Description: 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 lm)-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.

Description: 1. Different components of the climate system have been shown to affect temporal dynamics in natural plankton communities on scales varying from days to years. The seasonal dynamics in temperate lake plankton communities, with emphasis on both physical and biological forcing factors, were captured in the 1980s in a conceptual framework, the Plankton Ecology Group (PEG) model. 2. Taking the PEG model as our starting point, we discuss anticipated changes in seasonal and long-term plankton dynamics and extend this model to other climate regions, particularly polar and tropical latitudes. Based on our improved post-PEG understanding of plankton dynamics, we also evaluate the role of microbial plankton, parasites and fish in governing plankton dynamics and distribution. 3. In polar lakes, there is usually just a single peak in plankton biomass in summer. Lengthening of the growing season under warmer conditions may lead to higher and more prolonged phytoplankton productivity. Climate-induced increases in nutrient loading in these oligotrophic waters may contribute to higher phytoplankton biomass and subsequent higher zooplankton and fish productivity. 4. In temperate lakes, a seasonal pattern with two plankton biomass peaks – in spring and summer – can shift to one with a single but longer and larger biomass peak as nutrient loading increases, with associated higher populations of zooplanktivorous fish. Climate change will exacerbate these trends by increasing nutrient loading through increased internal nutrient inputs (due to warming) and increased catchment inputs (in the case of more precipitation). 5. In tropical systems, temporal variability in precipitation can be an important driver of the seasonal development of plankton. Increases in precipitation intensity may reset the seasonal dynamics of plankton communities and favour species adapted to highly variable environments. The existing intense predation by fish on larger zooplankters may increase further, resulting in a perennially low zooplankton biomass. 6. Bacteria were not included in the original PEG model. Seasonally, bacteria vary less than the phytoplankton but often follow its patterns, particularly in colder lakes. In warmer lakes, and with future warming, a greater influx of allochthonous carbon may obscure this pattern. 7. Our analyses indicate that the consequences of climate change for plankton dynamics are, to a large extent, system specific, depending on characteristics such as food-web structure and nutrient loading. Indirect effects through nutrient loading may be more important than direct effects of temperature increase, especially for phytoplankton. However, with warming a general picture emerges of increases in bacterivory, greater cyanobacterial dominance and smaller-bodied zooplankton that are more heavily impacted by fish predation.

Description: Changing seawater chemistry towards reduced pH as a result of increasing atmospheric carbon dioxide (CO2) is affecting oceanic organisms, particularly calcifying species. Responses of non-calcifying consumers are highly variable and mainly mediated through indirect ocean acidification effects induced by changing the biochemical content of their prey, as shown within single species and simple 2-trophic level systems. However, it can be expected that indirect CO2 impacts observed at the single species level are compensated at the ecosystem level by species richness and complex trophic interactions. A dampening of CO2-effects can be further expected for coastal communities adapted to strong natural fluctuations in pCO2, typical for productive coastal habitats. Here we show that a plankton community of the Kiel Fjord was tolerant to CO2 partial pressure (pCO2) levels projected for the end of this century (〈1400 µatm), and only subtle differences were observed at the extremely high value of 4000 µatm. We found similar phyto- and microzooplankton biomass and copepod abundance and egg production across all CO2 treatment levels. Stoichiometric phytoplankton food quality was minimally different at the highest pCO2 treatment, but was far from being potentially limiting for copepods. These results are in contrast to studies that include only a single species, which observe strong indirect CO2 effects for herbivores and suggest limitations of biological responses at the level of organism to community. Although this coastal plankton community was highly tolerant to high fluctuations in pCO2, increase in hypoxia and CO2 uptake by the ocean can aggravate acidification and may lead to pH changes outside the range presently experienced by coastal organisms.

Description: This article serves as an introduction to this special issue of Marine Biology, but also as a review of the key findings of the AQUASHIFT research program which is the source of the articles published in this issue. AQUASHIFT is an interdisciplinary research program targeted to analyze the response of temperate zone aquatic ecosystems (both marine and freshwater) to global warming. The main conclusions of AQUASHIFT relate to (a) shifts in geographic distribution, (b) shifts in seasonality, (c) temporal mismatch in food chains, (d) biomass responses to warming, (e) responses of body size, (f) harmful bloom intensity, (f), changes of biodiversity, and (g) the dependence of shifts to temperature changes during critical seasonal windows.

Description: The seasonal succession of plankton is an annually repeated process of community assembly during which all major external factors and internal interactions shaping communities can be studied. A quarter of a century ago, the state of this understanding was described by the verbal plankton ecology group (PEG) model. It emphasized the role of physical factors, grazing and nutrient limitation for phytoplankton, and the role of food limitation and fish predation for zooplankton. Although originally targeted at lake ecosystems, it was also adopted by marine plankton ecologists. Since then, a suite of ecological interactions previously underestimated in importance have become research foci: overwintering of key organisms, the microbial food web, parasitism, and food quality as a limiting factor and an extended role of higher order predators. A review of the impact of these novel interactions on plankton seasonal succession reveals limited effects on gross seasonal biomass patterns, but strong effects on species replacements.

Description: While there is a general sense that lakes can act as sentinels of climate change, their efficacy has not been thoroughly analyzed. We identified the key response variables within a lake that act as indicators of the effects of climate change on both the lake and the catchment. These variables reflect a wide range of physical, chemical, and biological responses to climate. However, the efficacy of the different indicators is affected by regional response to climate change, characteristics of the catchment, and lake mixing regimes. Thus, particular indicators or combinations of indicators are more effective for different lake types and geographic regions. The extraction of climate signals can be further complicated by the influence of other environmental changes, such as eutrophication or acidification, and the equivalent reverse phenomena, in addition to other land-use influences. In many cases, however, confounding factors can be addressed through analytical tools such as detrending or filtering. Lakes are effective sentinels for climate change because they are sensitive to climate, respond rapidly to change, and integrate information about changes in the catchment.

Description: Dynamics of prokaryotic and eukaryotic picophytoplankton were investigated over a 2-year time period using flow cytometry and combined with an in situ experiment in Lake Tahoe, USA to better characterize to which extent environmental factors control these communities. Pronounced seasonal patterns and clear temporal and spatial partitioning were observed between picocyanobacteria and picoeukaryotes. Picocyanobacteria dominated in the nutrient deficient upper water column during the stratified season, while picoeukaryotes reached maximum abundance during isothermal conditions and maintained high numbers in deep-water layers during the stratified season. Picocyanobacteria were more sensitive to high solar and UV radiation compared with picoeukaryotes, which were not affected by high solar radiation and nutrient enrichment stimulated their growth. The opposing response of these two populations is consistent with their vertical distribution: picocyanobacteria dominate below the 30% isolume and above the nitrocline depth, whereas picoeukaryotes increase in the vicinity of the nitrocline and thus increased nutrient concentration. This spatial separation of picophytoplankton groups along environmental gradients in Lake Tahoe is consistent with other deep-oligotrophic lakes and the marine environment, suggesting that these marine and freshwater organisms have similar ecophysiological requirements. These results highlight that the smallest photosynthetic communities show taxon-specific responses to mixing and resource availability, which affect the structure and dynamics of picophytoplankton.

Description: Lake Tahoe is an ultra-oligotrophic subalpine lake that is renowned for its clarity. The region experiences little cloud cover and is one of the most UV transparent lakes in the world. As such, it is an ideal environment to study the role of UV radiation in aquatic ecosystems. Long-term trends in Secchi depths showed that water transparency to visible light has decreased in recent decades, but limited data are available on the UV transparency of the lake. Here we examine how ultraviolet radiation varies relative to longer-wavelength photosynthetically active radiation (PAR, 400-700 nm, visible wavelengths) horizontally along inshore-offshore transects in the lake and vertically within the water column as well as temporally throughout 2007. UV transparency was more variable than PAR transparency horizontally across the lake and throughout the year. Seasonal patterns of Secchi transparency differed from both UV and PAR, indicating that different substances may be responsible for controlling transparency to UV, PAR, and Secchi. In surface waters, UVA (380 nm) often attenuated more slowly than PAR, a pattern visible in only exceptionally transparent waters with very low dissolved organic carbon. On many sampling dates, UV transparency decreased progressively with depth suggesting surface photobleaching, reductions in particulate matter, increasing chlorophyll a, or some combination of these increased during summer months. Combining these patterns of UV transparency with data on visible light provides a more comprehensive understanding of ecosystem structure, function, and effects of environmental change in highly transparent alpine and subalpine lakes such as Tahoe.

Description: Diatoms contribute to a substantial portion of primary production in the oceans and many lakes. Owing to their relatively heavy cell walls and high nutrient requirements, planktonic diatoms are expected to decrease with climate warming because of reduced nutrient redistribution and increasing sinking velocities. Using a historical dataset, this study shows that diatoms were able to maintain their biovolume with increasing stratification in Lake Tahoe over the last decades; however, the diatom community structure changed. Increased stratification and reduced nitrogen to phosphorus ratios selected for small-celled diatoms, particularly within the Cyclotella genus. An empirical model showed that a shift in phytoplankton species composition and cell size was consistent within different depth strata, indicating that altered nutrient concentrations were not responsible for the change. The increase in small-celled species was sufficient to decrease the average diatom size and thus sinking velocity, which strongly influences energy transfer through the food web and carbon cycling. Our results show that within the diverse group of diatoms, small-sized species with a high surface area to volume ratio were able to adapt to a decrease in mixing intensity, supporting the hypotheses that abiotic drivers affect the size structure of planktonic communities and that warmer climate favours small-sized diatom cells.

Description: Life history responses are expected to accompany climate warming, yet little is known how long-term effects of climate and environmental change affect the seasonal dynamics of planktonic organisms. We used an historical data set from Lake Washington (U.S.A.) to quantify population responses of a calanoid copepod (Leptodiaptomus ashlandi) to long-term changes in temperature and resource availability and explore potential mechanisms for the responses. Increasing water temperatures (annual mean increase of 1.5 degrees C in the upper 10-m water volume) and longer stratification periods (about 4 weeks) were observed between 1962 and 2005, coincident with a pronounced decline in Leptodiaptomus densities. However, production was maintained because of an increase in the production to biomass ratio and a life cycle shift in Leptodiaptomus from an annual to a 6-month cycle. Cross-wavelet analyses demonstrated that the annual thermal forcing of copepod recruitment observed during the first two decades of the study weakened substantially, leading to more stochastic population dynamics during the past two decades. This shift from one to two generations per year was most likely produced by a longer and warmer growing period combined with changing fluctuations in resource (phytoplankton) availability. Climate change can lead to higher-frequency voltinism in ectothermic organisms and to temporal reorganization of their population dynamics.