Abstract. A 38-year record of bottom-water dissolved oxygen concentrations in coastal marine ecosystems around Denmark (1965Denmark ( -2003 and a longer, partially reconstructed record of total nitrogen (TN) inputs were assembled with the purpose of describing longterm patterns in hypoxia and anoxia. In addition, interannual variations in bottom-water oxygen concentrations were analyzed in relation to various explanatory variables (bottom temperature, wind speed, advective transport, TN loading). Reconstructed TN loads peaked in the 1980s, with a gradual decline to the present, commensurate with a legislated nutrient reduction strategy. Mean bottom-water oxygen concentrations during summer have significantly declined in coastal marine ecosystems, decreasing substantially during the 1980s and were extremely variable thereafter. Despite decreasing TN loads, the worst hypoxic event ever recorded in open waters occurred in 2002. For estuaries and coastal areas, bottomwater oxygen concentrations were best described by TN input from land and wind speed in July-September, explaining 52% of the interannual variation in concentrations. For open sea areas, bottom-water oxygen concentrations were also modulated by TN input from land; however, additional significant variables included advective transport of water and Skagerrak surface-water temperature and explained 49% of interannual variations in concentrations. Reductions in the number of benthic species and alpha diversity were significantly related to the duration of the 2002 hypoxic event. Gradual decreases in diversity measures (number of species and alpha diversity) over the first 2-4 weeks show that the benthic community undergoes significant changes before the duration of hypoxia is severe enough to cause the community to collapse. Enhanced sediment-water fluxes of NH 4 þ and PO 4 3À occur with hypoxia, increasing nutrient concentrations in the water column and stimulating additional phytoplankton production. Repeated hypoxic events have changed the character of benthic communities and how organic matter is processed in sediments. Our data suggest that repeated hypoxic events lead to an increase in susceptibility of Danish waters to eutrophication and further hypoxia.

Much of the Baltic Sea is currently classified as 'affected by eutrophication'. The causes for this are twofold. First, current levels of nutrient inputs (nitrogen and phosphorus) from human activities exceed the natural processing capacity with an accumulation of nutrients in the Baltic Sea over the last 50-100 years. Secondly, the Baltic Sea is naturally susceptible to nutrient enrichment due to a combination of long retention times and stratification restricting ventilation of deep waters. Here, based on a unique data set collated from research activities and long-term monitoring programs, we report on the temporal and spatial trends of eutrophication status for the open Baltic Sea over a 112-year period using the HELCOM Eutrophication Assessment Tool (HEAT 3.0). Further, we analyse variation in the confidence of the eutrophication status assessment based on a systematic quantitative approach using coefficients of variation in the observations. The classifications in our assessment indicate that the first signs of eutrophication emerged in the mid-1950s and the central parts of the Baltic Sea changed from being unaffected by eutrophication to being affected. We document improvements in eutrophication status that are direct consequences of long-term efforts to reduce the inputs of nutrients. The reductions in both nitrogen and phosphorus loads have led to large-scale alleviation of eutrophication and to a healthier Baltic Sea. Reduced confidence in our assessment is seen more recently due to reductions in the scope of monitoring programs. Our study sets a baseline for implementation of the ecosystem-based management strategies and policies currently in place including the EU Marine Strategy Framework Directives and the HELCOM Baltic Sea Action Plan.

In the 1980s, Danish coastal waters suffered from eutrophication and several nutrient management plans have been implemented during the years to improve ecological status. This study aims at giving a holistic ecosystem perspective on 25 years of mitigation measures. We report trends of nutrient inputs and the responses to these in various chemical and biological components. Nutrient inputs from land were reduced by~50 % for nitrogen (N) and 56 % for phosphorus (P) since 1990. These reductions resulted in significant and parallel declines in nutrient concentrations, and initiated a shift in the dominance of primary producers towards reduced phytoplankton biomass (chlorophyll a concentration) and increased cover of macroalgae in deeper waters. In the last 5 years, eelgrass meadows have also expanded towards deeper waters, in response to improving water clarity. An expected improvement of bottom water oxygen conditions has not been observed, presumably because more frequent stratification and higher water temperatures have counteracted the expected positive effects of reduced nutrient inputs. The biomass of the benthic macrofauna decreased as expected, but it was composed of a drastic decline of filter feeders paralleled by a more moderate increase of deposit feeders. This shift was most likely induced by increasing stratification. The reduced benthic filtration along with the limited eelgrass cover probably kept relatively more particles in suspension, which can explain why improvements in the Secchi depths were modest. Overall, several ecosystem components demonstrated clear signs of improvement, suggesting that at least partial recovery is attainable. On this basis, we propose a conceptual scheme for recovery of shallow coastal ecosystems following marked reductions in nutrient inputs.

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Ecosystem functioning is threatened by an increasing number of anthropogenic stressors, creating a legacy of disturbance that undermines ecosystem resilience. However, few empirical studies have assessed to what extent an ecosystem can tolerate repeated disturbances and sustain its multiple functions. By inducing increasingly recurring hypoxic disturbances to a sedimentary ecosystem, we show that the majority of individual ecosystem functions experience gradual degradation patterns in response to repetitive pulse disturbances. The degradation in overall ecosystem functioning was, however, evident at an earlier stage than for single ecosystem functions and was induced after a short pulse of hypoxia (i.e., three days), which likely reduced ecosystem resistance to further hypoxic perturbations. The increasing number of repeated pulse disturbances gradually moved the system closer to a press response. In addition to the disturbance regime, the changes in benthic trait composition as well as habitat heterogeneity were important for explaining the variability in overall ecosystem functioning. Our results suggest that disturbance-induced responses across multiple ecosystem functions can serve as a warning signal for losses of the adaptive capacity of an ecosystem, and might at an early stage provide information to managers and policy makers when remediation efforts should be initiated.

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