Metacommunity Model Research Articles

Network topology mediates freshwater fish metacommunity response to loss of connectivity

AbstractChanges in connectivity regimes affect patterns of diversity and species richness. In riverine ecosystems, factors that vary through space and time, such as flow, the presence of barriers to movement, and network topology determine connectivity, and in turn shape patterns of diversity and richness. While the effects of network topology and changes in spatiotemporal connectivity regimes on patterns of species richness have been studied in isolation, they have not been studied simultaneously. We used a discrete‐time logistic growth metacommunity model to analyze the role of spatial and temporal functional connectivity in determining patterns of local (patch level) species richness in freshwater fish metacommunities. Our modeling suggests that: (1) the effect of spatial loss of connectivity on local species richness is mediated by network topology and where richness is measured in the system and (2) increasing temporal autocorrelation in connectivity results in increasing temporal autocorrelation in patch occupancy. Spatial and temporal loss of connectivity is a ubiquitous issue for river ecosystems, making understanding and predicting its effects of fundamental importance to both improving theory and guiding river management. Our findings highlight the importance of network topology (and hence context dependency) in shaping metacommunity response to changing connectivity regimes.

Multispecies coexistence in fragmented landscapes

Spatial dynamics have long been recognized as an important driver of biodiversity. However, our understanding of species' coexistence under realistic landscape configurations has been limited by lack of adequate analytical tools. To fill this gap, we develop a spatially explicit metacommunity model of multiple competing species and derive analytical criteria for their coexistence in fragmented heterogeneous landscapes. Specifically, we propose measures of niche and fitness differences for metacommunities, which clarify how spatial dynamics and habitat configuration interact with local competition to determine coexistence of species. We parameterize our model with a Bayesian approach using a 36-y time-series dataset of three Daphnia species in a rockpool metacommunity covering >500 patches. Our results illustrate the emergence of interspecific variation in extinction and recolonization processes, including their dependencies on habitat size and environmental temperature. We find that such interspecific variation contributes to the coexistence of Daphnia species by reducing fitness differences and increasing niche differences. Additionally, our parameterized model allows separating the effects of habitat destruction and temperature change on species extinction. By integrating coexistence theory and metacommunity theory, our study provides platforms to increase our understanding of species' coexistence in fragmented heterogeneous landscapes and the response of biodiversity to environmental changes.

Sharp thresholds limit the benefit of defector avoidance in cooperation on networks

Consider a cooperation game on a spatial network of habitat patches, where players can relocate between patches if they judge the local conditions to be unfavorable. In time, the relocation events may lead to a homogeneous state where all patches harbor the same relative densities of cooperators and defectors, or they may lead to self-organized patterns, where some patches become safe havens that maintain an elevated cooperator density. Here we analyze the transition between these states mathematically. We show that safe havens form once a certain threshold in connectivity is crossed. This threshold can be analytically linked to the structure of the patch network and specifically to certain network motifs. Surprisingly, a forgiving defector avoidance strategy may be most favorable for cooperators. Our results demonstrate that the analysis of cooperation games in ecological metacommunity models is mathematically tractable and has the potential to link topics such as macroecological patterns, behavioral evolution, and network topology.

Synchrony and Stability in Trophic Metacommunities: When Top Predators Navigate in a Heterogeneous World

Ecosystem stability strongly depends on spatial aspects since localized perturbations spread across an entire region through species dispersal. Assessing the synchrony of the response of connected populations is fundamental to understand stability at different scales because if populations fluctuate asynchronously, the risk of their simultaneous extinction is low, thus reducing the species' regional extinction risk. Here, we consider a metacommunity model consisting of two food chains connected by dispersal and we review the various mechanisms governing the transmission of small perturbations affecting populations in the vicinity of equilibrium. First, we describe how perturbations propagate vertically (i.e., within food chains through trophic interactions) and horizontally (i.e., between food chains through dispersal) in metacommunities. Then, we discuss the mechanisms susceptible to alter synchrony patterns such as density-depend dispersal or spatial heterogeneity. Density-dependent dispersal, which is the influence of prey or predator abundance on dispersal, has a major impact because the species with the highest coefficient of variation of biomass governs the dispersal rate of the dispersing species and determines the synchrony of its populations, thus bypassing the classic vertical transmission of perturbations. Spatial heterogeneity, which is a disparity between patches of the attack rate of predators on prey in our model, alters the vertical transmission of perturbations in each patch, thus making synchrony dependent on which patch is perturbed. Finally, by combining our understanding of the impact of each of these mechanisms on synchrony, we are able to full explain the response of realistic metacommunities such as the model developed by Rooney et al. (2006). By disentangling the main mechanisms governing synchrony, our metacommunity model provides a broad insight into the consequences of spacial aspects on food web stability.

Comparing management strategies for conserving communities of climate-threatened species with a stochastic metacommunity model.

Many species are shifting their ranges to keep pace with climate change, but habitat fragmentation and limited dispersal could impede these range shifts. In the case of climate-vulnerable foundation species such as tropical reef corals and temperate forest trees, such limitations might put entire communities at risk of extinction. Restoring connectivity through corridors, stepping-stones or enhanced quality of existing patches could prevent the extinction of several species, but dispersal-limited species might not benefit if other species block their dispersal. Alternatively, managers might relocate vulnerable species between habitats through assisted migration, but this is generally a species-by-species approach. To evaluate the relative efficacy of these strategies, we simulated the climate-tracking of species in randomized competitive metacommunities with alternative management interventions. We found that corridors and assisted migration were the most effective strategies at reducing extinction. Assisted migration was especially effective at reducing the extinction likelihood for short-dispersing species, but it often required moving several species repeatedly. Assisted migration was more effective at reducing extinction in environments with higher stochasticity, and corridors were more effective at reducing extinction in environments with lower stochasticity. We discuss the application of these approaches to an array of systems ranging from tropical corals to temperate forests.This article is part of the theme issue ‘Ecological complexity and the biosphere: the next 30 years’.

Metacommunity dynamics and the detection of species associations in co‐occurrence analyses: Why patch disturbance matters

Abstract Many statistical methods attempt to detect species associations—and so infer interspecific interactions—from species co‐occurrence patterns. Habitat heterogeneity and out‐of‐equilibrium colonization histories are well recognized as potentially causing species associations, even when interactions are absent. The potential for patch disturbance, a classical component of metacommunity dynamics, to also drive spurious species associations has however been overlooked. Using a new general metacommunity model, we derive mathematical predictions regarding how patch disturbance would affect the patterns of species associations detected in ‘null’ co‐occurrence matrices. We also conduct numerical simulations to test our predictions and to compare the performance of several widespread statistical methods, including direct tests of pairwise independence, matrix permutation approaches and joint species distribution modelling. We show how classical metacommunity dynamics can produce statistical associations, both positive and negative, even when species do not interact, when there is no habitat heterogeneity, and at equilibrium. This occurs as soon as there is some rate of patch disturbance (i.e. simultaneous extinction of several species in a patch) and/or a finite life span of patches, a common feature of a broad range of plant, animal or microbial systems. Patch disturbance can compromise species co‐occurrence analyses and cause the artefactual detection of species associations if not taken into account. Including patch age (i.e. the time since the last patch disturbance event) as a covariate in a joint species distribution model can resolve the artefact. However, this requires additional data that often are not available in practice. We argue that the consequences of patch disturbance should not be underestimated when analysing species distribution patterns in metacommunity‐like systems. Read the free Plain Language Summary for this article on the Journal blog.

Microbiome Transmission During Sexual Intercourse Appears Stochastic and Supports the Red Queen Hypothesis.

Microbes inhabit virtually everywhere on and/or in our bodies, including the seminal and vaginal fluids. They have significant importance in maintaining reproductive health and protecting hosts from diseases. The exchange of microbes during sexual intercourse is one of the most direct and significant microbial transmissions between men and women. Nevertheless, the mechanism of this microbial transmission was little known. Is the transmission mode stochastic, passive diffusion similar to the random walk of particles, or driven by some deterministic forces? What is the microbial transmission probability? What are the possible evolutionary implications, particularly from the perspective of sexual reproduction (selection)? We tackle these intriguing questions by leveraging the power of Hubbell’s unified neutral theory of biodiversity, specifically implemented as the HDP-MSN (hierarchical Dirichlet process approximated multi-site neutral model), which allows for constructing truly multi-site metacommunity models, simultaneously including vaginal and semen microbiomes. By reanalyzing the microbiome datasets of seminal and vaginal fluids from 23 couples both before and after sexual intercourses originally reported by Mändar and colleagues, we found that the microbial transmission between seminal and vaginal fluids is a stochastic, passive diffusion similar to the random walk of particles in physics, rather than driven by deterministic forces. The transmission probability through sexual intercourse seems to be approximately 0.05. Inspired by the results from the HDP-MSN model, we further conjecture that the stochastic drifts of microbiome transmissions during sexual intercourses can be responsible for the homogeneity between semen and vaginal microbiomes first identified in a previous study, which should be helpful for sexual reproduction by facilitating the sperm movement/survival and/or egg fertilization. This inference seems to be consistent with the classic Red Queen hypothesis, which, when extended to the co-evolutionary interactions between humans and their symbiotic microbiomes, would predict that the reproductive system microbiomes should support sexual reproduction.

Evolution of prudent predation in complex food webs.

Prudent predators catch sufficient prey to sustain their populations but not as much as to undermine their populations’ survival. The idea that predators evolve to be prudent has been dismissed in the 1970s, but the arguments invoked then are untenable in the light of modern evolution theory. The evolution of prudent predation has repeatedly been demonstrated in two‐species predator–prey metacommunity models. However, the vigorous population fluctuations that these models predict are not widely observed. Here we show that in complex model food webs prudent predation evolves as a result of consumer‐mediated (‘apparent’) competitive exclusion of resources, which disadvantages aggressive consumers and does not generate such fluctuations. We make testable predictions for empirical signatures of this mechanism and its outcomes. Then we discuss how these predictions are borne out across freshwater, marine and terrestrial ecosystems. Demonstrating explanatory power of evolved prudent predation well beyond the question of predator–prey coexistence, the predicted signatures explain unexpected declines of invasive alien species, the shape of stock–recruitment relations of fish, and the clearance rates of pelagic consumers across the latitudinal gradient and 15 orders of magnitude in body mass. Specific research to further test this theory is proposed.

Seed banks can help to maintain the diversity of interacting phytoplankton species

Seed formation is part of the reproductive cycle, leading to the accumulation of resistance stages that can withstand harsh environmental conditions for long periods of time. At the community level, multiple species with such long-lasting life stages can be more likely to coexist. While the implications of this process for biodiversity have been studied in terrestrial plants, seed banks are usually neglected in phytoplankton multispecies dynamic models, in spite of widespread empirical evidence for such seed banks. In this study, we build a metacommunity model of interacting phytoplankton species, including a resting stage supplying the seed bank. The model is parameterized with empirically-driven growth rate functions and field-based interaction estimates, which include both facilitative and competitive interactions. Exchanges between compartments (coastal pelagic cells, coastal resting cells on the seabed, and open ocean pelagic cells) are controlled by hydrodynamical parameters to which the sensitivity of the model is assessed. We consider two models, i.e., with and without a saturating effect of the interactions on the growth rates. Our results are consistent between models, and show that a seed bank allows to maintain all species in the community over 30 years. Indeed, a fraction of the species are vulnerable to extinction at specific times within the year, but this process is buffered by their survival in their resting stage. We thus highlight the potential role of the seed bank in the recurrent re-invasion of the coastal community, and of coastal environments in re-seeding oceanic regions. Moreover, the seed bank enables populations to tolerate stronger interactions within the community as well as more severe changes to the environment, such as those predicted in a climate change context. Our study therefore shows how resting stages may help phytoplanktonic diversity maintenance.

The internal structure of metacommunities

Current analyses of metacommunity data largely focus on global attributes across the entire metacommunity, such as mean alpha, beta, and gamma diversity, as well as the partitioning of compositional variation into single estimates of contributions of space and environmental effects and, more recently, possible contributions of species interactions. However, this view neglects the fact that different species and sites in the landscape can vary widely in how they contribute to these metacommunity‐wide attributes. We argue for a new conceptual framework with matched analytics with the goals of studying the complex and interactive relations between process and pattern in metacommunities that is focused on the variation among species and among sites which we call the ‘internal structure' of the metacommunity. To demonstrate how the internal structure could be studied, we create synthetic data using a process‐based colonization–extinction metacommunity model. We then use joint species distribution models to estimate how the contributions of space, environment, and biotic interactions driving metacommunity assembly differ among species and sites. We find that this approach to the internal structure of metacommunities provides useful information about the distinct ways that different species and different sites contribute to metacommunity structure. Although it has limitations, our work points at a more general approach to understand how other possible complexities might affect internal structure and might thus be incorporated into a more cohesive metacommunity theory.

Macroinvertebrate species occupancy frequency distribution patterns in eutrophic lakes

Metacommunity models describe species occupancy frequency distribution (hereinafter ‘SOFD’). Our goal is to present how the differences in eight macroinvertebrate orders dispersal ability affect SOFD patterns. A total of 293 species from eight macroinvertebrate orders were observed in 14 eutrophic lakes in southern Finland. Species occupancy ranged from 1 to 14. About 30% (89 out of 293) of the species were found in only one lake, yielding a surprisingly high number of rare species. So, there were few widely distributed common species and numerous rare species with a restricted distribution. Combined data from eight macroinvertebrate orders supported the bimodal truncated SOFD pattern. Similarly, the low dispersal ability orders, watermites and mayflies, fitted the bimodal truncated SOFD pattern. However, bimodal symmetric SOFD pattern also fitted relatively well to the dragonflies and damselflies with high dispersal ability. It seems that differences in dispersal ability among different macroinvertebrate orders may partly explain observed differences. Moreover, our results supported slightly more a niche-based model rather than a metapopulation dynamics model in eutrophic lakes littoral macroinvertebrate metacommunities. Our results highlight that the dispersal ability is important trait for species conservation in patchily distributed habitat.

Top predator introduction changes the effects of spatial isolation on freshwater community structure.

Current conceptual metacommunity models predict that the consequences of local selective pressures on community structure increase with spatial isolation when species favored by local conditions also have higher dispersal rates. This appears to be the case of freshwater insects in the presence of fish. The introduction of predatory fish can produce trophic cascades in freshwater habitats because fish tend to prey upon intermediate predatory taxa, such as predatory insects, indirectly benefiting herbivores and detritivores. Similarly, spatial isolation can limit dispersal and colonization rates of predatory insects more strongly than of herbivores and detritivores, thus generating similar cascading effects. Here we tested the hypothesis that the effect of introduced predatory fish on insect community structure increases with spatial isolation by conducting a field experiment in artificial ponds that manipulated the presence/absence of fish (the redbreast tilapia) at three different distances from a source wetland. Our results showed that fish have direct negative effects on the abundance of predatory insects but probably have variable net effects on the abundance of herbivores and detritivores because the direct negative effects of predation by fish may offset indirect positive ones. Spatial isolation also resulted in indirect positive effects on the abundance of herbivores and detritivores but this effect was stronger in the absence rather than in the presence of fish so that insect communities diverged more strongly between fish and fishless ponds at higher spatial isolation. We argue that an important additional mechanism, ignored in our initial hypothesis, was that as spatial isolation increases fish predation pressure upon herbivores and detritivores increases due to the relative scarcity of predatory insects, thus dampening the positive effect that spatial isolation confers to lower trophic levels. Our results highlight the importance of considering interspecific variation in dispersal and multiple trophic levels to better understand the processes generating community and metacommunity patterns.

Structure and Dynamics of the Breast Tissue Microbiomes Under Tumor Influences: An Approach With Neutral, Near-Neutral, and Niche-Neutral Hybrid Models.

The structure and dynamics of breast tissue bacteria can have far-reaching influences on women’s health, particularly on breast tumor development. However, there is little understanding on the ecological processes that shape the structure and dynamics of breast tissue bacteria. Here, we fill the gap by applying three metacommunity models for investigating the community assembly and diversity maintenance, including Sloan near neutral model, Harris et al. multisite neutral and Tang & Zhou niche-neutral hybrid models to reanalyze the 16S-rRNA sequencing datasets of 23 healthy, 12 benign tumor, and 33 malignant tumor tissue samples. First, we found that, at the community/metacommunity levels, the mechanisms of bacteria assembly and diversity maintenance of breast tissue bacteria were moderately influenced by stochastic drifts of bacteria demography (division, death, and dispersal of bacterial cells). At species level, on average, approximately 10 and 5% species were above (positively selected) and below (negatively selected) neutral, respectively. Furthermore, malignant tumor may raise the positively selected species up to 17%. Second, malignant tumor appears to inhibit microbial dispersal as evidenced by lowered migration rates, compared with the migration in normal and benign tumor tissues. These theoretic findings can be inspirational for further investigating the relationships between tissue bacteria and breast tumor progression/development.

Identifying robust strategies for assisted migration in a competitive stochastic metacommunity.

Assisted migration (AM) is the translocation of species beyond their historical range to locations that are expected to be more suitable under future climate change. However, a relocated population may fail to establish in its donor community if there is high uncertainty in decision‐making, climate, and interactions with the recipient ecological community. To quantify the benefit to persistence and risk of establishment failure of AM under different management scenarios (e.g., choosing target species, proportion of population to relocate, and optimal location to relocate), we built a stochastic metacommunity model to simulate several species reproducing, dispersing, and competing on a temperature gradient as temperature increases over time. Without AM, the species were vulnerable to climate change when they had low population sizes, short dispersal, and strong poleward competition. When relocating species that exemplified these traits, AM increased the long‐term persistence of the species most when relocating a fraction of the donor population, even if the remaining population was very small or rapidly declining. This suggests that leaving behind a fraction of the population could be a robust approach, allowing managers to repeat AM in case they move the species to the wrong place and at the wrong time, especially when it is difficult to identify a species’ optimal climate. We found that AM most benefitted species with low dispersal ability and least benefited species with narrow thermal tolerances, for which AM increased extinction risk on average. Although relocation did not affect the persistence of nontarget species in our simple competitive model, researchers will need to consider a more complete set of community interactions to comprehensively understand invasion potential.

Incorporating abiotic controls on animal movements in metacommunities.

Local dynamics are influenced by regional processes. Meta-ecology, or the study of spatial flows of energy, materials, and species between local systems, is becoming increasingly concerned with accurate depictions of species movements and the impacts of this movement on landscape-level ecosystem function. Indeed, incorporating diverse types of movement is a major frontier in metacommunity theory. Here, we synthesize literature to demonstrate that the movement of organisms between patches is governed by the interplay between both a species' ability to move and the combined effects of landscape structure and physical flows (termed abiotic controls), which together we refer to as abiotic-dependent species connectivity. For example, two lakes that share geographic proximity may be inaccessible for mobile fish species because they lack a river connecting them (landscape structure), but wind currents may disperse insects between them (physical flows). Empirical evidence suggests that abiotic controls, such as ocean currents, lead to abiotic-dependent species connectivity and that, in nature, this type of connectivity is the rule rather than the exception. Based on this empirical evidence, we introduce a novel mathematical framework to demonstrate how species movement capabilities and abiotic conditions, can interact to influence metacommunity stability. We apply this framework to predict how incorporating abiotic-dependent species connectivity applies to classic empirical examples of aquatic, aquatic-terrestrial, and terrestrial experimental metacommunities. We demonstrate that incorporating abiotic-dependent species connectivity into metacommunity models can lead to a much broader range of dynamics than models previously predicted, including a wider range of metacommunity stability. Our framework fills critical gaps in our basic understanding of organismal movement across landscapes and provides testable predictions for how such common natural phenomena impact landscape-level ecosystem function. Finally, we present future perspectives for further development of meta-ecological theory from questions about fragmentation to ecosystems. Anthropogenic change is not only leading to habitat loss from the damming of rivers to denuding the landscape, but altering the physical flows that have historically connected communities. Thus, recognizing the importance of these processes in tandem with species' movement abilities is critical for predicting and preserving the structure and function of ecological communities.

Large-scale interventions may delay decline of the Great Barrier Reef

On the iconic Great Barrier Reef (GBR), the cumulative impacts of tropical cyclones, marine heatwaves and regular outbreaks of coral-eating crown-of-thorns starfish (CoTS) have severely depleted coral cover. Climate change will further exacerbate this situation over the coming decades unless effective interventions are implemented. Evaluating the efficacy of alternative interventions in a complex system experiencing major cumulative impacts can only be achieved through a systems modelling approach. We have evaluated combinations of interventions using a coral reef meta-community model. The model consisted of a dynamic network of 3753 reefs supporting communities of corals and CoTS connected through ocean larval dispersal, and exposed to changing regimes of tropical cyclones, flood plumes, marine heatwaves and ocean acidification. Interventions included reducing flood plume impacts, expanding control of CoTS populations, stabilizing coral rubble, managing solar radiation and introducing heat-tolerant coral strains. Without intervention, all climate scenarios resulted in precipitous declines in GBR coral cover over the next 50 years. The most effective strategies in delaying decline were combinations that protected coral from both predation (CoTS control) and thermal stress (solar radiation management) deployed at large scale. Successful implementation could expand opportunities for climate action, natural adaptation and socioeconomic adjustment by at least one to two decades.

Coupled phase-amplitude dynamics in heterogeneous metacommunities

Spatial synchrony of population fluctuations is an important tool for predicting regional stability. Its application to natural systems is still limited by the complexity of ecological time series displaying great variation in the frequency and amplitude of their fluctuations, which are not fully resolved by current ecological theories of spatial synchrony. In particular, while environmental fluctuations and limited dispersal can each control the dynamics of frequency and amplitude of population fluctuations, ecological theories of spatial synchrony still need to resolve their role on synchrony and stability in heterogeneous metacommunities. Here, we adopt a heterogeneous predator–prey metacommunity model and study the response of dispersal-driven phase locking and frequency modulation to among-patch heterogeneity in carrying capacity. We find that frequency modulation occurs at intermediate values of dispersal and habitat heterogeneity. We also show how frequency modulation can emerge in metacommunities of autonomously oscillating populations as well as through the forcing of local communities at equilibrium. Frequency modulation was further found to produce temporal variation in population amplitudes, promoting local and regional stability through cyclic patterns of local and regional variability. Our results highlight the importance of approaching spatial synchrony as a non-stationary phenomenon, with implications for the assessment and interpretation of spatial synchrony observed in experimental and natural systems.

Habitat restoration in spatially explicit metacommunity models.

In a time of rapid habitat destruction threatening the existence of many species, restoration of degraded habitats plays a crucial role in hampering biodiversity decline and recovering ecosystem services. The goal of this study is to advance the understanding of the consequences of habitat restoration on metacommunities, which is of upmost importance for designing successful restoration projects. We approach habitat restoration from a theoretical perspective by analysing spatially explicit metacommunity models which have previously been essential to understanding the effects of habitat loss and fragmentation. We investigate the efficiency of various restoration strategies on metacommunities involving interactions ranging from pairwise competition, predation and mutualism to more complex three-trophic modules. Our novel approach for measuring the restoration efficiency enables direct comparison of the responses of species in different metacommunities. We show that species recovery is affected by the amount of habitat destroyed, and the restoration strategy. When habitat is restored by randomly selecting destroyed sites, species recovery becomes less efficient and more uncertain with increasing amount of previously destroyed habitat. However, when the destroyed sites are restored in clusters, minimising the effects of fragmentation, species recovery and the certainty of success are substantially improved. Furthermore, we demonstrate that the community structure and the types of interactions involved determine the most efficient restoration approach. Our findings highlight the importance of carefully planning the restoration process, especially in landscapes where a large proportion of habitat has been destroyed, and with species at the brink of extinction. Our results may be used as guidelines for designing habitat restoration projects.

Synchrony and Perturbation Transmission in Trophic Metacommunities

AbstractIn a world where natural habitats are ever more fragmented, the dynamics of metacommunities are essential to properly understand species responses to perturbations. If species' populations fluctuate asynchronously, the risk of their simultaneous extinction is low, thus reducing the species' regional extinction risk. However, identifying synchronizing or desynchronizing mechanisms in systems containing several species and when perturbations affect multiple species is challenging. We propose a metacommunity model consisting of two food chains connected by dispersal to study the transmission of small perturbations affecting populations in the vicinity of an equilibrium. In spite of the complex responses produced by such a system, two elements enable us to understand the key processes that rule the synchrony between populations: (1) knowing which species have the strongest response to perturbations and (2) the relative importance of dispersal processes compared with local dynamics for each species. We show that perturbing a species in one patch can lead to asynchrony between patches if the perturbed species is not the most affected by dispersal. The synchrony patterns of rare species are the most sensitive to the relative strength of dispersal to demographic processes, thus making biomass distribution critical to understanding the response of trophic metacommunities to perturbations. We further partition the effect of each perturbation on species synchrony when perturbations affect multiple trophic levels. Our approach allows disentangling and predicting the responses of simple trophic metacommunities to perturbations, thus providing a theoretical foundation for future studies considering more complex spatial ecological systems.

Tracking unstable states: ecosystem dynamics in a changing world

Ecological systems can show complex and sometimes abrupt responses to environmental change, with important implications for their resilience. Theories of alternate stable states have been used to predict regime shifts of ecosystems as equilibrium responses to sufficiently slow environmental change. The actual rate of environmental change is a key factor affecting the response, yet we are still lacking a non‐equilibrium theory that explicitly considers the influence of this rate of environmental change. We present a metacommunity model of predator–prey interactions displaying multiple stable states, and we impose an explicit rate of environmental change in habitat quality (carrying capacity) and connectivity (dispersal rate). We study how regime shifts depend on the rate of environmental change and compare the outcome with a stability analysis in the corresponding constant environment. Our results reveal that in a changing environment, the community can track states that are unstable in the constant environment. This tracking can lead to regime shifts, including local extinctions, that are not predicted by alternative stable state theory. In our metacommunity, tracking unstable states also controls the maintenance of spatial heterogeneity and spatial synchrony. Tracking unstable states can also lead to regime shifts that may be reversible or irreversible. Our study extends current regime shift theories to integrate rate‐dependent responses to environmental change. It reveals the key role of unstable states for predicting transient dynamics and long‐term resilience of ecological systems to climate change.