Description: Generalisations on the combined effects of consumers and resources on autotrophs in aquatic food webs largely rely on freshwater studies. In this study, we tested these general concepts with marine benthic microalgae, which are important components of coastal food webs. We manipulated nitrogen availability and herbivore presence in a factorial field experiment in the Western Baltic Sea. Moreover, we investigated how herbivore control varied among 3 sites and 2 seasons and tested for trophic cascades by enhancing demersal fish density at 2 sites. Nitrogen availability and herbivore presence had strong and antagonistic effects on microalgal biomass, species composition and diversity. Herbivores significantly reduced algal biomass, whereas nutrient enrichment led to an increase in biomass. Herbivore effects on microalgal biomass increased with increasing nitrogen availability, indicating a functional response of herbivores to nutrient enrichment. The response of microalgae at the species level suggested a trade-off between nutrient use and grazing resistance which appeared to be linked to algal growth form. Compared to other growth forms, large erect species were most responsive to both nitrogen loading and herbivory. Grazing reduced microalgal diversity at low nutrient supply, but enhanced it at high nutrient supply. Herbivore effects varied considerably among different sites and were stronger in spring than in summer. Manipulations of fish density during summer did not have any effects on microalgal community structure. In conclusion, our results demonstrate that herbivores and nutrients have strong and balancing effects on marine microbenthic community structure.

Description: The combined and interactive effects of climatic and ecological factors are rarely considered in marine communities. We designed a factorial field experiment to analyze (1) the interactive effects of ambient UV radiation and consumers; and (2) the effects of photosynthetically active radiation (PAR 400 to 700 nm), UVA (320 to 400 nm) and UVB (280 to 320 nm) radiation on a marine hard-bottom community in Nova Scotia, NW Atlantic. Species recruitment and succession on ceramic tiles were followed for 5 mo. We found strong negative UV effects on biomass and cover of the early colonizing macroalga Pilayella littoralis, whereas UVB was more harmful than UVA radiation. Consumers, mainly gammarid amphipods, increased P. littoralis biomass when UV was excluded, probably through fertilization. These initially strong and interacting UV and consumer effects on total biomass and cover diminished as species succession progressed. Species diversity was not affected by experimental treatments, but significant shifts in species composition occurred, especially at the recruitment stage. Red algae were most inhibited by UV, whereas sedentary invertebrates and some brown algae tended to increase under UV exposure. Consumers suppressed green and filamentous brown algae, but favored the other groups. Again, these effects diminished during the later stages of succession. We conclude that UV radiation can be a significant structuring force in early successional benthic communities, and that consumers can mediate its effects.

Description: Many coastal oceans experience not only increased loads of nutrients but also changes in the stoichiometry of nutrient supply. Excess supply of nitrogen and stable or decreased supply of silicon lower silicon to nitrogen (Si:N) ratios, which may decrease diatom proportion in phytoplankton. To examine how Si:N ratios affect plankton community composition and food web structure, we performed a mesocosm experiment where we manipulated Si:N ratios and copepod abundance in a Baltic Sea plankton community. In high Si:N treatments, diatoms dominated. Some of them were likely spared from grazing unexpectedly resulting in higher diatom biomass under high copepod grazing. With declining Si:N ratios, dinoflagellates became more abundant under low and picoplankton under high copepod grazing. This altered plankton food web structure: under high Si:N ratios, edible diatoms were directly accessible food for copepods, while under low Si:N ratios, microzooplankton and phago-mixotrophs (mixoplankton) were a more important food source for mesograzers. The response of copepods to changes in the phytoplankton community was complex and copepod density-dependent. We suggest that declining Si:N ratios favor microzoo- and mixoplankton leading to increased complexity of planktonic food webs. Consequences on higher trophic levels will, however, likely be moderated by edibility, nutritional value or toxicity of dominant phytoplankton species.

Description: Destructive macroalgal mass blooms threaten estuarine and coastal ecosystems worldwide. We asked which factors regulate macroalgal bloom intensity, distribution and species composition. In field experiments in the Baltic Sea, we analyzed the relative effects of nutrients, herbivores and algal propagule banks on population development and dominance patterns in two co-occurring bloom-forming macroalgae, Enteromorpha intestinalis and Pilayella littoralis. Both species were highly affected by the combined effects of a propagule bank, herbivory and nutrients. The magnitude of effects varied with season. The propagule bank was an important overwintering mechanism for both algae, and allowed for recruitment two months earlier than recruitment via freshly dispersed propagules. This provided a seasonal escape from intense herbivory and nutrient limitation later in the year. Favored by massive recruitment from the propagule bank, Enteromorpha was the superior space occupier in early spring, thereby reducing recruitment of Pilayella. Elimination of the propagule bank and recruitment via freshly dispersed propagules favored Pilayella. Strong and selective herbivory on Enteromorpha supported Pilayella in the presence, but not in the absence of the propagule bank. Nutrient enrichment in summer counteracted herbivore pressure on Enteromorpha, thereby negatively affecting Pilayella. Herbivore and nutrient effects were more pronounced for early life stages than adult algae. These results show that recruitment processes and forces affecting early life stages at the beginning of the vegetation period determine development and dominance patterns of macroalgal blooms. Herbivores naturally suppress blooms but increasing nutrient enrichment can override this important control mechanism. The propagule bank plays a previously unrecognized role for population and community dynamics.

Description: Diatoms often dominate phytoplankton in temperate, polar and upwelling regions. Decreases in silicate availability or silicon to nitrogen (Si:N) ratios may induce silicon limitation in diatoms and lower their proportion within phytoplankton communities. The effects of such changes on the nutritional quality of phytoplankton are not well understood. To examine how changing Si:N ratios affect plankton nutritional value, we applied a range of Si:N ratios on a natural plankton community and manipulated grazing pressure to assess top-down effects of copepod selective grazing. Diatom proportion in phytoplankton increased with increasing Si:N ratios and so did phytoplankton nutritional quality in terms of major fatty acid concentrations, such as polyunsaturated fatty acids, docosahexaenoic (DHA) and eicosapentaenoic (EPA) acids. However, stoichiometric quality (carbon to nitrogen and carbon to phosphorus ratios), DHA:EPA and omega 3:6 (omega 3:omega 6) ratios declined with increasing Si:N ratios, suggesting that proportions between essential compounds in copepod diet may be more favorable in lowered Si:N ratios. Copepods had a negative effect on DHA contents, DHA:EPA and omega 3:omega 6 ratios, indicating possible selective grazing on more nutritious plankton. Our findings show that declining silicate concentrations can affect stoichiometric and biochemical quality of phytoplankton, which copepods can also moderate by selective grazing.