Benthic-pelagic coupling is manifested as the exchange of energy, mass, or nutrients between benthic and pelagic habitats. It plays a prominent role in aquatic ecosystems, and it is crucial to functions from nutrient cycling to energy transfer in food webs. Coastal and estuarine ecosystem structure and function are strongly affected by anthropogenic pressures; however, there are large gaps in our understanding of the responses of inorganic nutrient and organic matter fluxes between benthic habitats and the water column. We illustrate the varied nature of physical and biological benthic-pelagic coupling processes and their potential sensitivity to three anthropogenic pressures -climate change, nutrient loading, and fishing -using the Baltic Sea as a case study and summarize current knowledge on the exchange of inorganic nutrients and organic material between habitats. Traditionally measured benthic-pelagic coupling processes (e.g., nutrient exchange and sedimentation of organic material) are to some extent quantifiable, but the magnitude and variability of biological processes are rarely assessed, preventing quantitative comparisons. Changing oxygen conditions will continue to have widespread effects on the processes that govern inorganic and organic matter exchange among habitats while climate change and nutrient load reductions may have large effects on organic matter sedimentation. Many biological processes (predation, bioturbation) are expected to be sensitive to anthropogenic drivers, but the outcomes for ecosystem function are largely unknown. We emphasize how improved empirical and experimental understanding of benthic-pelagic coupling processes and their variability are necessary to inform models that can quantify the feedbacks among processes and ecosystem responses to a changing world.

Since the days of Elton, population cycles have challenged ecologists and resource managers. Although the underlying mechanisms remain debated, theory holds that both density-dependent and density-independent processes shape the dynamics. One striking example is the large-scale fluctuations of sardine and anchovy observed across the major upwelling areas of the world. Despite a long history of research, the causes of these fluctuations remain unresolved and heavily debated, with significant implications for fisheries management. We here model the underlying causes of these fluctuations, using the California Current Ecosystem as a case study, and show that the dynamics, accurately reproduced since A.D. 1661 onward, are explained by interacting density-dependent processes (i.e., through species-specific life-history traits) and climate forcing. Furthermore, we demonstrate how fishing modifies the dynamics and show that the sardine collapse of the 1950s was largely unavoidable given poor recruitment conditions. Our approach provides unique insight into the origin of sardine-anchovy fluctuations and a knowledge base for sustainable fisheries management in the California Current Ecosystem and beyond.species replacement | population modeling | climate change | ecosystem-based management M arine fish typically show multidecadal fluctuations in abundance, mainly attributed to overexploitation (1), climate (2), or a combination of both (3). Furthermore, such low-frequency variability may arise from density-dependent processes (i.e., cohort-resonance) potentially masking external or anthropogenic effects (4), whereas fishing-induced demographic changes increase short-term variability (5-7). To understand these fluctuations, it is essential to disentangle anthropogenic forcing from internal processes, as well as understanding the way they interact (7). One of the most striking examples of population fluctuations is the alternating regimes of sardine and anchovy observed across the major upwelling areas of the world (8, 9). Although oceanatmosphere forcing is considered the main underlying driver (8-11), no generally accepted theory regarding sardine-anchovy fluctuations presently exists (12). Nevertheless, insight accumulated over a long history of research highlights that the key to understanding population cycles in general, and the sardine-anchovy puzzle in particular, lies not in identifying a single factor but a combination of interacting factors (12)(13)(14).In the California Current Ecosystem (CCE), the Pacific sardine (Sardinops sagax) and northern anchovy (Engraulis mordax) exhibit pronounced fluctuations, spanning several orders of magnitude in terms of biomass (15, 16). These fluctuations, occurring with a dominant periodicity of ∼60 y (15, 16), have been linked to changes in the strength and position of the Aleutian Low, as reflected by the Pacific Decadal Oscillation (PDO) index (17,18), and correlated with patterns of flow, upwelling, and physical/biotic conditions (10,12,19). Furthermore, fishing has been su...

Worldwide a number of fish stocks have collapsed because of overfishing and climate-induced ecosystem changes. Developing ecosystem-based fisheries management (EBFM) to prevent these catastrophic events in the future requires ecological models incorporating both internal food-web dynamics and external drivers such as fishing and climate. Using a stochastic food-web model for a large marine ecosystem (i.e., the Baltic Sea) hosting a commercially important cod stock, we were able to reconstruct the history of the stock. Moreover we demonstrate that in hindsight the collapse could only have been avoidable by adapting fishing pressure to environmental conditions and food-web interactions. The modeling approach presented here represents a significant advance for EBFM, the application of which is important for sustainable resource management in the future.ecosystem-based fisheries management ͉ sustainability

Collapses and regime changes are pervasive in complex systems (such as marine ecosystems) governed by multiple stressors. The demise of Atlantic cod ( Gadus morhua ) stocks constitutes a text book example of the consequences of overexploiting marine living resources, yet the drivers of these nearly synchronous collapses are still debated. Moreover, it is still unclear why rebuilding of collapsed fish stocks such as cod is often slow or absent. Here, we apply the stochastic cusp model, based on catastrophe theory, and show that collapse and recovery of cod stocks are potentially driven by the specific interaction between exploitation pressure and environmental drivers. Our statistical modelling study demonstrates that for most of the cod stocks, ocean warming could induce a nonlinear discontinuous relationship between fishing pressure and stock size, which would explain hysteresis in their response to reduced exploitation pressure. Our study suggests further that a continuing increase in ocean temperatures will probably limit productivity and hence future fishing opportunities for most cod stocks of the Atlantic Ocean. Moreover, our study contributes to the ongoing discussion on the importance of climate and fishing effects on commercially exploited fish stocks, highlighting the importance of considering discontinuous dynamics in holistic ecosystem-based management approaches, particularly under climate change.

Aim:The life history of a species is determined by trade-offs between growth, survival and reproduction to maximize fitness in a given environment. Following a theoretical model, we investigate whether the composition of marine fish communities can be understood in terms of a set of lifehistory strategies and whether the prevalence of the strategies follows specific spatial patterns that can be related to the environment.Location: European seas.Time period: 1980-2014.Major taxa studied: Fish.Methods: An extensive set of scientific bottom trawl surveys was collected to obtain the species composition of fish communities across European seas. We complemented these data with species-specific information regarding six life-history traits, reflecting reproductive, growth and feeding modes. We then calculated the optimal number of strategies needed to summarize the information contained in the traits by using archetypal analysis. The proportion of each obtained strategy in the communities and their spatial patterns were explained as a function of the environment and their temporal changes were investigated.Results: The species could be decomposed into a continuum of three life-history strategiesopportunistic, periodic and equilibrium-resulting from trade-offs between traits. The marked spatial patterns of these strategies could be explained by depth, temperature and its seasonality, chlorophyll and fishing effort. In recent years, opportunistic and equilibrium strategies significantly increased, probably due to an increase in temperature and decrease in fishing effort.Main conclusions: Our empirical analysis supports a theoretical framework outlining three lifehistory strategies of fish. The strategies vary predictably in space and time in response to the environment. This highlights the underlying process whereby fitness is optimized through trade-offs between growth, feeding and reproduction under different environmental conditions. Due to their response to the environment, life-history strategies provide a suitable tool for monitoring and understanding community changes in response to natural and anthropogenic stressors, including fishing and climate change. K E Y W O R D S archetypal analysis, community composition, depth, fecundity, life-history strategies, marine fish, offspring survival, size, temperature, trade-off, trait 812 |

Understanding the effects of cross-system fluxes is fundamental in ecosystem ecology and biological conservation. Source-sink dynamics and spillover processes may link adjacent ecosystems by movement of organisms across system boundaries. However, effects of temporal variability in these cross-system fluxes on a whole marine ecosystem structure have not yet been presented. Here we show, using 35 y of multitrophic data series from the Baltic Sea, that transitory spillover of the top-predator cod from its main distribution area produces cascading effects in the whole food web of an adjacent and semi-isolated ecosystem. At varying population size, cod expand/contract their distribution range and invade/retreat from the neighboring Gulf of Riga, thereby affecting the local prey population of herring and, indirectly, zooplankton and phytoplankton via top-down control. The Gulf of Riga can be considered for cod a "true sink" habitat, where in the absence of immigration from the source areas of the central Baltic Sea the cod population goes extinct due to the absence of suitable spawning grounds. Our results add a metaecosystem perspective to the ongoing intense scientific debate on the key role of top predators in structuring natural systems. The integration of regional and local processes is central to predict species and ecosystem responses to future climate changes and ongoing anthropogenic disturbances.ecosystem regulation | predator distribution | landscape ecology | exploited resources | cross-system management

Natural resource management requires approaches to understand and handle sources of uncertainty in future responses of complex systems to human activities. Here we present one such approach, the "biological ensemble modeling approach," using the Eastern Baltic cod (Gadus morhua callarias) as an example. The core of the approach is to expose an ensemble of models with different ecological assumptions to climate forcing, using multiple realizations of each climate scenario. We simulated the long-term response of cod to future fishing and climate change in seven ecological models ranging from single-species to food web models. These models were analyzed using the "biological ensemble modeling approach" by which we (1) identified a key ecological mechanism explaining the differences in simulated cod responses between models, (2) disentangled the uncertainty caused by differences in ecological model assumptions from the statistical uncertainty of future climate, and (3) identified results common for the whole model ensemble. Species interactions greatly influenced the simulated response of cod to fishing and climate, as well as the degree to which the statistical uncertainty of climate trajectories carried through to uncertainty of cod responses. Models ignoring the feedback from prey on cod showed large interannual fluctuations in cod dynamics and were more sensitive to the underlying uncertainty of climate forcing than models accounting for such stabilizing predator-prey feedbacks. Yet in all models, intense fishing prevented recovery, and climate change further decreased the cod population. Our study demonstrates how the biological ensemble modeling approach makes it possible to evaluate the relative importance of different sources of uncertainty in future species responses, as well as to seek scientific conclusions and sustainable management solutions robust to uncertainty of food web processes in the face of climate change.

The relationship between biodiversity and ecosystem functioning (BEF) is a topic of considerable interest to scientists and managers because a better understanding of its underlying mechanisms may help us mitigate the consequences of biodiversity loss on ecosystems. Our current knowledge of BEF relies heavily on theoretical and experimental studies, typically conducted on a narrow range of spatio-temporal scales, environmental conditions, and trophic levels. Hence, whether a relationship holds in the natural environment is poorly understood, especially in exploited marine ecosystems. Using large-scale observations of marine fish communities, we applied a structural equation modelling framework to investigate the existence and significance of BEF relationships across northwestern European seas. We find that ecosystem functioning, here represented by spatial patterns in total fish biomass, is unrelated to species richness—the most commonly used diversity metric in BEF studies. Instead, community evenness, differences in species composition, and abiotic variables are significant drivers. In particular, we find that high fish biomass is associated with fish assemblages dominated by a few generalist species of a high trophic level, who are able to exploit both the benthic and pelagic energy pathway. Our study provides a better understanding of the mechanisms behind marine ecosystem functioning and allows for the integration of biodiversity into management considerations.