Marine ecosystems such as the Baltic Sea are currently under strong atmospheric and anthropogenic pressure. Besides natural and human-induced changes in climate, major anthropogenic drivers such as overfishing and anthropogenic eutrophication are significantly affecting ecosystem structure and function. Recently, studies demonstrated the existence of alternative stable states in various terrestrial and aquatic ecosystems. These so-called ecosystem regime shifts have been explained mainly as a result of multiple causes, e.g. climatic regime shifts, overexploitation or a combination of both. The occurrence of ecosystem regime shifts has important management implications, as they can cause significant losses of ecological and economic resources. Because of hysteresis in ecosystem responses, restoring regimes considered as favourable may require drastic and expensive management actions. Also the Baltic Sea, the largest brackish water body in the world ocean, and its ecosystems are strongly affected by atmospheric and anthropogenic drivers. Here, we present results of an analysis of the state and development of the Central Baltic Sea ecosystem integrating hydroclimatic, nutrient, phyto-and zooplankton as well as fisheries data. Our analyses of 52 biotic and abiotic variables using multivariate statistics demonstrated a major reorganization of the ecosystem and identified two stable states between 1974 and 2005, separated by a transition period in 1988-1993. We show the change in Baltic ecosystem structure to have the characteristics of a discontinuous regime shift, initiated by climate-induced changes in the abiotic environment and stabilized by fisheries-induced feedback loops in the food web. Our results indicate the importance of maintaining the resilience of an ecosystem to atmospherically induced environmental change by reducing the anthropogenic impact.

Fisheries can have a large impact on marine ecosystems, because the effects of removing large predatory fish may cascade down the food web. The implications of these cascading processes on system functioning and resilience remain a source of intense scientific debate. By using field data covering a 30-year period, we show for the Baltic Sea that the underlying mechanisms of trophic cascades produced a shift in ecosystem functioning after the collapse of the top predator cod. We identified an ecological threshold, corresponding to a planktivore abundance of Ϸ17 ؋ 10 10 individuals, that separates 2 ecosystem configurations in which zooplankton dynamics are driven by either hydroclimatic forces or predation pressure. Abundances of the planktivore sprat above the threshold decouple zooplankton dynamics from hydrological circumstances. The current strong regulation by sprat of the feeding resources for larval cod may hinder cod recovery and the return of the ecosystem to a prior state. This calls for the inclusion of a food web perspective in management decisions.alternative dynamics ͉ ecological thresholds ͉ ecosystem resilience ͉ Baltic Sea ͉ climate versus top-down control

Anthropogenic disturbances intertwined with climatic changes can have a large impact on the upper trophic levels of marine ecosystems, which may cascade down the food web. So far it has been difficult to demonstrate multi-level trophic cascades in pelagic marine environments. Using field data collected during a 33-year period, we show for the first time a four-level community-wide trophic cascade in the open Baltic Sea. The dramatic reduction of the cod (Gadus morhua) population directly affected its main prey, the zooplanktivorous sprat (Sprattus sprattus), and indirectly the summer biomass of zooplankton and phytoplankton (top-down processes). Bottom-up processes and climate-hydrological forces had a weaker influence on sprat and zooplankton, whereas phytoplankton variation was explained solely by top-down mechanisms. Our results suggest that in order to dampen the occasionally harmful algal blooms of the Baltic, effort should be addressed not only to control anthropogenic nutrient inputs but also to preserve structure and functioning of higher trophic levels.

The index of the North Atlantic Oscillation, the dominant mode of climatic variability in the North Atlantic region, changed in the late 1980s (1987–1989) from a negative to a positive phase. This led to regime shifts in the ecology of the North Sea (NS) and the central Baltic Sea (CBS), which involved all trophic levels in the pelagial of these two neighbouring continental shelf seas. Increasing air and sea surface temperatures, which affected critical physical and biological processes, were the main direct and indirect driving forces. After 1987, phytoplankton biomass in both systems increased and the growing season was extended. The composition of phyto- and zooplankton communities in both seas changed conspicuously, e.g. dinoflagellate abundance increased and diatom abundance decreased in the CBS. Key copepod species that are essential in fish diets experienced pronounced changes in biomass. Abundance of Calanus finmarchicus (NS) and Pseudocalanus sp. (CBS) fell to low levels, whereas C. helgolandicus (NS) and Temora longicornis and Acartia spp. (CBS) were persistently abundant. These changes in biomass of different copepod species had dramatic consequences on biomass, fisheries, and landings of key fish species: North Sea cod declined, cod in the CBS remained at low levels, and CBS sprat reached unprecedented high biomass levels resulting in high yields. The synchronous regime shifts in NS and CBS resulted in profound changes in both marine ecosystems. However, the reaction of fish populations to the bottom-up mechanisms caused by the same climatic shift was very different for the three fish stocks.

Möllmann, C., Müller-Karulis, B., Kornilovs, G., and St John, M. A. 2008. Effects of climate and overfishing on zooplankton dynamics and ecosystem structure: regime shifts, trophic cascade, and feeback loops in a simple ecosystem. – ICES Journal of Marine Science, 65: 302–310. The Central Baltic Sea is the largest brackish waterbody in the world ocean, containing a highly productive but low-diversity ecosystem. Climate-induced changes in hydrography recently caused an ecosystem regime shift with changes at all trophic levels. The most pronounced changes in the ecosystem occurred at the zooplankton and fish trophic levels. In the zooplankton, dominance changed between the copepods Pseudocalanus acuspes and Acartia spp., a result of reduced salinities and increased temperatures. The change in hydrography also affected the reproductive success of the major fish species, resulting in a change in dominance from the piscivorous cod (Gadus morhua) to the planktivorous sprat (Sprattus sprattus). First, we investigate statistically the occurrence of regime shifts in time-series of key hydrographic variables and the biomass time-series of key species. Second, we demonstrate a three-level trophic cascade involving zooplankton. Finally, we model the ecosystem effects of the abiotic and biotic changes on copepod biomass and recruitment of fish stocks. Our results demonstrate the linkage between climate-induced zooplankton and fish regime changes, and how overfishing amplified the climate-induced changes at both trophic levels. Hence, our study demonstrates (i) the multiple pathways along which climatic and anthropogenic pressures can propagate through the foodweb; (ii) how both effects act synergistically to cause and stabilize regime changes; and (iii) the crucial role of zooplankton in mediating these ecosystem changes.

A unique dataset of stomach contents sampled between 1977 and 1999 in the central Baltic Sea was used to perform a comprehensive study of the feeding ecology of Central Baltic herring Clupea harengus and sprat Sprattus sprattus. Both fish species were mainly preying upon calanoid copepods with Pseudocalanus sp. dominating the diet of herring, whereas sprat generally preferred Temora longicornis. Sprat preyed upon older copepodite stages, indicating size-selective particulate feeding, whereas herring additionally fed on smaller copepodite stages, indicating occasional low food supply inducing filter-feeding. Additional food sources other than copepods were mysids in winter and autumn for medium to large herring, as well as cladocerans for sprat in spring and summer, determined by the seasonal occurrence of these plankton species. Seasonally the highest feeding activity of both fishes species occurred in spring and summer, the main reproductive periods of calanoid copepods. The most important food item for both predators in spring was Pseudocalanus sp. In summer sprat switched to T. longicornis and Acartia spp. Since the late 1970s, the total stomach fullness decreased and the fraction of empty stomachs increased. In parallel the amount of Pseudocalanus sp. in the diets of both fish species decreased. Further, a considerable dietary overlap between both species in spring indicated considerable competition for food resources, especially due to an enlarged sprat stock. The results of this study support the hypothesis that growth reductions observed in Baltic herring and sprat are due to combination of a change in food availability and an increase in density-dependent competition. # 2004 The Fisheries Society of the British Isles

The eastern Baltic (EB) cod (Gadus morhua) stock was depleted and overexploited for decades until the mid-2000s, when fishing mortality rapidly declined and biomass started to increase, as shown by stock assessments. These positive developments were partly assigned to effective management measures, and the EB cod was considered one of the most successful stock recoveries in recent times. In contrast to this optimistic view, the analytical stock assessment failed in 2014, leaving the present stock status unclear. Deteriorated quality of some basic input data for stock assessment in combination with changes in environmental and ecological conditions has led to an unusual situation for cod in the Baltic Sea, which poses new challenges for stock assessment and management advice. A number of adverse developments such as low nutritional condition and disappearance of larger individuals indicate that the stock is in distress. In this study, we (i) summarize the knowledge of recent changes in cod biology and ecosystem conditions, (ii) describe the subsequent challenges for stock assessment, and (iii) highlight the key questions where answers are urgently needed to understand the present stock status and provide scientifically solid support for cod management in the Baltic Sea.

A coupled hydrodynamic-trophodynamic individual-based model of drift and feeding was utilized to analyze the intra-and inter-annual variability in growth and survival of cod (Gadus morhua) larvae in the central Baltic Sea. Highly temporally and spatially resolved simulated flow fields were used to investigate the potential drift of larval cod from the centre of spawning effort in the Bornholm Basin towards their nursery areas through temporally resolved three-dimensional idealized prey fields. Stomach content analyses of larval cod from the Bornholm Basin revealed calanoid copepod nauplii and early copepodite stages to be the preferred prey organisms. The results of the model runs indicate that larval cod changed from a nonlimited to a food-limited stage because of the strong decrease in abundance of the calanoid copepod Pseudocalanus elongatus during the last two decades. The modeling study revealed retention and dispersal from the main spawning ground to be a key process influencing larval survival. When P. elongatus was available in the prey fields, high cod larval survival rates occurred in spring and early summer. In contrast, when P. elongatus was not available, hatched larvae had only high survival probabilities later in the year or if they were transported into shallower coastal regions.Résumé : Un modèle couplé hydrodynamique-trophodynamique basé sur l'individu qui décrit la dérive et l'alimentation nous a permis d'analyser la variabilité intra-et inter-annuelle de la croissance et de la survie des larves de la morue franche (Gadus morhua) dans le centre de la Baltique. Des champs à hautes résolutions temporelle et spatiale d'écoulement simulé ont servi à étudier la dérive potentielle des larves de morue à partir du point central où se produit l'effort maximal de fraie dans le bassin de Bornholm vers les zones de nourricerie à travers des champs idéalisés à trois dimensions et à haute résolution de prédateurs. L'analyse des contenus stomacaux des larves de morue du bassin de Bornholm révèle que les nauplies de copépodes calanoïdes et les premiers stades copépodites sont les proies préférées. À cause de l'important déclin de l'abondance du copépode calanoïde Pseudocalanus elongatus au cours des deux dernières décennies, les résultats de la modélisation indiquent que les larves de morue sont limitées par une pénurie de nourriture, alors qu'elles ne l'étaient pas auparavant. Le modèle démontre que la rétention et la dispersion à partir du site principal de fraie sont les processus déterminants qui influencent la survie des larves. Quand P. elongatus est disponible dans les champs de proies, les taux de survie des larves de morue au printemps et au début de l'été sont élevés. En revanche, lorsque P. elongatus n'est pas disponible, les larves à l'éclosion n'ont de fortes probabilités de survie que plus tard dans l'année et seulement si elles sont transportées vers les régions côtières moins profondes.[Traduit par la Rédaction] Hinrichsen et al. 1873