Description: Light and nutrients are essential resources for phytoplankton growth and considered to shape the size structure and other morphometric traits (surface:volume ratio, deviation from spherical shape) of phytoplankton communities. If morphometric traits influence the growth and resource use, shifts by one of the two factors should influence the capability to utilize the other factor. We performed a two-step experiment, where a natural phytoplankton community was first exposed to three different light levels (supposed to be limiting, saturating and slightly inhibiting for the majority of species) and grown until stationary phase. Then, the pre-conditioned communities were split into two nutrient treatments (control and saturating nutrient pulse) and again grown until stationary phase under the medium light intensity. During the experimental light phase, community mean cell-size increased with light, but surface:volume ratio and deviation from spherical shape decreased. Moreover, in response to the following nutrient pulse, the low light pre-conditioned communities showed the highest initial growth rates in response to the nutrient pulse. The high light pre-conditioned communities showed the highest conversion of the nutrient pulse into biomass during the stationary phase. These results demonstrate how the imprint of one environmental factor on trait distribution influences the ability to cope with another.

Description: Climate change has the potential to profoundly influence the community structure and function of marine ecosystems. Prior to testing the consequences of altered environmental conditions on ecosystem functioning, it is first necessary to better understand how the functioning of an ecosystem is affected by its structure. Using phytoplankton communities with 4 naturally co‑occurring coccolithophores including species of Emiliania, Gephyrocapsa, and Calcidiscus collected off the Azores, we experimentally tested whether varying initial dominance leads to different competitive outcomes and consequently affects community functioning, such as biomass and carbon accumulation. We manipulated initial community structure by creating 5 different dominance scenarios: (1) all species contributing evenly to total initial biomass, and (2–5) one of each species contributing 4× that of the remaining 3 species to total initial biomass. All 4 species were simultaneously grown in monocultures starting with the same total initial biomass as the communities. Monocultures differed significantly in total final biomass, particulate inorganic carbon, and particulate organic carbon content. Priority effects in the communities caused the initially dominant species to remain dominant during the stationary phase in 3 out of 4 cases. However, despite varying dominant species and different outcomes in the monocultures, community functioning was unaffected. We suggest that selective and facilitative effects are responsible for the equalization of community functioning. We conclude that monoculture experiments are not sufficient to predict whole-community responses, since species interactions can significantly alter the expected functional outcome.

Description: In an increasingly modified world, understanding and predicting the consequences of landscape alteration on biodiversity is a challenge for ecologists. To this end, metacommunity theory has developed to better understand the complexity of local and regional interactions that occur across larger landscapes. While metacommunity ecology has now provided several alternative models of species coexistence at different spatial scales, predictions regarding the consequences of landscape alteration have been done exclusively for the competition-colonization trade off model (CC). In this paper we investigate the effects of landscape perturbation on source-sink metacommunities. We show that habitat destruction perturbs the equilibria among species competitive effects within the metacommunity, driving both direct extinctions and an indirect extinction debt. As in CC models, we found a time lag for extinction following habitat destruction that varied in length depending upon the relative importance of direct and indirect effects. However, in contrast to CC models, we found that the less competitive species are more affected by habitat destruction. The best competitors can sometimes even be positively affected by habitat destruction, which corresponds well with the results of field studies. Our results are complementary to those results found in CC models of metacommunity dynamics. From a conservation perspective, our results illustrate that landscape alteration jeopardizes species coexistence in patchy landscapes through complex indirect effects and delayed extinctions patterns.

Description: We investigated the impacts of predicted ocean acidification and future warming on the quantity and nutritional quality of a natural phytoplankton autumn bloom in a mesocosm experiment. Since the effects of CO2-enrichment and temperature have usually been studied independently, we were also interested in the interactive effects of both aspects of climate change. Therefore, we used a factorial design with 2 temperature and 2 acidification levels in a mesocosm experiment with a Baltic Sea phytoplankton community. Our results show a significant time-dependent influence of warming on phytoplankton carbon, chlorophyll a, and particulate organic carbon. Phytoplankton carbon, for instance, decreased by more than half with increasing temperature at bloom time. Additionally, elemental carbon to phosphorus ratios (C:P) increased significantly, by approximately 5 to 8%, due to warming. Impacts of CO2 or synergetic effects of warming and acidification could not be detected. We suggest that stronger grazing pressure induced by temperature was responsible for the significant decline in phytoplankton biomass. Our results suggest that the biological effects of warming on Baltic Sea phytoplankton are considerable and will likely have fundamental consequences for trophic transfer in the pelagic food web

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: Ecological stoichiometry can be a powerful tool to understand food web consequences of altered biogeochemical cycles as well as consequences of biodiversity loss on biogeochemistry and has proved to be a suitable framework to predict effects of consumers on the nutrient content of their prey. However, predictions from ecological stoichiometry have mainly been tested using single consumer species, whereas in most natural ecosystems several consumer species coexist. We conducted 2 outdoor mesocosm experiments with marine rock pool communities to test whether species richness and species combination of benthic invertebrates affected the nutrient content of periphyton. We independently manipulated 12 different consumer combinations ranging from 0 to 6 (2004) or 0 to 4 (2005) grazer species and measured the biomass and nutrient content of the algae. Grazers included 3 gastropods and 3 crustaceans. In 2005, we additionally analyzed animal nutrient content and N excretion rate. Algal biomass and C:N ratios decreased in the presence of grazers in both years, indicating that the remaining algae had higher internal N content. Also, both biomass (2004 and 2005) and C:N ratios (only 2004) decreased even further when grazer richness increased. In 2004, significant net diversity effects of grazer richness on periphyton C:N ratios indicated that periphyton N content under multispecies grazing could not be predicted from the effect of single species. In 2005, significant net diversity effects on C:N ratios were rare, but periphyton C:N ratios consistently decreased with increasing grazer excretion rate, indicating that higher nitrogen regeneration by grazers led to higher N incorporation by algae. The effects of species richness were mainly affected by the presence of one efficient grazer, the gastropod Littorina littorea. Our experiments indicate that non-additive intraguild interactions may qualitatively alter the stoichiometric effects of multispecies consumer assemblages.

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: Phytoplankton cell size is important for a multitude of functional traits such as growth rates, storage capabilities, and resistance to grazing. Because these response traits are correlated, selective effects on mean community cell size of one environmental factor should impact the ability of phytoplankton to cope with other factors. Here, we experimentally apply expectations on the functional importance of phytoplankton cell size to the community level. We used a natural marine plankton community, and first altered the community’s cell size structure by exposing it to six different grazer densities. The size-shifted communities were then treated with a saturated nutrient pulse to test how the changes in community size structure influenced the mean community growth rate in the short-term (day 1–3) and nutrient storage capacity in the postbloom phase. Copepod grazing reduced the medium-sized phytoplankton and increased the share of the smallest (〈10 µm3 ) and the largest (〉100,000 µm3 ). Communities composed of on average small cells grew faster in response to the nutrient pulse, and thus confirmed the previously suggested growth advantage of small cells for the community level. In contrast, larger phytoplankton showed better storage capabilities, reflected in a slower post-bloom decline of communities that were on average composed of larger cells. Our findings underline that the easily measurable mean cell size of a taxonomically complex phytoplankton community can be used as an indicator trait to predict phytoplankton responses to sequential environmental changes.