Early warning indicators capture catastrophic transitions driven by explicit rates of environmental change.

In response to external changes, ecosystems can undergo catastrophic transitions. Early warning indicators aim to predict such transitions based on the phenomenon of critical slowing down at bifurcation points found under a constant environment. When an explicit rate of environmental change is considered, catastrophic transitions can become distinct phenomena from bifurcations, and result from a delayed response to noncatastrophic bifurcations. We use a trophic metacommunity model where transitions in time series and bifurcations of the system are distinct phenomena. We calculate early warning indicators from the time series of the continually changing system and show that they predict not the bifurcation of the underlying system but the actual catastrophic transition driven by the explicit rate of change. Predictions based on the bifurcation structure could miss catastrophic transitions that can still be captured by early warning signals calculated from time series. Our results expand the repertoire of mechanistic models used to anticipate catastrophic transitions to nonequilibrium ecological systems exposed to a constant rate of environmental change.

The significance of partial migration for food web and ecosystem dynamics.

Migration is ubiquitous and can strongly shape food webs and ecosystems. Less familiar, however, is that the majority of life cycle, seasonal and diel migrations in nature are partial migrations: only a fraction of the population migrates while the other individuals remain in their resident ecosystem. Here, we demonstrate different impacts of partial migration rendering it fundamental to our understanding of the significance of migration for food web and ecosystem dynamics. First, partial migration affects the spatiotemporal distribution of individuals and the food web and ecosystem-level processes they drive differently than expected under full migration. Second, whether an individual migrates or not is regularly correlated with morphological, physiological, and/or behavioural traits that shape its food-web and ecosystem-level impacts. Third, food web and ecosystem dynamics can drive the fraction of the population migrating, enabling the potential for feedbacks between the causes and consequences of migration within and across ecosystems. These impacts, individually and in combination, can yield unintuitive effects of migration and drive the dynamics, diversity and functions of ecosystems. By presenting the first full integration of partial migration and trophic (meta-)community and (meta-)ecosystem ecology, we provide a roadmap for studying how migration affects and is affected by ecosystem dynamics in a changing world.

Traits affecting nutrient recycling by mobile consumers can explain coexistence and spatially heterogeneous trophic regulation across a meta-ecosystem.

Ecosystems are linked through spatial flows of organisms and nutrients that impact their biodiversity and regulation. Theory has predominantly studied passive nutrient flows that occur independently of organism movement. Mobile organisms, however, commonly drive nutrient flows across ecosystems through nutrient recycling. Using a meta-ecosystem model where consumers move between ecosystems, we study how consumer recycling and traits related to feeding and sheltering preferences affect species diversity and trophic regulation. We show local effects of recycling can cascade across space, yielding spatially heterogeneous top-down and bottom-up effects. Consumer traits impact the direction and magnitude of these effects by enabling recycling to favour a single ecosystem. Recycling further modifies outcomes of competition between consumer species by creating a positive feedback on the production of one competitor. Our findings suggest spatial interactions between feeding and recycling activities of organisms are key to predicting biodiversity and ecosystem functioning across spatial scales.

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.

Seasonal food webs with migrations: multi-season models reveal indirect species interactions in the Canadian Arctic tundra.

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'.

Converting Ecological Currencies: Energy, Material, and Information Flows.

Understanding how the three currencies of life - energy, material, and information - interact is a key step towards synthesis in ecology and evolution. However, current theory focuses on the role of matter as a resource and energy, and typically ignores how the same matter can have other important effects as a carrier of information or modifier of the environment. Here we present the hypothesis that the dynamic conversion of matter by organisms among its three currencies mediates the structure and function of ecosystems, and that these effects can even supersede the effects of matter as a resource. Humans are changing the information in the environment and this is altering species interactions and flows of matter within and among ecosystems.

Author Correction: Effect of diversity on growth, mortality, and loss of resilience to extreme climate events in a tropical planted forest experiment.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Understanding Maladaptation by Uniting Ecological and Evolutionary Perspectives.

Evolutionary biologists have long trained their sights on adaptation, focusing on the power of natural selection to produce relative fitness advantages while often ignoring changes in absolute fitness. Ecologists generally have taken a different tack, focusing on changes in abundance and ranges that reflect absolute fitness while often ignoring relative fitness. Uniting these perspectives, we articulate various causes of relative and absolute maladaptation and review numerous examples of their occurrence. This review indicates that maladaptation is reasonably common from both perspectives, yet often in contrasting ways. That is, maladaptation can appear strong from a relative fitness perspective, yet populations can be growing in abundance. Conversely, resident individuals can appear locally adapted (relative to nonresident individuals) yet be declining in abundance. Understanding and interpreting these disconnects between relative and absolute maladaptation, as well as the cases of agreement, is increasingly critical in the face of accelerating human-mediated environmental change. We therefore present a framework for studying maladaptation, focusing in particular on the relationship between absolute and relative fitness, thereby drawing together evolutionary and ecological perspectives. The unification of these ecological and evolutionary perspectives has the potential to bring together previously disjunct research areas while addressing key conceptual issues and specific practical problems.

Causes of maladaptation.

Evolutionary biologists tend to approach the study of the natural world within a framework of adaptation, inspired perhaps by the power of natural selection to produce fitness advantages that drive population persistence and biological diversity. In contrast, evolution has rarely been studied through the lens of adaptation's complement, maladaptation. This contrast is surprising because maladaptation is a prevalent feature of evolution: population trait values are rarely distributed optimally; local populations often have lower fitness than imported ones; populations decline; and local and global extinctions are common. Yet we lack a general framework for understanding maladaptation; for instance in terms of distribution, severity, and dynamics. Similar uncertainties apply to the causes of maladaptation. We suggest that incorporating maladaptation-based perspectives into evolutionary biology would facilitate better understanding of the natural world. Approaches within a maladaptation framework might be especially profitable in applied evolution contexts - where reductions in fitness are common. Toward advancing a more balanced study of evolution, here we present a conceptual framework describing causes of maladaptation. As the introductory article for a Special Feature on maladaptation, we also summarize the studies in this Issue, highlighting the causes of maladaptation in each study. We hope that our framework and the papers in this Special Issue will help catalyze the study of maladaptation in applied evolution, supporting greater understanding of evolutionary dynamics in our rapidly changing world.

Phenotype-environment mismatch in metapopulations-Implications for the maintenance of maladaptation at the regional scale.

Maladaptation is widespread in natural populations. However, maladaptation has most often been associated with absolute population decline in local habitats rather than on a spectrum of relative fitness variation that can assist natural populations in their persistence at larger regional scales. We report results from a field experiment that tested for relative maladaptation between-pond habitats with spatial heterogeneity and (a)symmetric selection in pH. In the experiment, we quantified relative maladaptation in a copepod metapopulation as a mismatch between the mean population phenotype and the optimal trait value that would maximize mean population fitness under either stable or fluctuating pH environmental conditions. To complement the field experiment, we constructed a metapopulation model that addressed both relative (distance from the optimum) and absolute (negative population growth) maladaptation, with the aim of forecasting maladaptation to pH at the regional scale in relation to spatial structure (environmental heterogeneity and connectivity) and temporal environmental fluctuations. The results from our experiment indicated that maladaptation to pH at the regional scale depended on the asymmetry of the fitness surface at the local level. The results from our metapopulation model revealed how dispersal and (a)symmetric selection can operate on the fitness surface to maintain maladaptive phenotype-environment mismatch at local and regional scales in a metapopulation. Environmental stochasticity resulted in the maintenance of maladaptation that was robust to dispersal, but also revealed an interaction between the asymmetry in selection and environmental correlation. Our findings emphasize the importance of maladaptation for planning conservation strategies that can support adaptive potential in fragmented and changing landscapes.

Meta-ecosystem processes alter ecosystem function and can promote herbivore-mediated coexistence.

Herbivory and dispersal play roles in the coexistence of primary producers with shared resource limitation by imposing trade-offs either through apparent competition or dispersal limitation. These mechanisms of coexistence can further interact with meta-ecosystem effects, which results in spatial heterogeneity through the movement of herbivores and nutrients. Here, we investigate how herbivores influence autotroph coexistence through a meta-ecosystem effect, and how this effect couples mechanisms of coexistence to ecosystem structure and functioning. We articulate this framework through a parameterized one resource-k producer-one herbivore meta-ecosystem model. The results show that herbivore movement with nutrient recycling can generate spatial heterogeneity to allow coexistence where the well-mixed system predicts competitive exclusion. Furthermore, the presence of movement alters local and regional ecosystem functioning even when coexistence would occur without movement. These results highlight how meta-ecosystem theory can provide a mechanistic context for the observed complexity of biodiversity-ecosystem function relationships.

Ocean acidification causes mortality in the medusa stage of the cubozoan Carybdea xaymacana.

Ocean pH is decreasing due to anthropogenic activities, and the consequences of this acidification on marine fauna and ecosystems are the subject of an increasing number of studies. Yet, the impact of ocean acidification (OA) on several abundant and ecologically important taxa, such as medusozoans, is poorly documented. To date there have been no studies on the effect of post-2050 OA projections on the medusa stage of jellyfish. As medusae represent the reproductive stage of cnidarians, negative impacts on adult jellyfish could severely impact the long-term survival of this group. Using a laboratory experiment, we investigated the effect of 2300 OA projections (i.e. pH of 7.5) on the mortality rate of the medusa-stage of the cubozoan species Carybdea xaymacana, compared to ambient seawater pH conditions (i.e. pH of 8.1). After a 12-h exposure to OA, C. xaymacana medusae suffered higher mortality rates compared to ambient conditions. This study represents the first evidence of the potential lethal effects of post-2050 OA projections on jellyfish. The higher metabolic rates of cubozoans compared to other cnidarians might make box jellyfish more vulnerable to OA. A decrease in the density of cnidarians could lead to harmful ecological events, such as algal blooms.

Effect of diversity on growth, mortality, and loss of resilience to extreme climate events in a tropical planted forest experiment.

A pressing question is whether biodiversity can buffer ecosystem functioning against extreme climate events. However, biodiversity loss is expected to occur due to climate change with severe impacts to tropical forests. Using data from a ca. 15 year-old tropical planted forest, we construct models based on a bootstrapping procedure to measure growth and mortality among different species richness treatments in response to extreme climate events. In contrast to higher richness mixtures, in one-species plots we find growth is strongly regulated by climate events and we also find increasingly higher mortality during a consecutive four year dry event. Based on these results together with indicators of loss of resilience, we infer an effect of diversity on critical slowing down. Our work generates new methods, concepts, and applications for global change ecology and emphasises the need for research in the area of biodiversity-ecosystem functioning along environmental stress gradients.

The lateralization and distalization shoulder angles are important determinants of clinical outcomes in reverse shoulder arthroplasty.

BACKGROUND: Reverse shoulder arthroplasty (RSA) designs vary in the lateralization and distalization geometry, which may affect functional outcomes. The purpose was to determine the effect of RSA lateralization and distalization on final functional outcomes by using the "lateralization shoulder angle" (LSA) and the "distalization shoulder angle" (DSA). METHODS: Forty-six consecutive patients who underwent RSA for cuff tear arthropathy were retrospectively evaluated. Functional outcome and radiographs were evaluated at a minimum of 2 years postoperatively and compared between implants with or without glenoid lateralization and with or without humeral-sided lateralization. Anteroposterior shoulder radiographs were used to evaluate the LSA and DSA. RESULTS: Both angles showed substantial to almost perfect intrarater and inter-rater agreement. Higher LSA values were found in more lateralized RSAs (P = .027), and values between 75° and 95° were correlated with better active external rotation (quadratic regression analysis R2 = 0.553, P < .001). Postoperative active anterior elevation (R2 = 0.2, P = .008), Constant (rs = 0.29, P = .05), and Activities of Daily Living Requiring External Rotation scores (rs = 0.4 P = .007) had a positive correlation with the LSA. The quadratic regression analysis also showed that a DSA between 40° and 65° resulted in better active anterior elevation (R2 = 0.4, P < .001) and abduction (R2 = 0.4, P < .001). The negative correlation between the LSA and DSA (rs = -0.7, P < .001) revealed that, according to the implant used, the more distally the RSA is placed the less lateralization is achieved. CONCLUSIONS: The LSA and the DSA are reproducible measurements that may be used to estimate "lateralization and distalization" after RSA. These measurements are correlated with postoperative clinical outcomes.

Signatures of the collapse and incipient recovery of an overexploited marine ecosystem.

The Northwest Atlantic cod stocks collapsed in the early 1990s and have yet to recover, despite the subsequent establishment of a continuing fishing moratorium. Efforts to understand the collapse and lack of recovery have so far focused mainly on the dynamics of commercially harvested species. Here, we use data from a 33-year scientific trawl survey to determine to which degree the signatures of the collapse and recovery of the cod are apparent in the spatial and temporal dynamics of the broader groundfish community. Over this 33-year period, the groundfish community experienced four phases of change: (i) a period of rapid, synchronous biomass collapse in most species, (ii) followed by a regime shift in community composition with a concomitant loss of functional diversity, (iii) followed in turn by periods of slow compositional recovery, and (iv) slow biomass growth. Our results demonstrate how a community-wide perspective can reveal new aspects of the dynamics of collapse and recovery unavailable from the analysis of individual species or a combination of a small number of species. Overall, we found evidence that such community-level signals should be useful for designing more effective management strategies to ensure the persistence of exploited marine ecosystems.

Recent advances in metacommunities and meta-ecosystem theories.

Metacommunity theory has provided many insights into the general problem of local versus regional control of species diversity and relative abundance. The metacommunity framework has been extended from competitive interactions to whole food webs that can be described as spatial networks of interaction networks. Trophic metacommunity theory greatly contributed to resolving the community complexity-stability debate by predicting its dependence on the regional spatial context. The meta-ecosystem framework has since been suggested as a useful simplification of complex ecosystems to apply this spatial context to spatial flows of both individuals and matter. Reviewing the recent literature on metacommunity and meta-ecosystem theories suggests the importance of unifying theories of interaction strength into a meta-ecosystem framework that captures how the strength of spatial, species, and ecosystem fluxes are distributed across location and trophic levels. Such integration predicts important feedback between local and regional processes that drive the assembly of species, the stability of community, and the emergence of ecosystem functions, from limited spatial fluxes of individuals and (in)organic matter. These predictions are often mediated by the maintenance of environmental or endogenous fluctuations from local to regional scales that create important challenges and opportunities for the validation of metacommunity and meta-ecosystem theories and their application to conservation.

Reciprocal feedbacks between spatial subsidies and reserve networks in coral reef meta-ecosystems.

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.

Nonhierarchical Dispersal Promotes Stability and Resilience in a Tritrophic Metacommunity.

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.

Patterns and scales of connectivity: temporal stability and variation within a marine metapopulation.

Because many marine invertebrates have a dispersive planktonic phase, the spatial scale of demographic, connectivity among local populations remains a key, but elusive, parameter driving population and metapopulation dynamics. However, temporal variation in the scale of connectivity remains largely undocumented, despite its recognized importance for predicting population responses to environmental changes. To assess the temporal stability of metapopulation connectivity, we conducted a large-scale survey of a blue mussel (Mytilus spp.) metapopulation for five years along a 100-km section of coastline of the Gaspé Peninsula, Québec, Canada. For each year, we estimated the scale of demographic coupling among 27-29 sites within our study region, using the spatial cross-covariance between adult abundance and recruit density across sites. Despite large interannual variability in overall recruit abundance, our analysis revealed stationary spatial distributions of adult and recruit abundance. More importantly, our analysis revealed a consistent demographic coupling among populations at a distance ranging from 12 to 24 km in all but one of the five years studied. The scale of connectivity in this system is thus temporally stable, but can occasionally show irregular fluctuations, and our results provide evidence in support of the integration of time-varying connectivity to marine metapopulation and reserve network theories.