Climate change and resource exploitation have been shown to modify the importance of bottom-up and top-down forces in ecosystems. However, the resulting pattern of trophic control in complex food webs is an emergent property of the system and thus unintuitive. We develop a statistical nondeterministic model, capable of modeling complex patterns of trophic control for the heavily impacted North Sea ecosystem. The model is driven solely by fishing mortality and climatic variables and based on timeseries data covering >40 y for six plankton and eight fish groups along with one bird group (>20 y). Simulations show the outstanding importance of top-down exploitation pressure for the dynamics of fish populations. Whereas fishing effects on predators indirectly altered plankton abundance, bottom-up climatic processes dominate plankton dynamics. Importantly, we show planktivorous fish to have a central role in the North Sea food web initiating complex cascading effects across and between trophic levels. Our linked model integrates bottom-up and top-down effects and is able to simulate complex long-term changes in ecosystem components under a combination of stressor scenarios. Our results suggest that in marine ecosystems, pathways for bottomup and top-down forces are not necessarily mutually exclusive and together can lead to the emergence of complex patterns of control.trophic control | ecosystem modeling | marine food web functioning | wasp-waist | regime shifts T he question of whether food webs are resource-(bottom-up) or predation-(top-down) controlled is one of the most fundamental research questions in ecology (1-3). Marine ecosystems, originally thought to be mainly steered by bottom-up control, have recently been shown to exhibit periods of top-down control due to the extraction of large predators through fishing (4-7) or climate oscillations (8). Furthermore, experimental evidence shows climate warming may exert a host of indirect effects on aquatic food webs mediated through shifts in the magnitudes of top-down and bottom-up forcing (9, 10). However, for large marine ecosystems that are not amenable to experimentation studies, investigations of how interactions in their complex food webs mediate the influence of both top-down (e.g., fishing) and bottom-up (e.g., climate change) control are lacking or are based on aggregated species complexes. We model an extensive historical dataset for the North Sea (over 45 y) at the lowest possible resolution (often species) to determine key interactions between species and estimate their responses to pressures. The model reveals both simple (direct) and complex (indirect) pathways linking plankton to seabirds and can highlight the wider effects of climate change and potential actions by fishery managers.The North Sea is one of the most anthropogenically impacted marine ecosystem and is thought to be fundamentally driven from the bottom-up through climatic (temperature-related) influences on plankton, planktivorous fish, and the pelagic stages of demersal fish (11-13). Som...

It is well known that human activities, such as harvesting, have had major direct effects on marine ecosystems. However, it is far less acknowledged that human activities in the surroundings might have important effects on marine systems. There is growing evidence suggesting that major reorganization (i.e., a regime shift) is a common feature in the temporal evolution of a marine system. Here we show, and quantify, the interaction of human activities (nutrient upload) with a favourable climate (run-off) and its contribution to the eutrophication of the Black Sea in the 1980s. Based on virtual analysis of the bottom-up (eutrophication) vs. top-down (trophic cascades) effects, we found that an earlier onset of eutrophication could have counteracted the restructuring of the trophic regulation at the base of the food web that resulted from the depletion of top predators in the 1970s. These enhanced bottom-up effects would, however, not propagate upwards in the food web beyond the zooplankton level. Our simulations identified the removal of apex predators as a key element in terms of loss of resilience that inevitably leads to a reorganization. Once the food web has been truncated, the type and magnitude of interventions on the group replacing the apex predator as the new upper trophic level have no effect in preventing the trophic cascade. By characterizing the tipping point at which increased bottom-up forcing exactly counteracts the top-down cascading effects, our results emphasize the importance of a comprehensive analysis that take into account all structuring forces at play (including those beyond the marine system) at a given time.

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

Payne, M. R., Hatfield, E. M. C., Dickey-Collas, M., Falkenhaug, T., Gallego, A., Gröger, J., Licandro, P., Llope, M., Munk, P., Röckmann, C., Schmidt, J. O., and Nash, R. D. M. 2009. Recruitment in a changing environment: the 2000s North Sea herring recruitment failure. – ICES Journal of Marine Science, 66: 272–277. Environmentally induced change appears to be impacting the recruitment of North Sea herring (Clupea harengus). Despite simultaneously having a large adult population, historically low exploitation, and Marine Stewardship Council accreditation (implying sustainability), there have been an unprecedented 6 sequential years of poor juvenile production (recruitment). Analysis suggests that the poor recruitment arises during the larval overwintering phase, with recent survival rates greatly reduced. Contemporary warming of the North Sea has caused significant changes in the plankton community, and a recently identified regime shift around 2000 shows close temporal agreement with the reduced larval survival. It is, therefore, possible that we are observing the first consequences of this planktonic change for higher trophic levels. There is no indication of a recovery in recruitment in the short term. Fishing mortality is currently outside the agreed management plan, and forecasts show a high risk of the stock moving outside safe biological limits soon, potentially precipitating another collapse of the stock. However, bringing the realized fishing mortality back in line with the management plan would likely alleviate the problem. This illustrates again that recruitment is influenced by more than just spawning-stock biomass, and that changes in other factors can be of equal, or even greater, importance. In such dynamically changing environments, recent management success does not necessarily guarantee future sustainability.

In order to provide better fisheries management and conservation decisions, there is a need to discern the underlying relationship between the spawning stock and recruitment of marine fishes, a relationship which is influenced by the environmental conditions. Here, we demonstrate how the environmental conditions (temperature and the food availability for fish larvae) influence the stock–recruitment relationship and indeed what kind of stock–recruitment relationship we might see under different environmental conditions. Using unique zooplankton data from the Continuous Plankton Recorder, we find that food availability (i.e. zooplankton) in essence determines which model applies for the once large North Sea cod (Gadus morhua) stock. Further, we show that recruitment is strengthened during cold years and weakened during warm years. Our combined model explained 45 per cent of the total variance in cod recruitment, while the traditional Ricker and Beverton–Holt models only explained about 10 per cent. Specifically, our approach predicts that a full recovery of the North Sea cod stock might not be expected until the environment becomes more favourable.

[1] The southern Bay of Biscay (NW Spain) shows a very active hydrography due to the different origins of its Central Waters, the local modifications exerted on them by continental effects and the recurrence of mesoscale processes such as slope currents, upwellings and eddies. In order to assess the role of the different sources of variability we conducted a monthly series of CTD sampling in the central Cantabrian Sea along a coastal-oceanic transect, from 1993 to 2003. We analyzed the spatial variability of the hydrographic processes over different timescales. The thermohaline properties of Central Waters varied between those typical of the subpolar mode of the Eastern North Atlantic Central Water (ENACWsp) and a local mode, the Bay of Biscay Central Water (BBCW), though there has been a clear shift toward the BBCW prevalence in the last years. The Iberian Poleward Current (IPC) conveyed subtropical Central Waters (ENACWst) into the region almost every winter. This slope current may display a double-core structure during some extreme events. The upper layers of the ocean showed a long-term trend toward increasing temperature and decreasing salinity, and accordingly density was on the decrease. These patterns suggest an enhancement of the water column stratification. Coastal upwellings are an important source of inshore variability and counteract these long-term changes on the coast. However, their intensity seems to be decreasing and their seasonal pattern changing toward a general advancement of the upwelling-favorable season.

Overfishing of large predatory fish populations has resulted in lasting restructurings of entire marine food webs worldwide, with serious socio-economic consequences. Fortunately, some degraded ecosystems show signs of recovery. A key challenge for ecosystem management is to anticipate the degree to which recovery is possible. By applying a statistical food-web model, using the Baltic Sea as a case study, we show that under current temperature and salinity conditions, complete recovery of this heavily altered ecosystem will be impossible. Instead, the ecosystem regenerates towards a new ecological baseline. This new baseline is characterized by lower and more variable biomass of cod, the commercially most important fish stock in the Baltic Sea, even under very low exploitation pressure. Furthermore, a socio-economic assessment shows that this signal is amplified at the level of societal costs, owing to increased uncertainty in biomass and reduced consumer surplus. Specifically, the combined economic losses amount to approximately 120 million € per year, which equals half of today's maximum economic yield for the Baltic cod fishery. Our analyses suggest that shifts in ecological and economic baselines can lead to higher economic uncertainty and costs for exploited ecosystems, in particular, under climate change.

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