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.

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Nitrogen (N) fixation is fueling planktonic production in a multitude of aquatic environments. In meso-and poly-haline estuaries, however, the contribution of N by pelagic N 2 fixation is believed to be insignificant due to the high input of N from land and the presumed absence of active N 2 -fixing organisms. Here we report N 2 fixation rates, nifH gene composition and nifH gene transcript abundance for key diazotrophic groups over 1 year in two contrasting, temperate, estuarine systems: Roskilde Fjord (RF) and the Great Belt (GB) strait. Annual pelagic N 2 fixation rates averaged 17 and 61 mmol N m À 2 per year at the two sites, respectively. In RF, N 2 fixation was mainly accompanied by transcripts related to heterotrophic (for example, Pseudomonas sp.) and photoheterotrophic bacteria (for example, unicellular diazotrophic cyanobacteria group A). In the GB, the first of two N 2 fixation peaks coincided with a similar nifH-expressing community as in RF, whereas the second peak was synchronous with increased nifH expression by an array of diazotrophs, including heterotrophic organisms as well as the heterocystous cyanobacterium Anabaena. Thus, we show for the first time that significant planktonic N 2 fixation takes place in mesohaline, temperate estuaries and that the importance of heterotrophic, photoheterotrophic and photosynthetic diazotrophs is clearly variable in space and time.

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