Filamentous, nitrogen-fixing cyanobacteria form extensive summer blooms in the Baltic Sea. Their ability to fix dissolved N2 allows cyanobacteria to circumvent the general summer nitrogen limitation, while also generating a supply of novel bioavailable nitrogen for the food web. However, the fate of the nitrogen fixed by cyanobacteria remains unresolved, as does its importance for secondary production in the Baltic Sea. Here, we synthesize recent experimental and field studies providing strong empirical evidence that cyanobacterial nitrogen is efficiently assimilated and transferred in Baltic food webs via two major pathways: directly by grazing on fresh or decaying cyanobacteria and indirectly through the uptake by other phytoplankton and microbes of bioavailable nitrogen exuded from cyanobacterial cells. This information is an essential step toward guiding nutrient management to minimize noxious blooms without overly reducing secondary production, and ultimately most probably fish production in the Baltic Sea.Electronic supplementary materialThe online version of this article (doi:10.1007/s13280-015-0660-x) contains supplementary material, which is available to authorized users.

The link between biodiversity and ecosystem functioning is an important question that remains unresolved, particularly in marine systems, in which cycling of organic matter by benthic organisms is of global significance. Direct observations of specific resource use by each species in single- and multispecies communities, as quantified by stable isotopes, facilitates a mechanistic understanding of the importance of each species for ecosystem functioning. We tested the effects of altered biodiversity (species richness) of deposit-feeding macrofauna on incorporation and burial of phytodetritus in combinations of three species representing natural communities found in the sediments of the species-poor Baltic Sea. The three species, two amphipods and a bivalve, had different rates of incorporation and burial and different needs for carbon (C) and nitrogen (N). The amphipods exhibited clear resource partitioning in sympatry, as a result of vertical separation in the sediment and consequent differential use of food. Communities of several species incorporated more C and N than expected from the respective single-species treatments, due to higher incorporation by surface feeders in multispecies treatments. Community incorporation of N in the most diverse treatment even exceeded N incorporation by a single-species treatment of the best-performing species, showing transgressive over-yielding. This over-yielding was primarily due to positive complementarity in all treatments. Diverse soft bottoms are also likely to be more productive in the long run, as species-specific traits (subsurface feeding) preserve fresh phytodetritus by burying it to depths in the sediment at which the mineralization rate is low. The more diverse sediment communities showed more efficient trophic transfer of phytodetritus, a finding of general significance for understanding biological processes driving the transformation of nutrients and energy in benthic ecosystems.

Karlson, A. M. L., Almqvist, G., Skóra, K. E., and Appelberg, M. 2007. Indications of competition between non-indigenous round goby and native flounder in the Baltic Sea. – ICES Journal of Marine Science, 64: 479–486. The Ponto-Caspian round goby (Neogobius melanostomus) was introduced to the Gulf of Gdańsk, southern Baltic Sea, in the late 1980s, and it has now become the dominant demersal fish species in shallow water. This study aimed to assess diet preferences and the degree of diet overlap between the round goby and the native flounder (Platichthys flesus). Results from time-series of stomach contents and stable isotope analyses of wild-caught fish, together with prey preference experiments carried out in the laboratory, showed that the two species consumed similar species and sizes of prey. The similarities in diet suggest potential for food competition. Catch data showed both reverse depth distributions of round goby and flounder when round gobies were abundant and that the abundances of the two species were negatively correlated. The diet overlap between small flounders and round gobies was greatest when goby abundance was least, suggesting that abundance of round gobies may restrict flounder habitat utilization and, therefore, also food availability to the latter. Therefore, round gobies may have a negative influence on the commercially important flounder.

The potential for using stable isotope analysis in risk assessment of environmental contaminants is crucially dependent on the predictability of the trophic transfer of isotopes in food webs. The relationship between contaminant levels and trophic position of consumers is widely used to assess biomagnification properties of various pollutants by establishing trophic magnification factors (TMF). However, contaminant-induced variability of the isotopic composition in biota is poorly understood. Here, we investigated effects of toxic exposure on δ(15)N and δ(13)C values in a consumer, with a main hypothesis that these effects would be largely mediated via growth rate and metabolic turnover of the test animals. The cladoceran Daphnia magna was used in two experiments that were conducted to manipulate growth and body condition (assayed as C:N ratio) by food availability and temperature (Experiment 1) and by toxic exposure to the pesticide lindane (Experiment 2). We found a significant negative effect of growth rate and a positive effect of temperature on the consumer-diet discrimination factor for δ(15)N and δ(13)C, with no effects on the C:N ratio (Experiment 1). In lindane-exposed daphnids, a significant growth inhibition was observed, with concomitant increase in metabolic costs and significantly elevated size-specific δ(15)N and δ(13)C values. Moreover, a significantly higher incorporation of carbon relative to nitrogen, yet a concomitant decrease in C:N ratio was observed in the exposed animals. Together, these results have methodological implications for determining trophic positions and TMF in polluted environments, where elevated δ(15)N values would translate into overestimated trophic positions and underestimated TMF. Furthermore, altered δ(13)C values may lead to erroneous food-chain assignment of the consumer in question.

Ecosystem consequences of biodiversity change are often studied from a species loss perspective, while the effects of invasive species on ecosystem functions are rarely quantified. In this experimental study, we used isotope tracers to measure the incorporation and burial of carbon and nitrogen from a simulated spring phytoplankton bloom by communities of one to four species of deposit-feeding macrofauna found in the species-poor Baltic Sea. The recently invading polychaete Marenzelleriaarctia, which has spread throughout the Baltic Sea, grows more rapidly than the native species Monoporeia affinis, Pontoporeia femorata (both amphipods) and Macoma balthica (a bivalve), resulting in higher biomass increase (biomass production) in treatments including the polychaete. Marenzelleria incorporated and buried bloom material at rates similar to the native species. Multi-species treatments generally had higher isotope incorporation, indicative of utilization of bloom material, than expected from monoculture yields of the respective species. The mechanism behind this observed over-yielding was mainly niche complementarity in utilization of the bloom input, and was more evident in communities including the invader. In contrast, multi-species treatments had generally lower biomass increase than expected. This contrasting pattern suggests that there is little overlap in resource use of freshly deposited bloom material between Marenzelleria and the native species but it is likely that interference competition acts to dampen resulting community biomass. In conclusion, an invasive species can enhance incorporation and burial of organic matter from settled phytoplankton blooms, two processes fundamental for marine productivity.