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.

Ljunggren, L., Sandström, A., Bergström, U., Mattila, J., Lappalainen, A., Johansson, G., Sundblad, G., Casini, M., Kaljuste, O., and Eriksson, B. K. 2010. Recruitment failure of coastal predatory fish in the Baltic Sea coincident with an offshore ecosystem regime shift. – ICES Journal of Marine Science, 67: 1587–1595. The dominant coastal predatory fish in the southwestern Baltic Sea, perch and pike, have decreased markedly in abundance during the past decade. An investigation into their recruitment at 135 coastal sites showed that both species suffered from recruitment failures, mainly in open coastal areas. A detailed study of 15 sites showed that areas with recruitment problems were also notable for mortality of early-stage larvae at the onset of exogenous food-intake. At those sites, zooplankton abundance predicted 83 and 34% of the variation in young of the year perch and pike, respectively, suggesting that the declines were caused by recruitment failure attributable to zooplankton food limitation. Incidences of recruitment failure match in time an offshore trophic cascade that generated massive increases in planktivorous sprat and decreases in zooplankton biomass in the early 1990s. Therefore, sprat biomass explained 53% of the variation in perch recruitment from 1994 to 2007 at an open coastal site, where three-spined stickleback also increased exponentially after 2002. The results indicate that the dramatic change in the offshore ecosystem may have propagated to the coast causing declines of the dominating coastal predators perch and pike followed by an increase in the abundance of small-bodied fish.

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In this study, the combined influence of vegetation density and water turbidity on habitat utilization of a prey fish, 0þ year perch Perca fluviatilis, under predation risk (pike, Esox lucius) was investigated. The vegetated habitat was overall preferred over the open habitat in the presence of a predator. The level of turbidity, and to a lesser extent vegetation density, however, influenced the response of 0þ year perch. The use of the vegetated habitat was lower in very turbid than in clear and turbid conditions, suggesting reduced antipredator behaviour in very turbid water. The effect of vegetation density on antipredator behaviour was only present in clear water, where the use of a structural refuge decreased with increasing vegetation density. No such effect was observed in turbid and very turbid water. The results showed that the structuring role of vegetation or habitat complexity may diminish with increased turbidity. The observed masking effect of turbidity suggests that predator-prey interactions in vegetated habitats are more complex than what has generally been thought. # 2004 The Fisheries Society of the British Isles

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.

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.

Substrate spawning fish are believed to be selective in their choice of spawning habitat, yet few studies have shown the relative importance of different characteristics in terms of habitat quality. We used an extensive and detailed dataset to identify the factors that govern both large-scale (10 3 to 10 5 m) and local-scale (10 1 to 10 2 m) selection by a substrate-spawning fish, the Eurasian perch Perca fluviatilis L. Distribution of spawning habitat was strongly dependent on habitat characteristics defined by substrate, wave exposure, temperature and depth. The most important predictor was the type of spawning substrate, which generally consisted of different types of vegetation. Substrates providing rigidity and structural complexity were preferred, despite abundant presence of other substrate types. Shallow depth and sheltered areas were also selected habitat characteristics. The response to temperature was scale-dependent, with a stronger selection expressed at the local scale. The specific selectivity suggests that spawning patterns can be successfully modelled with sufficient detail using only a few fundamental environmental variables. Wave exposure and depth are readily available for large-scale spatial predictions, while temperature and substrate require further development in most coastal areas. The high specificity of the characteristics determining habitat quality suggests that it should be possible to apply this modelling approach for identification and conservation of spawning habitats of Eurasian perch and other substrate-spawning fishes in coastal waters.

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