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.

Abstract. Excessive amounts of nutrients and dissolved organic matter in freshwater bodies affect aquatic ecosystems. In this study, the spatial and temporal variability in nitrate (NO − 3 ), dissolved organic carbon (DOC) and soluble reactive phosphorus (SRP) was analyzed in the Selke (Germany) river continuum from three headwaters draining 1-3 km 2 catchments to two downstream reaches representing spatially integrated signals from 184-456 km 2 catchments. Three headwater catchments were selected as archetypes of the main landscape units (land use × lithology) present in the Selke catchment. Export regimes in headwater catchments were interpreted in terms of NO − 3 , DOC and SRP land-to-stream transfer processes. Headwater signals were subtracted from downstream signals, with the differences interpreted in terms of in-stream processes and contributions from point sources. The seasonal dynamics for NO − 3 were opposite those of DOC and SRP in all three headwater catchments, and spatial differences also showed NO − 3 contrasting with DOC and SRP. These dynamics were interpreted as the result of the interplay of hydrological and biogeochemical processes, for which riparian zones were hypothesized to play a determining role. In the two downstream reaches, NO − 3 was transported almost conservatively, whereas DOC was consumed and produced in the upper and lower river sections, respectively. The natural export regime of SRP in the three headwater catchments mimicked a point-source signal (high SRP during summer low flow), which may lead to overestimation of domestic contributions in the downstream reaches. Monitoring the river continuum from headwaters to downstream reaches proved effective to jointly investigate land-to-stream and instream transport, and transformation processes.

We independently manipulated mixing intensity (strong artificial mixing vs. background turbulence) and water-column depth (2 m, 4 m, 8 m, and 12 m) in order to explore their separate and combined effects in a field enclosure experiment. To accentuate the vertical light gradient, enclosures had black walls, resulting in a euphotic depth of only 3.7 m. All enclosures were placed in a well-mixed water bath to equalize temperature across treatments. Phytoplankton responded to an initial phosphorus pulse with a transient increase in biomass, which was highest in the shallowest, least light-limited water columns where dissolved mineral phosphorus subsequently became strongly limiting. As a consequence, the depth-averaged mineral phosphorus concentration increased and the seston carbon (C) : phosphorous (P) ratio decreased with increasing water-column depth. Low turbulence enclosures became quickly dominated by motile taxa (flagellates) in the upper water column, whereas mixed enclosures became gradually dominated by pennate diatoms, which resulted in higher average sedimentation rates in the mixed enclosures over the 35-d experimental period. Low turbulence enclosures showed pronounced vertical structure in water columns .4 m, where diversity was higher than in mixed enclosures, suggesting vertical niche partitioning. This interpretation is supported by a primary production assay, where phytoplankton originating from different water depths in low-turbulence treatments had the relatively highest primary productivity when incubated at their respective depths of origin.

In shallow aquatic systems, benthic and pelagic primary producers typically compete for light and nutrients along opposing vertical supply axes: pelagic algae shade the benthic habitat; conversely, benthic algae intercept the nutrient flux from the sediment to the pelagic habitat. We present a general framework for analyzing such spatially asymmetric resource competition across habitat boundaries using a mechanistic, dynamical model. We visualize the mechanisms determining the outcome of these cross-habitat interactions using zero-net-growth isoclines, resource supply points, and resource consumption vectors. In extensive invasion analyses, we characterize the abiotic and competitive persistence boundaries of pelagic and benthic primary producers, which are set by environmental factors determining nutrient and light supply and are modified by resource use by the competitor in the respective other habitat.We note several qualitative differences between cross-habitat and ''classical'' within-habitat resource competition. First, coexistence of cross-habitat competitors is facilitated by, but does not require niche differentiation with respect to, the utilization of resources. Because each species has a competitive edge for the resource that is supplied from ''its'' side of the system, a competitor that is inferior in utilizing both resources can sometimes coexist with, or even exclude, a superior competitor. Second, increasing the external supply of one resource (the nutrient) may initially favor both competitors, until a breakpoint is reached where the benthic producer goes abruptly extinct. Finally, whether a given pair of cross-habitat competitors coexist or shows alternative states may depend on the environment. Specifically, benthic and pelagic algae may coexist at low nutrient and light supply but produce alternative states at high nutrient and light supply. Alternative states are, in turn, promoted by any algal trait combination that increases the spatial asymmetry in resource consumption, i.e., leads to a higher nutrient consumption in the benthic habitat and/or a higher light consumption in the pelagic habitat.In a first empirical application, we show that predictions from our model give a good fit to published data on benthic and pelagic primary production in temperate and arctic lakes spanning a broad range of nutrient environments.

Phytoplankton-grazer dynamics are often characterized by long transients relative to the length of the growing season. Using a phytoplankton-grazer model parameterized for Daphnia pulex with either flexible or fixed algal carbon:nutrient stoichiometry, we explored how nutrient and light supply (the latter by varying depth of the mixed water column) affect the transient dynamics of the system starting from low densities. The system goes through an initial oscillation across nearly the entire light-nutrient supply space. With flexible (but not with fixed) algal stoichiometry, duration of the initial algal peak, timing and duration of the subsequent grazer peak, and timing of the algal minimum are consistently accelerated by nutrient enrichment but decelerated by light enrichment (decreasing mixing depth) over the range of intermediate to shallow mixing depths. These contrasting effects of nutrient vs. light enrichment are consequences of their opposing influences on food quality (algal nutrient content): algal productivity and food quality are positively related along a nutrient gradient but inversely related along a light gradient. Light enrichment therefore slows down grazer growth relative to algal growth, decelerating oscillatory dynamics; nutrient enrichment has opposite effects. We manipulated nutrient supply and mixing depth in a field enclosure experiment. The experimental results were qualitatively much more consistent with the flexible than with the fixed stoichiometry model. Nutrient enrichment increased Daphnia peak biomass, decreased algal minimum biomass, decreased the seston C:P ratio, and accelerated transient oscillatory dynamics. Light enrichment (decreasing mixing depth) produced the opposite patterns, except that Daphnia peak biomass increased monotonously with light enrichment, too. Thus, while the model predicts the possibility of the "paradox of energy enrichment" (a decrease in grazer biomass with light enrichment) at high light and low nutrient supply, this phenomenon did not occur in our experiment.

The Bode catchment (Germany) shows strong land use gradients from forested parts of the National Park (23% of total land cover) to agricultural (70%) and urbanised areas (7%). It is part of the Terrestrial Environmental Observatories of the German Helmholtz association. We performed a biogeochemical analysis of the entire river network. Surface water was sampled at 21 headwaters and at ten downstream sites, before (in early spring) and during the growing season (in late summer). Many parameters showed lower concentrations in headwaters than in downstream reaches, among them nutrients (ammonium, nitrate and phosphorus), dissolved copper and seston dry mass. Nitrate and phosphorus concentrations were positively related to the proportion of agricultural area within the catchment. Punctual anthropogenic loads affected some parameters such as chloride and arsenic. Chlorophyll a concentration and total phosphorus in surface waters were positively related. The concentration of dissolved organic carbon (DOC) was higher in summer than in spring, whereas the molecular size of DOC was lower in summer. The specific UV absorption at 254 nm, indicating the content of humic substances, was higher in headwaters than in downstream reaches and was positively related to the proportion of forest within the catchment. CO₂ oversaturation of the water was higher downstream compared with headwaters and was higher in summer than in spring. It was correlated negatively with oxygen saturation and positively with DOC concentration but negatively with DOC quality (molecular size and humic content). A principle component analysis clearly separated the effects of site (44%) and season (15%), demonstrating the strong effect of land use on biogeochemical parameters.

To counteract the severe consequences of eutrophication on water quality and ecosystem health, nutrient inputs have been reduced in many lakes and reservoirs during the last decades. Contrary to expectations, in some lakes phytoplankton biomass did not decrease in response to oligotrophication (nutrient reduction). The underlying mechanisms preventing a decrease in biomass in these lakes are the subject of ongoing discussion. We used a hitherto unpublished long‐term data set ranging from 1961 until 2016 from a German drinking water reservoir (Rappbode Reservoir) to investigate the underlying mechanisms preventing a decrease in biomass. Total phosphorus (TP) concentrations in the Rappbode Reservoir dropped abruptly in 1990 from 0.163 to 0.027 mg/L within three consecutive years, as a result of banning phosphate‐containing detergents. Despite substantial reductions in TP, total annual phytoplankton biomass did not decline in the long‐run, and therefore, the yield of total phytoplankton biomass per unit phosphorus largely increased. Regression analysis revealed a positive association between the yield and potentially phagotrophic mixotrophs (R2 = .465, p < .001). We infer that by ingesting bacteria, mixotrophic species were capable of exploiting additional P sources that are not accessible to obligate autotrophic phytoplankton, eventually preventing a decrease in algal biomass after TP reductions. Long‐term epilimnetic phosphorus concentrations during the winter mixing period decreased to a greater degree than summer phosphorus concentrations. Apparently, TP losses over the season were less intense. Spring diatom biomass also markedly decreased after oligotrophication. In fact, spring diatom biomass was positively related to the TP loss over the season suggesting diatoms play an important role in P reduction. However, this intraannual P processing was not the primary factor when focusing on the average yearly yield, which remained to be fully explained by mixotrophs. Our study demonstrates this ecosystem's ability to compensate for changes in resource availability through changes in phytoplankton community composition and functional strategies. We conclude that an increase in mixotrophy and the ability to make bacterial phosphorus available for phytoplankters were the main factors that allowed the phytoplankton community of the Rappbode Reservoir to adapt to lower nutrient levels without a loss in total biomass.

&lt;p&gt;&lt;strong&gt;Abstract.&lt;/strong&gt; Excessive amounts of nutrients and dissolved organic matter in freshwater bodies affect aquatic ecosystems. In this study, the spatial and temporal variability in nitrate (NO&lt;sub&gt;3&lt;/sub&gt;), dissolved organic carbon (DOC) and soluble reactive phosphorus (SRP) was analyzed in the Selke (Germany) river continuum from headwaters draining 1&amp;#8211;3&amp;#8201;km&lt;sup&gt;2&lt;/sup&gt; catchments to downstream reaches representing spatially integrated signals from 184&amp;#8211;456&amp;#8201;km&lt;sup&gt;2&lt;/sup&gt; catchments. Three headwater catchments were selected as archetypes of the main landscape units (land use x lithology) present in the Selke catchment. Export regimes in headwater catchments were interpreted in terms of NO&lt;sub&gt;3&lt;/sub&gt;, DOC and SRP land-to-stream transfer processes. Headwater signals were subtracted from downstream signals, with the differences interpreted in terms of in-stream processes and contribution of point-source emissions. The seasonal dynamics for NO&lt;sub&gt;3&lt;/sub&gt; were opposite those of DOC and SRP in all three headwater catchments, and spatial differences also showed NO&lt;sub&gt;3&lt;/sub&gt; contrasting with DOC and SRP. These dynamics were interpreted as the result of the interplay of hydrological and biogeochemical processes, for which riparian zones were hypothesized to play a determining role. In the two downstream reaches, NO&lt;sub&gt;3&lt;/sub&gt; was transported almost conservatively, whereas DOC was consumed and produced in the upper and lower river sections, respectively. The natural export regime of SRP in the three headwater catchments mimicked a point-source signal (high SRP during summer low flow), which may lead to overestimation of domestic contributions in the downstream reaches. Monitoring the river continuum from headwaters to downstream reaches proved effective to investigate jointly land-to-stream and in-stream transport and transformation processes.&lt;/p&gt;