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The potential of high-frequency profiling to assess vertical and seasonal patterns of phytoplankton dynamics in lakes: An extension of the Plankton Ecology Group (PEG) model

The use of high-frequency sensors on profiling buoys to investigate physical, chemical, and biological processes in lakes is increasing rapidly. Profiling buoys with automated winches and sensors that collect high-frequency chlorophyll fluorescence (ChlF) profiles in 11 lakes in the Global Lake Ecological Observatory Network (GLEON) allowed the study of the vertical and temporal distribution of ChlF, including the formation of subsurface chlorophyll maxima (SSCM). The effectiveness of 3 methods for sampling phytoplankton distributions in lakes, including (1) manual profiles, (2) single-depth buoys, and (3) profiling buoys were assessed. High-frequency ChlF surface data and profiles were compared to predictions from the Plankton Ecology Group (PEG) model. The depth-integrated ChlF dynamics measured by the profiling buoy data revealed a greater complexity that neither conventional sampling nor the generalized PEG model captured. Conventional sampling techniques would have missed SSCM in 7 of 11 study lakes. Although surface-only ChlF data underestimated average water column ChlF, at times by nearly 2-fold in 4 of the lakes, overall there was a remarkable similarity between surface and mean water column data. Contrary to the PEG model’s proposed negligible role for physical control of phytoplankton during the growing season, thermal structure and light availability were closely associated with ChlF seasonal depth distribution. Thus, an extension of the PEG model is proposed, with a new conceptual framework that explicitly includes physical metrics to better predict SSCM formation in lakes and highlight when profiling buoys are especially informative.

Inland Waters

Individual-based ecological particle tracking model (ECO-PTM) for simulating juvenile chinook salmon migration and survival through the Sacramento – San Joaquin Delta

Recovery of endangered salmon species in the Central Valley of California amidst prolonged drought and climate change necessitates innovative water management actions that balance species recovery and California's water demands. We describe an individual-based ecological particle tracking model (ECO-PTM) that can be used to assess the efficacy of proposed actions. Based on a random walk theory, the model tracks individual particles’ travel time, routing and survival in a flow field simulated by the Delta Simulation Model 2 hydrodynamic module (DSM2 HYDRO). The random walk particles are parameterized to have fish-like swimming behaviors, including upstream/downstream swimming, probabilistic holding behaviors, and stochastic swimming velocities. Particle routing at key junctions is based on well-established statistical models, and route-specific survival is calculated using the XT mean free-path length model. Behavioral parameters were estimated by fitting several competing models to a multiyear dataset of travel times from acoustic tagged juvenile salmon. The model’s baseline simulations under historical flow conditions from 1991 to 2016 successfully replicated essential relationships between salmon outmigration survival and hydrodynamic conditions, consistent with previous studies and the STARS (Survival Travel Time and Routing Simulation) statistical simulation model. Simulation results for management scenarios revealed multifaceted influences on fish survival, including Delta flow, flow at key junctions, route alterations, seasons, and water year characteristics. Importantly, these results highlight ECO-PTM’s potential to predict fish survival outcomes of proposed actions, serving as a foundation for informed future research, decision-making, and effective management strategies to enhance the survival prospects of out-migrating salmonids within the Sacramento-San Joaquin Delta ecosystem.

California

Structural Equation Modeling: Applications in ecological and evolutionary biology

This book presents an introduction to the methodology of structural equation modeling, illustrates its use, and goes on to argue that it has revolutionary implications for the study of natural systems. A major theme of this book is that we have, up to this point, attempted to study systems primarily using methods (such as the univariate model) that were designed only for considering individual processes. Understanding systems requires the capacity to examine simultaneous influences and responses. Structural equation modeling (SEM) has such capabilities. It also possesses many other traits that add strength to its utility as a means of making scientific progress. In light of the capabilities of SEM, it can be argued that much of ecological theory is currently locked in an immature state that impairs its relevance. It is further argued that the principles of SEM are capable of leading to the development and evaluation of multivariate theories of the sort vitally needed for the conservation of natural systems. Supplementary information can be found at the authors website, http://www.jamesbgrace.com/ . • Details why multivariate analyses should be used to study ecological systems • Exposes unappreciated weakness in many current popular analyses • Emphasizes the future methodological developments needed to advance our understanding of ecological systems.

Book

Translating stakeholder narratives for participatory modeling in landscape ecology

Context Engaging stakeholders in research is needed for many of the sustainability challenges that landscape ecologists address. Involving stakeholders’ perspectives through narratives in participatory modeling fosters better understanding of the problem and evaluation of the acceptability of tradeoffs and creates buy-in for management actions. However, stakeholder-driven inputs often take the form of complex qualitative descriptions, rather than model-ready numerical or categorical inputs. Objectives Translating narratives into models, model parameters, or scenarios is essential for leveraging stakeholder knowledge and engagement. Drawing from varied experiences to identify lessons learned and pitfalls, we address the practice of translating narratives into models and using those narratives to interpret and communicate results. Methods We drew from seven participatory landscape ecology projects across North America to synthesize lessons for the inclusion of stakeholder narratives in modeling studies. Results We offer 8 lessons as practical guidance for other landscape ecologists to move the science beyond a unilateral focus on ecological systems and to maximize the benefits of landscape sustainability science. Conclusions These lessons are starting points, as real projects are complex, nuanced, and sometimes contradictory. Translating narratives into models is important for addressing complex sustainability challenges; we hope that these starting points are helpful to those foraying into this type of research.

Landscape Ecology

Hydrologic modeling for flow-ecology science in the Southeastern United States and Puerto Rico

An understanding of the applicability and utility of hydrologic models is critical to support the effective management of water resources throughout the Southeastern United States (SEUS) and Puerto Rico (PR). Hydrologic models have the capacity to provide an estimate of the quantity of available water at ungauged locations (i.e., areas of the country where a U.S. Geological Survey [USGS] continuous record gauge is not installed) and provide the baseline flow information necessary to develop the linkages between water availability and characteristics of streamflow that support ecological communities (i.e., support the development of flow-ecology response models). This report inventories and then directly examines and compares a subset of hydrologic models used to estimate streamflow at a number of gauged basins across the SEUS and PR. This effort was designed to evaluate, quantify, and compare the magnitude of error and to investigate the potential causes of error associated with predicted streamflows from seven hydrologic models of varying complexity and calibration strategy. This was accomplished by computing and then comparing classical hydrologic model fit statistics (e.g., mean bias, coefficient of determination [R2], root mean squared error [RMSE], Nash-Sutcliffe Efficiency [NSE]) and understanding the bias in the prediction in these and a subset of ecologically relevant flow metrics (ERFMs). Additionally, streamflow predictions from a larger regional-scale hydrologic model were compared to those of several fine-scale hydrologic models under a range of hypothetical climate change scenarios to determine the range of predicted streamflow responses to fixed climate perturbations. A pilot study was conducted using predicted streamflow and boosted regression trees to develop a set of predictive flow-ecology response models to assess the potential change in fish species richness in the North Carolina Piedmont under several scenarios of water availability change. This report is intended to provide a general assessment of all the tools and techniques available to support hydrologic modeling for flow-ecology science in the SEUS and PR. It is our hope that the approach used herein to understand differences in streamflow predictions among a subset of hydrologic models that have been applied in the SEUS for developing flow-ecology response models will provide water resource managers and stakeholders with an informed pathway for developing the capacity to link streamflow and ecological response and an understanding of some of the limitations associated with these type of modeling efforts.

Alabama, Florida, Georgia, Mississippi, North Caro

Importance of the predator's ecological neighborhood in modeling predation on migrating prey

Most mathematical descriptions of predator-prey interactions fail to take into account the spatio-temporal structures of the populations, which can lead to errors or misinterpretations. For example, a compact pulse of prey migrating through a field of quasi-stationary predators may not be well described by standard predator-prey models, because the predators and prey are unlikely to be well mixed; that is, the prey may be exposed to only a fraction of the predator population at a time. This underscores the importance of properly accounting for the ecological neighborhood, or effective feeding range, of predators in models. We illustrate this situation with a series of models of salmon smolts migrating through a reservoir arrayed with predators. The reservoir is divided into a number of longitudinal compartments or spatial cells, the length of each cell representing the upstream-downstream range over which predators can forage. In this series of models a 100-km-long reservoir is divided, successively into 2, 5, 10, 25, 50, 100, 200, and 400 cells, with respective cell lengths of 50, 20, 10, 4, 2, 1, 0.5, and 0.25 km. We used a detailed individual-based simulation model at first, but to ensure robustness of results we supplemented this with a simple analytic model. Both models showed sharp differences in the predicted mortality to a compact pulse of smolt prey moving through the reservoir, depending on the number of spatial cells in the model. In particular, models with fewer than about 10 cells vastly overpredicted the amount of mortality due to predators with activity ranges of not more than a few kilometers. These results corroborate recent theoretical and simulation studies on the importance of spatial scale and behavior in modeling predator-prey dynamics.

Oikos

SAHM:VisTrails (Software for Assisted Habitat Modeling for VisTrails): training course

VisTrails is an open-source management and scientific workflow system designed to integrate the best of both scientific workflow and scientific visualization systems. Developers can extend the functionality of the VisTrails system by creating custom modules for bundled VisTrails packages. The Invasive Species Science Branch of the U.S. Geological Survey (USGS) Fort Collins Science Center (FORT) and the U.S. Department of the Interior’s North Central Climate Science Center have teamed up to develop and implement such a module—the Software for Assisted Habitat Modeling (SAHM). SAHM expedites habitat modeling and helps maintain a record of the various input data, the steps before and after processing, and the modeling options incorporated in the construction of an ecological response model. There are four main advantages to using the SAHM:VisTrails combined package for species distribution modeling: (1) formalization and tractable recording of the entire modeling process; (2) easier collaboration through a common modeling framework; (3) a user-friendly graphical interface to manage file input, model runs, and output; and (4) extensibility to incorporate future and additional modeling routines and tools. In order to meet increased interest in the SAHM:VisTrails package, the FORT offers a training course twice a year. The course includes a combination of lecture, hands-on work, and discussion. Please join us and other ecological modelers to learn the capabilities of the SAHM:VisTrails package.

Fact Sheet

Trait contributions to fish community assembly emerge from trophicinteractions in an individual-based model

Community ecology seeks to understand and predict the characteristics of communities that can develop under different environmental conditions, but most theory has been built on analytical models that are limited in the diversity of species traits that can be considered simultaneously. We address that limitation with an individual-based model to simulate assembly of fish communities characterized by life history and trophic interactions with multiple physiological tradeoffs as constraints on species performance. Simulation experiments were carried out to evaluate the distribution of 6 life history and 4 feeding traits along gradients of resource productivity and prey accessibility. These experiments revealed that traits differ greatly in importance for species sorting along the gradients. Body growth rate emerged as a key factor distinguishing community types and defining patterns of community stability and coexistence, followed by egg size and maximum body size. Dominance by fast-growing, relatively large, and fecund species occurred more frequently in cases where functional responses were saturated (i.e. high productivity and/or prey accessibility). Such dominance was associated with large biomass fluctuations and priority effects, which prevented richness from increasing with productivity and may have limited selection on secondary traits, such as spawning strategies and relative size at maturation. Our results illustrate that the distribution of species traits and the consequences for community dynamics are intimately linked and strictly dependent on how the benefits and costs of these traits are balanced across different conditions.

Ecological Modelling

An overview of agent-based models in plant biology and ecology

Agent-based modeling (ABM) has become an established methodology in many areas of biology, ranging from the cellular to the ecological population and community levels. In plant science, two different scales have predominated in their use of ABM. One is the scale of populations and communities, through the modeling of collections of agents representing individual plants, interacting with each other and with the environment. The other is the scale of the individual plant, through the modeling, by functional-structural plant models (FSPMs), of agents representing plant building blocks, or metamers, to describe the development of plant architecture and functions within individual plants. The purpose of this review is to show key results and parallels in ABM for growth, mortality, carbon allocation, competition, and reproduction across the scales from the plant organ to populations and communities on a range of spatial scale to the whole landscape. Several areas of application of ABMs are reviewed, showing that some issues are addressed by both population-level ABMs and FSPMs. Continued increase in the relevance of ABM to environmental science and management will be helped by greater integration of ABMs across these two scales.

Annals of Botany

Techniques to improve ecological interpretability of black box machine learning models

Statistical modeling of ecological data is often faced with a large number of variables as well as possible nonlinear relationships and higher-order interaction effects. Gradient boosted trees (GBT) have been successful in addressing these issues and have shown a good predictive performance in modeling nonlinear relationships, in particular in classification settings with a categorical response variable. They also tend to be robust against outliers. However, their black-box nature makes it difficult to interpret these models. We introduce several recently developed statistical tools to the environmental research community in order to advance interpretation of these black-box models. To analyze the properties of the tools, we applied gradient boosted trees to investigate biological health of streams within the contiguous USA, as measured by a benthic macroinvertebrate biotic index. Based on these data and a simulation study, we demonstrate the advantages and limitations of partial dependence plots (PDP), individual conditional expectation (ICE) curves and accumulated local effects (ALE) in their ability to identify covariate–response relationships. Additionally, interaction effects were quantified according to interaction strength (IAS) and Friedman’s H 2 "> H 2 statistic. Interpretable machine learning techniques are useful tools to open the black-box of gradient boosted trees in the environmental sciences. This finding is supported by our case study on the effect of impervious surface on the benthic condition, which agrees with previous results in the literature. Overall, the most important variables were ecoregion, bed stability, watershed area, riparian vegetation and catchment slope. These variables were also present in most identified interaction effects. In conclusion, graphical tools (PDP, ICE, ALE) enable visualization and easier interpretation of GBT but should be supported by analytical statistical measures. Future methodological research is needed to investigate the properties of interaction tests. Supplementary materials accompanying this paper appear on-line.

Journal of Agricultural, Biological, and Environme

Using structural equation modeling to investigate relationships among ecological variables

Structural equation modeling is an advanced multivariate statistical process with which a researcher can construct theoretical concepts, test their measurement reliability, hypothesize and test a theory about their relationships, take into account measurement errors, and consider both direct and indirect effects of variables on one another. Latent variables are theoretical concepts that unite phenomena under a single term, e.g., ecosystem health, environmental condition, and pollution (Bollen, 1989). Latent variables are not measured directly but can be expressed in terms of one or more directly measurable variables called indicators. For some researchers, defining, constructing, and examining the validity of latent variables may be the end task of itself. For others, testing hypothesized relationships of latent variables may be of interest. We analyzed the correlation matrix of eleven environmental variables from the U.S. Environmental Protection Agency's (USEPA) Environmental Monitoring and Assessment Program for Estuaries (EMAP-E) using methods of structural equation modeling. We hypothesized and tested a conceptual model to characterize the interdependencies between four latent variables-sediment contamination, natural variability, biodiversity, and growth potential. In particular, we were interested in measuring the direct, indirect, and total effects of sediment contamination and natural variability on biodiversity and growth potential. The model fit the data well and accounted for 81% of the variability in biodiversity and 69% of the variability in growth potential. It revealed a positive total effect of natural variability on growth potential that otherwise would have been judged negative had we not considered indirect effects. That is, natural variability had a negative direct effect on growth potential of magnitude -0.3251 and a positive indirect effect mediated through biodiversity of magnitude 0.4509, yielding a net positive total effect of 0.1258. Natural variability had a positive direct effect on biodiversity of magnitude 0.5347 and a negative indirect effect mediated through growth potential of magnitude -0.1105 yielding a positive total effects of magnitude 0.4242. Sediment contamination had a negative direct effect on biodiversity of magnitude -0.1956 and a negative indirect effect on growth potential via biodiversity of magnitude -0.067. Biodiversity had a positive effect on growth potential of magnitude 0.8432, and growth potential had a positive effect on biodiversity of magnitude 0.3398. The correlation between biodiversity and growth potential was estimated at 0.7658 and that between sediment contamination and natural variability at -0.3769.

Environmental and Ecological Statistics

Incorporating food web dynamics into ecological restoration: A modeling approach for river ecosystems

Restoration is frequently aimed at the recovery of target species, but also influences the larger food web in which these species participate. Effects of restoration on this broader network of organisms can influence target species both directly and indirectly via changes in energy flow through food webs. To help incorporate these complexities into river restoration planning we constructed a model that links river food web dynamics to in-stream physical habitat and riparian vegetation conditions. We present an application of the model to the Methow River, Washington (USA), a location of on-going restoration aimed at recovering salmon. Three restoration strategies were simulated: riparian vegetation restoration, nutrient augmentation via salmon carcass addition, and side-channel reconnection. We also added populations of nonnative aquatic snails and fish to the modeled food web to explore how changes in food web structure mediate responses to restoration. Simulations suggest that side-channel reconnection may be a better strategy than carcass addition and vegetation planting for improving conditions for salmon in this river segment. However, modeled responses were strongly sensitive to changes in the structure of the food web. The addition of nonnative snails and fish modified pathways of energy through the food web, which negated restoration improvements. This finding illustrates that forecasting responses to restoration may require accounting for the structure of food webs, and that changes in this structure—as might be expected with the spread of invasive species—could compromise restoration outcomes. Unlike habitat-based approaches to restoration assessment that focus on the direct effects of physical habitat conditions on single species of interest, our approach dynamically links the success of target organisms to the success of competitors, predators, and prey. By elucidating the direct and indirect pathways by which restoration affects target species, dynamic food web models can improve restoration planning by fostering a deeper understanding of system connectedness and dynamics.

Washington

South Fork Shenandoah River habitat-flow modeling to determine ecological and recreational characteristics during low-flow periods

The ecological habitat requirements of aquatic organisms and recreational streamflow requirements of the South Fork Shenandoah River were investigated by the U.S. Geological Survey in cooperation with the Central Shenandoah Valley Planning District Commission, the Northern Shenandoah Valley Regional Commission, and Virginia Commonwealth University. Physical habitat simulation modeling was conducted to examine flow as a major determinant of physical habitat availability and recreation suitability using field-collected hydraulic habitat variables such as water depth, water velocity, and substrate characteristics. Fish habitat-suitability criteria specific to the South Fork Shenandoah River were developed for sub-adult and adult smallmouth bass (Micropterus dolomieu), juvenile and sub-adult redbreast sunfish (Lepomis auritus), spotfin or satinfin shiner (Cyprinella spp), margined madtom (Noturus insignis),and river chub (Nocomis micropogon). Historic streamflow statistics for the summer low-flow period during July, August, and September were used as benchmark low-flow conditions and compared to habitat simulation results and water-withdrawal scenarios based on 2005 withdrawal data. To examine habitat and recreation characteristics during droughts, daily fish habitat or recreation suitability values were simulated for 2002 and other selected drought years. Recreation suitability during droughts was extremely low, because the modeling demonstrated that suitable conditions occur when the streamflows are greater than the 50th percentile flow for July, August, and September. Habitat availability for fish is generally at a maximum when streamflows are between the 75th and 25th percentile flows for July, August, and September. Time-series results for drought years, such as 2002, showed that extreme low-flow conditions less than the 5th percentile of flow for July, August, and September corresponded to below-normal habitat availability for both game and nongame fish in the upper section of the river. For the middle section near Luray, margined madtom and river chub habitat area were below normal, whereas adult and sub-adult smallmouth bass habitat area remained near the median expected available habitat. In the lower section near Front Royal, time-series results for adult smallmouth bass, sub-adult smallmouth bass, and margined madtom habitat were below normal when streamflows were below the 10th percentile flow for July, August, and September. All other species of fish had habitat availability within the normal range for July, August, and September. Water-conservation scenarios representing a 50 percent water-withdrawal reduction resulted in game fish habitat availability within the normal range for habitat in upper and middle river sections, instead of below normal conditions which were observed during the 2002 drought. The 50 percent water-withdrawal reduction had no measurable effect on recreation. For nongame fish such as river chub, a 20 percent withdrawal reduction resulted in habitat availability within the normal range for habitat in the upper and middle river sections. Increased water-use scenarios representing a 5 percent increase in water withdrawals resulted in a slight reduction in habitat availability; however, increased withdrawals of 20 and 50 percent resulted in habitat availability substantially less than the 25th habitat percentile, or below normal. Habitat reductions were more pronounced when flows were lower than the 10th percentile flow for July, August, and September. The results show that for normal or wet years, increased water withdrawals are not likely to correspond with extensive habitat loss for game fish or nongame fish. During drought years, however, a 20 to 50 percent increase in water withdrawals may result in below normal habitat availability for game fish throughout the river and nongame fish in the upper and middle sections of the river. These simulations of rare historic drought conditions, such as those observed in 2002, serve as a baseline for development of ecological flow thresholds for drought planning.

Virginia;West Virginia

Accounting for missing ticks: Use (or lack thereof) of hierarchical models in tick ecology studies

Ixodid (hard) ticks play important ecosystem roles and have significant impacts on animal and human health via tick-borne diseases and physiological stress from parasitism. Tick occurrence, abundance, activity, and key life-history traits are highly influenced by host availability, weather, microclimate, and landscape features. As such, changes in the environment can have profound impacts on ticks, their hosts, and the spread of diseases. Researchers recognize that spatial and temporal factors influence activity and abundance and attempt to account for both by conducting replicate sampling bouts spread over the tick questing period. However, common field methods notoriously underestimate abundance, and it is unclear how (or if) tick studies model the confounding effects of factors influencing activity and abundance. This step is critical as unaccounted variance in detection can lead to biased estimates of occurrence and abundance. We performed a descriptive review to evaluate the extent to which studies account for the detection process while modeling tick data. We also categorized the types of analyses that are commonly used to model tick data. We used hierarchical models (HMs) that account for imperfect detection to analyze simulated and empirical tick data, demonstrating that inference is muddled when detection probability is not accounted for in the modeling process. Our review indicates that only 5 of 412 (1 %) papers explicitly accounted for imperfect detection while modeling ticks. By comparing HMs with the most common approaches used for modeling tick data (e.g., ANOVA), we show that population estimates are biased low for simulated and empirical data when using non-HMs, and that confounding occurs due to not explicitly modeling factors that influenced both detection and abundance. Our review and analysis of simulated and empirical data shows that it is important to account for our ability to detect ticks using field methods with imperfect detection. Not doing so leads to biased estimates of occurrence and abundance which could complicate our understanding of parasite-host relationships and the spread of tick-borne diseases. We highlight the resources available for learning HM approaches and applying them to analyzing tick data.

Maine

Coupling ecological and social network models to assess “transmission” and “contagion” of an aquatic invasive species

Network analysis is used to address diverse ecological, social, economic, and epidemiological questions, but few efforts have been made to combine these field-specific analyses into interdisciplinary approaches that effectively address how complex systems are interdependent and connected to one another. Identifying and understanding these cross-boundary connections improves natural resource management and promotes proactive, rather than reactive, decisions. This research had two main objectives; first, adapt the framework and approach of infectious disease network modeling so that it may be applied to the socio-ecological problem of spreading aquatic invasive species, and second, use this new coupled model to simulate the spread of the invasive Chinese mystery snail ( Bellamya chinensis ) in a reservoir network in Southeastern Nebraska, USA. The coupled model integrates an existing social network model of how anglers move on the landscape with new reservoir-specific ecological network models. This approach allowed us to identify 1) how angler movement among reservoirs aids in the spread of B . chinensis , 2) how B . chinensis alters energy flows within individual-reservoir food webs, and 3) a new method for assessing the spread of any number of non-native or invasive species within complex, social-ecological systems.

Journal of Environmental Management

Assessing ecological uncertainty and simulation model sensitivity to evaluate an invasive plant species’ potential impacts to the landscape

Ecological forecasts of the extent and impacts of invasive species can inform conservation management decisions. Such forecasts are hampered by ecological uncertainties associated with non-analog conditions resulting from the introduction of an invader to an ecosystem. We developed a state-and-transition simulation model tied to a fire behavior model to simulate the spread of buffelgrass ( Cenchrus ciliaris ) in Saguaro National Park, AZ, USA over a 30-year period. The simulation models forecast the potential extent and impact of a buffelgrass invasion including size and frequency of fire events and displacement of saguaro cacti and other native species. Using simulation models allowed us to evaluate how model uncertainties affected forecasted landscape outcomes. We compared scenarios covering a range of parameter uncertainties including model initialization (landscape susceptibility to invasion) and expert-identified ecological uncertainties (buffelgrass patch infill rates and precipitation). Our simulations showed substantial differences in the amount of buffelgrass on the landscape and the size and frequency of fires for dry years with slow patch infill scenarios compared to wet years with fast patch infill scenarios. We identified uncertainty in buffelgrass patch infill rates as a key area for research to improve forecasts. Our approach could be used to investigate novel processes in other invaded systems.

Scientific Reports

Spatial autoregressive models for statistical inference from ecological data

Ecological data often exhibit spatial pattern, which can be modeled as autocorrelation. Conditional autoregressive (CAR) and simultaneous autoregressive (SAR) models are network‐based models (also known as graphical models) specifically designed to model spatially autocorrelated data based on neighborhood relationships. We identify and discuss six different types of practical ecological inference using CAR and SAR models, including: (1) model selection, (2) spatial regression, (3) estimation of autocorrelation, (4) estimation of other connectivity parameters, (5) spatial prediction, and (6) spatial smoothing. We compare CAR and SAR models, showing their development and connection to partial correlations. Special cases, such as the intrinsic autoregressive model (IAR), are described. Conditional autoregressive and SAR models depend on weight matrices, whose practical development uses neighborhood definition and row‐standardization. Weight matrices can also include ecological covariates and connectivity structures, which we emphasize, but have been rarely used. Trends in harbor seals ( Phoca vitulina ) in southeastern Alaska from 463 polygons, some with missing data, are used to illustrate the six inference types. We develop a variety of weight matrices and CAR and SAR spatial regression models are fit using maximum likelihood and Bayesian methods. Profile likelihood graphs illustrate inference for covariance parameters. The same data set is used for both prediction and smoothing, and the relative merits of each are discussed. We show the nonstationary variances and correlations of a CAR model and demonstrate the effect of row‐standardization. We include several take‐home messages for CAR and SAR models, including (1) choosing between CAR and IAR models, (2) modeling ecological effects in the covariance matrix, (3) the appeal of spatial smoothing, and (4) how to handle isolated neighbors. We highlight several reasons why ecologists will want to make use of autoregressive models, both directly and in hierarchical models, and not only in explicit spatial settings, but also for more general connectivity models.

Ecological Monographs

Adaptations in a hierarchical food web of southeastern Lake Michigan

Two issues in ecological network theory are: (1) how to construct an ecological network model and (2) how do entire networks (as opposed to individual species) adapt to changing conditions? We present a novel method for constructing an ecological network model for the food web of southeastern Lake Michigan (USA) and we identify changes in key system properties that are large relative to their uncertainty as this ecological network adapts from one time point to a second time point in response to multiple perturbations. To construct our food web for southeastern Lake Michigan, we followed the list of seven recommendations outlined in Cohen et al. [Cohen, J.E., et al., 1993. Improving food webs. Ecology 74, 252–258] for improving food webs. We explored two inter-related extensions of hierarchical system theory with our food web; the first one was that subsystems react to perturbations independently in the short-term and the second one was that a system's properties change at a slower rate than its subsystems’ properties. We used Shannon's equations to provide quantitative versions of the basic food web properties: number of prey, number of predators, number of feeding links, and connectance (or density). We then compared these properties between the two time-periods by developing distributions of each property for each time period that took uncertainty about the property into account. We compared these distributions, and concluded that non-overlapping distributions indicated changes in these properties that were large relative to their uncertainty. Two subsystems were identified within our food web system structure ( p < 0.001). One subsystem had more non-overlapping distributions in food web properties between Time 1 and Time 2 than the other subsystem. The overall system had all overlapping distributions in food web properties between Time 1 and Time 2. These results supported both extensions of hierarchical systems theory. Interestingly, the subsystem with more non-overlapping distributions in food web properties was the subsystem that contained primarily benthic taxa, contrary to expectations that the identified major perturbations (lower phosphorous inputs and invasive species) would more greatly affect the subsystem containing primarily pelagic taxa. Future food-web research should employ rigorous statistical analysis and incorporate uncertainty in food web properties for a better understanding of how ecological networks adapt.

Lake Michigan