Hypoxia, a growing worldwide problem, has been intermittently present in the modern Baltic Sea since its formation ca. 8000 cal. yr BP. However, both the spatial extent and intensity of hypoxia have increased with anthropogenic eutrophication due to nutrient inputs. Physical processes, which control stratification and the renewal of oxygen in bottom waters, are important constraints on the formation and maintenance of hypoxia. Climate controlled inflows of saline water from the North Sea through the Danish Straits is a critical controlling factor governing the spatial extent and duration of hypoxia. Hypoxia regulates the biogeochemical cycles of both phosphorus (P) and nitrogen (N) in the water column and sediments. Significant amounts of P are currently released from sediments, an order of magnitude larger than anthropogenic inputs. The Baltic Sea is unique for coastal marine ecosystems experiencing N losses in hypoxic waters below the halocline. Although benthic communities in the Baltic Sea are naturally constrained by salinity gradients, hypoxia has resulted in habitat loss over vast areas and the elimination of benthic fauna, and has severely disrupted benthic food webs. Nutrient load reductions are needed to reduce the extent, severity, and effects of hypoxia.

Science-based, multinational management of the Baltic Sea offers lessons on amelioration of highly disturbed marine ecosystems.

Hypoxia is a well-described phenomenon in the offshore waters of the Baltic Sea with both the spatial extent and intensity of hypoxia known to have increased due to anthropogenic eutrophication, however, an unknown amount of hypoxia is present in the coastal zone. Here we report on the widespread unprecedented occurrence of hypoxia across the coastal zone of the Baltic Sea. We have identified 115 sites that have experienced hypoxia during the period 1955–2009 increasing the global total to ca. 500 sites, with the Baltic Sea coastal zone containing over 20% of all known sites worldwide. Most sites experienced episodic hypoxia, which is a precursor to development of seasonal hypoxia. The Baltic Sea coastal zone displays an alarming trend with hypoxia steadily increasing with time since the 1950s effecting nutrient biogeochemical processes, ecosystem services, and coastal habitat.

Benthic-pelagic coupling is manifested as the exchange of energy, mass, or nutrients between benthic and pelagic habitats. It plays a prominent role in aquatic ecosystems, and it is crucial to functions from nutrient cycling to energy transfer in food webs. Coastal and estuarine ecosystem structure and function are strongly affected by anthropogenic pressures; however, there are large gaps in our understanding of the responses of inorganic nutrient and organic matter fluxes between benthic habitats and the water column. We illustrate the varied nature of physical and biological benthic-pelagic coupling processes and their potential sensitivity to three anthropogenic pressures -climate change, nutrient loading, and fishing -using the Baltic Sea as a case study and summarize current knowledge on the exchange of inorganic nutrients and organic material between habitats. Traditionally measured benthic-pelagic coupling processes (e.g., nutrient exchange and sedimentation of organic material) are to some extent quantifiable, but the magnitude and variability of biological processes are rarely assessed, preventing quantitative comparisons. Changing oxygen conditions will continue to have widespread effects on the processes that govern inorganic and organic matter exchange among habitats while climate change and nutrient load reductions may have large effects on organic matter sedimentation. Many biological processes (predation, bioturbation) are expected to be sensitive to anthropogenic drivers, but the outcomes for ecosystem function are largely unknown. We emphasize how improved empirical and experimental understanding of benthic-pelagic coupling processes and their variability are necessary to inform models that can quantify the feedbacks among processes and ecosystem responses to a changing world.

Invasive species and bottom-water hypoxia both constitute major global threats to the diversity and integrity of marine ecosystems. These stressors may interact with unexpected consequences, as invasive species that require an initial environmental disturbance to become established can subsequently become important drivers of ecological change. There is recent evidence that improved bottom-water oxygen conditions in coastal areas of the northern Baltic Sea coincide with increased abundances of the invasive polychaetes Marenzelleria spp. Using a reactive-transport model, we demonstrate that the long-term bioirrigation activities of dense Marenzelleria populations have a major impact on sedimentary phosphorus dynamics. This may facilitate the switch from a seasonally hypoxic system back to a normoxic system by reducing the potential for sediment-induced eutrophication in the upper water column. In contrast to short-term laboratory experiments, our simulations, which cover a 10-year period, show that Marenzelleria has the potential to enhance long-term phosphorus retention in muddy sediments. Over time bioirrigation leads to a substantial increase in the iron-bound phosphorus content of sediments while reducing the concentration of labile organic carbon. As surface sediments are maintained oxic, iron oxyhydroxides are able to persist and age into more refractory forms. The model illustrates mechanisms through which Marenzelleria can act as a driver of ecological change, although hypoxic disturbance or natural population declines in native species may be needed for them to initially become established. Invasive species are generally considered to have a negative impact; however, we show here that one of the main recent invaders in the Baltic Sea may provide important ecosystem services. This may be of particular importance in low-diversity systems, where disturbances may dramatically alter ecosystem services due to low functional redundancy. Thus, an environmental problem in one region may be either exacerbated or alleviated by a single species from another region, with potentially ecosystem-wide consequences.

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In the northern Baltic Sea, occurrences of benthic drifting macroalgal mats have become an ~ncreasing problem. Accumulations of algae induce hypoxia and anoxia in zoobenthos above the halocline on shallow sandy bottoms. The immediate and temporal responses of a macrozoobenthic community to accumulat~ons of drifting algal mats were studied in a field experiment on an exposed shallow sandy bottom (7.5 m depth) in the northern Baltic Sea. Experimental algal plots (50 X 50 cm) corresponding to amounts recorded in the field (440 g dwt m-') were enclosed in net-bags and attached to the bottom. Changes in zoobenthic community structure under the algae were compared with ambient control plots for a period of 5 wk with sampling every seventh day. Community parameters (species, abundance and biomass) in the control community remained stable. Structural differences were recorded after 9 d, and community breakdown after 16 to 21 d of algal cover. Only opportunists and species tolerant to hypoxia remained under the algae. High particulate organic C/N ratios in the sediment under algal plots were recorded after 4 wk, indicating deposition of organic material from the algae to the sediment. Init~al recovery 15 d ) after termmated algal stress was rapid and dominated by a mass invasion ot hydrobud snatls, poss~bly attracted by enl-~chment of the sediment. Laboratory tests on algal degradation showed a 10-fold increase In phosphorus In the water due to nutrient leakdye during hypoxia (20'% Oz). Algal cover, and the ~nduction of hypoxla through degradation of the algae, exhibited severe effects on zoobenthlc community structure and a potential to accelerate local eutrophication. We therefore [eel that the escalating amounts of drifting algae recorded in the f~e l d are a significant threat to the coastal biota.

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