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In many regions across the globe, extreme weather events such as storms have increased in frequency, intensity, and duration due to climate change. Ecological theory predicts that such extreme events should have large impacts on ecosystem structure and function. High winds and precipitation associated with storms can affect lakes via short‐term runoff events from watersheds and physical mixing of the water column. In addition, lakes connected to rivers and streams will also experience flushing due to high flow rates. Although we have a well‐developed understanding of how wind and precipitation events can alter lake physical processes and some aspects of biogeochemical cycling, our mechanistic understanding of the emergent responses of phytoplankton communities is poor. Here we provide a comprehensive synthesis that identifies how storms interact with lake and watershed attributes and their antecedent conditions to generate changes in lake physical and chemical environments. Such changes can restructure phytoplankton communities and their dynamics, as well as result in altered ecological function (e.g., carbon, nutrient and energy cycling) in the short‐ and long‐term. We summarize the current understanding of storm‐induced phytoplankton dynamics, identify knowledge gaps with a systematic review of the literature, and suggest future research directions across a gradient of lake types and environmental conditions., Our understanding of ecosystem‐scale responses to storm events is fragmented, and terminological variability in defining a “storm” hinders resolution of generalizable impacts. We provide a comprehensive synthesis of the interactions between physical properties and biological responses in lakes that demonstrates the context‐dependency of storm impacts; lake/watershed attributes and their antecedent conditions mediate the extent to which storms impact lake ecosystems. We develop a framework which conceptualizes how abrupt, storm‐induced changes in the lake environment influence phytoplankton community dynamics through functional traits, identifies current limitations, and highlights several avenues of research to narrow knowledge gaps in synthetic and interdisciplinary ways.

Highlights • Modest brownification did not affect primary production, but increased bacterial production. • Concentration of inorganic nitrogen was the primary driver for the phytoplankton development. • Brownification benefitted picophytoplankton. Climate change is projected to cause brownification of some coastal seas due to increased runoff of terrestrially derived organic matter. We carried out a mesocosm experiment (15 d) to test the effect of this on the planktonic ecosystem expecting reduced primary production and shifts in the phytoplankton community composition. The experiment was set up in 2.2 m3 mesocosm bags using four treatments, each with three replicates: control (Contr) without any manipulation, organic carbon additive HuminFeed (Hum; 2 mg L−1), inorganic nutrients (Nutr; 5.7 μM NH4 and 0.65 μM PO4), and combined Nutr and Hum (Nutr + Hum) additions. Measured variables included organic and inorganic nutrient pools, chlorophyll a (Chla), primary and bacterial production and particle counts by flow cytometry. The bags with added inorganic nutrients developed a phytoplankton bloom that depleted inorganic N at day 6, followed by a rapid decline in Chla. Brownification did not reduce primary production at the tested concentration. Bacterial production was lowest in the Contr, but similar in the three treatments receiving additions likely due to increased carbon available for heterotrophic bacteria. Picoeukaryotes clearly benefited by brownification after inorganic N depletion, which could be due to more effective nutrient recycling, nutrient affinity, light absorption, or alternatively lower grazing pressure. In conclusion, brownification shifted the phytoplankton community composition towards smaller species with potential effects on carbon fluxes, such as sinking rates and export to the sea floor.

Functional traits are becoming more common in the analysis of marine zooplankton community dynamics associated with environmental change. We used zooplankton groups with common functional properties to assess long‐term trends in the zooplankton caused by certain environmental conditions in a highly eutrophicated gulf.Time series of zooplankton traits have been collected since the 1960s in the Gulf of Riga, Baltic Sea, and were analyzed using a combination of multivariate methods (principal coordinate analysis) and generalized additive models.One of the most significant changes was the considerable increase in the amount of the zooplankton functional groups (FGR) in coastal springtime communities, and dominance shifts from more complex to simpler organism groups—cladocerans and rotifers.The results also show that functional trait organism complexity (body size) decreased considerably due to cladoceran and rotifer increase following elevated water temperature. Salinity and oxygen had negligible effects on the zooplankton community., Time series of zooplankton traits were collected since the 1960s in the Gulf of Riga, Baltic Sea, and were analysed using general additive model, principal component analysis, and multivariate model. One of the most significant changes was the considerable increase in the amount of the zooplankton functional groups (FGR) in coastal springtime communities, and dominance shifts from more complex to simpler organism groups—cladocerans and rotifers. The results also show that the functional trait organism complexity (body size) decreased considerably due to cladoceran and rotifer increase following elevated water temperature.

1. Global pressures on freshwater ecosystems are high and rising. Viewed primarily as a resource for humans, current practices of water use have led to catastrophic declines in freshwater species and the degradation of freshwater ecosystems, including their genetic and functional diversity. Approximately three-quarters of the world's inland wetlands have been lost, one-third of the 28 000 freshwater species assessed for the International Union for Conservation of Nature (IUCN) Red List are threatened with extinction, and freshwater vertebrate populations are undergoing declines that are more rapid than those of terrestrial and marine species. This global loss continues unchecked, despite the importance of freshwater ecosystems as a source of clean water, food, livelihoods, recreation, and inspiration. 2. The causes of these declines include hydrological alterations, habitat degradation and loss, overexploitation, invasive species, pollution, and the multiple impacts of climate change. Although there are policy initiatives that aim to protect freshwater life, these are rarely implemented with sufficient conviction and enforcement. Policies that focus on the development and management of fresh waters as a resource for people almost universally neglect the biodiversity that they contain. 3. Here we introduce the Alliance for Freshwater Life, a global initiative, uniting specialists in research, data synthesis, conservation, education and outreach, and policymaking. This expert network aims to provide the critical mass required for the effective representation of freshwater biodiversity at policy meetings, to develop solutions balancing the needs of development and conservation, and to better convey the important role freshwater ecosystems play in human well-being. Through this united effort we hope to reverse this tide of loss and decline in freshwater biodiversity. We introduce several short- and medium-term actions as examples for making positive change, and invite individuals, organizations, authorities, and governments to join the Alliance for Freshwater Life.

About 60 years ago, the critical depth hypothesis was proposed to describe the occurrence of spring phytoplankton blooms and emphasized the role of stratification for the timing of onset. Since then, several alternative hypotheses appeared focusing on the role of grazing and mixing processes such as turbulent convection or wind activity. Surprisingly, the role of community composition—and thus the distribution of phytoplankton traits—for bloom formation has not been addressed. Here, we discuss how trait variability between competing species might influence phytoplankton growth during the onset of the spring bloom. We hypothesize that the bloom will only occur if there are species with a combination of traits fitting to the environmental conditions at the respective location and time. The basic traits for formation of the typical spring bloom are high growth rates and photoadaptation to low light conditions, but other traits such as nutrient kinetics and grazing resistance might also be important. We present concise ideas on how to test our theoretical considerations experimentally. Furthermore, we suggest that future models of phytoplankton blooms should include both water column dynamics and variability of phytoplankton traits to make realistic projections instead of treating the phytoplankton bloom as an aggregate community phenomenon.

The combined effects of warming and overwintering copepod densities on the spring succession of Baltic Sea plankton were investigated using indoor mesocosms. Three zooplankton (1.5, 4 and 10 copepods L-1) and two temperature levels called ∆0°C and ∆6°C (0°C and 6°C above the present day temperature scenario for Kiel Bight) were chosen. Both, the timing and the duration of the protozooplankton (PZP) bloom were significantly affected by temperature, but not by copepod density. In contrast, the bloom intensity of PZP was highly affected by the factors temperature and copepod density and its interaction. This suggests that at elevated temperature conditions PZP grows faster but, at the same time, are subject to higher top-down control by copepods. At low temperatures and low copepod densities, PZP in turn fully escaped from copepod predation. Further changes in the overwintering copepod densities resulted in a strong ciliate suppression of which small-sized ciliates (

Ecological research in recent decades revealed that species loss has a predominantly negative effect on ecosystem functioning and stability. Most of these studies were based on random species loss scenarios, but extinctions in nature are not random. Recent experimental studies using macroscopic communities largely advanced knowledge about the effects of non-random species loss. However, in microscopic communities like the phytoplankton, implementing realistic species loss scenarios is challenging and experimental data are scarce. Creating more realistic experiments to study the role of phytoplankton diversity for ecosystem functioning is particularly important, as they provide up to 50% of global primary productivity, form the basis of all pelagic food webs, and are important for biogeochemical cycling. In this study, we experimentally tested and evaluated three methods for non-random species loss in a natural marine phytoplankton community. Dilution, filtration, and heat stress removed the targeted rare, large, and sensitive species, respectively. All these species groups are extremely vulnerable to extinction in future climate scenarios and play important roles in the communities. Dilution and filtration with a fine mesh additionally decreased initial biomass, which increased the variability of species left in the respective replicates. The methods tested in this study can be used to non-randomly manipulate phytoplankton species diversity in communities used for experiments. However, in studies where species identities are more important than species richness, the dilution and filtration methods should be modified to eliminate the effect of decreasing initial biomass.

In this chapter, the effects of temperature change—as a main aspect of climate change—on marine biodiversity are assessed. Starting from a general discussion of species responses to temperature, the chapter presents how species respond to warming. These responses comprise adaptation and phenotypic plasticity as well as range shifts. The observed range shifts show more rapid shifts at the poleward range edge than at the equator-near edge, which probably reflects more rapid immigration than extinction in a warming world. A third avenue of changing biodiversity is change in species interactions, which can be altered by temporal and spatial shifts in interacting species. We then compare the potential changes in biodiversity to actual trends recently addressed in empirical synthesis work on local marine biodiversity, which lead to conceptual issues in quantifying the degree of biodiversity change. Finally we assess how climate change impacts the protection of marine environments.

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.

Numerous studies show that increasing species richness leads to higher ecosystem productivity. This effect is often attributed to more efficient portioning of multiple resources in communities with higher numbers of competing species, indicating the role of resource supply and stoichiometry for biodiversity–ecosystem functioning relationships. Here, we merged theory on ecological stoichiometry with a framework of biodiversity–ecosystem functioning to understand how resource use transfers into primary production. We applied a structural equation model to define patterns of diversity–productivity relationships with respect to available resources. Meta-analysis was used to summarize the findings across ecosystem types ranging from aquatic ecosystems to grasslands and forests. As hypothesized, resource supply increased realized productivity and richness, but we found significant differences between ecosystems and study types. Increased richness was associated with increased productivity, although this effect was not seen in experiments. More even communities had lower productivity, indicating that biomass production is often maintained by a few dominant species, and reduced dominance generally reduced ecosystem productivity. This synthesis, which integrates observational and experimental studies in a variety of ecosystems and geographical regions, exposes common patterns and differences in biodiversity–functioning relationships, and increases the mechanistic understanding of changes in ecosystems productivity.

A comparative analysis of data, obtained during four indoor-mesocosm experiments with natural spring plankton communities from the Baltic Sea, was conducted to investigate whether biogeochemical cycling is affected by an increase in water temperature of up to 6 °C above present-day conditions. In all experiments, warming stimulated in particular heterotrophic bacterial processes and had an accelerating effect on the temporal development of phytoplankton blooms. This was also mirrored in the build-up and partitioning of organic matter between particulate and dissolved phases. Thus, warming increased both the magnitude and rate of dissolved organic carbon (DOC) build-up, whereas the accumulation of particulate organic carbon (POC) and phosphorus (POP) decreased with rising temperature. In concert, the observed temperature-mediated changes in biogeochemical components suggest strong shifts in the functioning of marine pelagic food webs and the ocean’s biological carbon pump, hence providing potential feedback mechanisms to Earth’s climate system.

Indoor mesocosm experiments were conducted to test for potential climate change effects on the spring succession of Baltic Sea plankton. Two different temperature (Delta 0 A degrees C and Delta 6 A degrees C) and three light scenarios (62, 57 and 49 % of the natural surface light intensity on sunny days), mimicking increasing cloudiness as predicted for warmer winters in the Baltic Sea region, were simulated. By combining experimental and modeling approaches, we were able to test for a potential dietary mismatch between phytoplankton and zooplankton. Two general predator-prey models, one representing the community as a tri-trophic food chain and one as a 5-guild food web were applied to test for the consequences of different temperature sensitivities of heterotrophic components of the plankton. During the experiments, we observed reduced time-lags between the peaks of phytoplankton and protozoan biomass in response to warming. Microzooplankton peak biomass was reached by 2.5 day A degrees C-1 earlier and occurred almost synchronously with biomass peaks of phytoplankton in the warm mesocosms (Delta 6 A degrees C). The peak magnitudes of microzooplankton biomass remained unaffected by temperature, and growth rates of microzooplankton were higher at Delta 6 A degrees C (mu(a dagger 0 A degrees C) = 0.12 day(-1) and mu(a dagger 6 A degrees C) = 0.25 day(-1)). Furthermore, warming induced a shift in microzooplankton phenology leading to a faster species turnover and a shorter window of microzooplankton occurrence. Moderate differences in the light levels had no significant effect on the time-lags between autotrophic and heterotrophic biomass and on the timing, biomass maxima and growth rate of microzooplankton biomass. Both models predicted reduced time-lags between the biomass peaks of phytoplankton and its predators (both microzooplankton and copepods) with warming. The reduction of time-lags increased with increasing Q(10) values of copepods and protozoans in the tritrophic food chain. Indirect trophic effects modified this pattern in the 5-guild food web. Our study shows that instead of a mismatch, warming might lead to a stronger match between protist grazers and their prey altering in turn the transfer of matter and energy toward higher trophic levels.

Through the use of mesocosm experiments, we show that an unusually early spring phytoplankton bloom can be induced by intermittent high-light periods. We performed mesocosm experiments where plankton assemblages from Kiel Bight (Western Baltic Sea) received a light regime based on the natural seasonal irradiance dimmed to 43% of surface irradiance of cloudless days, starting with irradiance levels of mid-January (6 mesocosms) and mid-February (6 mesocosms). After 6 d, half of the mesocosms received a ca. 2-fold increase in irradiance. In the January mesocosms, a phytoplankton bloom developed only in the treatments with the high-light episode, whereas in the February mesocosms a phytoplankton bloom also developed in the controls. Phytoplankton net growth rates, production:biomass ratios and biomass at the end of the high irradiance episodes were positively correlated to the daily light dose. The relative biomass of diatoms increased with increasing light, whereas the relative biomass of cryptophytes decreased. A bottom-up transmission to mesozooplankton (mainly copepods of the genera Acartia and Oithona) was evident by increased densities of copepod nauplii and egg production under higher light conditions, whereas copepodids and adults showed no responses during the experimental period. The taxonomic composition of the nauplii was shifted to the advantage of Acartia/Centropages (not distinguished at the naupliar stage) under higher light conditions.

Indoor mesocosms were used to study the combined effect of warming and of different densities of overwintering mesozooplankton (mainly copepods) on the spring development of phytoplankton in shallow, coastal waters. Similar to previous studies, warming accelerated the spring phytoplankton peak by ca. 1 day 1C1 whereas zooplankton did not significantly influence timing. Phytoplankton biomass during the experimental period decreased with warming and with higher densities of overwintering zooplankton. Similarly, average cell size and average effective particle size (here: colony size) decreased both with zooplankton density and warming. A decrease in phytoplankton particle size is generally considered at typical footprint of copepod grazing. We conclude that warming induced changes in the magnitude and structure of the phytoplankton spring bloom cannot be understood without considering grazing by overwintering zooplankton.

Rising ocean temperature is expected to change the balance between production and degradation of organic matter due to different temperature sensitivities of auto- and heterotrophic processes. Copepods are the most prominent zooplankton group, and elevated temperature increases their growth and grazing rates. So far, it is unknown to what extent copepods affect the partitioning and stoichiometry of organic matter in a warmer surface ocean. We therefore conducted a mesocosm experiment with 3 copepod densities and 2 temperature scenarios to determine effects on the pools of dissolved and particulate organic matter and their C:N:P ratios. Here we show that particulate organic C (POC) concentrations decreased with increasing copepod abundance. This effect was more pronounced at elevated temperature, yielding a decrease in the POC to particulate nitrogen ratio (POC:PN) from 26 to 13 and in the POC:particulate organic phosphorus (POP) ratio from 567 to 257, from low to high copepod density. Dissolved organic carbon (DOC) accumulation was positively affected by temperature. However, increasing copepod abundance decreased the accumulation of DOC at elevated temperature. Copepod grazing and egestion enhanced the recycling of N and P, thereby increasing the availability of these nutrients for autotrophs. In concert with temperature-induced shifts in the phytoplankton community composition and size, changes in copepod abundance may therefore have contributed to altering the elemental composition of seston. Our findings suggest combined effects of zooplankton grazing and temperature on the composition and recycling of organic matter that should be taken into account when simulating biogeochemical cycles in a future ocean.

Ocean warming has been implicated in the observed decline of oceanic phytoplankton biomass. Some studies suggest a physical pathway of warming via stratification and nutrient flux, and others a biological effect on plankton metabolic rates; yet the relative strength and possible interaction of these mechanisms remains unknown. Here, we implement projections from a global circulation model in a mesocosm experiment to examine both mechanisms in a multi-trophic plankton community. Warming treatments had positive direct effects on phytoplankton biomass, but these were overcompensated by the negative effects of decreased nutrient flux. Zooplankton switched from phytoplankton to grazing on ciliates. These results contrast with previous experiments under nutrient-replete conditions, where warming indirectly reduced phytoplankton biomass via increased zooplankton grazing. We conclude that the effect of ocean warming on marine plankton depends on the nutrient regime, and provide a mechanistic basis for understanding global change in marine ecosystems.

Ecosystem functioning is simultaneously affected by changes in community composition and environmental change such as increasing atmospheric carbon dioxide (CO2 ) and subsequent ocean acidification. However, it largely remains uncertain how the effects of these factors compare to each other. Addressing this question, we experimentally tested the hypothesis that initial community composition and elevated CO2 are equally important to the regulation of phytoplankton biomass. We full-factorially exposed three compositionally different marine phytoplankton communities to two different CO2 levels and examined the effects and relative importance (ω(2) ) of the two factors and their interaction on phytoplankton biomass at bloom peak. The results showed that initial community composition had a significantly greater impact than elevated CO2 on phytoplankton biomass, which varied largely among communities. We suggest that the different initial ratios between cyanobacteria, diatoms, and dinoflagellates might be the key for the varying competitive and thus functional outcome among communities. Furthermore, the results showed that depending on initial community composition elevated CO2 selected for larger sized diatoms, which led to increased total phytoplankton biomass. This study highlights the relevance of initial community composition, which strongly drives the functional outcome, when assessing impacts of climate change on ecosystem functioning. In particular, the increase in phytoplankton biomass driven by the gain of larger sized diatoms in response to elevated CO2 potentially has strong implications for nutrient cycling and carbon export in future oceans.

Shifts in the timing and magnitude of the spring plankton bloom in response to climate change have been observed across a wide range of aquatic systems. We used meta-analysis to investigate phenological responses of marine and freshwater plankton communities in mesocosms subjected to experimental manipulations of temperature and light intensity. Systems differed with respect to the dominant mesozooplankton (copepods in seawater and daphnids in freshwater). Higher water temperatures advanced the bloom timing of most functional plankton groups in both marine and freshwater systems. In contrast to timing, responses of bloom magnitudes were more variable among taxa and systems and were influenced by light intensity and trophic interactions. Increased light levels increased the magnitude of the spring peaks of most phytoplankton taxa and of total phytoplankton biomass. Intensified size-selective grazing of copepods in warming scenarios affected phytoplankton size structure and lowered intermediate (20–200 μm)-sized phytoplankton in marine systems. In contrast, plankton peak magnitudes in freshwater systems were unaffected by temperature, but decreased at lower light intensities, suggesting that filter feeding daphnids are sensitive to changes in algal carrying capacity as mediated by light supply. Our analysis confirms the general shift toward earlier blooms at increased temperature in both marine and freshwater systems and supports predictions that effects of climate change on plankton production will vary among sites, depending on resource limitation and species composition.

The response of the Baltic Sea spring bloom was studied in mesocosm experiments, where temperatures were elevated up to 6°C above the present-day sea surface temperature of the spring bloom season. Four of the seven experiments were carried out at different light levels (32–202 Wh m −2 at the start of the experiments) in the different experimental years. In one further experiment, the factors light and temperature were crossed, and in one experiment, the factors density of overwintering zooplankton and temperature were crossed. Overall, there was a slight temporal acceleration of the phytoplankton spring bloom, a decline of peak biomass and a decline of mean cell size with warming. The temperature influence on phytoplankton bloom timing, biomass and size structure was qualitatively highly robust across experiments. The dependence of timing, biomass, and size structure on initial conditions was tested by multiple regression analysis of the y-temperature regressions with the candidate independent variables initial light, initial phytoplankton biomass, initial microzooplankton biomass, and initial mesozooplankton (=copepod) biomass. The bloom timing predicted for mean temperatures (5.28°C) depended on light. The peak biomass showed a strong positive dependence on light and a weaker negative dependence on initial copepod density. Mean phytoplankton cell size predicted for the mean temperature responded positively to light and negatively to copepod density. The anticipated mismatch between phytoplankton supply and food demand by newly hatched copepod nauplii occurred only under the combination of low light and warm temperatures. The analysis presented here confirms earlier conclusions about temperature responses that are based on subsets of our experimental series. However, only the comprehensive analysis across all experiments highlights the importance of the factor light.

We examined the simultaneous effect of climate warming and light availability on the phytoplankton spring bloom using 1400 l (1 m depth) indoor mesocosms. The timing of the spring bloom was advanced both by warming and higher light intensity, but the influence of temperature on the phytoplankton community was stronger than the light effect. Warming affected phytoplankton directly and indirectly via enhanced grazing pressure at higher temperatures. Warming resulted in markedly lower phytoplankton biomass and a shift towards smaller cell sizes. It also led to changes in the community structure of phytoplankton and zooplankton. Among phytoplankton, large-celled diatoms were most negatively affected by warming. Overwintering zooplankton species (Oithona, Pseudocalanus) remained dominant in the cold treatments, while they were replaced by late spring or summer species (Acartia, Centropages, Temora) in the warmed treatments. Our results show that understanding food web interactions might be very important to the study of the effects of climate warming on pelagic ecosystems.