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Current knowledge on the seasonality of benthic nitrate reduction pathways in the aphotic, density stratified coastal zone of the Baltic Sea is largely based on data from muddy sediments, neglecting the potential contribution of sandy sediments. To gain a more comprehensive understanding of seasonality in this part of the Baltic Sea coast, we measured rates of benthic denitrification, anammox and dissimilatory nitrate reduction to ammonium (DNRA) monthly in the ice-free period of 2016 in both sandy and muddy aphotic sediments, northwestern Gulf of Finland. No anammox was observed. The seasonal cycle of denitrification in both sediment types was related to the hydrography-driven development of bottom water temperature. The seasonal cycle of DNRA was less clear and likely connected to a combination of bottom water temperature, carbon to nitrogen ratio, and substrate competition with denitrification. Denitrification and DNRA rates were 50-80 and 20% lower in the sandy than in the muddy sediment. The share of DNRA in total nitrate reduction, however, was higher in the sandy than in the muddy sediment, being (by similar to 50%) the highest DNRA share in sandy sediments so far measured. Our data add to the small pool of published studies showing significant DNRA in both cold and/or sandy sediments and suggest that DNRA is currently underestimated in the Baltic coastal nitrogen filter. Our results furthermore emphasize that the various environmental conditions of a coastal habitat (light regime, hydrography, and geomorphology) affect biogeochemical element cycling and thus need to be considered in data interpretation.

The coastal zone of the Baltic Sea is diverse with strong regional differences in the physico-chemical setting. This diversity is also reflected in the importance of different biogeochemical processes altering nutrient and organic matter fluxes on the passage from land to sea. This review investigates the most important processes for removal of nutrients and organic matter, and the factors that regulate the efficiency of the coastal filter. Nitrogen removal through denitrification is high in lagoons receiving large inputs of nitrate and organic matter. Phosphorus burial is high in archipelagos with substantial sedimentation, but the stability of different burial forms varies across the Baltic Sea. Organic matter processes are tightly linked to the nitrogen and phosphorus cycles. Moreover, these processes are strongly modulated depending on composition of vegetation and fauna. Managing coastal ecosystems to improve the effectiveness of the coastal filter can reduce eutrophication in the open Baltic Sea.

Benthic macrofaunal communities have a profound impact on organic matter turnover and nutrient cycling in marine sediments. Their activities are of particular importance in the coastal filter, where nutrients and organic matter from land are transformed and/or retained before reaching the open sea. The benthic fauna modify the coastal filter directly (through consumption, respiration, excretion and biomass production) and indirectly (through bioturbation). It is hard to experimentally quantify faunal contribution to the coastal filter over large spatial and temporal scales that encompass significant environmental and biological heterogeneity. However, estimates can be obtained with biological trait analyses. By using benthic biological traits, we explored how the potential contribution of macrofaunal communities to the coastal filter differ between inner and outer sites in an extensive archipelago area and examine the generality of the observed pattern across contrasting coastal areas of the entire Baltic Sea. Estimates of benthic bioturbation, longevity and size (i.e. ‘stability’) and total energy and nutrient contents differed between coastal areas and inner versus outer sites. Benthic traits indicative of an enhanced nutrient turnover but a decreased capacity for temporal nutrient retention dominated inner sites, while outer sites were often dominated by larger individuals, exhibiting traits that are likely to enhance nutrient uptake and retention. The overarching similarities in benthic trait expression between more eutrophied inner vs. less affected outer coastal sites across the Baltic Sea suggest that benthic communities might contribute in a similar manner to nutrient recycling and retention in the coastal filter over large geographical scales. peerReviewed

During three cruises to the Bay of Gdansk, Baltic Sea, the fauna, porewater and bottom water were sampled at stations parallel to the shore and along a transect offshore. Diffusive porewater fluxes were calculated and related to the total net fluxes (TNF) of nutrients. The TNF comprise all nutrients that reach the bottom water from the sediment including diffusive nutrient efflux, discharge from macrozoobenthos and microbial activity. They were determined during in situ incubations using a benthic chamber lander, which is rarely done in coastal research. The lander restricts the physical influence of currents and waves on the sediments and only allows nutrient fluxes due to bioturbation by natural communities. Strong benthic-pelagic coupling in the shallow coastal zone suggested a crucial filter function for the bioturbated coastal sediments, which are separated from muddy deep sediments with little or no fauna at a depth of 50 m; in between is a small intermediate zone. While diffusive fluxes were highest at intermediate and offshore stations, TNF were highest at sandy coastal stations, where reservoirs of dissolved nutrients were small and sediments almost devoid of organic material. The greatest impact of macrofauna on sedimentary fluxes was found at stations whose communities were dominated by deep-burrowing polychaetes. The largest TNF were measured directly at the mouth of the Vistula River, where riverine food and nutrients supplies were highest. Macrofauna communities and sediment variables can thus serve as descriptive indicator to estimate the extent of the coastal filter. Finally, based on the total areal size of the different sediment types, annual efflux for the complete coastal zone of the Gdansk Bay was estimated to be 6.9 kt N, 19 kt Si, and 0.9 kt P. Compared to the muddy offshore area, which is twice as large, these amounts were similar for P and threefold higher for N and Si.

Estuaries are often seen as natural filters of riverine nitrate, but knowledge of this nitrogen sink in oligotrophic systems is limited. We measured spring and summer dinitrogen production (denitrification, anammox) in muddy and non-permeable sandy sediments of an oligotrophic estuary in the northern Baltic Sea, to estimate its function in mitigating the riverine nitrate load. Both sediment types had similar denitrification rates, and no anammox was detected. In spring at high nitrate loading, denitrification was limited by likely low availability of labile organic carbon. In summer, the average denitrification rate was similar to 138 mu mol N m(-2) d(-1). The corresponding estuarine nitrogen removal for August was similar to 1.2 t, of which similar to 93% was removed by coupled nitrification-denitrification. Particulate matter in the estuary was mainly phytoplankton derived (> 70% in surface waters) and likely based on the riverine nitrate which was not removed by direct denitrification due to water column stratification. Subsequently settling particles served as a link be tween the otherwise uncoupled nitrate in surface waters and benthic nitrogen removal. We suggest that the riverine nitrate brought into the oligotrophic estuary during the spring flood is gradually, and with a time delay, removed by benthic denitrification after being temporarily ` trapped' in phytoplankton particulate matter. The oligotrophic system is not likely to face eutrophication from increasing nitrogen loading due to phosphorus limitation. In response, coupled nitrification-denitrification rates are likely to stay constant, which might increase the future export of nitrate to the open sea and decrease the estuary's function as a nitrogen sink relative to the load.