Description: Highlights: • We used indoor mesocosms to test the impact of warming on plankton communities. • Different stages of phytoplankton bloom were analysed. • Increased temperature and zooplankton grazing had similar effects on phytoplankton. • Warming and increased zooplankton density decreased phytoplankton richness. • Warming and increased zooplankton density increased phytoplankton evenness. Recent climate warming is expected to affect phytoplankton biomass and diversity in marine ecosystems. Temperature can act directly on phytoplankton (e.g. rendering physiological processes) or indirectly due to changes in zooplankton grazing activity. We tested experimentally the impact of increased temperature on natural phytoplankton and zooplankton communities using indoor mesocosms and combined the results from different experimental years applying a meta-analytic approach. We divided our analysis into three bloom phases to define the strength of temperature and zooplankton impacts on phytoplankton in different stages of bloom development. Within the constraints of an experiment, our results suggest that increased temperature and zooplankton grazing have similar effects on phytoplankton diversity, which are most apparent in the post-bloom phase, when zooplankton abundances reach the highest values. Moreover, we observed changes in zooplankton composition in response to warming and initial conditions, which can additionally affect phytoplankton diversity, because changing feeding preferences of zooplankton can affect phytoplankton community structure. We conclude that phytoplankton diversity is indirectly affected by temperature in the post-bloom phase through changing zooplankton composition and grazing activities. Before and during the bloom, however, these effects seem to be overruled by temperature enhanced bottom-up processes such as phytoplankton nutrient uptake.

Description: Landscape connectivity can increase the capacity of communities to maintain their function when environments change by promoting the immigration of species or populations with adapted traits. However, high immigration may also restrict fine tuning of species compositions to local environmental conditions by homogenizing the community. Here we demonstrate that dispersal generates such a tradeoff between maximizing local biomass and the capacity of model periphyton metacommunities to recover after a simulated heat wave. In non-disturbed metacommunities, dispersal decreased the total biomass by preventing differentiation in species composition between the local patches making up the metacommunity. On the contrary, in metacommunities exposed to a realistic summer heat wave, dispersal promoted recovery by increasing the biomass of heat tolerant species in all local patches. Thus, the heat wave reorganized the species composition of the metacommunities and after an initial decrease in total biomass by 38.7%, dispersal fueled a full recovery of biomass in the restructured metacommunities. Although dispersal may decrease equilibrium biomass, our results highlight that connectivity is a key requirement for the response diversity that allows ecological communities to adapt to climate change through species sorting.

Description: Ecosystem functioning is affected by horizontal (within trophic groups) and vertical (across trophic levels) biodiversity. Theory predicts that the effects of vertical biodiversity depend on consumer specialization. In a microcosm experiment, we investigated ciliate consumer diversity and specialization effects on algal prey biovolume, evenness and composition, and on ciliate biovolume production. The experimental data was complemented by a process-based model further analyzing the ecological mechanisms behind the observed diversity effects. Overall, increasing consumer diversity had no significant effect on prey biovolume or evenness. However, consumer specialization affected the prey community. Specialist consumers showed a stronger negative impact on prey biovolume and evenness than generalists. The model confirmed that this pattern was mainly driven by a single specialist with a high per capita grazing rate, consuming the two most productive prey species. When these were suppressed, the prey assemblage became dominated by a less productive species, consequently decreasing prey biovolume and evenness. Consumer diversity increased consumer biovolume, which was stronger for generalists than for specialists and highest in mixed combinations, indicating that consumer functional diversity, i.e. more diverse feeding strategies, increased resource use efficiency. Overall, our results indicate that consumer diversity effects on prey and consumers strongly depend on species-specific growth and grazing rates, which may be at least equally important as consumer specialization in driving consumer diversity effects across trophic levels.

Description: Conceptual models predict counteractive effects of herbivores and nutrient enrichment on plant diversity and reversed effects of grazers under different nutrient regimes. I tested these hypotheses in 11 field experiments with periphyton communities in three different aquatic habitats (a highly eutrophic lake, an meso-eutrophic lake, and an meso-eutrophic part of the Baltic Sea coast) and in different seasons. Grazer access and nutrient supply were manipulated in a factorial design. Species richness and evenness were chosen as response variables. Both manipulated factors had significant and contrasting effects on diversity, with variable effect strength between sites and seasons. From the two aspects of diversity, evenness well reflected the changes in community composition. Fertilization tended to increase the dominance of few species and thus to decrease evenness, whereas grazers counteracted these effects by removing dominant life forms. The response of species richness was not as expected, since grazers decreased richness throughout, whereas nutrients had weaker effects but tended to increase richness. Species richness rather reflected changes in periphyton architecture. Grazers reduced algal richness presumably by co-consumption of rare species in the tightly connected periphyton assemblages, whereas enrichment may increase richness by providing more structure via increased dominance of filamentous species. Although grazer and nutrient effects on richness and evenness were opposing, there was no change in the effect of one factor by manipulation of the other.

Description: Climatic warming is a primary driver of change in ecosystems worldwide. Here, we synthesize responses of species richness and evenness from 187 experimental warming studies in a quantitative meta-analysis. We asked 1) whether effects of warming on diversity were detectable and consistent across terrestrial, freshwater and marine ecosystems, 2) if effects on diversity correlated with intensity, duration, and experimental unit size of temperature change manipulations, and 3) whether these experimental effects on diversity interacted with ecosystem types. Using multilevel mixed linear models and model averaging, we also tested the relative importance of variables that described uncontrolled environmental variation and attributes of experimental units. Overall, experimental warming reduced richness across ecosystems (mean log-response ratio = –0.091, 95% bootstrapped CI: –0.13, –0.05) representing an 8.9% decline relative to ambient temperature treatments. Richness did not change in response to warming in freshwater systems, but was more strongly negative in terrestrial (–11.8%) and marine (–10.5%) experiments. In contrast, warming impacts on evenness were neutral overall and in aquatic systems, but weakly negative on land (7.6%). Intensity and duration of experimental warming did not explain variation in diversity responses, but negative effects on richness were stronger in smaller experimental units, particularly in marine systems. Model-averaged parameter estimation confirmed these main effects while accounting for variation in latitude, ambient temperature at the sites of manipulations, venue (field versus lab), community trophic type, and whether experiments were open or closed to colonization. These analyses synthesize extensive experimental evidence showing declines in local richness with increased temperature, particularly in terrestrial and marine communities. However, the more variable effects of warming on evenness were better explained by the random effect of site identity, suggesting that effects on species’ relative abundances were contingent on local species composition.

Description: The factors regulating species diversity have received increasing attention in the face of the global biodiversity loss, but are not well understood for unicellular organisms. We conducted in situ experiments in Kiel Fjord in order to analyze the response of microalgal diversity to colonization time and to artificial eutrophication. Diversity decreased throughout colonization time (maximum: 12 weeks), whereas species richness initially increased to about 25 species before it leveled off. The proposed unimodal time course of diversity during succession could not be detected for diversity or species richness. The rapid decrease of evenness indicated a greater importance of algal growth on the substrata compared to the arrival of new species. Artificial eutrophication led to an decrease of diversity, which could be correlated to the supply concentrations of the limiting nutrient: P in spring, N in summer and Si in the presence of high concentrations of N and P. The decrease was due to an increased dominance of few species (i.e. reduced evenness), whereas species richness was not or positively correlated to nutrient supply. Species richness was negatively correlated to evenness and diversity measures. Thus, species diversity indices are useful response variable to measure environmental effects on local periphyton communities

Description: In order to examine the effects of warming and diversity changes on primary productivity, we conducted a meta-analysis on six independent indoor mesocosm experiments with a natural plankton community from the Baltic Sea. Temperature effects on primary productivity changed with light intensity and zooplankton density and analysed pathways between temperature, diversity and productivity, elucidating direct and indirect effects of warming on primary productivity during the spring phytoplankton bloom. Our findings indicate that warming directly increased carbon specific primary productivity, which was more pronounced under low grazing pressure. On the other hand, primary productivity per unit water volume did not respond to increased temperature, because of a negative temperature effect on phytoplankton biomass. Moreover, primary productivity response to temperature changes depended on light limitation. Using path analysis, we tested whether temperature effects were direct or mediated by warming effects on phytoplankton diversity. Although phytoplankton species richness had a positive impact on both net primary productivity and carbon specific primary productivity – and evenness had a negative effect on net primary productivity – both richness and evenness were not affected by temperature. Thus, we suggest that diversity effects on primary productivity depended mainly on other factors than temperature like grazing, sinking or nutrient limitation, which themselves are temperature dependent. Highlights ► Impact of warming on primary productivity and diversity–productivity relationship. ► Meta-analysis on indoor mesocosm experiments with a natural plankton community. ► Temperature has a direct impact on specific productivity, not on net productivity. ► Species richness increases and evenness decreases net primary productivity. ► Temperature does not directly affect diversity–productivity relationship.

Description: Recent experiments, mainly in terrestrial environments, have provided evidence of the functional importance of biodiversity to ecosystem processes and properties. Compared to terrestrial systems, aquatic ecosystems are characterised by greater propagule and material exchange, often steeper physical and chemical gradients, more rapid biological processes and, in marine systems, higher metazoan phylogenetic diversity. These characteristics limit the potential to transfer conclusions derived from terrestrial experiments to aquatic ecosystems whilst at the same time provide opportunities for testing the general validity of hypotheses about effects of biodiversity on ecosystem functioning. Here, we focus on a number of unique features of aquatic experimental systems, propose an expansion to the scope of diversity facets to be considered when assessing the functional consequences of changes in biodiversity and outline a hierarchical classification scheme of ecosystem functions and their corresponding response variables. We then briefly highlight some recent controversial and newly emerging issues relating to biodiversity-ecosystem functioning relationships. Based on lessons learnt from previous experimental and theoretical work, we finally present four novel experimental designs to address largely unresolved questions about biodiversity-ecosystem functioning relationships. These include (1) investigating the effects of non-random species loss through the manipulation of the order and magnitude of such loss using dilution experiments; (2) combining factorial manipulation of diversity in interconnected habitat patches to test the additivity of ecosystem functioning between habitats; (3) disentangling the impact of local processes from the effect of ecosystem openness via factorial manipulation of the rate of recruitment and biodiversity within patches and within an available propagule pool; and (4) addressing how non-random species extinction following sequential exposure to different stressors may affect ecosystem functioning. Implementing these kinds of experimental designs in a variety of systems will, we believe, shift the focus of investigations from a species richness-centred approach to a broader consideration of the multifarious aspects of biodiversity that may well be critical to understanding effects of biodiversity changes on overall ecosystem functioning and to identifying some of the potential underlying mechanisms involved.

Description: Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the P-content of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N- and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.

Description: Field experiments were conducted to investigate the effects of grazing and nutrient supply on sediment microflora in a freshwater habitat (Lake Erken, Sweden) and at the brackish Baltic Sea coast (Väddö, Sweden). The two sites were of similar productivity, but had contrasting herbivore composition. In a full-factorial experiment design, closed cages excluded macrozoobenthos (〉1 mm) from sediment patches, whereas open cages allowed grazer access. The cage design applied here proved to successfully prevent in- and epifauna to access the sediment in closed cages. In half of the treatments, nutrients were added to the water-column by a slow-release fertilizer. The experiments were seasonally replicated four times at Väddö and two times in Lake Erken. After 4–5 weeks, sediment cores were sampled and analyzed for chlorophyll, carbon, nitrogen and phosphorus. The benthic microalgae showed strong seasonal variation in biomass and internal nutrient content. At Väddö, neither grazing nor nutrients affected the algal biomass significantly, but significant grazer effects were detected on C:N:P ratios. In Lake Erken, grazer presence reduced algal biomass by ca. 50%, whereas nutrients were without effect on biomass or on nutrient content. Compared to results from hard substrata at the same sites, sediment microflora was less affected by nutrients and grazing. This may be due to the harsh physico-chemical environment on sediments, to low grazer density at the coastal site and to low availability of water column nutrients to sediment microalgae. In our experiments, sand-dwelling microphytobenthic communities represented a highly dynamic assemblage, which, however, is less structured by biotic interactions than epilithic periphyton