecosystem function: Implications for conservation ecology

We evaluate the empirical and theoretical support for the hypothesis that a large proportion of native species richness is required to maximize ecosystem stability and sustain function. This assessment is important for conservation strategies because sustenance of ecosystem functions has been used as an argument for the conservation of species. If ecosystem functions are sustained at relatively low species richness, then arguing for the conservation of ecosystem function, no matter how important in its own right, does not strongly argue for the conservation of species. Additionally, for this to be a strong conservation argument the link between species diversity and ecosystem functions of value to the human community must be clear. We review the empirical literature to quantify the support for two hypotheses: (1) species richness is positively correlated with ecosystem function, and (2) ecosystem functions do not saturate at low species richness relative to the observed or experimental diversity. Few empirical studies demonstrate improved function at high levels of species richness. Second, we analyze recent theoretical models in order to estimate the level of species richness required to maintain ecosystem function. Again we find that, within a single trophic level, most mathematical models predict saturation of ecosystem function at a low proportion of local species richness. We also analyze a theoretical model linking species number to ecosystem stability. This model predicts that species richness beyond the first few species does not typically increase ecosystem stability. One reason that high species richness may not contribute significantly to function or stability is that most communities are characterized by strong dominance such that a few species provide the vast majority of the community biomass. Rapid turnover of species may rescue the concept that diversity leads to maximum function and stability. The role of turnover in ecosystem function and stability has not been investigated. Despite the recent rush to embrace the linkage between biodiversity and ecosystem function, we find little support for the hypothesis that there is a strong dependence of ecosystem function on the full complement of diversity within sites. Given this observation, the conservation community should take a cautious view of endorsing this linkage as a model to promote conservation goals.

Oecologia

Stochastic ecological network occupancy (SENO) models: a new tool for modeling ecological networks across spatial scales

Stochastic ecological network occupancy (SENO) models predict the probability that species will occur in a sample of an ecological network. In this review, we introduce SENO models as a means to fill a gap in the theoretical toolkit of ecologists. As input, SENO models use a topological interaction network and rates of colonization and extinction (including consumer effects) for each species. A SENO model then simulates the ecological network over time, resulting in a series of sub-networks that can be used to identify commonly encountered community modules. The proportion of time a species is present in a patch gives its expected probability of occurrence, whose sum across species gives expected species richness. To illustrate their utility, we provide simple examples of how SENO models can be used to investigate how topological complexity, species interactions, species traits, and spatial scale affect communities in space and time. They can categorize species as biodiversity facilitators, contributors, or inhibitors, making this approach promising for ecosystem-based management of invasive, threatened, or exploited species.

Theoretical Ecology

Magnitude and direction of stream–forest community interactions change with timescale

Networks of direct and indirect biotic interactions underpin the complex dynamics and stability of ecological systems, yet experimental and theoretical studies often yield conflicting evidence regarding the direction (positive or negative) or magnitude of these interactions. We revisited pioneering data sets collected at the deciduous forested Horonai Stream and conducted ecosystem‐level syntheses to demonstrate that the direction of direct and indirect interactions can change depending on the timescale of observation. Prior experimental studies showed that terrestrial prey that enter the stream from the adjacent forest caused positive indirect effects on aquatic invertebrates during summer by diverting fish consumption. Seasonal and annual estimates of secondary production and organic matter flows along food web pathways demonstrate that this seasonal input of terrestrial invertebrate prey increases production of certain fish species, reversing the indirect effect on aquatic invertebrates from positive at the seasonal timescale to negative at the annual timescale. Even though terrestrial invertebrate prey contributed 54% of the annual organic matter flux to fishes, primarily during summer, fish still consumed 98% of the aquatic invertebrate annual production, leading to top‐down control that is not revealed in short‐term experiments and demonstrating that aquatic prey may be a limiting resource for fishes. Changes in the direction or magnitude of interactions may be a key factor creating nonlinear or stabilizing feedbacks in complex systems, and these dynamics can be revealed by merging experimental and comparative approaches at different scales.

Ecology

Effects of diffusion on total biomass in heterogeneous continuous and discrete-patch systems

Theoretical models of populations on a system of two connected patches previously have shown that when the two patches differ in maximum growth rate and carrying capacity, and in the limit of high diffusion, conditions exist for which the total population size at equilibrium exceeds that of the ideal free distribution, which predicts that the total population would equal the total carrying capacity of the two patches. However, this result has only been shown for the Pearl-Verhulst growth function on two patches and for a single-parameter growth function in continuous space. Here, we provide a general criterion for total population size to exceed total carrying capacity for three commonly used population growth rates for both heterogeneous continuous and multi-patch heterogeneous landscapes with high population diffusion. We show that a sufficient condition for this situation is that there is a convex positive relationship between the maximum growth rate and the parameter that, by itself or together with the maximum growth rate, determines the carrying capacity, as both vary across a spatial region. This relationship occurs in some biological populations, though not in others, so the result has ecological implications.

Theoretical Ecology

Toward a social-ecological theory of forest macrosystems for improved ecosystem management

The implications of cumulative land-use decisions and shifting climate on forests, require us to integrate our understanding of ecosystems, markets, policy, and resource management into a social-ecological system. Humans play a central role in macrosystem dynamics, which complicates ecological theories that do not explicitly include human interactions. These dynamics also impact ecological services and related markets, which challenges economic theory. Here, we use two forest macroscale management initiatives to develop a theoretical understanding of how management interacts with ecological functions and services at these scales and how the multiple large-scale management goals work either in consort or conflict with other forest functions and services. We suggest that calling upon theories developed for organismal ecology, ecosystem ecology, and ecological economics adds to our understanding of social-ecological macrosystems. To initiate progress, we propose future research questions to add rigor to macrosystem-scale studies: (1) What are the ecosystem functions that operate at macroscales, their necessary structural components, and how do we observe them? (2) How do systems at one scale respond if altered at another scale? (3) How do we both effectively measure these components and interactions, and communicate that information in a meaningful manner for policy and management across different scales?

Forests

Complexity and integration of recreational fisheries

Recreational fisheries are interconnected, complex, adaptive systems characterized by multiple direct and indirect interactions among ecological and human subsystems. This is important for many reasons, including that feedbacks between the social and ecological dimensions lead to difficult-to-predict, often entirely unexpected, outcomes and because many management and governance systems have multiple objectives that can involve social (e.g. fisher satisfaction), economic (e.g. license revenue), and ecological (e.g. fish conservation) dimensions. Embracing a social-ecological-system perspective can usher in an improved era of recreational fisheries science and management. Interdisciplinary approaches that unite experts across disciplines (e.g. social and ecological sciences) to create a unique theoretical, conceptual, and methodological identity are needed to gather crucial information from recreational fishers, quantify and predict fisher behaviours and outcomes from these behaviours, and integrate these findings into fisheries management. In this chapter, we lay the conceptual foundation for recreational fisheries as coupled social-ecological systems that are also complex and adaptive, and discuss the interdisciplinary approach to operationalize this book’s vision.

Book chapter

More than a meal: integrating non-feeding interactions into food webs

Organisms eating each other are only one of many types of well documented and important interactions among species. Other such types include habitat modification, predator interference and facilitation. However, ecological network research has been typically limited to either pure food webs or to networks of only a few (<3) interaction types. The great diversity of non-trophic interactions observed in nature has been poorly addressed by ecologists and largely excluded from network theory. Herein, we propose a conceptual framework that organises this diversity into three main functional classes defined by how they modify specific parameters in a dynamic food web model. This approach provides a path forward for incorporating non-trophic interactions in traditional food web models and offers a new perspective on tackling ecological complexity that should stimulate both theoretical and empirical approaches to understanding the patterns and dynamics of diverse species interactions in nature.

Ecology Letters

Numerical computation of hurricane effects on historic coastal hydrology in Southern Florida

Introduction Numerical models are critical for assessing the effects of sea level rise (SLR), hurricanes, and storm surge on vegetation change in the Everglades National Park. The model must be capable of representing short-timescale hydrodynamics, salinity transport, and groundwater interaction. However, there is also a strong need to adapt these numerical models to hindcast past conditions in order to examine long-term effects on the distribution of vegetation that cannot be determined using only the modern record. Methods Based on parameters developed for a numerical model developed for the recent 1996 to 2004 period, a hindcast model was developed to represent sea level and water management for the period of 1926 to 1932, constrained by the limited hydrology and meteorology data available from the historic past. Realistic hurricane-wind and storm surge representations, required for the hindcast model, are based on information synthesized from modern storm data. A series of simulation scenarios with various hurricane representations inserted into both hindcast and recent numerical models were used to assess the utility of the storm representation in the model and compare the two simulations. Results The comparison of the hindcast and recent models showed differences in the hydrology patterns that are consistent with known differences in water delivery systems and sea level rise. A 30x lower-resolution spatially variable wind grid for the hindcast produced similar results to the original high-resolution full wind grid representation of the recent simulation. Storm effects on hydrologic patterns demonstrated with the simulations show hydrologic processes that could have a long-term effect on vegetation change. Conclusions The hindcast simulation estimated hydrologic processes for the 1926 to 1932 period. It shows promise as a simulator in long-term ecological studies to test hypotheses based on theoretical or empirical-based studies at larger landscape scales.

Florida

On the specification of structural equation models for ecological systems

The use of structural equation modeling (SEM) is often motivated by its utility for investigating complex networks of relationships, but also because of its promise as a means of representing theoretical concepts using latent variables. In this paper, we discuss characteristics of ecological theory and some of the challenges for proper specification of theoretical ideas in structural equation models (SE models). In our presentation, we describe some of the requirements for classical latent variable models in which observed variables (indicators) are interpreted as the effects of underlying causes. We also describe alternative model specifications in which indicators are interpreted as having causal influences on the theoretical concepts. We suggest that this latter nonclassical specification (which involves another variable type—the composite) will often be appropriate for ecological studies because of the multifaceted nature of our theoretical concepts. In this paper, we employ the use of meta‐models to aid the translation of theory into SE models and also to facilitate our ability to relate results back to our theories. We demonstrate our approach by showing how a synthetic theory of grassland biodiversity can be evaluated using SEM and data from a coastal grassland. In this example, the theory focuses on the responses of species richness to abiotic stress and disturbance, both directly and through intervening effects on community biomass. Models examined include both those based on classical forms (where each concept is represented using a single latent variable) and also ones in which the concepts are recognized to be multifaceted and modeled as such. To address the challenge of matching SE models with the conceptual level of our theory, two approaches are illustrated, compositing and aggregation. Both approaches are shown to have merits, with the former being preferable for cases where the multiple facets of a concept have widely differing effects in the system and the latter being preferable where facets act together consistently when influencing other parts of the system. Because ecological theory characteristically deals with concepts that are multifaceted, we expect the methods presented in this paper will be useful for ecologists wishing to use SEM.

Ecological Monographs

Roosting and foraging social structure of the endangered Indiana bat ( Myotis sodalis )

Social dynamics are an important but poorly understood aspect of bat ecology. Herein we use a combination of graph theoretic and spatial approaches to describe the roost and social network characteristics and foraging associations of an Indiana bat ( Myotis sodalis ) maternity colony in an agricultural landscape in Ohio, USA. We tracked 46 bats to 50 roosts (423 total relocations) and collected 2,306 foraging locations for 40 bats during the summers of 2009 and 2010. We found the colony roosting network was highly centralized in both years and that roost and social networks differed significantly from random networks. Roost and social network structure also differed substantially between years. Social network structure appeared to be unrelated to segregation of roosts between age classes. For bats whose individual foraging ranges were calculated, many shared foraging space with at least one other bat. Compared across all possible bat dyads, 47% and 43% of the dyads showed more than expected overlap of foraging areas in 2009 and 2010 respectively. Colony roosting area differed between years, but the roosting area centroid shifted only 332 m. In contrast, whole colony foraging area use was similar between years. Random roost removal simulations suggest that Indiana bat colonies may be robust to loss of a limited number of roosts but may respond differently from year to year. Our study emphasizes the utility of graphic theoretic and spatial approaches for examining the sociality and roosting behavior of bats. Detailed knowledge of the relationships between social and spatial aspects of bat ecology could greatly increase conservation effectiveness by allowing more structured approaches to roost and habitat retention for tree-roosting, socially-aggregating bat species.

Ohio

Spatial ecology of refuge selection by an herbivore under risk of predation

Prey species use structures such as burrows to minimize predation risk. The spatial arrangement of these resources can have important implications for individual and population fitness. For example, there is evidence that clustered resources can benefit individuals by reducing predation risk and increasing foraging opportunity concurrently, which leads to higher population density. However, the scale of clustering that is important in these processes has been ignored during theoretical and empirical development of resource models. Ecological understanding of refuge exploitation by prey can be improved by spatial analysis of refuge use and availability that incorporates the effect of scale. We measured the spatial distribution of pygmy rabbit (Brachylagus idahoensis) refugia (burrows) through censuses in four 6-ha sites. Point pattern analyses were used to evaluate burrow selection by comparing the spatial distribution of used and available burrows. The presence of food resources and additional overstory cover resources was further examined using logistic regression. Burrows were spatially clustered at scales up to approximately 25 m, and then regularly spaced at distances beyond ~40 m. Pygmy rabbit exploitation of burrows did not match availability. Burrows used by pygmy rabbits were likely to be located in areas with high overall burrow density (resource clusters) and high overstory cover, which together minimized predation risk. However, in some cases we observed an interaction between either overstory cover (safety) or understory cover (forage) and burrow density. The interactions show that pygmy rabbits will use burrows in areas with low relative burrow density (high relative predation risk) if understory food resources are high. This points to a potential trade-off whereby rabbits must sacrifice some safety afforded by additional nearby burrows to obtain ample forage resources. Observed patterns of clustered burrows and non-random burrow use improve understanding of the importance of spatial distribution of refugia for burrowing herbivores. The analyses used allowed for the estimation of the spatial scale where subtle trade-offs between predation avoidance and foraging opportunity are likely to occur in a natural system.

Ecosphere

Are all prey created equal? A review and synthesis of differential predation on prey in substandard condition

Our understanding of predator-prey interactions in fishes has been influenced largely by research assuming that the condition of the participants is normal. However, fish populations today often reside in anthropogenically altered environments and are subjected to many kinds of stressors, which may reduce their ecological performance by adversely affecting their morphology, physiology, or behaviour. One consequence is that either the predator or prey, or both, may be in a substandard condition at the time of an interaction. We reviewed the literature on predator-prey interactions in fishes where substandard prey were used as experimental groups. Although most of this research indicates that such prey are significantly more vulnerable to predation, prey condition has rarely been considered in ecological theory regarding predator-prey interactions. The causal mechanisms for increased vulnerability of substandard prey to predation include a failure to detect predators, lapses in decision-making, poor fast-start performance, inability to shoal effectively, and increased prey conspicuousness. Despite some problems associated with empirical predator-prey studies using substandard prey, their results can have theoretical and applied uses, such as in ecological modelling or justification of corrective measures to be implemented in the wild. There is a need for more corroborative field experimentation, a better understanding of the causal mechanisms behind differential predation, and increased incorporation of prey condition into the research of predator-prey modellers and theoreticians. If the concept of prey condition is considered in predator-prey interactions, our understanding of how such interactions influence the structure and dynamics of fish communities is likely to change, which should prove beneficial to aquatic ecosystems.

Journal of Fish Biology

Taming wildlife disease: bridging the gap between science and management

1.Parasites and pathogens of wildlife can threaten biodiversity, infect humans and domestic animals, and cause significant economic losses, providing incentives to manage wildlife diseases. Recent insights from disease ecology have helped transform our understanding of infectious disease dynamics and yielded new strategies to better manage wildlife diseases. Simultaneously, wildlife disease management (WDM) presents opportunities for large-scale empirical tests of disease ecology theory in diverse natural systems. 2.To assess whether the potential complementarity between WDM and disease ecology theory has been realized, we evaluate the extent to which specific concepts in disease ecology theory have been explicitly applied in peer-reviewed WDM literature. 3.While only half of WDM articles published in the past decade incorporated disease ecology theory, theory has been incorporated with increasing frequency over the past 40 years. Contrary to expectations, articles authored by academics were no more likely to apply disease ecology theory, but articles that explain unsuccessful management often do so in terms of theory. 4.Some theoretical concepts such as density-dependent transmission have been commonly applied, whereas emerging concepts such as pathogen evolutionary responses to management, biodiversity–disease relationships and within-host parasite interactions have not yet been fully integrated as management considerations. 5.Synthesis and applications. Theory-based disease management can meet the needs of both academics and managers by testing disease ecology theory and improving disease interventions. Theoretical concepts that have received limited attention to date in wildlife disease management could provide a basis for improving management and advancing disease ecology in the future.

Journal of Applied Ecology

Biodiversity and Habitat Markets—Policy, Economic, and Ecological implications of Market-Based Conservation

This report is a primer on market-like and market-based mechanisms designed to conserve biodiversity and habitat. The types of markets and market-based approaches that were implemented or are emerging to benefit biodiversity and habitat in the United States are examined. The central approaches considered in this report include payments for ecosystem services, conservation banks, habitat exchanges, and eco-labels. Based on literature reviews and input from experts and practitioners, the report characterizes each market-based approach including policy context and structure; the theoretical basis for applying market-based approaches; the ecological effectiveness of practices and tools for measuring performance; and the future outlook for biodiversity and habitat markets. This report draws from previous research and serves as a summary of pertinent information associated with biodiversity and habitat markets while providing references to materials that go into greater detail on specific topics.

Circular

Ecological diffusion models are still useful in ecology

Context Connecting animal movement across scales—both among spatial scales and between individual and population level processes—is a central theme in ecology, including the sub-fields of movement and landscape ecology. However, most modeling frameworks operate at either the individual (Lagrangian) or population (Eulerian) level. Objectives We aimed to review how a partial differential equation (PDE) known as the ecological diffusion equation (EDE) offers connections among the scales of animal movement and how it can be embedded in commonly used statistical frameworks (e.g., hierarchical models) and fit to multiple types of data, such as count, presence-only, presence-absence, and telemetry data. In doing so, we also highlight emerging and promising avenues of research related to statistical EDE models. Methods We present how the EDE is derived from a simple set of first principles, can be used to represent both individual animal movement and spatiotemporal population dynamics, and can be fit to data by being applied in a hierarchical statistical framework. Finally, we review literature on recent applications and advances related to EDE models. Results In presenting the derivation of the EDE and reviewing recent advances and applications of EDE models, we established that the EDE can enable a mechanistic understanding of animal movement across scales, as well as the integration of multiple types of data. We also identified a number of topics where EDE models can be advanced to gain additional ecological insight. Conclusions Ecological diffusion is a process that emerges naturally from first principles governing animal movement and can be used to model both individual and population-level spatiotemporal dynamics. Statistical EDE models are related to other modeling frameworks, such as species distribution models, occupancy models, and step-selection analysis. The flexibility of hierarchical statistical modeling allows combining different types of data collected on movement and population dynamics, and the theoretical underpinnings of the EDE allow it to connect ecological processes occurring at fine and coarse spatial scales, as well as individual animal movement and population dynamics.

Landscape Ecology

Moving forward in circles: Challenges and opportunities in modeling population cycles

Population cycling is a widespread phenomenon, observed across a multitude of taxa in both laboratory and natural conditions. Historically, the theory associated with population cycles was tightly linked to pairwise consumer–resource interactions and studied via deterministic models, but current empirical and theoretical research reveals a much richer basis for ecological cycles. Stochasticity and seasonality can modulate or create cyclic behaviour in non-intuitive ways, the high-dimensionality in ecological systems can profoundly influence cycling, and so can demographic structure and eco-evolutionary dynamics. An inclusive theory for population cycles, ranging from ecosystem-level to demographic modelling, grounded in observational or experimental data, is therefore necessary to better understand observed cyclical patterns. In turn, by gaining better insight into the drivers of population cycles, we can begin to understand the causes of cycle gain and loss, how biodiversity interacts with population cycling, and how to effectively manage wildly fluctuating populations, all of which are growing domains of ecological research.

Ecology Letters