Environmental abundances of the non-native round goby Neogobius melanostomus influence feeding of native fish predators.

The authors assessed the importance of the round goby Neogobius melanostomus as prey for three native predatory fish species, Atlantic cod Gadus morhua, European perch Perca fluviatilis and northern pike Esox lucius, in a northern and southern area of the Baltic Proper, using a combination of visual analysis and DNA metabarcoding of predator stomach contents. To explore the influence of environmental abundances of N. melanostomus on predation, they related the occurrence of N. melanostomus in predator diets to its abundance in survey fishing. Gadus morhua and E. lucius in the southern area showed the highest tendency to feed on N. melanostomus when it was abundant, as N. melanostomus occurred in up to 100% of stomachs and constituted up to 88% of the total diet volume proportion. The diet contribution of N. melanostomus was associated with N. melanostomus abundances for G. morhua and E. lucius, and when N. melanostomus was abundant, these predators exhibited lower prey richness and a higher degree of piscivory. G. morhua and P. fluviatilis also fed less on crustacean prey when N. melanostomus was abundant. The high importance of N. melanostomus in diets of native fish predators may modify indirect interactions between N. melanostomus and native prey species in invaded coastal communities.

External nutrient loading from land, sea and atmosphere to all 656 Swedish coastal water bodies.

Identifying the main sources of nutrient loading is a key factor for efficient mitigation of eutrophication. This study has investigated the pathways of external nutrient loading to 656 coastal water bodies along the entire Swedish coastline. The studied water bodies have been delineated to meet requirements in the European Union's Water Framework Directive, and recent status assessments have shown that 57% of them fail to attain good or high ecological status with respect to nutrients. The analysis in the study was performed on data from mass-balance based nutrient budgets computed using the modelling framework Vattenwebb. The external nutrient contribution from the sea to the water bodies was highly variable, ranging from about 1% to nearly 100%, but the median contribution was >99% of the total external loading regarding both nitrogen and phosphorus. External loading from the atmosphere and local catchment area played a minor role in general. However, 45 coastal water bodies received >25% of the external nitrogen and phosphorus from their catchments. Loading from land typically peaked in April following ice-break and snow melting and was comparatively low during summer. The results indicate that for many eutrophicated Swedish coastal water bodies, nutrient abatement is likely to be optimally effective when potential measures in all of the catchment area of the concerned sea basin are considered. Local-scale mitigation in single water bodies will likely be locally effective only in the small proportion of areas where water and thereby also nutrient input from the catchment is high compared to the influx from the sea. Future studies should include nutrient reduction scenarios in order to refine these conclusions and to identify relevant spatial scales for coastal eutrophication mitigation measures from a water body perspective.

Factors controlling phosphorus release from sediments in coastal archipelago areas.

In coastal archipelago areas of the northern Baltic Sea, significantly higher phosphate concentrations (6.0±4.5µmol/l, mean±SD) were measured in water samples close to the sediment surface compared with those from 1m above the seafloor (1.6±2.0µmol/l). The results indicated notable phosphate release from sediments under the bottom water oxygen concentrations of up to 250µmol/l, especially in areas that had experienced recent temporal fluctuation between oxic and hypoxic/anoxic conditions. No single factor alone was found to control the elevated PO4-P concentrations in the near-bottom water. In addition to the oxygen in the water, the contents of potentially mobile phosphorus fractions, grain-size, the organic content at the sediment surface, and the water depth were all important factors controlling the internal loading of phosphorus. The complexity of this process needs to be accounted for in assessments of the internal loading of phosphorus and in potential mitigation plans.

Testing the potential for predictive modeling and mapping and extending its use as a tool for evaluating management scenarios and economic valuation in the Baltic Sea (PREHAB).

We evaluated performance of species distribution models for predictive mapping, and how models can be used to integrate human pressures into ecological and economic assessments. A selection of 77 biological variables (species, groups of species, and measures of biodiversity) across the Baltic Sea were modeled. Differences among methods, areas, predictor, and response variables were evaluated. Several methods successfully predicted abundance and occurrence of vegetation, invertebrates, fish, and functional aspects of biodiversity. Depth and substrate were among the most important predictors. Models incorporating water clarity were used to predict increasing cover of the brown alga bladderwrack Fucus vesiculosus and increasing reproduction area of perch Perca fluviatilis, but decreasing reproduction areas for pikeperch Sander lucioperca following successful implementation of the Baltic Sea Action Plan. Despite variability in estimated non-market benefits among countries, such changes were highly valued by citizens in the three Baltic countries investigated. We conclude that predictive models are powerful and useful tools for science-based management of the Baltic Sea.

Pikeperch (Sander lucioperca (L.)) in decline: high mortality of three populations in the northern Baltic Sea.

The development of three pikeperch (Sander lucioperca (L.)) populations in the northern Baltic Sea was monitored using standardized multimesh gillnets in 1995-2009. Declining trends in the abundances of pikeperch over 40 cm total length, low numbers of individuals older than 6 years, and high mortality rates were observed in all three populations. In the site with the largest commercial catches per unit area and a rapidly increased colony of great cormorants (Phalacrocorax carbo sinensis Blumenbach 1798), also the abundance of pikeperch below 40 cm total length and year-class strength showed declining trends. The adverse population level changes did not correlate with changes in water quality or eutrophication status. Together, the results suggest that in all study sites fisheries are harvesting a large proportion of the pikeperch soon after or even before reaching the maturity, and that predation from great cormorants may increase mortality of juveniles. Pikeperch is important not only for fisheries but also for ecosystem functioning, and our results point at the need for further management measures to ensure viable populations in the areas studied.

Differential habitat use and antipredator response of juvenile roach (Rutilus rutilus) to olfactory and visual cues from multiple predators.

The indirect, behavioral effects of predation and predator-predator interactions can significantly alter the trophic ecology of many communities. In numerous instances, the strength of these effects may be determined by the ability of prey to identify predation risk through predator-specific cues and respond accordingly to avoid capture. We exposed juvenile roach (Rutilus rutilus), a common forage fish in many brackish and freshwater environments, to vision and/or olfactory cues from two predators with different hunting methods: northern pike (Esox lucius, an ambush predator) and European perch (Perca fluviatilis, a roving predator). Our results demonstrated that responses of roach to perceived risk (as evidenced by their selection of structured or open-water habitats) were highly dependent on cue type and predator identity. For instance, roach responded to olfactory cues of pike by entering open-water habitat, but entered structured habitat when presented with a vision cue of this predator. Opposite responses were elicited from roach for both olfactory and visual cues of perch. Interestingly, roach defaulted to selection of structured habitat when presented with vision + olfaction cues of either predator. Moreover, when presented individual cues of both predators together, roach responded by choosing open-water habitat. Upon being presented with vision + olfaction cues of both predators, however, roach strongly favored structured habitat. Differences in habitat selection of roach were likely in response to the alternative foraging strategies of the two predators, and suggest that prey species may not always use structured habitats as protection. This appears particularly true when a threat is perceived, but cannot immediately be located. These results provide insight to the complex and variable nature by which prey respond to various cues and predators, and offer a mechanistic guide for how behaviorally mediated and predator-predator interactions act as structuring processes in aquatic systems.

Declines in predatory fish promote bloom-forming macroalgae.

In the Baltic Sea, increased dominance of ephemeral and bloom-forming algae is presently attributed to increased nutrient loads. Simultaneously, coastal predatory fish are in strong decline. Using field data from nine areas covering a 700-km coastline, we examined whether formation of macroalgal blooms could be linked to the composition of the fish community. We then tested whether predator or nutrient availability could explain the field patterns in two small-scale field experiments, by comparing joint effects on algal net production from nutrient enrichment with agricultural fertilizer and exclusion of larger predatory fish with cages. We also manipulated the presence of invertebrate grazers. The abundance of piscivorous fish had a strong negative correlation with the large-scale distribution of bloom-forming macroalgae. Areas with depleted top-predator communities displayed massive increases in their prey, small-bodied fish, and high covers of ephemeral algae. Combining the results from the two experiments showed that excluding larger piscivorous fish: (1) increased the abundance of small-bodied predatory fish; (2) changed the size distribution of the dominating grazers, decreasing the smaller gastropod scrapers; and (3) increased the net production of ephemeral macroalgae. Effects of removing top predators and nutrient enrichment were similar and additive, together increasing the abundance of ephemeral algae many times. Predator effects depended on invertebrate grazers; in the absence of invertebrates there were no significant effects of predator exclusion on algal production. Our results provide strong support for regional declines of larger predatory fish in the Baltic Sea promoting algal production by decreasing invertebrate grazer control. This highlights the importance of trophic interactions for ecosystem responses to eutrophication. The view emerges that to achieve management goals for water quality we need to consider the interplay between top-down and bottom-up processes in future ecosystem management of marine resources.

The relative importance of food and shelter for seagrass-associated invertebrates: a latitudinal comparison of habitat choice by isopod grazers.

The generality of mechanisms affecting habitat choice and grazing in seagrass meadows was evaluated in a latitudinal comparison of seagrass grazers from the temperate (60°N) Baltic Sea and the subtropical (30°N) Gulf of Mexico. Using similar habitat choice experiment set-ups in Finland and the USA, the role of food type, habitat complexity and predation hazard on habitat choice of the isopods Idotea baltica (Pallas) and Erichsonella attenuata Harger were tested. When shelter was provided by both living and artificial seagrass, epiphytic food resources on artificial vegetation were clearly preferred by both species, although Idotea was attracted to epiphyte-free seagrass when no alternative food was present. When choosing between food and shelter, both species preferred epiphytic food over shelter. However, under predation hazard of fish, Erichsonella clearly switched to the habitat offering shelter, while the presence of a predatory fish produced no preference for shelter by Idotea. Food type may be considered as an universal mechanism that partly determines the presence of grazers in seagrass habitats and is, in the absence of a predator, more important than shelter. Predation risk affected the behaviour of the grazers, but the response varied between species possibly due to varying importance of fish predation in the areas studied.