No abstract

Investigating the factors regulating fish condition is crucial in ecology and the management of exploited fish populations. The body condition of cod (Gadus morhua) in the Baltic Sea has dramatically decreased during the past two decades, with large implications for the fishery relying on this resource. Here, we statistically investigated the potential drivers of the Baltic cod condition during the past 40 years using newly compiled fishery-independent biological data and hydrological observations. We evidenced a combination of different factors operating before and after the ecological regime shift that occurred in the Baltic Sea in the early 1990s. The changes in cod condition related to feeding opportunities, driven either by density-dependence or food limitation, along the whole period investigated and to the fivefold increase in the extent of hypoxic areas in the most recent 20 years. Hypoxic areas can act on cod condition through different mechanisms related directly to species physiology, or indirectly to behaviour and trophic interactions. Our analyses found statistical evidence for an effect of the hypoxia-induced habitat compression on cod condition possibly operating via crowding and density-dependent processes. These results furnish novel insights into the population dynamics of Baltic Sea cod that can aid the management of this currently threatened population.

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.

Five decades of stomach content data allowed insight into the development of consumption, diet composition, and resulting somatic growth of Gadus morhua (Atlantic cod) in the eastern Baltic Sea. We show a recent reversal in feeding level over body length. Present feeding levels of small cod indicate severe growth limitation and increased starvation-related mortality. For young cod, the low growth rate and the high mortality rate are manifested through a reduction in size-at-age. The low feeding levels are likely the result of a decrease in benthic prey abundance due to increased hypoxic areas, while decreasing abundances of pelagic species in the area of cod distribution have prevented a compensatory shift in diet. Our study emphasizes that environmental forcing and the decline in pelagic prey caused changes in consumption and growth rates of small cod. The food reduction is amplified by stunted growth leading to high densities of cod of smaller size competing for the scarce resources. The average growth rate is negative, and only individuals with feeding levels well above average will survive, though growing slowly. These results suggest that the relation between consumption rate, somatic growth and predatorprey population densities is strongly environmentally mediated.

Individual behaviour of Atlantic cod Gadus morhua in the presence of hypoxic water was measured in situ in the vertically stratified Bornholm Basin of the Baltic Sea. Considering all recaptured individuals, the use of hypoxic habitat was comparable to data derived by traditional survey data, but some G. morhua had migrated towards the centre of the c.100 m deep basin and spent about a third of their time at oxygen saturation <50%, possibly to forage on zoobenthos. Maximal residence time per visit in such hypoxic water was limited to a few hours, allowing for the digestion of consumed prey items in waters with sufficient dissolved oxygen.

The Baltic Sea comprises a heterogeneous oceanographic environment influencing the spatial and temporal potential for reproductive success of cod (Gadus morhua) and sprat (Sprattus sprattus) in the different spawning basins. Hence, to quantify stock and recruitment dynamics, it is necessary to resolve species-specific regional reproductive success in relation to size, structure, and distribution of the spawning stock. Furthermore, as species and fisheries interactions vary between areas, it is necessary to include these interactions on an area-specific basis. Therefore, area-disaggregated multispecies virtual population analyses (MSVPA) were performed for interacting species cod, herring (Clupea harengus), and sprat in the different subdivisions of the Central Baltic. The MSVPA runs revealed distinct spatial trends in population abundance, spawning biomass, recruitment, and predation-induced mortality. Results, when evaluated with respect to trends in population sizes from research surveys, were similar for the cod and sprat stocks but different for herring. Horizontal distributions from MSVPA runs and research surveys indicate that cod and sprat undergo migrations between basins during different life stages. This is an observation potentially influencing estimates for the different stock components but not affecting the overall stock sizes.

Theory behind ecosystem-based management (EBM) and ecosystem-based fisheries management (EBFM) is now well developed. However, the implementation of EBFM exemplified by fisheries management in Europe is still largely based on single-species assessments and ignores the wider ecosystem context and impact. The reason for the lack or slow implementation of EBM and specifically EBFM is a lack of a coherent strategy. Such a strategy is offered by recently developed integrated ecosystem assessments (IEAs), a formal synthesis tool to quantitatively analyse information on relevant natural and socio-economic factors, in relation to specified management objectives. Here, we focus on implementing the IEA approach for Baltic Sea fish stocks. We combine both tactical and strategic management aspects into a single strategy that supports the present Baltic Sea fish stock advice, conducted by the International Council for the Exploration of the Sea (ICES). We first review the state of the art in the development of IEA within the current management framework. We then outline and discuss an approach that integrates fish stock advice and IEAs for the Baltic Sea. We intentionally focus on the central Baltic Sea and its three major fish stocks cod (Gadus morhua), herring (Clupea harengus), and sprat (Sprattus sprattus), but emphasize that our approach may be applied to other parts and stocks of the Baltic, as well as other ocean areas.

Cod is the top piscivore predator in the Baltic Sea ecosystem. Based on stomach content data from 62 427 cod collected during 1977-1994 and food consumption rates, cannibalism in the Eastern and Western Baltic cod stocks has been quantified using multispecies virtual population analysis. In the Eastern Baltic stock, depending on model assumptions, an average of 25-38% of the 0-group and 11-17% of the 1-group were removed by predation by adults. Thus, between age 0 and age 2 a year class may lose on average about 31% and 44% of the initial number as a result of cannibalism. Cannibalism is lower in the Western Baltic. On average, 19% of the 0-group and 9% of the 1-group are consumed per year, i.e. 24% of the initial cohort is eaten before reaching age 2. Predation was most intense in 1978-1984, a period with high juvenile abundance and large adult stock sizes in both areas. Subsequently, stock, recruitment, and cannibalism declined steadily until the early 1990s and then increased again. Problems identified in relation to data compilation and estimation procedure are discussed with respect to their impact on estimates of cannibalism and stockrecruitment relationships.