Description: By factorial field experiments we analyzed the relative effects of increased nutrient (N+P) loading and natural grazing pressure on species composition, carbon storage, and nitrogen retention in the Baltic Sea littoral food web, composed of macroalgae, grazers (snails, isopods, amphipods), and predators (shrimps, crabs, fish). Nitrogen was depleted relative to phosphorus throughout most of the year. Increasing nitrogen (6–200% over ambient concentrations) enhanced algal productivity and cover of fast-growing annual algae, grazer, and predator densities, suggesting a three-level bottom-up effect. With increasing nitrogen loading, annual algae increasingly blocked perennial algal recruitment (65–98% decrease) and growth. Grazers counteracted the effects of nutrient enrichment on algal species composition through selective consumption of annual algae. Grazer exclusion had equivalent negative effects on perennial recruitment as a 85% increase in nitrogen loading. Nutrient enrichment increased algal nitrogen content and decreased tissue C: N ratios in spring and summer but not in fall. Carbon storage and nitrogen retention, measured as C and N retained in plant biomass at the end of the growth season, were increased by grazers (C: 39%, N: 24%) but decreased with increasing nitrogen loading (C: -71%, N: -74%). Our results emphasize the important role of grazers in buffering moderate eutrophication effects and illustrate how food web interactions and shifts in species composition are tightly linked to coastal ecosystem function

Description: In contrast to most pelagic primary producers, benthic macrophytes pass through morphologically distinct life stages, which can be subject to different ecological controls. Using factorial field experiments, we investigated how grazing pressure (three levels) and nutrient supply (four levels) interact in controlling the passage of marine macroalgae through an apparent recruitment bottleneck at the germling stage. In comparative experiments, we asked whether relative bottom-up and top-down effects on early life stages (〈4 week germlings) vary (1) between the eutrophic Baltic Sea and the oligotrophic NW Atlantic, (2) across seasons in the NW Atlantic, and (3) among annual and perennial macroalgae. In both systems nutrient enrichment favored and grazers suppressed recruitment of green and brown annual algae; however, enrichment effects were much more pronounced in the Baltic, whereas grazer effects dominated in the NW Atlantic. Grazers induced a shift from grazer-susceptible green to more resistant brown algae in the Baltic without reducing total germling density. In the NW Atlantic, grazers strongly reduced overall recruitment rate throughout all seasons. Effects on perennials were similar in both systems with moderate losses to grazing and no effects of nutrient enrichment. Recruit densities and species composition shifted with season in the NW Atlantic. We conclude that the relative effects of grazers and nutrient enrichment depended on the nutrient status of the system, algal life history strategy, and season. Strong bottom-up and top-down controls shape benthic community composition before macroalgae reach visible size

Description: Microzooplankton have received increased attention as an important trophic link between the microbial loop and calanoid copepods. On the basis of food size spectra overlap in some microzooplankton groups and calanoid copepods, however, such microzooplankton could function as competitors rather than as food for calanoid copepods (intraguild prey). Mixotrophic flagellates presumably represent a link between the microbial loop and the micro and mesozooplankton. We investigated the effects of microzooplankton and mixotrophy by altering the presence of a heterotrophic dinoflagellate and of a mixotrophic nanoflagellate in artificial food webs with calanoid copepods as terminal consumers. Overall system productivity was manipulated by two levels of nutrient enrichment. The heterotrophic dinoflagellate drastically reduced the nanophytoplankton and enhanced the reproduction of the copepods, suggesting that its role as a competitor is negligible compared to its function as a trophic link. In spite of the presence of heterotrophic nanoflagellates, the mixotroph had a strong negative effect on the picophytoplankton and (presumably) on bacterial biomass. At the same time, the mixotroph enhanced the atomic C:N ratio of the seston biomass, indicating a higher efficiency in overall primary production. Copepod reproduction was enhanced in the presence of the mixotrophic nanoflagellate. Results did not support predictions of the intraguild predation theory: The ratios of the intraguild predators and their preys were not affected by overall system productivity

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: Ecosystem resistance to a single stressor relies on tolerant species that can compensate for sensitive competitors and maintain ecosystem processes, such as primary production. We hypothesize that resistance to additional stressors depends increasingly on species tolerances being positively correlated (i.e. positive species co-tolerance). Initial exposure to a stressor combined with positive species co-tolerance should reduce the impacts of other stressors, which we term stress-induced community tolerance. In contrast, negative species co-tolerance is expected to result in additional stressors having pronounced additive or synergistic impacts on biologically impoverished functional groups, which we term stress-induced community sensitivity. Therefore, the sign and strength of the correlation between species sensitivities to multiple stressors must be considered when predicting the impacts of global change on ecosystem functioning as mediated by changes in biodiversity.