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'.

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.

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.

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.

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.

Emergence of nutrient co-limitation through movement in stoichiometric meta-ecosystems.

Evidence that ecosystems and primary producers are limited in their productivity by multiple nutrients has caused the traditional nutrient limitation framework to include multiple limiting nutrients. The models built to mimic these responses have invoked local mechanisms at the level of the primary producers. In this paper, we explore an alternative explanation for the emergence of co-limitation by developing a simple, stoichiometrically explicit meta-ecosystem model with two limiting nutrients, autotrophs and herbivores. Our results show that differences in movement rates for the nutrients, autotrophs and herbivores can allow for nutrient co-limitation in biomass response to emerge despite no local mechanisms of nutrient co-limitation. Furthermore, our results provide an explanation to why autotrophs show positive growth responses to nutrients despite 'nominal' top-down control by herbivores. These results suggest that spatial processes can be mechanisms for nutrient co-limitation at local and regional scales, and can help explain anomalous results in the co-limitation literature.

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.

Fidelity drive: a mechanism for chaperone proteins to maintain stable mutation rates in prokaryotes over evolutionary time.

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.

How robust is dispersal-induced spatial synchrony?

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.

The paradox of enrichment in metaecosystems.

The paradox of enrichment has been studied almost exclusively within communities or metacommunities, without explicit nutrient dynamics. Yet local recycling of materials from enriched ecosystems may affect the stability of connected ecosystems. Here we study the effect of nutrient, detritus, producer, and consumer spatial flows-combined with changes in regional enrichment-on the stability of a metaecosystem model. We considered both spatially homogeneous and heterogeneous enrichment. We found that nutrient and detritus spatial flows are destabilizing, whereas producer or consumer spatial flows are either neutral or stabilizing. We noticed that detritus spatial flows have only a weak impact on stability. Our study reveals that heterogeneity no longer stabilizes well-connected systems when accounting for explicit representation of nutrient dynamics. We also found that intermediate consumer diffusion could lead to multiple equilibria in strongly enriched metaecosystems. Stability can emerge from a top-down control allowing the storage of materials into inorganic form, a mechanism never documented before. In conclusion, local enrichment can be stabilized if spatial flows are strong enough to efficiently redistribute the local excess of enrichment to unfertile ecosystems. However, high regional enrichment can be dampened only by intermediate consumer diffusion rates.

Meta-ecosystem dynamics and functioning on finite spatial networks.

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.

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.

Designing effective reserve networks for nonequilibrium metacommunities.

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.

Ecological processes can synchronize marine population dynamics over continental scales.

Determining the relative importance of local and regional processes for the distribution of population abundance is a fundamental but contentious issue in ecology. In marine systems, classical theory holds that the influence of demographic processes and dispersal is confined to local populations whereas the environment controls regional patterns of abundance. Here, we use spatial synchrony to compare the distribution of population abundance of the dominant mussel Mytilus californianus observed along the West Coast of the United States to that predicted by dynamical models undergoing different dispersal and environmental treatments to infer the relative influence of local and regional processes. We reveal synchronized fluctuations in the abundance of mussel populations across a whole continent despite limited larval dispersal and strong environmental forcing. We show that dispersal among neighboring populations interacts with local demographic processes to generate characteristic patterns of spatial synchrony that can govern the dynamic distribution of mussel abundance over 1,800 km of coastline. Our study emphasizes the importance of dispersal and local dynamics for the distribution of abundance at the continental scale. It further highlights potential limits to the use of "climate envelope" models for predicting the response of large-scale ecosystems to global climate change.

Pairing dynamics and the origin of species.

Whether sexual selection alone can drive the evolution of assortative mating in the presence of gene flow is a long-standing question in evolutionary biology. Here, we report a role for pairing dynamics of individuals when mate choice is mutual, which is sufficient for the evolution of assortative mating by sexual selection alone in the presence of gene flow. Through behavioural observation, individual-based simulation and population genetic analysis, we evaluate the pairing dynamics of coral reef fish in the genus Hypoplectrus (Serranidae), and the role these dynamics can play for the evolution of assortative mating. When mate choice is mutual and the stability of mating pairs is critical for reproductive success, the evolution of assortative mating in the presence of gene flow is not only possible, but is also a robust evolutionary outcome.

Disentangling invasion processes in a dynamic shipping-boating network.

The relative importance of multiple vectors to the initial establishment, spread and population dynamics of invasive species remains poorly understood. This study used molecular methods to clarify the roles of commercial shipping and recreational boating in the invasion by the cosmopolitan tunicate, Botryllus schlosseri. We evaluated (i) single vs. multiple introduction scenarios, (ii) the relative importance of shipping and boating to primary introductions, (iii) the interaction between these vectors for spread (i.e. the presence of a shipping-boating network) and (iv) the role of boating in determining population similarity. Tunicates were sampled from 26 populations along the Nova Scotia, Canada, coast that were exposed to either shipping (i.e. ports) or boating (i.e. marinas) activities. A total of 874 individuals (c. 30 per population) from five ports and 21 marinas was collected and analysed using both mitochondrial cytochrome c oxidase subunit I gene (COI) and 10 nuclear microsatellite markers. The geographical location of multiple hotspot populations indicates that multiple invasions have occurred in Nova Scotia. A loss of genetic diversity from port to marina populations suggests a stronger influence of ships than recreational boats on primary coastal introductions. Population genetic similarity analysis reveals a dependence of marina populations on those that had been previously established in ports. Empirical data on marina connectivity because of boating better explains patterns in population similarities than does natural spread. We conclude that frequent primary introductions arise by ships and that secondary spread occurs gradually thereafter around individual ports, facilitated by recreational boating.