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Dominance by cyanobacteria hampers human use of lakes and reservoirs worldwide. Previous studies indicate that excessive nutrient loading and warmer conditions promote dominance by cyanobacteria, but evidence from global scale field data has so far been scarce. Our analysis, based on a study of 143 lakes along a latitudinal transect ranging from subarctic Europe to southern South America, shows that although warmer climates do not result in higher overall phytoplankton biomass, the percentage of the total phytoplankton biovolume attributable to cyanobacteria increases steeply with temperature. Our results also reveal that the percent cyanobacteria is greater in lakes with high rates of light absorption. This points to a positive feedback because restriction of light availability is often a consequence of high phytoplankton biovolume, which in turn may be driven by nutrient loading. Our results indicate a synergistic effect of nutrients and climate. The implications are that in a future warmer climate, nutrient concentrations may have to be reduced substantially from present values in many lakes if cyanobacterial dominance is to be controlled.

1.A logical way of distinguishing functional groups of phytoplankton is to cluster species according to their functional traits, such as growth rate and nutrient assimilation constants. However, data for such an approach are lacking for the vast majority of the species. 2. In this study, we show that a classification based on simple morphological traits may capture much of the variability in functional properties among the phytoplankton. We used information on more than 700 freshwater species, from more than 200 lakes situated in climate zones ranging from subpolar to tropical. 3. Morphological characteristics correlated well with functional properties, such as growth rate and sinking rate, and also with the population size and biomass attained in the field. This suggests that morphology is a good predictor of the functional characteristics of species. 4. Cluster analysis was used to define seven species groups based on morphology. Although some of the clusters are taxonomically homogeneous, others include species of several separate divisions. Functional traits (not used for the classification) differed significantly among the clusters, suggesting that the clusters may indeed represent meaningful functional groups. 5. Advantages of our morphological approach to classification include its objectivity, its independence from taxonomic affiliations, and the relative ease of its application to the majority of species for which physiological traits are unknown and are not readily determined.

Summary 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.

Summary 1. The hypothesis that cyanobacteria have higher optimum growth temperatures and higher growth rates at the optimum as compared to chlorophytes was tested by running a controlled experiment with eight cyanobacteria species and eight chlorophyte species at six different temperatures (20–35 °C) and by performing a literature survey. 2. In the experiment, all organisms except the chlorophyte Monoraphidium minutum grew well up to 35 °C. The chlorophyte Chlamydomonas reinhardtii was the fastest‐growing organism over the entire temperature range (20–35 °C). 3. Mean optimum growth temperatures were similar for cyanobacteria (29.2 °C) and chlorophytes (29.2 °C). These results are concordant with published data, yielding slightly higher mean optimum growth temperatures for cyanobacteria (27.2 °C) than for chlorophytes (26.3 °C). 4. Mean growth rates of cyanobacteria at 20 °C (0.42 day−1) were significantly lower than those of chlorophytes at 20 °C (0.62 day−1). However, at all other temperatures, there were no differences between mean growth rates of cyanobacteria and chlorophytes. 5. Mean growth rates at the optimum temperature were similar for cyanobacteria (0.92 day−1) and chlorophytes (0.96 day−1). However, analysis of published data revealed that growth rates of cyanobacteria (0.65 day−1) were significantly lower than those of chlorophytes (0.93 day−1) at their optimum temperatures. 6. Although climate warming will probably lead to an intensification of cyanobacterial blooms, our results indicate that this might not be as a result of higher growth rates of cyanobacteria compared with their chlorophyte competitors. The competitive advantage of cyanobacteria can more likely be attributed to their ability to migrate vertically and prevent sedimentation in warmer and more strongly stratified waters and to their resistance to grazing, especially when warming reduces zooplankton body size.

The cyanobacteria Planktothrix agardhii and Cylindrospermopsis raciborskii are bloom-forming species common in eutrophic freshwaters. These filamentous species share certain physiological traits which imply that they might flourish under similar environmental conditions. We compared the distribution of the two species in a large database (940 samples) covering different climatic regions and the Northern and Southern hemispheres, and carried out laboratory experiments to compare their morphological and physiological responses. The environmental ranges of the two species overlapped with respect to temperature, light and total phosphorus (TP); however, they responded differently to environmental gradients; C. raciborskii biovolume changed gradually while P. agardhii shifted sharply from being highly dominated to a rare component of the phytoplankton. As expected, P. agardhii dominates the phytoplankton with high TP and low light availability conditions. Contrary to predictions, C. raciborskii succeeded in all climates and at temperatures as low as 11 °C. Cylindrospermopsis raciborskii had higher phenotypic plasticity than P. agardhii in terms of pigments, individual size and growth rates. We conclude that the phenotypic plasticity of C. raciborskii could explain its ongoing expansion to temperate latitudes and suggest its future predominance under predicted climate-change scenarios.

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