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.
By mainly targeting larger predatory fish, commercial fisheries have indirectly promoted rapid increases in densities of their prey; smaller predatory fish like sprat, stickleback and gobies. This process, known as mesopredator release, has effectively transformed many marine offshore basins into mesopredator-dominated ecosystems. In this article, we discuss recent indications of trophic cascades on the Atlantic and Baltic coasts of Sweden, where increased abundances of mesopredatory fish are linked to increased nearshore production and biomass of ephemeral algae. Based on synthesis of monitoring data, we suggest that offshore exploitation of larger predatory fish has contributed to the increase in mesopredator fish also along the coasts, with indirect negative effects on important benthic habitats and coastal water quality. The results emphasize the need to rebuild offshore and coastal populations of larger predatory fish to levels where they regain their control over lower trophic levels and important links between offshore and coastal systems are restored.
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.
Spawning habitat of pike (Esox lucius) in the Baltic Sea include brackish water bays, brooks and rivers. Elevated salinity concentrations are one of several stressors that might increase the use and importance of freshwater habitats for spawning. In the Baltic Sea, one of the largest brackish seas in the world, freshwater species like pike, perch (Perca fluviatilis), whitefish (Coregonus sp), bream (Abramis brama), ide (Leuciscus idus), roach (Rutilus rutilus) and burbot (Lota lota) all undertake spawning migrations to freshwater. However, over the last decades populations densities of these species have declined, and recruitment failure has been argued to be at least part of the problem. The importance of brooks and rivers as spawning areas for these species have not been quantified and set in relation to spawning success in brackish bays. In this study, we collected 175 adult pike (Esox lucius) on their foraging grounds in the sea. Fish were collected in two regions on the Baltic coast, more than 600 km apart. Subsequently we determined their origin (freshwater or marine) using otolith chemistry. Sagittal otoliths were analysed for strontium using the PIXE-method. The results show that 80 of the 175 pike were recruited in freshwater, and several of the larger specimens showed reoccurring migration behaviour. Data show that freshwater is an important recruitment habitat for Baltic Sea pike, suggesting that habitat improvements in rivers entering the Baltic Sea might significantly contribute to population restoration.
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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