Designing More Informative Multiple-Driver Experiments.

For decades, multiple-driver/stressor research has examined interactions among drivers that will undergo large changes in the future: temperature, pH, nutrients, oxygen, pathogens, and more. However, the most commonly used experimental designs-present-versus-future and ANOVA-fail to contribute to general understanding or predictive power. Linking experimental design to process-based mathematical models would help us predict how ecosystems will behave in novel environmental conditions. We review a range of experimental designs and assess the best experimental path toward a predictive ecology. Full factorial response surface, fractional factorial, quadratic response surface, custom, space-filling, and especially optimal and sequential/adaptive designs can help us achieve more valuable scientific goals. Experiments using these designs are challenging to perform with long-lived organisms or at the community and ecosystem levels. But they remain our most promising path toward linking experiments and theory in multiple-driver research and making accurate, useful predictions.

PERFICT: A Re-imagined foundation for predictive ecology.

Making predictions from ecological models-and comparing them to data-offers a coherent approach to evaluate model quality, regardless of model complexity or modelling paradigm. To date, our ability to use predictions for developing, validating, updating, integrating and applying models across scientific disciplines while influencing management decisions, policies, and the public has been hampered by disparate perspectives on prediction and inadequately integrated approaches. We present an updated foundation for Predictive Ecology based on seven principles applied to ecological modelling: make frequent Predictions, Evaluate models, make models Reusable, Freely accessible and Interoperable, built within Continuous workflows that are routinely Tested (PERFICT). We outline some benefits of working with these principles: accelerating science; linking with data science; and improving science-policy integration.

Energy-mediated responses to changing prey size and distribution in marine top predator movements and population dynamics.

Climate change is modifying the structure of marine ecosystems, including that of fish communities. Alterations in abiotic and biotic conditions can decrease fish size and change community spatial arrangement, ultimately impacting predator species which rely on these communities. To conserve predators and understand the drivers of observed changes in their population dynamics, we must advance our understanding of how shifting environmental conditions can impact populations by limiting food available to individuals. To investigate the impacts of changing fish size and spatial aggregation on a top predator population, we applied an existing agent-based model parameterized for harbour porpoises Phocoena phocoena which represents animal energetics and movements in high detail. We used this framework to quantify the impacts of shifting prey size and spatial aggregation on porpoise movement, space use, energetics and population dynamics. Simulated individuals were more likely to switch from area-restricted search to transit behaviour with increasing prey size, particularly when starving, due to elevated resource competition. In simulations with highly aggregated prey, higher prey encounter rates counteracted resource competition, resulting in no impacts of prey spatial aggregation on movement behaviour. Reduced energy intake with decreasing prey size and aggregation level caused population decline, with a 15% decrease in fish length resulting in total population collapse Increasing prey consumption rates by 42.8 ± 4.5% could offset population declines; however, this increase was 21.3 ± 12.7% higher than needed to account for changes in total energy availability alone. This suggests that animals in realistic seascapes require additional energy to locate smaller prey which should be considered when assessing the impacts of decreased energy availability. Changes in prey size and aggregation influenced movements and population dynamics of simulated harbour porpoises, revealing that climate-induced changes in prey structure, not only prey abundance, may threaten predator populations. We demonstrate how a population model with realistic animal movements and process-based energetics can be used to investigate population consequences of shifting food availability, such as those mediated by climate change, and provide a mechanistic explanation for how changes in prey structure can impact energetics, behaviour and ultimately viability of predator populations.

Toward a function-first framework to make soil microbial ecology predictive.

Soil microbial communities perform vital ecosystem functions, such as the decomposition of organic matter to provide plant nutrition. However, despite the functional importance of soil microorganisms, attribution of ecosystem function to particular constituents of the microbial community has been impeded by a lack of information linking microbial function to community composition and structure. Here, we propose a function-first framework to predict how microbial communities influence ecosystem functions. We first view the microbial community associated with a specific function as a whole and describe the dependence of microbial functions on environmental factors (e.g., the intrinsic temperature dependence of bacterial growth rates). This step defines the aggregate functional response curve of the community. Second, the contribution of the whole community to ecosystem function can be predicted, by combining the functional response curve with current environmental conditions. Functional response curves can then be linked with taxonomic data in order to identify sets of "biomarker" taxa that signal how microbial communities regulate ecosystem functions. Ultimately, such indicator taxa may be used as a diagnostic tool, enabling predictions of ecosystem function from community composition. In this paper, we provide three examples to illustrate the proposed framework, whereby the dependence of bacterial growth on environmental factors, including temperature, pH, and salinity, is defined as the functional response curve used to interlink soil bacterial community structure and function. Applying this framework will make it possible to predict ecosystem functions directly from microbial community composition.

Soil enzymes as indicators of soil function: A step toward greater realism in microbial ecological modeling.

Soil carbon (C) and nitrogen (N) cycles and their complex responses to environmental changes have received increasing attention. However, large uncertainties in model predictions remain, partially due to the lack of explicit representation and parameterization of microbial processes. One great challenge is to effectively integrate rich microbial functional traits into ecosystem modeling for better predictions. Here, using soil enzymes as indicators of soil function, we developed a competitive dynamic enzyme allocation scheme and detailed enzyme-mediated soil inorganic N processes in the Microbial-ENzyme Decomposition (MEND) model. We conducted a rigorous calibration and validation of MEND with diverse soil C-N fluxes, microbial C:N ratios, and functional gene abundances from a 12-year CO2  × N grassland experiment (BioCON) in Minnesota, USA. In addition to accurately simulating soil CO2 fluxes and multiple N variables, the model correctly predicted microbial C:N ratios and their negative response to enriched N supply. Model validation further showed that, compared to the changes in simulated enzyme concentrations and decomposition rates, the changes in simulated activities of eight C-N-associated enzymes were better explained by the measured gene abundances in responses to elevated atmospheric CO2 concentration. Our results demonstrated that using enzymes as indicators of soil function and validating model predictions with functional gene abundances in ecosystem modeling can provide a basis for testing hypotheses about microbially mediated biogeochemical processes in response to environmental changes. Further development and applications of the modeling framework presented here will enable microbial ecologists to address ecosystem-level questions beyond empirical observations, toward more predictive understanding, an ultimate goal of microbial ecology.

Predicting insect outbreaks using machine learning: A mountain pine beetle case study.

Planning forest management relies on predicting insect outbreaks such as mountain pine beetle, particularly in the intermediate-term future, e.g., 5-year. Machine-learning algorithms are potential solutions to this challenging problem due to their many successes across a variety of prediction tasks. However, there are many subtle challenges in applying them: identifying the best learning models and the best subset of available covariates (including time lags) and properly evaluating the models to avoid misleading performance-measures. We systematically address these issues in predicting the chance of a mountain pine beetle outbreak in the Cypress Hills area and seek models with the best performance at predicting future 1-, 3-, 5- and 7-year infestations. We train nine machine-learning models, including two generalized boosted regression trees (GBM) that predict future 1- and 3-year infestations with 92% and 88% AUC, and two novel mixed models that predict future 5- and 7-year infestations with 86% and 84% AUC, respectively. We also consider forming the train and test datasets by splitting the original dataset randomly rather than using the appropriate year-based approach and show that this may obtain models that score high on the test dataset but low in practice, resulting in inaccurate performance evaluations. For example, a k-nearest neighbor model with the actual performance of 68% AUC, scores the misleadingly high 78% on a test dataset obtained from a random split, but the more accurate 66% on a year-based split. We then investigate how the prediction accuracy varies with respect to the provided history length of the covariates and find that neural network and naive Bayes, predict more accurately as history-length increases, particularly for future 1- and 3-year predictions, and roughly the same holds with GBM. Our approach is applicable to other invasive species. The resulting predictors can be used in planning forest and pest management and planning sampling locations in field studies.

From theory to practice in pattern-oriented modelling: identifying and using empirical patterns in predictive models.

To robustly predict the effects of disturbance and ecosystem changes on species, it is necessary to produce structurally realistic models with high predictive power and flexibility. To ensure that these models reflect the natural conditions necessary for reliable prediction, models must be informed and tested using relevant empirical observations. Pattern-oriented modelling (POM) offers a systematic framework for employing empirical patterns throughout the modelling process and has been coupled with complex systems modelling, such as in agent-based models (ABMs). However, while the production of ABMs has been rising rapidly, the explicit use of POM has not increased. Challenges with identifying patterns and an absence of specific guidelines on how to implement empirical observations may limit the accessibility of POM and lead to the production of models which lack a systematic consideration of reality. This review serves to provide guidance on how to identify and apply patterns following a POM approach in ABMs (POM-ABMs), specifically addressing: where in the ecological hierarchy can we find patterns; what kinds of patterns are useful; how should simulations and observations be compared; and when in the modelling cycle are patterns used? The guidance and examples provided herein are intended to encourage the application of POM and inspire efficient identification and implementation of patterns for both new and experienced modellers alike. Additionally, by generalising patterns found especially useful for POM-ABM development, these guidelines provide practical help for the identification of data gaps and guide the collection of observations useful for the development and verification of predictive models. Improving the accessibility and explicitness of POM could facilitate the production of robust and structurally realistic models in the ecological community, contributing to the advancement of predictive ecology at large.

Range-wide variations in common milkweed traits and their effect on monarch larvae.

PREMISE: Leaf economic spectrum (LES) theory has historically been employed to inform vegetation models of ecosystem processes, but largely neglects intraspecific variation and biotic interactions. We attempt to integrate across environment-plant trait-herbivore interactions within a species at a range-wide scale. METHODS: We measured traits in 53 populations spanning the range of common milkweed (Asclepias syriaca) and used a common garden to determine the role of environment in driving patterns of intraspecific variation. We used a feeding trial to determine the role of plant traits in monarch (Danaus plexippus) larval development. RESULTS: Trait-trait relationships largely followed interspecific patterns in LES theory and persisted in a common garden when individual traits change. Common milkweed showed intraspecific variation and biogeographic clines in traits. Clines did not persist in a common garden. Larvae ate more and grew larger when fed plants with more nitrogen. A longitudinal environmental gradient in precipitation corresponded to a resource gradient in plant nitrogen, which produces a gradient in larval performance. CONCLUSIONS: Biogeographic patterns in common milkweed traits can sometimes be predicted from LES, are largely driven by environmental conditions, and have consequences for monarch larval performance. Changes to nutrient dynamics of landscapes with common milkweed could potentially influence monarch population dynamics. We show how biogeographic trends in intraspecific variation can influence key ecological interactions, especially in common species with large distributions.

Two decades of functional-structural plant modelling: now addressing fundamental questions in systems biology and predictive ecology.

BACKGROUND: Functional-structural plant models (FSPMs) explore and integrate relationships between a plant's structure and processes that underlie its growth and development. In the last 20 years, scientists interested in functional-structural plant modelling have expanded greatly the range of topics covered and now handle dynamical models of growth and development occurring from the microscopic scale, and involving cell division in plant meristems, to the macroscopic scales of whole plants and plant communities. SCOPE: The FSPM approach occupies a central position in plant science; it is at the crossroads of fundamental questions in systems biology and predictive ecology. This special issue of Annals of Botany features selected papers on critical areas covered by FSPMs and examples of comprehensive models that are used to solve theoretical and applied questions, ranging from developmental biology to plant phenotyping and management of plants for agronomic purposes. Altogether, they offer an opportunity to assess the progress, gaps and bottlenecks along the research path originally foreseen for FSPMs two decades ago. This review also allows discussion of current challenges of FSPMs regarding (1) integration of multidisciplinary knowledge, (2) methods for handling complex models, (3) standards to achieve interoperability and greater genericity and (4) understanding of plant functioning across scales. CONCLUSIONS: This approach has demonstrated considerable progress, but has yet to reach its full potential in terms of integration and heuristic knowledge production. The research agenda of functional-structural plant modellers in the coming years should place a greater emphasis on explaining robust emergent patterns, and on the causes of possible deviation from it. Modelling such patterns could indeed fuel both generic integration across scales and transdisciplinary transfer. In particular, it could be beneficial to emergent fields of research such as model-assisted phenotyping and predictive ecology in managed ecosystems.

Predicting ecosystem state changes in shallow lakes using an aquatic ecosystem model: Lake Hinge, Denmark, an example.

In recent years, considerable efforts have been made to restore turbid, phytoplankton-dominated shallow lakes to a clear-water state with high coverage of submerged macrophytes. Various dynamic lake models with simplified physical representations of vertical gradients, such as PCLake, have been used to predict external nutrient load thresholds for such nonlinear regime shifts. However, recent observational studies have questioned the concept of regime shifts by emphasizing that gradual changes are more common than sudden shifts. We investigated if regime shifts would be more gradual if the models account for depth-dependent heterogeneity of the system by including the possibility of vertical gradients in the water column and sediment layers for the entire depth. Hence, bifurcation analysis was undertaken using the 1D hydrodynamic model GOTM, accounting for vertical gradients, coupled to the aquatic ecosystem model PCLake, which is implemented in the framework for aquatic biogeochemical modeling (FABM). First, the model was calibrated and validated against a comprehensive data set covering two consecutive 7-yr periods from Lake Hinge, a shallow, eutrophic Danish lake. The autocalibration program Auto-Calibration Python (ACPy) was applied to achieve a more comprehensive adjustment of model parameters. The model simulations showed excellent agreement with observed data for water temperature, total nitrogen, and nitrate and good agreement for ammonium, total phosphorus, phosphate, and chlorophyll a concentrations. Zooplankton and macrophyte coverage were adequately simulated for the purpose of this study, and in general the GOTM-FABM-PCLake model simulations performed well compared with other model studies. In contrast to previous model studies ignoring depth heterogeneity, our bifurcation analysis revealed that the spatial extent and depth limitation of macrophytes as well as phytoplankton chlorophyll-a responded more gradually over time to a reduction in the external phosphorus load, albeit some hysteresis effects still appeared. In a management perspective, our study emphasizes the need to include depth heterogeneity in the model structure to more correctly determine at which external nutrient load a given lake changes ecosystem state to a clear-water condition.

Foreseeing the future of mutualistic communities beyond collapse.

Changing conditions may lead to sudden shifts in the state of ecosystems when critical thresholds are passed. Some well-studied drivers of such transitions lead to predictable outcomes such as a turbid lake or a degraded landscape. Many ecosystems are, however, complex systems of many interacting species. While detecting upcoming transitions in such systems is challenging, predicting what comes after a critical transition is terra incognita altogether. The problem is that complex ecosystems may shift to many different, alternative states. Whether an impending transition has minor, positive or catastrophic effects is thus unclear. Some systems may, however, behave more predictably than others. The dynamics of mutualistic communities can be expected to be relatively simple, because delayed negative feedbacks leading to oscillatory or other complex dynamics are weak. Here, we address the question of whether this relative simplicity allows us to foresee a community's future state. As a case study, we use a model of a bipartite mutualistic network and show that a network's post-transition state is indicated by the way in which a system recovers from minor disturbances. Similar results obtained with a unipartite model of facilitation suggest that our results are of relevance to a wide range of mutualistic systems.

Linking Microbial Community Structure to Trait Distributions and Functions Using Salinity as an Environmental Filter.

The structure and function of microbial communities vary along environmental gradients; however, interlinking the two has been challenging. In this study, salinity was used as an environmental filter to study how it could shape trait distributions, community structures, and the resulting functions of soil microbes. The environmental filter was applied by salinizing nonsaline soil (0 to 22 mg NaCl g-1). Our targeted community trait distribution (salt tolerance) was determined with dose-response relationships between bacterial growth and salinity. The bacterial community structure responses were resolved with Illumina 16S rRNA gene amplicon sequencing, and the microbial functions determined were respiration and bacterial and fungal growth. Salt exposure quickly resulted in filtered trait distributions, and stronger filters resulted in larger shifts. The filtered trait distributions correlated well with community composition differences, suggesting that trait distribution shifts were driven at least partly by species turnover. While salt exposure decreased respiration, microbial growth responses appeared to be characterized by competitive interactions. Fungal growth was highest when bacterial growth was inhibited by the highest salinity, and it was lowest when the bacterial growth rate peaked at intermediate salt levels. These findings corroborated a higher potential for fungal salt tolerance than bacterial salt tolerance for communities derived from a nonsaline soil. In conclusion, by using salt as an environmental filter, we could interlink the targeted trait distribution with both the community structure and resulting functions of soil microbes.IMPORTANCE Understanding the role of ecological communities in maintaining multiple ecosystem processes is a central challenge in ecology. Soil microbial communities perform vital ecosystem functions, such as the decomposition of organic matter to provide plant nutrition. However, despite the functional importance of soil microorganisms, attribution of ecosystem function to particular constituents of the microbial community has been impeded by a lack of information linking microbial processes to community composition and structure. Here, we apply a conceptual framework to determine how microbial communities influence ecosystem processes, by applying a "top-down" trait-based approach. By determining the dependence of microbial processes on environmental factors (e.g., the tolerance to salinity), we can define the aggregate response trait distribution of the community, which then can be linked to the community structure and the resulting function performed by the microbial community.

Transferability of biotic interactions: Temporal consistency of arctic plant-rodent relationships is poor.

Variability in biotic interaction strength is an integral part of food web functioning. However, the consequences of the spatial and temporal variability of biotic interactions are poorly known, in particular for predicting species abundance and distribution. The amplitude of rodent population cycles (i.e., peak-phase abundances) has been hypothesized to be determined by vegetation properties in tundra ecosystems. We assessed the spatial and temporal predictability of food and shelter plants effects on peak-phase small rodent abundance during two consecutive rodent population peaks. Rodent abundance was related to both food and shelter biomass during the first peak, and spatial transferability was mostly good. Yet, the temporal transferability of our models to the next population peak was poorer. Plant-rodent interactions are thus temporally variable and likely more complex than simple one-directional (bottom-up) relationships or variably overruled by other biotic interactions and abiotic factors. We propose that parametrizing a more complete set of functional links within food webs across abiotic and biotic contexts would improve transferability of biotic interaction models. Such attempts are currently constrained by the lack of data with replicated estimates of key players in food webs. Enhanced collaboration between researchers whose main research interests lay in different parts of the food web could ameliorate this.

Indicators of transitions in biological systems.

In the face of global biodiversity declines, predicting the fate of biological systems is a key goal in ecology. One popular approach is the search for early warning signals (EWSs) based on alternative stable states theory. In this review, we cover the theory behind nonlinearity in dynamic systems and techniques to detect the loss of resilience that can indicate state transitions. We describe the research done on generic abundance-based signals of instability that are derived from the phenomenon of critical slowing down, which represent the genesis of EWSs research. We highlight some of the issues facing the detection of such signals in biological systems - which are inherently complex and show low signal-to-noise ratios. We then document research on alternative signals of instability, including measuring shifts in spatial autocorrelation and trait dynamics, and discuss potential future directions for EWSs research based on detailed demographic and phenotypic data. We set EWSs research in the greater field of predictive ecology and weigh up the costs and benefits of simplicity vs. complexity in predictive models, and how the available data should steer the development of future methods. Finally, we identify some key unanswered questions that, if solved, could improve the applicability of these methods.

Predicting biotic interactions and their variability in a changing environment.

Global environmental change is altering the patterns of biodiversity worldwide. Observation and theory suggest that species' distributions and abundances depend on a suite of processes, notably abiotic filtering and biotic interactions, both of which are constrained by species' phylogenetic history. Models predicting species distribution have historically mostly considered abiotic filtering and are only starting to integrate biotic interaction. However, using information on present interactions to forecast the future of biodiversity supposes that biotic interactions will not change when species are confronted with new environments. Using bacterial microcosms, we illustrate how biotic interactions can vary along an environmental gradient and how this variability can depend on the phylogenetic distance between interacting species.

The emergence and promise of functional biogeography.

Understanding, modeling, and predicting the impact of global change on ecosystem functioning across biogeographical gradients can benefit from enhanced capacity to represent biota as a continuous distribution of traits. However, this is a challenge for the field of biogeography historically grounded on the species concept. Here we focus on the newly emergent field of functional biogeography: the study of the geographic distribution of trait diversity across organizational levels. We show how functional biogeography bridges species-based biogeography and earth science to provide ideas and tools to help explain gradients in multifaceted diversity (including species, functional, and phylogenetic diversities), predict ecosystem functioning and services worldwide, and infuse regional and global conservation programs with a functional basis. Although much recent progress has been made possible because of the rising of multiple data streams, new developments in ecoinformatics, and new methodological advances, future directions should provide a theoretical and comprehensive framework for the scaling of biotic interactions across trophic levels and its ecological implications.

Simple relationships to predict attributes of fish assemblages in patches of submerged macrophytes

Submerged macrophytes play an important role in structuring habitats and, therefore, in determining patterns of aquatic biodiversity. Because these plants are widespread in shallow areas of many Neotropical reservoirs, the present work investigated if variables related to habitat structure, measured in patches of submerged macrophytes (Egeria densa and E. najas), can be used to predict fish assemblage attributes (fish density and species richness). Based on patch characteristics at fine spatial extents (macrophyte patches within reservoir arms), we considered plant biomass, volume and proportional volume (i.e. percentage of macrophyte volume in the water column) as potential predictors. Fish and macrophytes were sampled with a 1-m² throw trap in littoral habitats of Rosana Reservoir, Paranapanema River, and simple correlation analyses were performed. Fish richness and abundance were highly correlated with all variables (R = 0.53 to 0.90), a relationship consistently observed in all sites. When compared to biomass, plant volume and proportional volume did not yield stronger correlations. We observed stronger correlations when E. densa and E. najas patches were analyzed separately (mono-specificity), probably because particular effects of each macrophyte on habitat structuring were removed (e.g. unnoticed morphological differences or unknown effects on habitat quality). The high R values observed in all pairwise relationships are uncommon in ecological studies, highlighting the predictive potential of variables related to habitat structure. These results suggest that, at small spatial extents, macrophyte biomass may represent an interesting predictor of fish density and richness in reservoirs with extensive colonization of submerged plants.

As macrófitas aquáticas submersas desempenham um importante papel na estruturação de hábitats e, por isso, determinam padrões gerais de biodiversidade. Como essas plantas colonizam muitos reservatórios neotropicais, o presente trabalho investigou simples relações capazes de predizer atributos de assembléias de peixes (densidade e riqueza) a partir de variáveis relacionadas à estruturação do habitat, medidas em manchas de macrófitas submersas (Egeria densa e E. najas). Baseando-se em características das manchas em pequenas escalas espaciais (bancos de macrófitas em braços do reservatório), nós consideramos a biomassa de plantas, volume e volume proporcional (percentual de volume de macrófitas na coluna d'água) como preditores em potencial. Peixes e macrófitas foram amostrados com uma armadilha de arremesso (1-m²) em hábitats litorâneos do reservatório de Rosana, rio Paranapanema, e correlação simples foram utilizadas para analisar as relações. Todas as variáveis apresentaram correlação positiva com a riqueza e densidade de peixes (R = 0.53 to 0.90), uma relação consistentemente observada nos três locais amostrados. Com relação à performance individual de cada variável, o volume e o volume proporcional não aumentaram a magnitude das correlações quando comparados à biomassa de macrófitas. Correlações mais fortes foram observadas quando as manchas de E. densa e E. najas foram analisadas separadamente (mono-especificidade), provavelmente porque efeitos particulares de cada espécie na estruturação dos hábitats foram removidos (e.g. pequenas diferenças morfológicas ou efeitos na qualidade da água). Os altos valores de correlação (R) observados em todas as relações são incomuns em estudos ecológicos, e enfatizam o potencial preditivo de variáveis relacionadas à estrutura espacial dos hábitats. Esses resultados sugerem que, em pequenas escalas espaciais, a biomassa de macrófitas pode representar um interessante preditor da densidade e riqueza...