SummaryChytridiomycota, often referred to as chytrids, can be virulent parasites with the potential to inflict mass mortalities on hosts, causing e.g. changes in phytoplankton size distributions and succession, and the delay or suppression of bloom events. Molecular environmental surveys have revealed an unexpectedly large diversity of chytrids across a wide range of aquatic ecosystems worldwide. As a result, scientific interest towards fungal parasites of phytoplankton has been gaining momentum in the past few years. Yet, we still know little about the ecology of chytrids, their life cycles, phylogeny, host specificity and range. Information on the contribution of chytrids to trophic interactions, as well as coevolutionary feedbacks of fungal parasitism on host populations is also limited. This paper synthesizes ideas stressing the multifaceted biological relevance of phytoplankton chytridiomycosis, resulting from discussions among an international team of chytrid researchers. It presents our view on the most pressing research needs for promoting the integration of chytrid fungi into aquatic ecology.

We studied the effects of water column mixing depth and background turbidity on phytoplankton biomass, light climate, and nutrients in two field enclosure experiments designed to test predictions of a dynamical model. In 1997 and 1998, we created gradients of mixing depth by enclosing the 100-m-filtered phytoplankton community of a phosphorus-deficient lake in cylindrical plastic bags of varying depth (1.5-15 m) which were continuously mixed. To mimic different levels of background turbidity, we surrounded the transparent enclosure walls with a layer of opaque white (1997) or black (1998) plastic. The experiments were run for 4 wk (1997) and 6 wk (1998). The results supported two key assumptions of the model: specific production and specific sedimentation losses both decreased with increasing mixing depth. At all mixing depths, fast-sinking diatoms dominated the communities. In accordance with model predictions, algal biomass concentration and standing stock (summed over the mixed layer) showed a unimodal relationship to mixing depth when background turbidity was high (1998). When background turbidity was lower (1997), only the ascending limbs of the corresponding relationships were found, which supports the prediction that the mixing depth at which biomass peaks (i.e., becomes predominantly limited by light) increases with decreasing background turbidity. Also in accordance with predictions, light intensity at the bottom of the mixed layer decreased with increasing mixing depth and with increasing background turbidity. Finally, the data supported only the ascending limbs of the predicted inverse unimodal relationships among mixing depth and dissolved inorganic and total water column phosphorus. The absence of descending limbs in these relationships at low mixing depths was probably due to deviations of the experimental systems from two model assumptions. First, the remineralization rate of sedimented phosphorus may have been too slow to equilibrate with sedimentation losses over the experimental periods. Second, biomass yield per unit nutrient (the ratio of seston carbon to phosphorus) was not constant, but decreased with increasing mixing depth. To our knowledge, these are the first field experiments in which the effects of mixing depth on phytoplankton and its resources have been investigated systematically along a large gradient.

In many lakes, the most conspicuous seasonal events are the phytoplankton spring bloom and the subsequent clear-water phase, a period of low-phytoplankton biomass that is frequently caused by mesozooplankton (Daphnia) grazing. In Central European lakes, the timing of the clear-water phase is linked to large-scale climatic forcing, with warmer winters being followed by an earlier onset of the clear-water phase. Mild winters may favour an early build-up of Daphnia populations, both directly through increased surface temperatures and indirectly by reducing light limitation and enhancing algal production, all being a consequence of earlier thermal stratification. We conducted a field experiment to disentangle the separate impacts of stratification depth (affecting light supply) and temperature on the magnitude and timing of successional events in the plankton. We followed the dynamics of the phytoplankton spring bloom, the clear-water phase and the spring peak in Daphnia abundance in response to our experimental manipulations. Deeper mixing delayed the timing of all spring seasonal events and reduced the magnitudes of the phytoplankton bloom and the subsequent Daphnia peak. Colder temperatures retarded the timing of the clear-water phase and the subsequent Daphnia peak, whereas the timing of the phytoplankton peak was unrelated to temperature. Most effects of mixing depth (light) and temperature manipulations were independent, effects of mixing depth being more prevalent than effects of temperature. Because mixing depth governs both the light climate and the temperature regime in the mixed surface layer, we propose that climate-driven changes in the timing and depth of water column stratification may have far-reaching consequences for plankton dynamics and should receive increased attention.

We studied the effects of water column mixing depth and background turbidity on phytoplankton biomass, light climate, and nutrients in two field enclosure experiments designed to test predictions of a dynamical model. In 1997 and 1998, we created gradients of mixing depth by enclosing the 100-m-filtered phytoplankton community of a phosphorus-deficient lake in cylindrical plastic bags of varying depth (1.5-15 m) which were continuously mixed. To mimic different levels of background turbidity, we surrounded the transparent enclosure walls with a layer of opaque white (1997) or black (1998) plastic. The experiments were run for 4 wk (1997) and 6 wk (1998). The results supported two key assumptions of the model: specific production and specific sedimentation losses both decreased with increasing mixing depth. At all mixing depths, fast-sinking diatoms dominated the communities. In accordance with model predictions, algal biomass concentration and standing stock (summed over the mixed layer) showed a unimodal relationship to mixing depth when background turbidity was high (1998). When background turbidity was lower (1997), only the ascending limbs of the corresponding relationships were found, which supports the prediction that the mixing depth at which biomass peaks (i.e., becomes predominantly limited by light) increases with decreasing background turbidity. Also in accordance with predictions, light intensity at the bottom of the mixed layer decreased with increasing mixing depth and with increasing background turbidity. Finally, the data supported only the ascending limbs of the predicted inverse unimodal relationships among mixing depth and dissolved inorganic and total water column phosphorus. The absence of descending limbs in these relationships at low mixing depths was probably due to deviations of the experimental systems from two model assumptions. First, the remineralization rate of sedimented phosphorus may have been too slow to equilibrate with sedimentation losses over the experimental periods. Second, biomass yield per unit nutrient (the ratio of seston carbon to phosphorus) was not constant, but decreased with increasing mixing depth. To our knowledge, these are the first field experiments in which the effects of mixing depth on phytoplankton and its resources have been investigated systematically along a large gradient.

According to a recent dynamical model, the depth of a well-mixed water column should have contrasting effects on the abundances of sinking and nonsinking phytoplankton taxa. Because of increasing light limitation, nonsinking taxa should decline monotonically with increasing mixing depth, and because of sinking loss limitation at low mixing depths, sinking taxa should peak at intermediate mixing depths. Along a gradient of mixing depths, the position of this maximum should increase with increasing taxon-specific sinking velocity and decrease with increasing background turbidity. In two field-enclosure experiments, we investigated the effects of mixing depth and background turbidity on a variety of sinking and nonsinking phytoplankton taxa. We exposed the natural, 100-mscreened phytoplankton community of a clear, unproductive, but silica-rich lake to a gradient of mixing depths (1.5-15 m) during 4-6 weeks. To mimic two different background turbidities, the transparent enclosure walls were surrounded by either white or black foliage. Although diatoms suffered from high sedimentation losses at low mixing depths, they dominated biomass at all mixing depths throughout both experiments. Results were largely in accordance with model predictions. Specific gross growth rates of most common taxa were negatively related to mixing depth. In both experiments, the abundances of most sinking taxa showed a unimodal pattern along the mixing depth gradient, while two of three motile taxa declined monotonically with mixing depth. The depths where these taxa reached their maximal abundances were positively related to taxon-specific sinking velocity and negatively related to background turbidity.

In deep temperate lakes, the beginning of the growing season is triggered by thermal stratification, which alleviates light limitation of planktonic producers in the surface layer and prevents heat loss to deeper strata. The sequence of subsequent phenological events (phytoplankton spring bloom, grazer peak, clearwater phase) results in part from coupled phytoplankton-grazer interactions. Disentangling the separate, direct effects of correlated climatic drivers (stratificationdependent underwater light climate vs. water temperature) from their indirect effects mediated through trophic feedbacks is impossible using observational field data, which challenges our understanding of global warming effects on seasonal plankton dynamics. We therefore manipulated water temperature and stratification depth independently in experimental field mesocosms containing ambient microplankton and inocula of the resident grazer Daphnia hyalina. Higher light availability in shallower surface layers accelerated primary production, warming accelerated consumption and growth of Daphnia, and both factors speeded up successional dynamics driven by trophic feedbacks. Specifically, phytoplankton peaked and decreased earlier and Daphnia populations increased and peaked earlier at both shallower stratification and higher temperature. The timing of ciliate dynamics was unrelated to both factors. Volumetric peak densities of phytoplankton, ciliates and Daphnia in the surface layer were also unaffected by temperature but declined with stratification depth in parallel with light availability. The latter relationship vanished, however, when population sizes were integrated over the entire water column. Overall our results suggest that, integrated over the entire water column of a deep lake, surface warming and shallower stratification independently speed up spring successional events, whereas the magnitudes of phytoplankton and zooplankton spring peaks are less sensitive to these factors. Therefore, accelerated dynamics under warming need not lead to a trophic mismatch (given similar grazer inocula at the time of stratification). We emphasize that entire water column dynamics must be studied to estimate global warming effects on lake ecosystems.

We derive from a dynamic model that light availability, phytoplankton density, and the carbon : nutrient ratio of phytoplankton biomass should all be negatively related to mixed surface layer depth, whereas the areal standing stock of phytoplankton should show a unimodal, and total and dissolved nutrients a horizontal or increasing, relationship to mixing depth. These predictions agree closely with data from 65 central European lakes during summer stratification. In addition, zooplankton biomass was strongly negatively related to mixing depth in a subset of lakes. A decrease in mixing depth is thus a form of enrichment with light of the mixed surface layer, the effects of which could propagate to higher trophic levels.

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.Climatic warming is a major driver of change in ecosystems worldwide. Shifts in species ranges, phenology, and relative abundances, with resulting changes in species interactions, are predicted to alter local biodiversity within ecosystems (Dawson et al. 2011) OIKOSA global research priority is to understand the consequences of climate change for biodiversity. A growing number of experimental studies have manipulated climatic drivers, in particular changes in temperature, in local communities. In the first quantitative meta-analysis of community-level studies across freshwater, marine and terrestrial experiments, species richness declined consistently with experimental warming. This effect was insensitive to warming intensity, duration, and multiple environmental and procedural covariates. However, evenness responses were weakly negative only in terrestrial systems and more variable across ecosystem types. Linear mixed model analyses ...