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Ecosystem-based management requires understanding of food webs. Consequently, assessment of food web status is mandatory according to the European Union’s Marine Strategy Framework Directive (MSFD) for EU Member States. However, how to best monitor and assess food webs in practise has proven a challenging question. Here, we review and assess the current status of food web indicators and food web models, and discuss whether the models can help addressing current shortcomings of indicator-based food web assessments, using the Baltic Sea as an example region. We show that although the MSFD food web assessment was designed to use food web indicators alone, they are currently poorly fit for the purpose, because they lack interconnectivity of trophic guilds. We then argue that the multiple food web models published for this region have a high potential to provide additional coherence to the definition of good environmental status, the evaluation of uncertainties, and estimates for unsampled indicator values, but we also identify current limitations that stand in the way of more formal implementation of this approach. We close with a discussion of which current models have the best capacity for this purpose in the Baltic Sea, and of the way forward towards the combination of measurable indicators and modelling approaches in food web assessments.

Reproductive rate of grey seal (Halichoerus grypus) females has fluctuated during the 2000s, although reproductive disturbances, which occurred earlier, are rare at present. Here we aimed to study especially the food web factors possibly affecting the birth rate of Baltic grey seals and whether birth rate can be used as an indicator of the changes in the Baltic food web. Our results showed that the birth rate of grey seals was significantly related to herring (Clupea harengus membras) and sprat (Sprattus sprattus) quality (weight) which, in turn were influenced by sprat and cod (Gadus morhua) abundance, as well as zooplankton biomass and plankter size. This suggests strong trophic coupling over three trophic levels. We thus conclude that the birth rate of grey seals can be used as an indicator of the status of the Baltic food web. Based on this, we suggest a threshold value for good food web status for a stable, non-growing seal population. Highlights • Birth rate of grey seals was related to changes in the quality of clupeids. • The quality of clupeids reflected changes in the Baltic food web. • Birth rate can thus be used as an indicator of the status of the Baltic food web. • Different threshold values of birth rate for a growing and stable seal population should be determined.

Marine ecosystems are under high demand for human use, giving concerns about how pressures from human activities may affect their structure, function, and status. In Europe, recent developments in mapping of marine habitats and human activities now enable a coherent spatial evaluation of potential combined effects of human activities. Results indicate that combined effects from multiple human pressures are spread to 96% of the European marine area, and more specifically that combined effects from physical disturbance are spread to 86% of the coastal area and 46% of the shelf area. We compare our approach with corresponding assessments at other spatial scales and validate our results with European-scale status assessments for coastal waters. Uncertainties and development points are identified. Still, the results suggest that Europe’s seas are widely disturbed, indicating potential discrepancy between ambitions for Blue Growth and the objective of achieving good environmental status within the Marine Strategy Framework Directive.

The EU Water Framework and Marine Strategy Framework Directives require that macroalgae are used in status assessments of marine areas. Thus, practical macroalgal indicators are needed. We tested the functionality of cumulative algal cover, covers of late-successional and opportunistic algae, and fraction of opportunistic algae as indicators of eutrophication in the brackish northern Baltic Sea. We found a positive response of cumulative algal cover to increasing Secchi depth that accords with results from earlier studies. However, the increase in cumulative cover was mainly due to an increase in the cover of opportunistic algae, usually considered as a sign of proceeding eutrophication. Therefore, the use of increasing cumulative cover of algae as an indicator of a better status of the marine area is not supported. The effects of increasing Secchi depth on late-successional algae showed positive responses to Secchi depth only at deeper strata, suggesting suitability of deeper areas for monitoring, if late-successional algal cover is used as an indicator. Nitrogen concentration had no effects on algal cover, but decreased the fraction of opportunistics. In general, the fraction of opportunistics was high. Also environmental factors not related to eutrophication had strong effects on macroalgal indicators: the algal cover (both late-successionals and opportunistics) decreased clearly with depth, increased with exposure and we also found a latitudinal gradient within our study area, with increasing algal covers towards higher latitudes. More studies on species responses to eutrophication, enabling a more precise division to eutrophication sensitive vs. tolerant species, could improve the indicators tested.

Eutrophication is a serious threat to aquatic ecosystems globally with pronounced negative effects in the Baltic and other semi-enclosed estuaries and regional seas, where algal growth associated with excess nutrients causes widespread oxygen free “dead zones” and other threats to sustainability. Decades of policy initiatives to reduce external (land-based and atmospheric) nutrient loads have so far failed to control Baltic Sea eutrophication, which is compounded by significant internal release of legacy phosphorus (P) and biological nitrogen (N) fixation. Farming and harvesting of the native mussel species (Mytilus edulis/trossulus) is a promising internal measure for eutrophication control in the brackish Baltic Sea. Mussels from the more saline outer Baltic had higher N and P content than those from either the inner or central Baltic. Despite their relatively low nutrient content, harvesting farmed mussels from the central Baltic can be a cost-effective complement to land-based measures needed to reach eutrophication status targets and is an important contributor to circularity. Cost effectiveness of nutrient removal is more dependent on farm type than mussel nutrient content, suggesting the need for additional development of farm technology. Furthermore, current regulations are not sufficiently conducive to implementation of internal measures, and may constitute a bottleneck for reaching eutrophication status targets in the Baltic Sea and elsewhere. Highlights • Mussel farming is a viable internal measure to address Baltic Sea eutrophication. • Rates of nutrient removal depend on salinity at the regional scale and food availability at the local scale. • Cost effectiveness of nutrient removal by mussel farming depends also on farm type. • Total farm area needed for achieving HELCOM nutrient reduction targets is realistic.

Contamination status of coastal areas of Finland (northern Baltic Sea) markedly affected by anthropogenic activities (harbours, shipyards and maritime activity, industry, municipal and agricultural inputs, legacy contamination) was assessed for the first time using the weight of evidence (WOE) approach. The key element of the study was the caging (transplantation) of Baltic mussels (Mytilus trossulus) for the measurement of tissue accumulation of polycyclic aromatic hydrocarbons (PAHs) and applying a suite of biomarkers of biological effects of contaminants. Additional variables included in the assessment were trace metals in seawater, macrozoobenthos, near-bottom oxygen levels and eutrophication indicators. The chemical parameters were supported by passive sampling of PAHs and organotins at the study sites. The integrated approach combining all the line of evidence (LOE) variables into the WOE showed separation of some sites as more affected by hazardous substances than others, with the most contaminated areas found around harbour and ship yard areas. The contaminant levels measured in the different matrices were not alarmingly high at none of the areas compared to many other areas within or outside the Baltic Sea under more heavy anthropogenic impact, rarely exceeding any given threshold values for Good Environmental Status of the EU Marine Strategy Framework Directive. However, significant biological effects were recorded in mussels in the most contaminated sites, signifying that the combined effects caused by the contaminants and other environmental factors are disturbing the health of marine organisms in the area. The results of this successful combined application based on the mussel transplantation method and the WOE approach are highly encouraging for further trials in developing the monitoring of chemical contamination in the Baltic Sea.

Ecological patterns are inherently scale-dependent and driven by the interplay of abiotic gradients and biotic processes. Despite the fundamental importance of such gradients, there are many gaps in our understanding of how abiotic stress gradients interplay with biotic processes and how these collectively affect species distributions. Using a hierarchical design, we sampled two communities separated by depth along wave exposure and salinity gradients to elucidate how these two gradients affect species composition in habitats formed by the foundation species Mytilus trossulus and Fucus vesiculosus. Specifically, we looked at the impacts of regional salinity and temperature, local wave exposure, and site-dependent facilitation effects on the associated community composition. Wave exposure was the best predictor for species assembly structure, which was also affected by Mytilus biomass and by salinity and water temperature. While the tested variables provided robust explanations for community structure and density, they did not provide conclusive explanations for variation in species richness or evenness. Mytilus biomass had a stronger effect on the associated community with increasing wave exposure at the deeper depth, but the patterns were less obvious at the shallower depth. The latter was also the case for Fucus. These findings comply partly with theoretical predictions suggesting stronger facilitation effects in physically harsh environments. Our results indicate that environmental drivers are the main structuring forces that affect species assembly structure, but also foundation species are important. Thus, predicting changes in species distributions and biodiversity requires the simultaneous consideration of environmental gradients, as well as the structure and composition of foundation species and the interplay between these factors. This work advances our understanding of the processes that modulate species distributions in a marginal marine area and broadens the knowledge of how biological and environmental factors interplay and have an influence on hard-bottom community structure in brackish water seas. peerReviewed

Efforts to attain good environmental status in the marine realm require decisions which cannot be done without knowledge of effects of different management measures. Given the wide diversity of marine ecosystems, multitude of pressures affecting it and the still poor understanding on linkages between those, there are likely no models available to give all the required answers. Hence, several separate approaches can be used in parallel to give support for management measures. We tested three completely different methods – a spatial impact index, a food web model and a Bayesian expert method. We found that a large uncertainty existed regarding the ecosystem response to the management scenarios, and that the three different modelling approaches complemented each other. The models indicated that in order to reach an improved overall state of the ecosystem nutrient reductions are the more effective of the two management variables explored, and that cumulative effects of the management of nutrient inputs and fishing mortality are likely to exist.

We studied the spatio-temporal impacts of physical and chemical environmental variables (depth, sediment type, salinity, temperature, oxygen, total phosphorus, and total nitrogen) on soft-sediment zoobenthos in the open coastal Gulf of Finland during 2001–2015. The study included 55 sampling-stations covering the east-west gradient of the Finnish coastal zone. The chosen environmental variables significantly influenced the distribution of the species in space and over time. Some zoobenthic taxa formed assemblages with each other, occurring in similar environmental conditions, while Gammarus spp. and Chironomidae clearly differed from other taxa in regards to ecological requirements. We showed the critical influence of oxygen (normoxia, hypoxia, anoxia) on individual species, some better adapted to low oxygen conditions (e.g., Chironomidae) than others (e.g., Monoporeia affinis). The nutrient concentrations in the surface sediment also significantly affected the benthic assemblage patterns. The number of species in space and time increased with increasing oxygen concentrations. This study clearly shows that in order to maintain healthy marine communities, it is essential to counteract excess nutrient inputs and their indirect effects on sufficient O2 conditions for the benthic habitats.

The representativeness of aquatic ecosystem monitoring and the precision of the assessment results are of high importance when implementing the EU’s Water Framework Directive that aims to secure a good status of waterbodies in Europe. However, adapting monitoring designs to answer the objectives and allocating the sampling resources effectively are seldom practiced. Here, we present a practical solution how the sampling effort could be re-allocated without decreasing the precision and confidence of status class assignment. For demonstrating this, we used a large data set of 272 intensively monitored Finnish lake, coastal, and river waterbodies utilizing an existing framework for quantifying the uncertainties in the status class estimation. We estimated the temporal and spatial variance components, as well as the effect of sampling allocation to the precision and confidence of chlorophyll-a and total phosphorus. Our results suggest that almost 70% of the lake and coastal waterbodies, and 27% of the river waterbodies, were classified without sufficient confidence in these variables. On the other hand, many of the waterbodies produced unnecessary precise metric means. Thus, reallocation of sampling effort is needed. Our results show that, even though the studied variables are among the most monitored status metrics, the unexplained variation is still high. Combining multiple data sets and using fixed covariates would improve the modeling performance. Our study highlights that ongoing monitoring programs should be evaluated more systematically, and the information from the statistical uncertainty analysis should be brought concretely to the decision-making process. peerReviewed

Oceans around the world are threatened by human pressures. Ecological indicators are useful tools in understanding complex systems and their changes caused by human pressures, and the information they offer is also needed for ecosystem-based management. Integrated assessments combine information produced by several indicators at different spatial scales and thus offer a more holistic view of the status of the ecosystem. In this study, we evaluate the integration of biodiversity indicators at different spatial scales in two study areas in the Baltic Sea: Gulf of Finland and Bothnian Sea. By producing time series of the indicators and integrated assessments, we study the historical changes in the overall marine biodiversity status, and the impact of data availability, indicator selection, and choice of spatial assessment units on the status assessment. The integrated assessments are produced using the Biodiversity Assessment Tool (BEAT 3.0) and following the procedure of the HELCOM integrated assessment of biodiversity. The analysis shows that the results of the integrated assessment depend strongly on which indicators are available for the assessment, and on the chosen spatial assessment units. While the integrated assessments are a strong communication tool, their interpretation needs to be accompanied by information of indicators to avoid misleading conclusions about the marine ecosystem status.

The European Marine Strategy Framework Directive requires the development of suitable indicators for regular reporting on the environmental state and achievement of a good environmental status of EU’s marine waters by 2020. The development of indicators for determining seafloor integrity and its possible disturbance by human activities have so far largely ignored the geological properties of seafloor. This paper presents a study of Vuosaari and Uusikaupunki-D offshore dumping sites in Finland, the northern Baltic Sea. Full coverage multibeam bathymetry and relative backscatter data, and a number of sediment cores were collected over the sites. The areas covered by dumped dredge spoil stand out in the multibeam images because of their irregular surface and elevated backscatter. The short gravity cores were studied for lithology, and in 1-cm slices for 137Cs activity, organic content, and grain size distribution. The dumped material is represented in the cores by the gravelly mud lithofacies with massive texture and angular coarse particles. The dumped material is coarser, less sorted and has higher kurtosis compared to natural sediment due to the admixing of blasted rock during the dredging activities, and limited sorting during fall through the water column upon dumping. Dispersed dredge spoil, which was suspended in the water column during the dumping activities or reworked from the dumped material mounds and redistributed along the seafloor soon thereafter, was deposited over a wide area as a thin layer that is not necessarily readily identifiable by visual inspection in the cores. Cesium activity helped distinguish the dumped material from the 137Cs-enriched natural sediments deposited after the 1986 Chernobyl disaster. Considering that the dumped material at many of the coring sites in the Vuosaari dumping area is covered by natural sediment, it probably is largely stable. In contrast, dumped material at the shallower Uusikaupunki-D site has slumped down to an adjacent channel and is likely being redistributed by near-bottom currents. Based on the findings of the study, a protocol for the assessment of the geological integrity of seafloor, its anthropogenic change due to dumping, and its potential recovery is proposed, as required by the Marine Strategy Framework Directive.

Evaluating the state of benthic communities has played an important role in water quality assessments. Indices incorporating species sensitivities, richness and densities are commonly applied. In Europe, the importance of benthic indices has increased in the last years with the implementation of the European Marine Strategy Framework Directive (MSFD) which at the same time demands the applicability of an index across regional scales. To date, environmental variability is rarely considered in benthic indices and most sensitivity rankings have the disadvantages of static values (i.e. the same value in all areas), expert judgement and a limited geographical range. This study presents species sensitivity values calculated along environmental gradients for the Baltic Sea. Sensitivities were calculated according to the procedure of the Benthic Quality Index (BQI). We created a matrix of subregions, classes of salinity, depth and gear to identify comparable subsets for data analysis. Altogether, 19 subsets were defined within the Baltic Sea basins. Sensitivity values were calculated for 329 species out of a total of 678 species that were recorded in this study. Sensitivity values of taxa vary between subsets as it was expected for different environmental conditions. Most sensitivity values can be assigned to species occurring in euhaline and polyhaline waters. Distribution of species with high and low sensitivity values differed along the salinity gradient. In euhaline waters more species with high sensitivity values occurred than species with low sensitivity values, while in mesohaline waters the ratio of high and low sensitivity values among species was almost equal. In oligohaline waters more species with lower sensitivity values were present. For the first time, sensitivity values were calculated for a large number of species using the same method for the entire Baltic Sea. This results in a Baltic-wide comprehensive set of sensitivity values based on a dataset across subregional borders, and divided along environmental gradients and gear type. The same principles can be applied to transient waters from rivers to coastal lagoons as well as to other environments with gradients of, e.g. hydrodynamic characteristics. Publicly available sensitivity values will increase transparency and support the improvement of state assessments under the MSFD.

Using remote sensing methods to identify valuable underwater habitats Seagrasses are submerged plants found all over the world from brackish bays to salty continental shelves. Seagrasses create high-productive habitats and are a vital part of the marine ecosystems, providing shelter, nurseries, food, and habitats for other species. Seagrasses can be seen as indicators of environmental changes and ecosystem heath as they are sensitive to water quality. Accurate and detailed spatial information of seagrasses would be important in assessments of threatened habitats, establishing MPAs and evaluating ecosystem state. However, spatially comprehensive species information is usually lacking and the knowledge of seagrass habitats around the world relies on scarce field observations and models based on inventory records. Because field work is expensive and time consuming, alternative ways of acquiring information of seagrass habitats are needed. Therefore, aerial and satellite images are useful as they are provided in different resolution, times, and prices depending on the purpose of use. We tested how aerial and high-resolution satellite images could be utilized in finding seagrass habitats in two different environments; in the brackish Finnish coast and in the clear tropical waters of Zanzibar, Tanzania. We did supervised classification to identify seagrasses of pre-processed aerial and satellite images. For the satellite image of the coast of Zanzibar, we also calculated water column correction. Both ways of the method worked, as the seagrass areas were found, although in the Finnish coast poor water visibility hinders light penetration below 4 meter. In developing countries, where marine inventories are usually non-existent, our methodology of utilizing low-cost images for identifying valuable habitats is the only reasonable way of acquiring marine biodiversity data.

The Baltic Sea suffers from extensive blooms of the toxic cyanobacteria Nodularia spumigena that produces nodularin toxin (NOD). Additionally, intensification of oil transportation and related activities in the area increase the risk of oil spills. The current experiment was designed to mimic a situation where an oil spill occurs during a cyanobacterial bloom by exposing the amphipod Gammarus oceanicus to a NOD-rich cyanobacterial extract and the polycyclic aromatic hydrocarbon compound benzo[a]pyrene (B[a]P), a common constituent of oil. The activity of the antioxidant enzymes glutathione S-transferase (GST), glutathione peroxidase (GPx), catalase (CAT) and superoxide dismutase (SOD) were examined after 48 and 96 h of exposure. Exposure to low and high levels of the NOD-rich extract produced a time-dependent activation of GST, GPx and SOD. CAT levels were elevated only by high NOD treatment. Also the toxicity of B[a]P was indicated by significantly elevated antioxidant response. In the combined exposures treatment-dependent additive increases in the activity of GPx and SOD were observed as well as inhibitory (antagonistic) effects on GST, CAT and GPx. Rapid concentration-dependent accumulation of NOD by G. oceanicus was observed. The addition of B[a]P reduced the accumulation of NOD and resulted in different biomarker response patterns compared to single exposures demonstrating the effects of mixture toxicity.

Ever more extensive use of marine space by human activities and greater demands for marine natural resources has led to increases in both duration and spatial extent of pressures on the marine environment. In parallel, the global crisis of decreasing biodiversity and loss of habitats has revitalized scientific research on human impacts and lead to methodological development of cumulative pressure and impact assessments (CPIA). In Europe alone, almost 20 CPIAs have been published in the past 10 years and some more in other sea regions of the world. In this review, we have analyzed 40 recent marine CPIAs and focused on their methodological approaches. We were especially interested in uncovering methodological similarities, identifying best practices and analysing whether the CPIAs have addressed the recent criticism. The review results showed surprisingly similar methodological approaches in half of the studies, raising hopes for finding coherence in international assessment efforts. Although the CPIA methods showed relatively few innovative approaches for addressing the major caveats of previous CPIAs, the most recent studies indicate that improved approaches may be soon found.

Traditional and emerging human activities are increasingly putting pressures on marine ecosystems and impacting their ability to sustain ecological and human communities. To evaluate the health status of marine ecosystems we need a science-based, integrated Ecosystem Approach, that incorporates knowledge of ecosystem function and services provided that can be used to track how management decisions change the health of marine ecosystems. Although many methods have been developed to assess the status of single components of the ecosystem, few exist for assessing multiple ecosystem components in a holistic way. To undertake such an integrative assessment, it is necessary to understand the response of marine systems to human pressures. Hence, innovative monitoring is needed to obtain data to determine the health of large marine areas, and in an holistic way. Here we review five existing methods that address both of these needs (monitoring and assessment): the Ecosystem Health Assessment Tool; a method for the Marine Strategy Framework Directive in the Bay of Biscay; the Ocean Health Index (OHI); the Marine Biodiversity Assessment Tool, and the Nested Environmental status Assessment Tool. We have highlighted their main characteristics and analyzing their commonalities and differences, in terms of: use of the Ecosystem Approach; inclusion of multiple components in the assessment; use of reference conditions; use of integrative assessments; use of a range of values to capture the status; weighting ecosystem components when integrating; determine the uncertainty; ensure spatial and temporal comparability; use of robust monitoring approaches, and address pressures and impacts. Ultimately, for any ecosystem assessment to be effective it needs to be: transparent and repeatable and, in order to inform marine management, the results should be easy to communicate to wide audiences, including scientists, managers, and policymakers.

The eutrophication status of the entire Baltic Sea is classified using a multi-metric indicator- based assessment tool. A total of 189 areas are assessed using indicators where information on reference conditions (RefCon), and acceptable devi- ation (AcDev) from reference condition could be combined with national monitoring data from the period 2001-2006. Most areas (176) are classified as 'affected by eutrophication' and only two open water areas and 11 coastal areas are classified as 'unaf- fected by eutrophication'. The classification is made by application of the recently developed HELCOM Eutrophication Assessment Tool (HEAT), which is described in this paper. The use of harmonized

Closely related species may occupy very similar niches but are often found to diverge by one or more traits when they share the same habitat. Five indigenous and sympatric Gammarus species are characteristic for the Baltic rocky littoral ecosystem. Yet, the species-specific distribution of these sympatric gammarids has not been well studied in the northern Baltic Sea. This study was undertaken to assess the spatial distribution of gammarid amphipods along wave exposure and depth gradients to study whether they show segregation in their microhabitat use. We sampled 12 rocky sublittoral shores along a wave exposure gradient over a period of 5 years. Samples differed with respect to depth and macroalgal type. Three of the five gammarid species occurred mainly in different depth zones and among different macroalgae at the exposed shores. In contrast, on protected shores, where algal zonation is weaker, a link to zonation and macroalgal type was almost absent. Moreover, the microhabitat use was strongest during the reproductive seasons of the species. The observed microhabitats of the three gammarid species fit well to their species-specific mean body sizes.

Coastal eutrophication has resulted in the reduced abundance of the perennial macroalga Fucus vesiculosus L. due to negative effects of increased turbidity, spatial competition, and grazing. We conducted a factorial field experiment to explore the main and interactive effects of three factors on the recruitment success of F. vesiculosus. We manipulated (1) substrate occupancy by offering unoccupied substrates and substrates occupied by a natural assembly of macroalgal spores, periphyton and loose material, (2) nutrient availability by exposing substrates to ambient and enriched nutrient conditions, and (3) herbivory by exclosing the natural herbivore community and enclosing either small (Hydrobia spp.) or large (Theodoxus fluviatilis L.) snails. Microrecruit density of F. vesiculosus depended on the density of parent algae in ambient nutrient availability, whereas, under enriched conditions, such dependency disappeared. Microrecruit density was lower on previously occupied than on unoccupied substrates and at enriched rather than ambient nutrient levels. The lowest recruitment success resulted from a combination of these two treatment levels. The microrecruit density of F. vesiculosus decreased sharply with the density of filamentous algae indicating that competition with opportunistic species contributed to poor recruitment success. Microrecruit density did not differ between the grazer exclosure and the natural grazer community treatment level reflecting locally low grazer densities and consequent weak grazing pressure in the study site. However, the enclosure of snails at higher densities that still naturally occur in the nearby areas affected the recruitment success of F. vesiculosus. This effect was dependent on nutrient availability; at ambient levels, snail grazers had only a minor negative effect on microrecruit density whereas under enriched conditions, both snail species negatively affected recruitment success. Thus, instead of reducing competition and thereby benefiting the perennial slow-growing F. vesiculosus, negative grazing effects by both snail species increased under nutrient-enriched conditions.

Rocky macroalgal assemblages are typically composed of patches differing in age and species composition and grazing is generally a very important modifier of such assemblages. We hypothesized that patch colonization time determines its algal community and that grazing effects depend on the colonization time and vary with depth. We created patches by placing empty substrates at two sublittoral depths over five consecutive months, manipulated grazer entry and determined the algal species composition in each patch in the next growing season. Distinct algal colonization periods resulted in different algal assemblages. Although algal communities in our study area consist mainly of opportunistic species, thus being highly dynamic, the resulting macroalgal assemblages differed in species richness, diversity, composition, and total biomass even a year after first colonization. Substrates close to the water-surface supported a higher species richness and diversity than those in the deeper littoral. The community characteristics, total density, total biomass and species richness were only slightly, if at all affected by grazing. However, individual algal species or taxa showed varying and even contrasting responses to grazing, often differently between depths and depending on colonization time. In the deeper littoral, but not close to the water surface, grazing increased the density of filamentous brown algae while reducing the green alga Cladophora glomerata. In these taxa, grazing effects were strongest in patches colonized during the early growing season. Grazing at the colonization stage had lasting consequences for the density of several individual species.

Grazing pressure on macroalgae in littoral communities may vary with algal species, the age of an algal individual and grazer identity. Previous studies on alga–grazer interactions have shown that grazer preference for an algal species may release another one from interspecific competition. We measured the impacts of four common grazer taxa and the natural grazer guild on macroalgal communities at both their colonization and adult stages, and compared the impacts to grazer exclosures. The grazer effects were stronger on colonizing than on adult macroalgae; grazers did not reduce the total density of adult algae. Grazers both feed on propagules and indirectly facilitate other algae, depending on the grazer or algal species. Hydrobia species increased the settlement of spores of the red alga Ceramium tenuicorne. Similarly, the gastropod Theodoxus fluviatilis tended to facilitate one crustose algal species, but decreased the propagule density of annual filamentous algae, suggesting a preference for one species to the advantage of another. Effects of crustacean mesograzers on the studied macroalgae were weak. These results indicate that northern Baltic macroalgae are limited to grazing mainly during their colonization stage.

The invasive amphipod Gammarus tigrinus was found for the first time and in abundant populations in shallow-water habitats in coastal areas of the city of Helsinki (Gulf of Finland, Baltic Sea). In summer-autumn 2007 the species occurred at four out of seven study sites, dominating almost exclusively at one site and comprising a progressively increasing share of the local gammarid community at another site with a reducing portion of the native Gammarus zaddachi during the study period. The species did not occur at the least polluted archipelago stations, which are also most exposed to wave action. With only single previous observations of this species in Finland the current findings show that G. tigrinus has now firmly established itself into the northern Baltic littoral ecosystem. As this exceedingly omnivoric species is able to outcompete and replace native herbivorous Gammarus species its environmentally detrimental effects may include lesser consumption of e.g. filamentous macroalgae, which are already highly abundant in the study region due to eutrophication.

We report the development and application of a prototype tool for integrated assessment of chemical status in aquatic environments based on substance- and matrix-specific environmental assessment criteria (thresholds). The Chemical Status Assessment Tool (CHASE) integrates data on hazardous substances in water, sediments and biota as well as bio-effect indicators and is based on a substance- or bio-effect-specific calculation of a 'contamination ratio' being the ratio between an observed concentration and a threshold value. Values1.0 indicate areas potentially 'unaffected', while values1.0 indicate areas potentially 'affected'. These ratios are combined within matrices, i.e. for water, sediment and biota and for biological effects. The overall assessment used a 'one out, all out principle' with regard to each matrix. The CHASE tool was tested in the Baltic Sea and the North Sea in 376 assessment units. In the former, the chemical status was1.0 in practically all areas indicating that all areas assessed were potentially affected. The North Sea included areas classified as unaffected or affected. The CHASE tool can in combination with temporal trend assessments of individual substances be advantageous for use in remedial action plans and, in particular, for the science-based evaluation of the status and for determining which specific substances are responsible for a status as potentially affected.

Cyanobacteria of the Baltic Sea have multiple effects on organisms that influence the food chain dynamics on several trophic levels. Cyanobacteria contain several bioactive compounds, such as alkaloids, peptides, and lipopolysaccharides. A group of nonribosomally produced oligopeptides, namely microcystins and nodularin, are tumor promoters and cause oxidative stress in the affected cells. Zooplankton graze on cyanobacteria, and when ingested, the hepatotoxins (nodularin) decrease the egg production of, for example, copepods. However, the observed effects are very variable, because many crustaceans are tolerant to nodularin and because cyanobacteria may complement the diet of grazers in small amounts. Cyanobacterial toxins are transferred through the food web from one trophic level to another. The transfer rate is relatively low in the pelagic food web, but reduced feeding and growth rates of fish larvae have been observed. In the benthic food web, especially in blue mussels, nodularin concentrations are high, and benthic feeding juvenile flounders have been observed to disappear from bloom areas. In the littoral ecosystem, gammarids have shown increased mortality and weakening of reproductive success under cyanobacterial exposure. In contrast, mysid shrimps seem to be tolerant to cyanobacterial exposure. In fish larvae, detoxication of nodularin poses a metabolic cost that is reflected as decreased growth and condition, which may increase their susceptibility to predation. Cyanobacterial filaments and aggregates also interfere with both hydromechanical and visual feeding of planktivores. The feeding appendages of mysid shrimps may clog, and the filaments interfere with prey detection of pike larvae. On the other hand, a cyanobacterial bloom may provide a refuge for both zooplankton and small fish. As the decaying bloom also provides an ample source of organic carbon and nutrients for the organisms of the microbial loop, the zooplankton species capable of selective feeding may thrive in bloom conditions. Cyanobacteria also compete for nutrients with other primary producers and change the nitrogen (N): phosphorus (P) balance of their environment by their N-fixation. Further, the bioactive compounds of cyanobacteria directly influence other primary producers, favoring cyanobacteria, chlorophytes, dinoflagellates, and nanoflagellates and inhibiting cryptophytes. As the selective grazers also shift the grazing pressure on other species than cyanobacteria, changes in the structure and functioning of the Baltic Sea communities and ecosystems are likely to occur during the cyanobacterial bloom season.

On marine rocky shores, macroalgal herbivory is often intense, such that the cascading effects of fish predation may contribute to the control of algal communities. To estimate the magnitudes of top–down and bottom–up control on a macroalgal community, we manipulated the access of carnivorous fish to macroalgal colonization substrates, as well as nutrient availability, at two sub-littoral depths. There were three levels of fish manipulation: natural fish community, no fish and the enclosure of one common species, the perch, Perca fluviatilis . We found a clear cascade effect of fish predation on both the total density and several individual species of macroalgae, which was more pronounced in deep than shallow water. The density of the dominant grazers, i.e. snails, increased in nutrient-enriched conditions; perch were inefficient in controlling herbivores, and had therefore no cascading effect on algal densities under such conditions. Although nutrients enhanced the growth of opportunistic algae, herbivores, in the absence of fish, inhibited this response. While algal diversity was higher in shallow than in deep water, the enrichment effect was opposite at the two depths with lowered diversity in the shallows and increased at depth. Our results indicate that fish predation is an efficient regulator of meso-herbivores and that its effect thereby cascades onto the producer trophic level such that both perennial and opportunistic algae benefit from the presence of fish. This cascade effect is probably stronger at depth where predation efficiency is less disturbed by wave motion.

Many studies have tried to explain spatial and temporal variations in biodiversity status of marine areas from a single-issue perspective, such as fishing pressure or coastal pollution, yet most continental seas experience a wide range of human pressures. Cumulative impact assessments have been developed to capture the consequences of multiple stressors for biodiversity, but the ability of these assessments to accurately predict biodiversity status has never been tested or ground-truthed. This relationship has similarly been assumed for the Baltic Sea, especially in areas with impaired status, but has also never been documented. Here we provide a first tentative indication that cumulative human impacts relate to ecosystem condition, i.e. biodiversity status, in the Baltic Sea. Thus, cumulative impact assessments offer a promising tool for informed marine spatial planning, designation of marine protected areas and ecosystem-based management, and may prove useful for setting limits on allowable levels of human impact on ecosystems. Many studies have tried to explain spatial and temporal variations in biodiversity status of marine areas from a single-issue perspective, such as fishing pressure or coastal pollution, yet most continental seas experience a wide range of human pressures. Cumulative impact assessments have been developed to capture the consequences of multiple stressors for biodiversity, but the ability of these assessments to accurately predict biodiversity status has never been tested or ground-truthed. This relationship has similarly been assumed for the Baltic Sea, especially in areas with impaired status, but has also never been documented. Here we provide a first tentative indication that cumulative human impacts relate to ecosystem condition, i.e. biodiversity status, in the Baltic Sea. Thus, cumulative impact assessments offer a promising tool for informed marine spatial planning, designation of marine protected areas and ecosystem-based management, and may prove useful for setting limits on allowable levels of human impact on ecosystems.

Toxic cyanobacteria can have harmful or fatal impacts on aquatic organisms. In the archipelagos of the northern Baltic Sea, the open sea blooms often drift into littoral areas, where they decompose and release toxins and other chemical compounds in the water. However, the effects of cyanobacteria on the littoral organisms have not previously been investigated. We studied the effects of three cyanobacteria species (toxic Nodularia spumigena, non-toxic N. sphaerocarpa and non-toxic Aphanizomenon flos-aquae) and purified dissolved nodularin (produced by N. spumigena) on a common littoral amphipod Gammarus zaddachi. Nodularin was transferred to eggs, juveniles and adults of G. zaddachi, but no significant negative effects of dissolved nodularin were detected on adults, eggs or juveniles. However, survival of adults decreased by the exposure to toxic N. spumigena cells. The egg hatching rate and juvenile survival were not affected when exposed to the three cyanobacteria species. In contrast, a weak decrease in the egg production and an increased abortion of embryos from the brood pouch of females was observed, the later indicating a failure in parental care. Further, a decrease in grazing rate on the filamentous green alga Enteromorpha intestinalis was observed. The results suggest that toxic cyanobacteria blooms are not extremely fatal, but may have, in high concentrations, negative effects on the adult survival, fecundity, and feeding behaviour of gammarids inhabiting the littoral zone.

Integrated assessment of the status of marine biodiversity is and has been problematic compared to, for example, assessments of eutrophication and contamination status, mostly as a consequence of the fact that monitoring of marine habitats, communities and species is expensive, often collected at an incorrect spatial scale and/or poorly integrated with existing marine environmental monitoring efforts. The objective of this Method Paper is to introduce and describe a simple tool for integrated assessment of biodiversity status based on the HELCOM Biodiversity Assessment Tool (BEAT), where interim biodiversity indicators are grouped by themes: broad-scale habitats, communities, and species as well as supporting non-biodiversity indicators. Further, we report the application of an initial indicator-based assessment of biodiversity status of Danish marine waters where we have tentatively classified the biodiversity status of Danish marine waters. The biodiversity status was in no areas classified as “unaffected by human activities.” In all the 22 assessment areas, the status was classified as either “moderately affected by human activities” or “significantly affected by human activities.” Spatial variations in the biodiversity status were in general related to the eutrophication status as well as fishing pressure. &nbsp

The European seas are under anthropogenic pressures impacting the state of water quality, benthic habitats and species. The EU Marine Strategy Framework Directive (MSFD) requires the Member States to assess the impacts of pressures and make a programme of measures leading to good environmental status (GES) by 2020. This study presents a method for assessing the quantity and distribution of anthropogenic impacts on benthic habitats in the Baltic Sea by using spatial data of human pressures and benthic habitats. The southern sub-basins were more extensively impacted than the northern sub-basins. Over the entire sea area, deep sea habitats were more impacted than shallower infralittoral and circalittoral habitats. Sand and coarse sediments were the seabed types relatively most impacted in the Baltic Sea scale. A comparison against tentative thresholds for GES showed that in the sub-basin scale only one third of the habitat types was in GES.

The Baltic Sea countries have been quite successful in preventing major pollution spills from shipping, and establishing a system to monitor ship traffic and detect illegal oil spills using aerial and satellite surveillance. The regional cooperation is carried out in the framework of HELCOM, an intergovernmental organization of the nine coastal states and the European Union. Ships operating in the Baltic Sea have to follow strict global and regional anti-discharge regulations, and the number of illegal, deliberate oil discharges has decreased since 1980s substantially. While the risk of large accidental spills is constantly present, requiring that sufficient response capacities are available in the region, smaller oil discharges also pose a threat to, and have an impact on the marine environment of the Baltic Sea.

The EU Marine Strategy Framework Directive requires Member States to estimate the level of human impacts on their marine waters. We report the first attempt to quantify the magnitude and distribution of cumulative impacts of anthropogenic pressures for an entire regional sea, the Baltic Sea. We used a method which takes account of the sensitivity of different ecosystem components and gives scores for potential impacts in 5 km × 5 km areas. Our quantification of impacts was based on data layers of anthropogenic pressures and ecosystem components. The classification of the anthropogenic pressures follows the MSFD and the outcome of the index was targeted to facilitate the implementation of the directive. The study presents the cumulative impacts over the entire sea area and shows that the highest estimated impacts were in the southern and south-western sea areas and in the Gulf of Finland. The lowest index values were found in the Gulf of Bothnia. The results coincide with the population densities of the adjacent catchment areas. Fishing, inputs of nutrients and organic matter and inputs of hazardous substances comprised 25%, 30% and 30%, respectively, of the total cumulative impact. The approach used is transparent and the results are useful in regard to ecosystem-based management, e.g. for area-based management and assessments. Examples of uses are given together with analysis of the strengths and weaknesses of the approach.

The brown alga Fucus vesiculosus is a foundation species in the Baltic Sea littoral, hosting a rich faunal community. We compared the species composition and diversity of invertebrate macrofauna living on F. vesiculosus between sites differing in their eutrophication status and exposure to waves at three different times during a season. We determined the size, nitrogen and phlorotannin content of the alga. The invertebrate community differed substantially between sites near fish farms and those in more pristine environment. Snails and bivalves were more abundant on the Fucus stands near fish farms than on control stands, where crustaceans were more abundant. The abundance of molluscs decreased with the increasing shore exposure, while gammaridean amphipods dominated on the exposed shores. Abundance of several taxa increased during the proceeding growing season. The density of the most important herbivore of F. vesiculosus, Idotea balthica, varied 100-fold during the season being the lowest in June and the highest in August when the generation born in the summer started to feed on Fucus. Thus, the diversity and composition of Fucus-associated invertebrate fauna varies both with environmental conditions of the stand and seasonally. Although the negative effects of eutrophication on distribution and abundance of Fucus stands are well documented, a moderate increase of nutrients was found to increase the species richness of Fucus-associated fauna in early summer.

We studied the interacting roles of nutrient availability and herbivory in determining the macroalgal community in a rocky littoral environment. We conducted a factorial field experiment where we manipulated nutrient levels and herbivory at two sublittoral depths and measured macroalgal colonization and the following young assemblage during the growing season. At the community level, grazing reduced algal colonization, though the effect varied with depth and its interaction with nutrient availability varied in time. In shallow water, the total density of macroalgae increased in response to nutrient enrichment, but the ability of grazers to reduce macroalgal density also increased with the nutrient enrichment, and thus, the community could not escape from the top-down control. In deep water, the algal density was lower, except in July when nutrient enrichment caused a very dense algal growth. Grazing at the greater depth, though effective, was generally of smaller magnitude, and in July it could not limit algal recruitment and growth. Species richness peaked at the intermediate nutrient level in deep but not in shallow water during most of the growing season. Grazing had no effect on diversity of the algal community at either depth and only a minor effect on species richness at the greater depth. Opportunistic and ephemeral algae benefited from the nutrient enrichment but were also grazed to very low densities. Slowly growing and/or perennial species colonized poorly in the nutrient enriched treatments, and depending on the species, either suffered or indirectly benefited from herbivory. For all species, effects of nutrients on colonization depended on depth; usually both nutrient and herbivory effects were more pronounced at the shallow depth. We conclude that grazers are able to reduce macroalgae over a large range of nutrient availabilities, up to 12-fold nutrient enrichment in the current experiment, and that the sublittoral depth gradient generates variation in the algal community control exerted by both herbivory and nutrient availability. Thus temporal and spatial variability in both top-down and bottom-up control and in their interaction, especially along the depth gradient, may be crucially important for producer diversity and for the successional dynamic in a rocky sublittoral environment.

In diverse littoral communities, biotic interactions play an important role in community regulation. This article reviews how eutrophication modifies biotic interactions in littoral macroalgal communities. Eutrophication causes blooms of opportunistic algae, increases epibiotism, and affects regulation by grazers. Opportunistic algae and epibionts harm colonization and growth of perennial algae. Grazing regulates the density and species composition of macroalgal communities, especially at the early stage of algal colonization. Eutrophication supports higher grazer densities by increasing the availability and quality of algae to grazers. This may, on the one hand, enhance the capability of grazers to regulate and counteract the increase of harmful, bloom-forming macroalgae; on the other hand, it may increase grazing pressure on perennial species, with a poor tolerance of grazing. In highly eutrophic conditions, bloom-forming algae may also escape grazing control and accumulate. Increasing epibiotism and grazing threaten in particular the persistence of habitat-forming perennials such as the bladderwrack. An interesting property of biotic interactions is that they do not remain fixed but are able to evolve, as the traits of the interacting species adapt to each other and to abiotic conditions. The potential of plants and grazers to adapt is crucial to their chances to survive in changing environment.