Seasonal variability in ecosystem functions: quantifying the contribution of invasive species to nutrient cycling in coastal ecosystems

Benthic ecosystems at temperate and high latitudes experience marked seasonal variation in the environmental factors affecting nutrient remineralization processes both directly and indirectly through their effects on the benthic communities. The invasive polychaete genus Marenzelleria represents new functionality in Baltic Sea sediments through its deep burrowing and extensive gallery formation, thus possibly greatly affecting benthic oxygen and nutrient fluxes. We assessed the seasonal contribution of Marenzelleria spp. to fluxes of solutes in monthly field measurements at 2 sites, 10 and 33 m deep, in the northern Baltic Proper over 1 yr. In general, the fluxes of inorganic nutrients and oxygen were higher during summer than during winter, and the seasonal variation was more pronounced at the deeper, more biologically active site. By using variation partitioning, we were able to demonstrate that Marenzelleria and other macrofauna could account for up to 92% of the variation in the fluxes depending on the site and season. Fauna was the most important in predicting the fluxes in spring when the sediment organic content and the abundance of juvenile Marenzelleria spp. were highest, while during e.g. winter, the influence of Marenzelleria spp., even though abundant, on solute fluxes was negligible. The results from this study have implications for management, and, importantly, for the modelling of nutrient budgets often based on values from studies conducted during the summer period only, thus possibly greatly miscalculating the annual nutrient fluxes.