Climate and land-use change drive a suite of stressors that shape ecosystems and interact to yield complex ecological responses, i.e. additive, antagonistic and synergistic effects.Currently we know little about the spatial scale relevant for the outcome of such interactions and about effect sizes. This knowledge gap needs to be filled to underpin future land management decisions or climate mitigation interventions, for protecting and restoring freshwater ecosystems. The study combines data across scales from 33 mesocosm experiments with those from 14 river basins and 22 cross-basin studies in Europe producing 174 combinations of paired-stressor effects on a biological response variable. Generalised linear models showed that only one of the two stressors had a significant effect in 39% of the analysed cases, 28% of the paired-stressor combinations resulted in additive and 33% in interactive (antagonistic, synergistic, opposing or reversal) effects. For lakes the frequency of additive and interactive effects was similar for all spatial scales addressed, while for rivers this frequency increased with scale. Nutrient enrichment was the overriding stressor for lakes, generally exceeding those of secondary stressors. For rivers, the effects of nutrient enrichment were dependent on the specific stressor combination and biological response variable. These results vindicate the traditional focus of lake restoration and management on nutrient stress, while highlighting that river management requires more bespoke management solutions.

The importance of nitrogen (N) versus phosphorus (P) in explaining total cyanobacterial biovolume, the biovolume of specific cyanobacterial taxa, and the incidence of cyanotoxins was determined for 102 north German lakes, using methods to separate the effects of joint variation in N and P concentration from those of differential variation in N versus P. While the positive relationship between total cyanobacteria biovolume and P concentration disappeared at high P concentrations, cyanobacteria biovolume increased continually with N concentration, indicating potential N limitation in highly P enriched lakes. The biovolumes of all cyanobacterial taxa were higher in lakes with above average joint NP concentrations, although the relative biovolumes of some Nostocales were higher in less enriched lakes. Taxa were found to have diverse responses to differential N versus P concentration, and the differences between taxa were not consistent with the hypothesis that potentially N 2 -fixing Nostocales taxa would be favoured in low N relative to P conditions. In particular Aphanizomenon gracile and the subtropical invasive species Cylindrospermopsis raciborskii often reached their highest biovolumes in lakes with high nitrogen relative to phosphorus concentration. Concentrations of all cyanotoxin groups increased with increasing TP and TN, congruent with the biovolumes of their likely producers. Microcystin concentration was strongly correlated with the biovolume of Planktothrix agardhii but concentrations of anatoxin, cylindrospermopsin and paralytic shellfish poison were not strongly related to any individual taxa. Cyanobacteria should not be treated as a single group when considering the potential effects of changes in nutrient loading on phytoplankton community structure and neither should the N 2 -fixing Nostocales. This is of particular importance when considering the occurrence of cyanotoxins, as the two most abundant potentially toxin producing Nostocales in our study were found in lakes with high N relative to P enrichment.

Summary 1.A safe, clean water supply is critical for sustaining many important ecosystem services provided by freshwaters. The development of cyanobacterial blooms in lakes and reservoirs has a major impact on the provision of these services, particularly limiting their use for recreation and water supply for drinking and spray irrigation. Nutrient enrichment is thought to be the most important pressure responsible for the widespread increase in cyanobacterial blooms in recent decades. Quantifying how nutrients limit cyanobacterial abundance in lakes is, therefore, a key need for setting robust targets for the management of freshwaters. 2. Using a data set from over 800 European lakes, we highlight the use of quantile regression modelling for understanding the maximum potential capacity of cyanobacteria in relation to total phosphorus (TP) and the use of a range of quantile responses, alongside World Health Organisation (WHO) health alert thresholds for recreational waters, for setting robust phosphorus targets for lake management in relation to water use. 3. The analysis shows that cyanobacteria exhibit a nonlinear response to phosphorus with the sharpest increase in cyanobacterial abundance occurring in the TP range from about 20 lg L À1 up to about 100 lg L À1 .4. The likelihood of exceeding the World Health Organisation (WHO) 'low health alert' threshold increases from about 5% exceedance at 16 lg L À1 to 40% exceedance at 54 lg L À1 . About 50% of the studied lakes remain below this WHO health alert threshold, irrespective of high summer TP concentrations, highlighting the importance of other factors affecting cyanobacteria population growth and loss processes, such as high flushing rate. 5. Synthesis and applications. Developing a more quantitative understanding of the effect of nutrients on cyanobacterial abundance in freshwater lakes provides important knowledge for restoring and sustaining a safe, clean water supply for multiple uses. Our models can be used to set nutrient targets to sustain recreational services and provide different levels of precaution that can be chosen dependent on the importance of the service provision.Key-words: algal bloom, blue-green algae, ecosystem services, freshwater, lake, nutrient, quantile regression, WHO *Correspondence author. E-mail: laca@ceh.ac.uk † On secondment from CEH 2 to JRC 1 .

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Data on phytoplankton, macrophytes, benthic invertebrates and fish from more than 2000 lakes in 22 European countries were used to develop and test metrics for assessing the ecological status of European lakes as required by the Water Framework Directive. The strongest and most sensitive of the 11 metrics responding to eutrophication pressure were phytoplankton chlorophyll a, a taxonomic composition trophic index and a functional traits index, the macrophyte intercalibration taxonomic composition metric and a Nordic lake fish index. Intermediate response was found for a cyanobacterial bloom intensity index (Cyano), the Ellenberg macrophyte index and a multimetric index for benthic invertebrates. The latter also responded to hydromorphological pressure. The metrics provide information on primary and secondary impacts of eutrophication in the pelagic and the littoral zone of lakes. Several of these metrics were used as common metrics in the intercalibration of national assessment systems or have been incorporated directly into the national systems. New biological metrics have been developed Hydrobiologia (2013) 704:57-74 DOI 10.1007/s10750-012-1436 to assess hydromorphological pressures, based on aquatic macrophyte responses to water level fluctuations, and on macroinvertebrate responses to morphological modifications of lake shorelines. These metrics thus enable the quantification of biological impacts of hydromorphological pressures in lakes.