New insights into the recent collapse of Eastern Baltic cod from historical data on stock health.

The Eastern Baltic cod (Gadus morhua) stock is currently in a very poor state, with low biomass and adverse trends in several life history and demographic parameters. This raises concern over whether and to what level recovery is possible. Here, we look for new insights from a historical perspective, extending the time series of various stock health indicators back to the 1940s, i.e. to the beginning of intensive exploitation of the Eastern Baltic cod. The historical data confirm that the stock deterioration in recent years is unprecedented, as all indicators are presently in their worst states on record. Cod body condition and energy reserves were equally low in the 1940s-1950s, accompanied by high parasitic liver worm infection, comparable to that measured in recent years. However, other stock parameters (size structure, size at maturity, stock distribution) are currently in their worst states over the past 80 years. In contrast, the state of cod in the 1970s to early 1990s that is often perceived as a desirable target, was exceptional, with the most favorable indicator levels in the time series. Long-term observation data reveal concurrent or asynchronous trends in different indicators of stock health and to what extent these have coincided with changes in possible external drivers. In this way, the extended time series contribute to ongoing research on understanding the collapse of the cod and its recovery potential.

Genetic analyses reveal complex dynamics within a marine fish management area.

Genetic data have great potential for improving fisheries management by identifying the fundamental management units-that is, the biological populations-and their mixing. However, so far, the number of practical cases of marine fisheries management using genetics has been limited. Here, we used Atlantic cod in the Baltic Sea to demonstrate the applicability of genetics to a complex management scenario involving mixing of two genetically divergent populations. Specifically, we addressed several assumptions used in the current assessment of the two populations. Through analysis of 483 single nucleotide polymorphisms (SNPs) distributed across the Atlantic cod genome, we confirmed that a model of mechanical mixing, rather than hybridization and introgression, best explained the pattern of genetic differentiation. Thus, the fishery is best monitored as a mixed-stock fishery. Next, we developed a targeted panel of 39 SNPs with high statistical power for identifying population of origin and analyzed more than 2,000 tissue samples collected between 2011 and 2015 as well as 260 otoliths collected in 2003/2004. These data provided high spatial resolution and allowed us to investigate geographical trends in mixing, to compare patterns for different life stages and to investigate temporal trends in mixing. We found similar geographical trends for the two time points represented by tissue and otolith samples and that a recently implemented geographical management separation of the two populations provided a relatively close match to their distributions. In contrast to the current assumption, we found that patterns of mixing differed between juveniles and adults, a signal likely linked to the different reproductive dynamics of the two populations. Collectively, our data confirm that genetics is an operational tool for complex fisheries management applications. We recommend focussing on developing population assessment models and fisheries management frameworks to capitalize fully on the additional information offered by genetically assisted fisheries monitoring.

Testing spatial heterogeneity with stock assessment models.

This paper describes a methodology that combines meta-population theory and stock assessment models to gain insights about spatial heterogeneity of the meta-population in an operational time frame. The methodology was tested with stochastic simulations for different degrees of connectivity between sub-populations and applied to two case studies, North Sea cod (Gadus morua) and Northeast Atlantic sardine (Sardina pilchardus). Considering that the biological components of a population can be partitioned into discrete spatial units, we extended this idea into a property of additivity of sub-population abundances. If the additivity results hold true for putative sub-populations, then assessment results based on sub-populations will provide information to develop and monitor the implementation of finer scale/local management. The simulation study confirmed that when sub-populations are independent and not too heterogeneous with regards to productivity, the sum of stock assessment model estimates of sub-populations' SSB is similar to the SSB estimates of the meta-population. It also showed that a strong diffusion process can be detected and that the stronger the connection between SSB and recruitment, the better the diffusion process will be detected. On the other hand it showed that weak to moderate diffusion processes are not easy to identify and large differences between sub-populations productivities may be confounded with weak diffusion processes. The application to North Sea cod and Atlantic sardine exemplified how much insight can be gained. In both cases the results obtained were sufficiently robust to support the regional analysis.

Has eutrophication promoted forage fish production in the Baltic Sea?

Reducing anthropogenic nutrient inputs is a major policy goal for restoring good environmental status of coastal marine ecosystems. However, it is unclear to what extent reducing nutrients would also lower fish production and fisheries yields. Empirical examples of changes in nutrient loads and concurrent fish production can provide useful insights to this question. In this paper, we investigate to what extent a multi-fold increase in nutrient loads from the 1950s to 1980s enhanced forage fish production in the Baltic Sea. We use monitoring data on fish stock dynamics covering the period of the nutrient increase, combined with nutrient concentrations from a 3-dimensional coupled physical-biogeochemical ocean model. The results suggest that nutrient enrichment enhanced the biomass level of forage fish by up to 50 % in some years and areas due to increased body weight of fish. However, the trends in fish biomasses were generally decoupled from changes in nutrient concentrations.

Impact of climate change on fish population dynamics in the Baltic sea: a dynamical downscaling investigation.

Understanding how climate change, exploitation and eutrophication will affect populations and ecosystems of the Baltic Sea can be facilitated with models which realistically combine these forcings into common frameworks. Here, we evaluate sensitivity of fish recruitment and population dynamics to past and future environmental forcings provided by three ocean-biogeochemical models of the Baltic Sea. Modeled temperature explained nearly as much variability in reproductive success of sprat (Sprattus sprattus; Clupeidae) as measured temperatures during 1973-2005, and both the spawner biomass and the temperature have influenced recruitment for at least 50 years. The three Baltic Sea models estimate relatively similar developments (increases) in biomass and fishery yield during twenty-first century climate change (ca. 28 % range among models). However, this uncertainty is exceeded by the one associated with the fish population model, and by the source of global climate data used by regional models. Knowledge of processes and biases could reduce these uncertainties.

Methodological challenges in assessing the environmental status of a marine ecosystem: case study of the Baltic Sea.

Assessments of the environmental status of marine ecosystems are increasingly needed to inform management decisions and regulate human pressures to meet the objectives of environmental policies. This paper addresses some generic methodological challenges and related uncertainties involved in marine ecosystem assessment, using the central Baltic Sea as a case study. The objectives of good environmental status of the Baltic Sea are largely focusing on biodiversity, eutrophication and hazardous substances. In this paper, we conduct comparative evaluations of the status of these three segments, by applying different methodological approaches. Our analyses indicate that the assessment results are sensitive to a selection of indicators for ecological quality objectives that are affected by a broad spectrum of human activities and natural processes (biodiversity), less so for objectives that are influenced by a relatively narrow array of drivers (eutrophications, hazardous substances). The choice of indicator aggregation rule appeared to be of essential importance for assessment results for all three segments, whereas the hierarchical structure of indicators had only a minor influence. Trend-based assessment was shown to be a useful supplement to reference-based evaluation, being independent of the problems related to defining reference values and indicator aggregation methodologies. Results of this study will help in setting priorities for future efforts to improve environmental assessments in the Baltic Sea and elsewhere, and to ensure the transparency of the assessment procedure.

Could seals prevent cod recovery in the Baltic Sea?

Fish populations are increasingly affected by multiple human and natural impacts including exploitation, eutrophication, habitat alteration and climate change. As a result many collapsed populations may have to recover in ecosystems whose structure and functioning differ from those in which they were formerly productive and supported sustainable fisheries. Here we investigate how a cod (Gadus morhua) population in the Baltic Sea whose biomass was reduced due to a combination of high exploitation and deteriorating environmental conditions might recover and develop in the 21st century in an ecosystem that likely will change due to both the already started recovery of a cod predator, the grey seal Halichoerus grypus, and projected climate impacts. Simulation modelling, assuming increased seal predation, fishing levels consistent with management plan targets and stable salinity, shows that the cod population could reach high levels well above the long-term average. Scenarios with similar seal and fishing levels but with 15% lower salinity suggest that the Baltic will still be able to support a cod population which can sustain a fishery, but biomass and yields will be lower. At present knowledge of cod and seal interactions, seal predation was found to have much lower impact on cod recovery, compared to the effects of exploitation and salinity. These results suggest that dual management objectives (recovery of both seal and cod populations) are realistic but success in achieving these goals will also depend on how climate change affects cod recruitment.

Multi-decadal responses of a cod (Gadus morhua) population to human-induced trophic changes, fishing, and climate.

Understanding how human impacts have interacted with natural variability to affect populations and ecosystems is required for sustainable management and conservation. The Baltic Sea is one of the few large marine ecosystems worldwide where the relative contribution of several key forcings to changes in fish populations can be analyzed with empirical data. In this study we investigate how climate variability and multiple human impacts (fishing, marine mammal hunting, eutrophication) have affected multi-decadal scale dynamics of cod in the Baltic Sea during the 20th century. We document significant climate-driven variations in cod recruitment production at multi-annual timescales, which had major impacts on population dynamics and the yields to commercial fisheries. We also quantify the roles of marine mammal predation, eutrophication, and exploitation on the development of the cod population using simulation analyses, and show how the intensity of these forcings differed over time. In the early decades of the 20th century, marine mammal predation and nutrient availability were the main limiting factors; exploitation of cod was still relatively low. During the 1940s and subsequent decades, exploitation increased and became a dominant forcing on the population. Eutrophication had a relatively minor positive influence on cod biomass until the 1980s. The largest increase in cod biomass occurred during the late 1970s, following a long period of hydrographically related above-average cod productivity coupled to a temporary reduction in fishing pressure. The Baltic cod example demonstrates how combinations of different forcings can have synergistic effects and consequently dramatic impacts on population dynamics. Our results highlight the potential and limitations of human manipulations to influence predator species and show that sustainable management can only be achieved by considering both anthropogenic and naturally varying processes in a common framework.