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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., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Models incorporating seasonality are necessary to fully assess the impact of global warming on Arctic communities. Seasonal migrations are a key component of Arctic food webs that still elude current theories predicting a single community equilibrium. We develop a multi-season model of predator-prey dynamics using a hybrid dynamical systems framework applied to a simplified tundra food web (lemming-fox-goose-owl). Hybrid systems models can accommodate multiple equilibria, which is a basic requirement for modelling food webs whose topology changes with season. We demonstrate that our model can generate multi-annual cycling in lemming dynamics, solely from a combined effect of seasonality and state-dependent behaviour. We compare our multi-season model to a static model of the predator-prey community dynamics and study the interactions between species. Interestingly, including seasonality reveals indirect interactions between migrants and residents not captured by the static model. Further, we find that the direction and magnitude of interactions between two species are not necessarily accurate using only summer time-series. Our study demonstrates the need for the development of multi-season models and provides the tools to analyse them. Integrating seasonality in food web modelling is a vital step to improve predictions about the impacts of climate change on ecosystem functioning. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Competing Interests: The author declares no conflict of interest.

Community interactions (e.g., predation, competition) can be characterized by two factors: their strengths and how they are structured between and within species. Both factors play a role in determining community dynamics. In addition to trophic interactions, dispersal acts as an interaction between separate populations. As with other interactions, the structure of dispersal can affect the stability of a system. However, the primary structure that has been studied in consumer-resource models has been hierarchical dispersal, where between-patch dispersal rates increase with trophic level. Here we use analytical, numerical, and simulation approaches on a two-patch, three-species metacommunity model to investigate the relationship between structure and community stability and resilience. We show that metacommunity stability is greater in systems with both weak and strong dispersal rates. Our system is stabilized by the formation of patterns when predators disperse frequently and herbivores disperse rarely, and via asynchrony when both predators and herbivores disperse infrequently. Our results show how interaction strengths within both trophic and spatial networks shape metacommunity stability.

Top-down processes such as predation and herbivory have been shown to control the dynamics of communities across a range of ecosystems by generating trophic cascades. However, theory is only beginning to describe how these local trophic processes interact with spatial subsidies in the form of material (nutrient, detritus) transport and organismal dispersal to (1) shape the structure of interconnected (meta-) ecosystems and (2) determine their optimal management via reserve networks. Here, we develop a meta-ecosystem model to understand how the reciprocal feedbacks between spatial subsidies and reserve networks modulate the importance of top-down control in a simple herbivorous fish-macroalgae-coral system. We show that in large and isolated reserve networks where connectivity between protected and unprotected areas is limited, spatial subsidies remain largely confined to reserves. This retention of spatial subsidies promotes the top-down control of corals and macroalgae by herbivores inside reserves but reduces it outside reserves. Conversely, in small and aggregated reserves where connectivity between protected and unprotected areas is high, the spillover of spatial subsidies causes a reduction in top-down control of corals and macroalgae by herbivores inside reserves and an increase in the strength of top-down control outside reserves. In addition, we demonstrate that there is a trade-off between local and regional conservation objectives when designing reserve networks: small and aggregated reserves based on the extent of dispersal maximize the abundance of corals and herbivores regionally, whereas large and isolated reserves always maximize the abundance of corals within reserves, regardless of the extent of dispersal. The existence of such "conservation traps," which arise from the fulfillment of population-level objectives within local reserves at the cost of community-level objectives at regional scales, suggests the importance of adopting a more holistic strategy to manage complex and interconnected ecosystems.

Many biological populations fluctuate in synchrony over large geographic regions. This behavior may increase the chance of extinction. The combination of time-scale separation between interacting species and weak spatial linear diffusive coupling is one mechanism that can generate synchrony; however, accounting for travel time between habitat patches may destabilize this synchrony. Here, we show that ubiquitous behavioral aspects of dispersal (e.g., predator avoidance), implemented as nonlinear diffusive coupling, may also destabilize synchrony. In addition, these aspects interact with travel-time delays and amplify mechanisms that destroy synchrony. Our work suggests that dispersal-induced synchrony is more rare than typically assumed.

We show a mechanism by which chaperone proteins can play a key role in maintaining the long-term evolutionary stability of mutation rates in prokaryotes with perfect genetic linkage. Since chaperones can reduce the phenotypic effects of mutations, higher mutation rate, by affecting chaperones, can increase the phenotypic effects of mutations. This in turn leads to greater mutation effect among the proteins that control mutation repair and DNA replication, resulting in large changes in mutation rate. The converse of this is that when mutation rate is low and chaperones are functioning well, then the rate of change in mutation rate will also be low, leading to low mutation rates being evolutionarily frozen. We show that the strength of this recursion is critical to determining the long-term evolutionary patterns of mutation rate among prokaryotes. If this recursion is weak, then mutation rates can grow without bound, leading to the extinction of the lineage. However, if this recursion is strong, then we can reproduce empirical patterns of prokaryotic mutation rates, where mutation rates remain stable over evolutionary time, and where most mutation rates are low, but with a significant fraction of high mutators., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Ecological systems show tremendous variability across temporal and spatial scales. It is this variability that ecologists try to predict and that managers attempt to harness in order to mitigate risk. However, the foundations of ecological science and its mainstream agenda focus on equilibrium dynamics to describe the balance of nature. Despite a rich body of literature on non-equilibrium ecological dynamics, we lack a well-developed set of predictions that can relate the spatiotemporal heterogeneity of natural systems to their underlying ecological processes. We argue that ecology needs to expand its current toolbox for the study of non-equilibrium ecosystems in order to both understand and manage their spatiotemporal variability. We review current approaches and outstanding questions related to the study of spatial dynamics and its application to natural ecosystems, including the design of reserves networks. We close by emphasizing the importance of ecosystem function as a key component of a non-equilibrium ecological theory, and of spatial synchrony as a central phenomenon for its inference in natural systems., (Copyright © 2014 Elsevier Inc. All rights reserved.)

The addition of spatial structure to ecological concepts and theories has spurred integration between sub-disciplines within ecology, including community and ecosystem ecology. However, the complexity of spatial models limits their implementation to idealized, regular landscapes. We present a model meta-ecosystem with finite and irregular spatial structure consisting of local nutrient-autotrophs-herbivores ecosystems connected through spatial flows of materials and organisms. We study the effect of spatial flows on stability and ecosystem functions, and provide simple metrics of connectivity that can predict these effects. Our results show that high rates of nutrient and herbivore movement can destabilize local ecosystem dynamics, leading to spatially heterogeneous equilibria or oscillations across the meta-ecosystem, with generally increased meta-ecosystem primary and secondary production. However, the onset and the spatial scale of these emergent dynamics depend heavily on the spatial structure of the meta-ecosystem and on the relative movement rate of the autotrophs. We show how this strong dependence on finite spatial structure eludes commonly used metrics of connectivity, but can be predicted by the eigenvalues and eigenvectors of the connectivity matrix that describe the spatial structure and scale. Our study indicates the need to consider finite-size ecosystems in meta-ecosystem theory.

The proliferation of efficient fishing practices has promoted the depletion of commercial stocks around the world and caused significant collateral damage to marine habitats. Recent empirical studies have shown that marine reserves can play an important role in reversing these effects. Equilibrium metapopulation models predict that networks of marine reserves can provide similar benefits so long as individual reserves are sufficiently large to achieve self-sustainability, or spaced based on the extent of dispersal of the target species in order to maintain connectivity between neighboring reserves. However, these guidelines have not been tested in nonequilibrium metacommunity models that exhibit the kinds of complex spatiotemporal dynamics typically seen in natural marine communities. Here, we used a spatially explicit predator-prey model whose predictions have been validated in a marine system to show that current guidelines are not optimal for metacommunities. In equilibrium metacommunities, there is a community-level trade-off for designing effective reserves: Networks whose size and spacing are smaller than the extent of dispersal maximize global predator abundance but minimize global prey abundance because of trophic cascades, whereas the converse is true for reserve networks whose size and spacing are larger than the extent of dispersal. In nonequilibrium metacommunities, reserves whose size and spacing match the extent of spatial autocorrelation in adult abundance (i.e., the extent of patchiness) escape this community-level trade-off by maximizing global abundance and persistence of both the prey and the predator. Overall, these results suggest that using the extent of adult patchiness instead of the extent of larval dispersal as the size and spacing of reserve networks is critical for designing community-based management strategies. By emphasizing patchiness over dispersal distance, our results show how the apparent complexity of nonequilibrium communities can actually simplify management guidelines and reduce uncertainty associated with the assessment of dispersal in marine environments.

The contribution of deterministic and stochastic processes to species coexistence is widely debated. With the introduction of powerful statistical techniques, we can now better characterise different sources of uncertainty when quantifying niche differentiation. The theoretical literature on the effect of stochasticity on coexistence, however, is often ignored by field ecologists because of its technical nature and difficulties in its application. In this review, we examine how different sources of variability in population dynamics contribute to coexistence. Unfortunately, few general rules emerge among the different models that have been studied to date. Nonetheless, we believe that a greater understanding is possible, based on the integration of coexistence and population extinction risk theories. There are two conditions for coexistence in the presence of environmental and demographic variability: (1) the average per capita growth rates of all coexisting species must be positive when at low densities, and (2) these growth rates must be strong enough to overcome negative random events potentially pushing densities to extinction. We propose that critical tests for species coexistence must account for niche differentiation arising from this variability and should be based explicitly on notions of stability and ecological drift., (© 2011 Blackwell Publishing Ltd/CNRS.)

Dispersal of organisms has large effects on the dynamics and stability of populations and communities. However, current metacommunity theory largely ignores how the flows of limiting nutrients across ecosystems can influence communities. We studied a meta-ecosystem model where two autotroph-consumer communities are spatially coupled through the diffusion of the limiting nutrient. We analyzed regional and local stability, as well as spatial and temporal synchrony to elucidate the impacts of nutrient recycling and diffusion on trophic dynamics. We show that nutrient diffusion is capable of inducing asynchronous local destabilization of biotic compartments through a diffusion-induced spatiotemporal bifurcation. Nutrient recycling interacts with nutrient diffusion and influences the susceptibility of the meta-ecosystem to diffusion-induced instabilities. This interaction between nutrient recycling and transport is further shown to depend on ecosystem enrichment. It more generally emphasizes the importance of meta-ecosystem theory for predicting species persistence and distribution in managed ecosystems., (Copyright 2010 Elsevier Ltd. All rights reserved.)

We add an ecosystem perspective to spatially structured communities subject to colonization-extinction dynamics. We derive a plant-based metaecosystem model to analyze how the spatial flows of the biotic and abiotic forms of a limiting nutrient affect persistence and coexistence. We show that the proportion of patches supporting plants in a region has a considerable impact on local nutrient dynamics. Then we explicitly couple nutrient dynamics to patch dynamics. Our model shows strong feedback between local and regional dynamics mediated by nutrient flows. We find that nutrient flows can have either positive or negative effects on species persistence and coexistence. The essential feature of this local-regional coupling is the net direction of the nutrient flows between occupied and empty patches. A net flow of nutrients from occupied to empty patches leads to indirect facilitative interactions, such as an inferior competitor promoting the persistence of a superior competitor. We show that nutrient flows affect the potential diversity of the metaecosystem and key features of plant community dynamics, such as the shape of the competition-colonization trade-off and successional sequences. Our analysis revealed that integrating ecosystem and spatial dynamics can lead to various indirect interactions that contribute significantly to community organization.

We present a theory extending the source-sink concept with an ecosystem perspective. We analyze a model for meta-ecosystem dynamics in a heterogeneous environment to study how the spatial flows of materials such as inorganic nutrients and nutrients sequestered into producers, herbivores, and detritus affect the community dynamics. We show that spatial flows of an inorganic nutrient (direct nutrient flow) and organic matter (indirect nutrient flow) through detritus, producer, or herbivore compartments can reverse the source-sink dynamics of a local ecosystem. More precisely, the balance between such direct and indirect nutrient flows determines the net direction of nutrient flows between two ecosystems of contrasted productivities. It allows a sink to turn into a source and vice versa. This effect of nutrient flows on source and sink dynamics is robust to the ecosystem structure (with and without herbivores) and to specific ecosystem compartments contributing to nutrient flows (primary producers, herbivores, or detritus). Ecosystems in distinct localities thus interact together with the structure at one place influencing that of the other. In meta-ecosystems, the source-sink dynamics of an organism is not only constrained by its dispersal from the source to the sink, but also by the fertility and community composition in the neighborhood responsible for spatial flows of nutrients and energy. The meta-ecosystem perspective provides a powerful theoretical framework to address novel questions in spatial ecosystem ecology.

Synchrony has fundamental but conflicting implications for the persistence and stability of food webs at local and regional scales. In a constant environment, compensatory dynamics between species can maintain food web stability, but factors that synchronize population fluctuations within and between communities are expected to be destabilizing. We studied the dynamics of a food web in a metacommunity to determine how environmental variability and dispersal affect stability by altering compensatory dynamics and average species abundance. When dispersal rate is high, weak correlated environmental fluctuations promote food web stability by reducing the amplitude of compensatory dynamics. However, when dispersal rate is low, weak environmental fluctuations reduce food web stability by inducing intraspecific synchrony across communities. Irrespective of dispersal rate, strong environmental fluctuations disrupt compensatory dynamics and decrease stability by inducing intermittent correlated fluctuations between consumers in local food webs, which reduce both total consumer abundance and predator abundance. Strong correlated environmental fluctuations lead to (i) spatially asynchronous and highly correlated local consumer dynamics when dispersal is low and (ii) spatially synchronous but intermediate local consumer correlation when dispersal is high. By controlling intraspecific synchrony, dispersal mediates the capacity of strong environmental fluctuations to disrupt compensatory dynamics at both local and metacommunity scales.

The relative importance of extrinsic and intrinsic causes of variability is among the oldest unresolved problems in ecology. However, the interaction between large-scale intrinsic variability in species abundance and environmental heterogeneity is still unknown. We use a metacommunity model with disturbance-recovery dynamics to resolve the interaction between scales of environmental heterogeneity, biotic processes and of intrinsic variability. We explain how population density increases with environmental variability only when its scale matches that of intrinsic patterns of abundance, through their ability to develop in heterogeneous environments. Succession dynamics reveals how the strength of local species interactions, through its control of intrinsic variability, can in turn control the scale of metapopulation response to environmental scales. Our results show that the environment and species density might fail to show any correlation despite their strong causal association. They more generally suggest that the spatial scale of ecological processes might not be sufficient to build a predictive framework for spatially heterogeneous habitats, including marine reserve networks.

Weak trophic interactions have been shown to promote the stability of ecological food webs characterized by perfect mixing. However, their importance at the landscape level and response to enrichment has not been extensively examined. In this paper we examine the food-web model explored by McCann et al. [1998. Weak trophic interactions and the balance of nature. Nature 395, 794-798]. The model is expanded into a metacommunity construct where local communities are coupled through global or local dispersal. We analyze global and local stability, as well as spatial synchrony in relation to trophic interaction strength and dispersal regimes. Results reveal that weak interactions can operate through two scale-dependent mechanisms: (i) under low local dispersal regimes, local stabilization of each community under weak interactions directly scales-up to global stability. (ii) Under high local dispersal, asynchronous local destabilization associated with weak interactions proves the driver behind global stability. In the face of enrichment, weak trophic interactions are shown to be instrumental in promoting global stability when dispersal is local. These results demonstrate how the importance of weak trophic interactions can be generalized at the landscape level despite contrary local predictions.

Species interactions and connectivity are both central to explaining the stability of ecological communities and the problem of species extinction. Yet, the role of species interactions for the stability of spatially subdivided communities still eludes ecologists. Ecological models currently address the problem of stability by exploring the role of interaction strength in well mixed habitats, or of connectivity in subdivided communities. Here I propose a unification of interaction strength and connectivity as mechanisms explaining regional community stability. I introduce a metacommunity model based on succession dynamics in coastal ecosystems, incorporating limited dispersal and facilitative interactions. I report a sharp transition in regional stability and extinction probability at intermediate interaction strength, shown to correspond to a phase transition that generates scale-invariant distribution and high regional stability. In contrast with previous studies, stability results from intermediate interaction strength only in subdivided communities, and is associated with large-scale (scale-invariant) synchrony. These results can be generalized to other systems exhibiting phase transitions to show how local interaction strength can be used to resolve the link between regional community stability and pattern formation.

I present a model based on benthic populations by first defining a competition landscape where competitive interactions are confined to small areas, defined by the distribution of resource renewal from a global pool. Using truncated kernels and results from 1-dimensional percolation theory, I then derive an approximation allowing the scaling up of local interactions among individuals to patch and landscape levels. This approximation is used to explore how spatially structured habitats can affect competition intensity. The competition landscape is then compared to a simulation of mussel colonization under various flow velocity patterns, and is shown to predict the qualitative effect of the scale of patchiness in habitat quality on total competition intensity. The interaction between large scale hydrodynamic forcing and local competition is also shown to provide an alternate hypothesis explaining the observed among-year reversal of the relationship between topographic scale and mussel abundance in some intertidal communities. The model more generally highlights the nonlinear scaling up of competitive interactions in competition landscapes, and provides a framework for the study of self-organization in heterogeneous ecosystems.

Three different lattice-based models for antagonistic ecological interactions, both nonlinear and stochastic, exhibit similar power-law scalings in the geometry of clusters. Specifically, cluster size distributions and perimeter-area curves follow power-law scalings. In the coexistence regime, these patterns are robust: their exponents, and therefore the associated Korcak exponent characterizing patchiness, depend only weakly on the parameters of the systems. These distributions, in particular the values of their exponents, are close to those reported in the literature for systems associated with self-organized criticality (SOC) such as forest-fire models; however, the typical assumptions of SOC need not apply. Our results demonstrate that power-law scalings in cluster size distributions are not restricted to systems for antagonistic interactions in which a clear separation of time-scales holds. The patterns are characteristic of processes of growth and inhibition in space, such as those in predator-prey and disturbance-recovery dynamics. Inversions of these patterns, that is, scalings with a positive slope as described for plankton distributions, would therefore require spatial forcing by environmental variability.