Description: Despite management efforts, anthropogenic nutrient enrichments continue to enhance phytoplankton blooms worldwide. Release of nitrogen and phosphorus compounds not only provides surplus of nutrients but also disbalances their stoichiometry. Declines in the relative availability of dissolved silicon might induce limitation in diatoms, major primary producers with silicified shells. We studied experimentally how nutrient enrichment and resulting decline in dissolved silicon to nitrogen ratios (Si:N) affect the structure and functioning of natural plankton communities. Nitrate was added to create a range of Si:N ratios and phosphate was supplied in Redfield ratio to nitrogen. We also manipulated copepod abundance to understand the top-down effects on communities experiencing nutrient enrichment. Nitrogen and phosphorus additions resulted in a steep phytoplankton biomass increase, followed by a post-bloom decline. Phytoplankton bloom biomass was higher in high nitrogen treatments but during the post-bloom period this trend switched. Biomass was sustained longer in high Si:N treatments, indicating that silicon limitation terminates the bloom. Many diatom species did not benefit from nitrogen and phosphorus enrichment and diatom dominance ceased below Si:N of 0.4:1. Under high grazing pressure, silicate was taken up faster suggesting that silicification is important in diatom defense. Copepods shaped plankton communities via feeding on dinoflagellates, chlorophytes and the diatom Skeletonema costatum but there was no significant effect of nitrogen and phosphorus enrichment on copepod abundance. Our results, combined with previous studies, show that while nutrient concentrations define the total phytoplankton bloom biomass, resource ratios are important in sustaining biomass and determining community structure and composition.

Description: The plea for using more “realistic,” community‐level, investigations to assess the ecological impacts of global change has recently intensified. Such experiments are typically more complex, longer, more expensive, and harder to interpret than simple organism‐level benchtop experiments. Are they worth the extra effort? Using outdoor mesocosms, we investigated the effects of ocean warming (OW) and acidification (OA), their combination (OAW), and their natural fluctuations on coastal communities of the western Baltic Sea during all four seasons. These communities are dominated by the perennial and canopy‐forming macrophyte Fucus vesiculosus—an important ecosystem engineer Baltic‐wide. We, additionally, assessed the direct response of organisms to temperature and pH in benchtop experiments, and examined how well organism‐level responses can predict community‐level responses to the dominant driver, OW. OW affected the mesocosm communities substantially stronger than acidification. OW provoked structural and functional shifts in the community that differed in strength and direction among seasons. The organism‐level response to OW matched well the community‐level response of a given species only under warm and cold thermal stress, that is, in summer and winter. In other seasons, shifts in biotic interactions masked the direct OW effects. The combination of direct OW effects and OW‐driven shifts of biotic interactions is likely to jeopardize the future of the habitat‐forming macroalga F. vesiculosus in the Baltic Sea. Furthermore, we conclude that seasonal mesocosm experiments are essential for our understanding of global change impact because they take into account the important fluctuations of abiotic and biotic pressures.

Description: Ecological theory suggests that a combination of local and regional factors regulate biodiversity and community functioning in metacommunities. The relative importance of different factors structuring communities likely changes over successional time, but to date this concept is scarcely documented. In addition, the few studies describing successional dynamics in metacommunity regulation have only focused on a single group of organisms. Here, we report results of an experimental study testing the effect size of initial local community composition and dispersal between local patches on community dynamics of benthic microalgae and their associated bacteria over community succession. Our results show that over time dispersal outweighed initial effects of community composition on microalgal evenness and biomass, microalgal β-diversity, and the ratio of bacteria to microalgae. At the end of the experiment (ca. 20 microalgae generations), dispersal significantly decreased microalgal evenness and β-diversity by promoting one regionally superior competitor. Dispersal also decreased the ratio of bacteria to microalgae, while it significantly increased microalgal biomass. These results suggest that the dispersal-mediated establishment of a dominant and superior microalgae species prevented bacteria from gaining competitive advantage over the autotrophs in these metacommunities, ultimately maintaining the provision of autotrophic biomass. Our study emphasizes the importance of time for dispersal to be a relevant community-structuring mechanism. Moreover, we highlight the need for considering multiple competitors in complex metacommunity systems to properly pinpoint the consequences of local change in dominance through dispersal for metacommunity function.

Description: How ecological and evolutionary processes interact and together determine species and community responses to climate change is poorly understood. We studied long-term dynamics (over approximately 200 asexual generations) in two phytoplankton species, a coccolithophore (Emiliania huxleyi), and a diatom (Chaetoceros affinis), to increased CO2 growing alone, or competing with one another in co-occurrence. To allow for rapid evolutionary responses, the experiment started with a standing genetic variation of nine genotypes in each of the species. Under co-occurrence of both species, we observed a dominance shift from C. affinis to E. huxleyi after about 120 generations in both CO2 treatments, but more pronounced under high CO2. Associated with this shift, we only found weak adaptation to high CO2 in the diatom and none in the coccolithophore in terms of species’ growth rates. In addition, no adaptation to interspecific competition could be observed by comparing the single to the two-species treatments in reciprocal assays, regardless of the CO2 treatment. Nevertheless, highly reproducible genotype sorting left only one genotype remaining for each of the species among all treatments. This strong evolutionary selection coincided with the dominance shift from C. affinis to E. huxleyi. Since all other conditions were kept constant over time, the most parsimonious explanation for the dominance shift is that the strong evolutionary selection was driven by the experimental nutrient conditions, and in turn potentially altered competitive ability of the two species. Thus, observed changes in the simplest possible two-species phytoplankton “community” demonstrated that eco-evolutionary interactions can be critical for predicting community responses to climate change in rapidly dividing organisms such as phytoplankton.

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: While the effect of the global biodiversity crisis on local species loss is still debated, there is empirical evidence for major changes in local biodiversity attributed to increased species turnover. In communities exposed to a climate stressor, species turnover can lead to increased dominance of well-adapted species and consequently to an overall decline in species diversity. Despite the known importance of species turnover for community dynamics and functioning, experimental results on the connection between biodiversity loss and species turnover are scarce. We still do not fully understand which specific factors increase the rate of change in species composition, especially when considering natural compared to artificially lab assembled communities. In the present study, we experimentally tested whether a heatwave and dispersal increased species turnover and decreased species diversity in natural benthic diatom communities with different initial species compositions. We found that on the local scale, dispersal had overall positive effects on species richness while the relationship between exposure to the heatwave, species turnover, and diversity depended on initial community composition. However, on the regional (i.e. metacommunity) scale, exposure to the heatwave and dispersal both increased turnover and decreased Shannon diversity by almost 50%. Turnover in these metacommunities was not caused by a loss of species, but rather by a change in dominance patterns leading to homogenization, and consequently decreased diversity. Our study shows that climate change can destabilize community composition and degrade species diversity, but still after ca. 15 generations does not decrease the number of species in the community, demonstrating that the response of species diversity and richness to changing conditions can be fundamentally decoupled on ecological time scales.

Description: Previous studies with Baltic Sea phytoplankton combining elevated seawater temperature with CO2 revealed the importance of size trait‐based analyses, in particular dividing the plankton into edible (〉5 and 〈100 μm) and inedible ([removed]100 μm) size classes for mesozoopankton grazers. While the edible phytoplankton responded predominantly negative to warming and the inedible group stayed unaffected or increased, independent from edibility most phytoplankton groups gained from CO2. Because the ratio between edible and inedible taxa changes profoundly over seasons, we investigated if community responses can be predicted according to the prevailing composition of edible and inedible groups. We experimentally explored the combined effects of elevated temperatures and CO2 concentrations on a late‐summer Baltic Sea community. Total phytoplankton significantly increased in response to elevated CO2 in particular in combination with temperature, driven by a significant gain of the inedible 〈5 μm fraction and large filamentous cyanobacteria. Large flagellates disappeared. The edible group was low as usual in summer and decreased with both factors due to enhanced copepod grazing and overall decline of small flagellates. Our results emphasize that the responses of summer communities are complex, but can be predicted by the composition and dominance of size classes and groups.

Description: Intraspecific diversity is a substantial part of biodiversity, yet little is known about its maintenance. Understanding mechanisms of intraspecific diversity shifts provides realistic detail about how phytoplankton communities evolve to new environmental conditions, a process especially important in times of climate change. Here, we aimed to identify factors that maintain genotype diversity and link the observed diversity change to measured phytoplankton morpho-functional traits Vmax and cell size of the species and genotypes. In an experimental setup, the two phytoplankton species Emiliania huxleyi and Chaetoceros affinis, each consisting of nine genotypes, were cultivated separately and together under different fluctuation and nutrient regimes. Their genotype composition was assessed after 49 and 91 days, and Shannon’s diversity index was calculated on the genotype level. We found that a higher intraspecific diversity can be maintained in the presence of a competitor, provided it has a substantial proportion to total biovolume. Both fluctuation and nutrient regime showed species-specific effects and especially structured genotype sorting of C. affinis. While we could relate species sorting with the measured traits, genotype diversity shifts could only be partly explained. The observed context dependency of genotype maintenance suggests that the evolutionary potential could be better understood, if studied in more natural settings including fluctuations and competition.

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.

Description: Shifts in microbial communities and their functioning in response to environmental change result from contemporary interspecific and intraspecific diversity changes. Interspecific changes are driven by ecological shifts in species composition, while intraspecific changes are here assumed to be dominated by evolutionary shifts in genotype frequency. Quantifying the relative contributions of interspecific and intraspecific diversity shifts to community change thus addresses the essential, yet understudied question as to how important ecological and evolutionary contributions are to total community changes. This debate is to date practically constrained by (a) a lack of studies integrating across organizational levels and (b) a mismatch between data requirements of existing partitioning metrics and the feasibility to collect such data, especially in microscopic organisms like phytoplankton. We experimentally assessed the relative ecological and evolutionary contributions to total phytoplankton community changes using a new design and validated its functionality by comparisons to established partitioning metrics. We used a community of coexisting Emiliania huxleyi and Chaetoceros affinis with initially nine genotypes each. First, we exposed the community to elevated CO2 concentration for 80 days (~50 generations) to induce interspecific and intraspecific diversity changes and a total abundance change. Second, we independently manipulated the induced interspecific and intraspecific diversity changes in an assay to quantify the corresponding ecological and evolutionary contributions to the total change. Third, we applied existing partitioning metrics to our experimental data and compared the outcomes. Total phytoplankton abundance declined to one-fifth in the high CO2 exposed community compared to ambient conditions. Consistently across all applied partitioning metrics, the abundance decline could predominantly be explained by ecological shifts and to a low extent by evolutionary changes. We discuss potential consequences of the observed community changes on ecosystem functioning. Furthermore, we explain that the low evolutionary contributions likely resulted of intraspecific diversity changes that occurred irrespectively of CO2. We discuss how the assay could be upscaled to more realistic settings, including more species and drivers. Overall, the presented calculations of eco-evolutionary contributions to phytoplankton community changes constitute another important step towards understanding future phytoplankton shifts, and eco-evolutionary dynamics in general.