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We investigate the "macronutrient-access hypothesis", which states that the balance between stoichiometric macronutrient demand and accessible macronutrients controls nutrient assimilation by aquatic heterotrophs. Within this hypothesis, we consider bioavailable dissolved organic carbon (bDOC), reactive nitrogen (N) and reactive phosphorus (P) to be the macronutrients accessible to heterotrophic assimilation. Here, reactive N and P are the sums of dissolved inorganic N (nitrate-N, nitrite-N, ammonium-N), soluble-reactive P (SRP), and bioavailable dissolved organic N (bDON) and P (bDOP). Previous data from various freshwaters suggests this hypothesis, yet clear experimental support is missing. We assessed this hypothesis in a proof-of-concept experiment for waters from four small agricultural streams. We used seven different bDOC:reactive N and bDOC:reactive P ratios, induced by seven levels of alder leaf leachate addition. With these treatments and a stream-water specific bacterial inoculum, we conducted a 3-day experiment with three independent replicates per combination of stream water, treatment, and sampling occasion. Here, we extracted dissolved organic matter (DOM) fluorophores by measuring excitation-emission matrices with subsequent parallel factor decomposition (EEM-PARAFAC). We assessed the true bioavailability of DOC, DON, and the DOM fluorophores as the concentration difference between the beginning and end of each experiment. Subsequently, we calculated the bDOC and bDON concentrations based on the bioavailable EEM-PARAFAC fluorophores, and compared the calculated bDOC and bDON concentrations to their true bioavailability. Due to very low DOP concentrations, the DOP determination uncertainty was high, and we assumed DOP to be a negligible part of the reactive P. For bDOC and bDON, the true bioavailability measurements agreed with the same fractions calculated indirectly from bioavailable EEM-PARAFAC fluorophores (bDOC r2 = 0.96, p 2 = 0.77, p 2 = 0.80 and 0.83). To define zones of C:N:P (co-)limitation of heterotrophic assimilation, we used a novel ternary-plot approach combining our data with literature data on C:N:P ranges of bacterial biomass. Here, we found a zone of maximum reactive N uptake (C:N:P approx. > 114: 170:21: 204:14:1). The “macronutrient-access hypothesis” links ecological stoichiometry and biogeochemistry, and may be of importance for nutrient uptake in many freshwater ecosystems. However, this experiment is only a starting point and this hypothesis needs to be corroborated by further experiments for more sites, by in-situ studies, and with different DOC sources.

Background Aggregations of cyanobacteria in lakes and reservoirs are commonly associated with surface blooms, but may also occur in the metalimnion as subsurface or deep chlorophyll maxima. Metalimnetic cyanobacteria blooms are of great concern when potentially toxic species, such as Planktothrix rubescens, are involved. Metalimnetic blooms of P. rubescens have apparently increased in frequency and severity in recent years, so there is a strong need to identify reservoir management options to control it. We hypothesized that P. rubescens blooms in reservoirs can be suppressed using selective withdrawal to maximize its export from the reservoir. We also expect that altering the light climate can affect the dynamics of this species. We tested our hypothesis in Rappbode Reservoir (the largest drinking water reservoir in Germany) by establishing a series of withdrawal and light scenarios based on a calibrated water quality model (CE-QUAL-W2). Results The novel withdrawal strategy, in which water is withdrawn from a certain depth below the surface within the metalimnion instead of at a fixed elevation relative to the dam wall, significantly reduced P. rubescens biomass in the reservoir. According to the simulation results, we defined an optimal withdrawal volume to control P. rubescens blooms in the reservoir as approximately 10 million m3 (10% of the reservoir volume) during its bloom phase. The results also illustrated that P. rubescens growth can be most effectively suppressed if the metalimnetic withdrawal is applied in the early stage of its rapid growth, i.e., before the bloom occurs. In addition, our study showed that P. rubescens biomass gradually decreased with increasing light extinction and nearly disappeared when the extinction coefficient exceeded 0.55 m−1. Conclusions Our study indicates the rise in P. rubescens biomass can be effectively offset by selective withdrawal as well as by reducing light intensity beneath the water surface. Considering the widespread occurrence of P. rubescens in stratified lakes and reservoirs worldwide, we believe the results will be helpful for scientists and managers working on other water bodies to minimize the negative impacts of this harmful cyanobacteria. Our model may serve as a transferable tool to explore local dynamics in other standing waters.

Model ensembles have several benefits compared to single-model applications but are not frequently used within the lake modelling community. Setting up and running multiple lake models can be challenging and time consuming, despite the many similarities between the existing models (forcing data, hypsograph, etc.). Here we present an R package, LakeEnsemblR, that facilitates running ensembles of five different one-dimensional hydrodynamic lake models (FLake, GLM, GOTM, Simstrat, MyLake). The package requires input in a standardised format and a single configuration file. LakeEnsemblR formats these files to the input files required by each model, and provides functions to run and calibrate the models. The outputs of the different models are compiled into a single file, and several post-processing operations are supported. LakeEnsemblR’s workflow standardisation can simplify model benchmarking, sharing of output files, and improve collaborations between aquatic scientists. We showcase the successful application of LakeEnsemblR for two different lakes.

Empirical evidence demonstrates that lakes and reservoirs are warming across the globe. Consequently, there is an increased need to project future changes in lake thermal structure and resulting changes in lake biogeochemistry in order to plan for the likely impacts. Previous studies of the impacts of climate change on lakes have often relied on a single model forced with limited scenario-driven projections of future climate for a relatively small number of lakes. As a result, our understanding of the effects of climate change on lakes is fragmentary, based on scattered studies using different data sources and modelling protocols, and mainly focused on individual lakes or lake regions. This has precluded identification of the main impacts of climate change on lakes at global and regional scales and has likely contributed to the lack of lake water quality considerations in policy-relevant documents, such as the Assessment Reports of the Intergovernmental Panel on Climate Change (IPCC). Here, we describe a simulation protocol developed by the Lake Sector of the Inter-Sectoral Impact Model Intercomparison Project (ISIMIP) for simulating climate change impacts on lakes using an ensemble of lake models and climate change scenarios. The protocol prescribes lake simulations driven by climate forcing from gridded observations and different Earth system models under various Representative Greenhouse Gas Concentration Pathways, all consistently bias-corrected on a 0.5° × 0.5° global grid. In ISIMIP phase 2, 11 lake models were forced with these data to project the thermal structure of 62 well-studied lakes where data were available for calibration under historical conditions, and for nearly 17,500 lakes using uncalibrated models and forcing data from the global grid where lakes are present. In ISIMIP phase 3, this approach was expanded to consider more lakes, more models, and more processes. The ISIMIP Lake Sector is the largest international effort to project future water temperature, thermal structure, and ice phenology of lakes at local and global scales and paves the way for future simulations of the impacts of climate change on water quality and biogeochemistry in lakes.

Light pollution is an environmental stressor of global extent that is growing exponentially in area and intensity. Artificial skyglow, a form of light pollution with large range, is hypothesized to have environmental impact at ecosystem level. However, testing the impact of skyglow at large scales and in a controlled fashion under in situ conditions has remained elusive so far. Here we present the first experimental setup to mimic skyglow at ecosystem level outdoors in an aquatic environment. Spatially diffuse and homogeneous surface illumination that is adjustable between 10 mlx and 10 lx, resembling rural to urban skyglow levels, was achieved with white light-emitting diodes at a large-scale lake enclosure facility. The illumination system was enabled by optical modeling with Monte-Carlo raytracing and validated by measurements. Our method can be adapted to other outdoor and indoor skyglow experiments, urgently needed to understand the impact of skyglow on ecosystems.

The Qinghai-Tibet Plateau possesses the largest alpine lake system, which plays a crucial role in the land-atmosphere interaction. We report first observations on the thermal and radiation regime u...

One of the most important physical characteristics driving lifecycle events in lakes is stratification. Already subtle variations in the timing of stratification onset and break-up (phenology) are known to have major ecological effects, mainly by determining the availability of light, nutrients, carbon and oxygen to organisms. Despite its ecological importance, historic and future global changes in stratification phenology are unknown. Here, we used a lake-climate model ensemble and long-term observational data, to investigate changes in lake stratification phenology across the Northern Hemisphere from 1901 to 2099. Under the high-greenhouse-gas-emission scenario, stratification will begin 22.0 ± 7.0 days earlier and end 11.3 ± 4.7 days later by the end of this century. It is very likely that this 33.3 ± 11.7 day prolongation in stratification will accelerate lake deoxygenation with subsequent effects on nutrient mineralization and phosphorus release from lake sediments. Further misalignment of lifecycle events, with possible irreversible changes for lake ecosystems, is also likely., Stratification has a considerable influence on lake ecology, but there is little understanding of past or future changes in its seasonality. Here, the authors use modelling and empirical data to determine that between 1901–2099, climate change causes stratification to start earlier and end later.

Evaporation of surface water is critical to the basic functioning of lakes. It directly and, in some cases, substantially modifies the hydrologic, chemical, and energy budgets, making evaporation one of the most important physical controls on lake ecosystems. Predicting lake evaporation response to climate change is, therefore, of paramount importance. Most studies that simulate climate change impacts on lake evaporation have utilised only a single mechanistic model. Whilst such studies have merit, the advantage of applying multiple, independently developed models (i.e., an ensemble approach), is that some of the inherent uncertainties in the individual lake models due to, for example, different model structures, can be reduced thus enabling increased robustness of historic and future projections. In this study, we present results from the Inter-Sectoral Impact Model Intercomparison Project phase 2b (ISIMIP) Lake Sector, where lake evaporation responses to 20th and 21st century (1901-2099) climate change has been simulated with a suite of independently developed lake models under different climate change scenarios (Representative Concentration Pathways, RCP, 2.6, 6.0 and 8.5). Our study focuses on Lake Kinneret (Israel), a sub-tropical monomictic lake of socioeconomic importance. Our simulations are validated during the historic period with bulk evaporation estimates calculated from high frequency meteorological and in-lake observations. Our results demonstrate that the lake models provide an accurate representation of historical variability in lake evaporation, with promising comparisons of the magnitude, timing and seasonality of evaporative water loss. Future evaporation projections at Lake Kinneret show that evaporation anomalies will increase by the end of the century. We show that multi-model projections of lake evaporation can accurately represent the historic period and hence provide reliable future projections that will be vital for water management.

We assess the “macronutrient-balance hypothesis,” which we define as: “Aquatic heterotrophic nutrient assimilation is controlled by the balance between the bioavailable DOC : reactive macronutrient stoichiometry and the microbial stoichiometric macronutrient demand.” Here we define the reactive macronutrients as the sum of dissolved inorganic nitrogen, soluble-reactive phosphorus (SRP), and dissolved bioavailable organic N (bDON) & P (bDOP). A global meta-analysis of monitoring data from various freshwaters suggests this hypothesis, yet clear experimental support is missing.We assessed this hypothesis in a proof-of-concept experiment for waters from four different small agricultural streams. We used seven different bioavailable DOC (bDOC) : reactive N and bDOC : reactive P ratios, induced by seven different levels of alder leaf leachate addition. With these treatments and a stream-water specific bacterial inoculum, we conducted a separate 3-day experiment, with three independent replicates per combination of stream water, treatment and sampling occasion. Here, we extracted dissolved organic matter (DOM) fluorophores by measuring excitation-emission matrices with subsequent parallel factor decomposition (EEM-PARAFAC). We assessed the true bioavailability of DOC, DON, and the DOM fluorophores as solute concentration difference between the beginning and end of each experiment. Separately, we predicted bDOC and bDON concentrations based on the bioavailable fluorophores, which we compared to their true bioavailability measured before. Due to very low DOP concentrations, the DOP determination uncertainty was high, and we had to neglect DOP as part of the reactive P.For bDOC and bDON, the bioavailability measurements agreed with the same fractions calculated indirectly from bioavailable DOM fluorophores (bDOC r² = 0.96, p In summary, the results of our simple laboratory experiment provide first proof that the bDOC : reactive macronutrient ratio strongly controls heterotrophic reactive macronutrient uptake. Combined with the previous large-scale monitoring evidence, our study implies that the “macronutrient-balance hypothesis” holds in many aquatic ecosystems. However, this hypothesis needs to be corroborated by further experiments with different DOC sources and assessments of changes in bDOC : reactive macronutrient ratios on freshwater carbon and nutrient cycles.

Aggregations of cyanobacteria in lakes and reservoirs are commonly associated with surface blooms, but may also occur in the metalimnion as subsurface or deep chlorophyll maxima. Metalimnetic cyanobacteria blooms are of great concern when potentially toxic species, such as Planktothrix rubescens (P. rubescens), are involved. Apparently, metalimnetic blooms of P. rubescens have increased in frequency and severity in recent years so there is a strong need to establish the external factors controlling its growth. We hypothesized that P. rubescens blooms in reservoirs can be managed by modifying the water withdrawal strategy and altering the light climate experienced by the algae. We tested our hypothesis in Rappbode Reservoir by establishing a series of withdrawal and light scenarios based on a calibrated water quality model (CE-QUAL-W2). Our scenarios demonstrated that metalimnetic water withdrawal reduced P. rubescens biomass in the reservoir. According to the simulation results we defined an optimal withdrawal volume to control P. rubescens blooms in the reservoir as approximately 10 million m3 during its blooming period. The numerical results also indicated that P. rubescens growth can be most effectively suppressed if the metalimnetic withdrawal is applied in the early stage of its rapid growth (i.e. before the occurrence of blooms). Additionally, the results showed that P. rubescens biomass gradually decreased with increasing light extinction and nearly disappeared when the extinction coefficient exceeded 0.55 m-1. Our results indicated that close linkages among in situ measurements, model simulations, empirical growth rate and flushing rate calculations could inform management strategies to minimise the harmful impacts of P. rubescens in water supplies. Such a strategy could be used in reservoir operational strategies as an adaptation way to offset the rise in P. rubescens populations that has been linked to climate change.

The thermal structure in reservoirs affects the development of aquatic ecosystems and is substantially influenced by changing climate conditions. At the same time, reservoir management strategies can also affect the thermal structure of the water body and may enable adaptation strategies in a warmer world. We applied a two-dimensional hydrodynamicmodel to explore the response of the thermal structure in Germany's largest drinking water reservoir, Rappbode Reservoir, to future climate projections and different water withdrawal strategies. We used projections for representative concentration pathways (RCP) 2.6, 6.0 and 8.5 from an ensemble of 4 different global climate models taken from the ISIMIP project. Simulation results showed that epilimnetic water temperatures in the reservoir strongly increased under all three climate scenarios while the magnitude of warming directly corresponds to the increase in air temperatures. Hypolimnetic temperatures remained rather constant under RCP 2.6 and RCP 6.0 but increased markedly under RCP 8.5. Under the intense warming in RCP 8.5, hypolimnion temperatures were projected to rise from 5 °C to 8 °C by the end of the century. Moreover, the results suggested that surface withdrawal can be an effective adaptation strategy under strong climate warming (RCP 8.5) to reduce surface warming and even avoid hypolimnetic warming. This study documents how global scale climate projections can be translated into site-specific climate impacts to derive adaptation strategies for reservoir operation. Moreover, our results illustrate that the most intense warming scenario, i.e. RCP 8.5, demands far-reaching climate adaptation while the mitigation scenario (RCP 2.6) does not require adaptation of reservoir management before 2100.

Lake ecosystems, and the organisms that live within them, are vulnerable to temperature change1–5, including the increased occurrence of thermal extremes6. However, very little is known about lake heatwaves—periods of extreme warm lake surface water temperature—and how they may change under global warming. Here we use satellite observations and a numerical model to investigate changes in lake heatwaves for hundreds of lakes worldwide from 1901 to 2099. We show that lake heatwaves will become hotter and longer by the end of the twenty-first century. For the high-greenhouse-gas-emission scenario (Representative Concentration Pathway (RCP) 8.5), the average intensity of lake heatwaves, defined relative to the historical period (1970 to 1999), will increase from 3.7 ± 0.1 to 5.4 ± 0.8 degrees Celsius and their average duration will increase dramatically from 7.7 ± 0.4 to 95.5 ± 35.3 days. In the low-greenhouse-gas-emission RCP 2.6 scenario, heatwave intensity and duration will increase to 4.0 ± 0.2 degrees Celsius and 27.0 ± 7.6 days, respectively. Surface heatwaves are longer-lasting but less intense in deeper lakes (up to 60 metres deep) than in shallower lakes during both historic and future periods. As lakes warm during the twenty-first century7,8, their heatwaves will begin to extend across multiple seasons, with some lakes reaching a permanent heatwave state. Lake heatwaves are likely to exacerbate the adverse effects of long-term warming in lakes and exert widespread influence on their physical structure and chemical properties. Lake heatwaves could alter species composition by pushing aquatic species and ecosystems to the limits of their resilience. This in turn could threaten lake biodiversity9 and the key ecological and economic benefits that lakes provide to society. Modelling and remote sensing show that by the end of the twenty-first century, lake heatwaves will be several degrees Celsius warmer and some will be months longer, with potentially major adverse consequences for lake ecosystems.

In temperate lakes, it is generally assumed that light rather than temperature constrains phytoplankton growth in winter. Rapid winter warming and increasing observations of winter blooms warrant more investigation of these controls. We investigated the mechanisms regulating a massive winter diatom bloom in a temperate lake. High frequency data and process-based lake modeling demonstrated that phytoplankton growth in winter was dually controlled by light and temperature, rather than by light alone. Water temperature played a further indirect role in initiating the bloom through ice-thaw, which increased light exposure. The bloom was ultimately terminated by silicon limitation and sedimentation. These mechanisms differ from those typically responsible for spring diatom blooms and contributed to the high peak biomass. Our findings show that phytoplankton growth in winter is more sensitive to temperature, and consequently to climate change, than previously assumed. This has implications for nutrient cycling and seasonal succession of lake phytoplankton communities. The present study exemplifies the strength in integrating data analysis with different temporal resolutions and lake modeling. The new lake ecological model serves as an effective tool in analyzing and predicting winter phytoplankton dynamics for temperate lakes.

The surface mixed layer in lakes is where phytoplankton grow and where most of the primary production occurs. Knowledge of the thickness of the mixed layer is essential to estimate for instance primary productivity and to interpret remote sensing measurements, because it determines the mean light supply and indicates how homogeneous the water column is. Modelling studies, primarily in the ocean, have concluded that the mixed layer will shoal as a result of climate warming, but the empirical evidence does not support this. Here we seek to determine how climate change affects the mixed layer thickness and mean underwater irradiance in lakes. We use an ensemble modelling approach to simulate mixed layer depth in 3 warming scenarios (RCP2.6, 6.0, 8.5) in about 50 lakes across the globe using the hydrodynamic model Flake forced by four downscaled global climate models. Results indicate that warming has little direct effect on the mixed layer depth. Mean underwater light in the mixed layer was nevertheless projected to increase as a result of the global radiation increases in the global climate models.

Metalimnetic oxygen minima are observed in many lakes and reservoirs, but the mechanisms behind this phenomenon are not well understood. Thus, we simulated the metalimnetic oxygen minimum (MOM) in the Rappbode Reservoir (Germany) with a well-established two-dimensional water quality model (CE-QUAL-W2) to systematically quantify the chain of events leading to its formation. We used high-resolution measured data to calibrate the model, which accurately reproduced the physical (e.g. water level and water temperature), biogeochemical (e.g. nutrient and oxygen dynamics) and ecological (e.g. algal community dynamics) features of the reservoir, particularly the spatial and temporal extent of the MOM. The results indicated that around 60% of the total oxygen consumption rate in the MOM layer originated from benthic processes whereas the remainder originated from pelagic processes. The occurrence of the cyanobacterium Planktothrix rubescens in the metalimnion delayed and slightly weakened the MOM through photosynthesis, although its decaying biomass ultimately induced the MOM. Our research also confirmed the decisive role of water temperature in the formation of the MOM since the water temperatures, and thus benthic and pelagic oxygen consumption rates, were higher in the metalimnion than in the hypolimnion. Our model is not only providing novel conclusions about the drivers of MOM development and their quantitative contributions, it is also a new tool for understanding and predicting ecological and biogeochemical water quality dynamics.

The thermal structure in reservoirs affects the development of aquatic ecosystems, and can be substantially influenced by climate change and management strategies. We applied a two-dimensional hydrodynamic model to explore the response of the thermal structure in Germany's largest drinking water reservoir, Rappbode Reservoir, to future climate projections and different water withdrawal strategies. We used projections for representative concentration pathways (RCP) 2.6, 6.0 and 8.5 from an ensemble of 4 different global climate models. Simulation results showed that epilimnetic water temperatures in the reservoir strongly increased under all three climate scenarios. Hypolimnetic temperatures remained rather constant under RCP 2.6 and RCP 6.0 but increased markedly under RCP 8.5. Under the intense warming in RCP 8.5, hypolimnion temperatures were projected to rise from 5 °C to 8 °C by the end of the century. Stratification in the reservoir was projected to be more stable under RCP 6.0 and RCP 8.5, but did not show significant changes under RCP 2.6. Similar results were found with respect to the light intensity within the mixed-layer. Moreover, the results suggested that surface withdrawal can be an effective adaptation strategy under strong climate warming (RCP 8.5) to reduce surface warming and avoid hypolimnetic warming. This study documents how global scale climate projections can be translated into site-specific climate impacts to derive adaptation strategies for reservoir operation. Moreover, our results illustrate that the most intense warming scenario, i.e. RCP 8.5, demands far-reaching climate adaptation while the mitigation scenario (RCP 2.6) does not require adaptation of reservoir management before 2100.

Climate forecasts project a global increase in extreme weather events, but information on the consequences for ecosystems is scarce. Of particular significance for lakes are severe storms that can influence biogeochemical processes and biological communities by disrupting the vertical thermal structure during periods of stratification. An exceptional storm passing over northern Germany in July 2011 provided an opportunity to assess the consequences and underlying mechanisms of such extreme events on the interplay between the physics and ecological characteristics of a deep, nutrient-poor lake. Wind speeds were among the most extreme on record. A suite of variables measured throughout the event consistently indicates that a cascade of processes pushed the clear-water lake into an exceptionally turbid state. Specifically, thermocline deepening by the storm-entrained cyanobacteria of a deep chlorophyll maximum located at about 8 m depth into the surface mixed layer. Released from light limitation, intense photosynthesis of the cyanobacteria boosted primary production, increased algal biomass, raised the pH and thus induced massive calcite precipitation to a level never observed within three decades of lake monitoring. As a consequence, water transparency dropped from 6.5 to 2.1 m, the minimum on record for 40 years, and the euphotic zone shrank by about 8 m for several weeks. These results show that cyanobacterial blooms not only are promoted by climate warming, but can also be triggered by extreme storms. Clear-water lakes developing a deep chlorophyll maximum appear to be particularly at risk in the future, if such events become more intense or frequent.

Metalimnetic oxygen minima are observed in many lakes and reservoirs, but the mechanisms behind this phenomena are not well understood. Thus, we simulated the metalimnetic oxygen minimum (MOM) in the Rappbode Reservoir with a well-established two-dimensional water quality model (CE-QUAL-W2) to systematically quantify the chain of events leading to its formation. We used high-resolution measured data to calibrate the model, which accurately reproduced the physical (e.g. water level and water temperature), biogeochemical (e.g. nutrient and oxygen dynamics) and ecological (e.g. algal community dynamics) features of the reservoir, particularly the spatial and temporal extent of the MOM. The results indicated that around 60% of the total oxygen consumption rate in the MOM layer originated from benthic processes whereas the remainder originated from pelagic processes. The occurrence of the cyanobacterium Planktothrix rubescens in the metalimnion delayed and slightly weakened the MOM through photosynthesis, although its decaying biomass ultimately induced the MOM. Our research also confirmed the decisive role of water temperature in the formation of the MOM since the water temperatures, and thus benthic and pelagic oxygen consumption rates, were higher in the metalimnion than in the hypolimnion. Our model is not only providing novel conclusions about the drivers of MOM development and their quantitative contributions, it is also a new tool for understanding and predicting ecological and biogeochemical water quality dynamics.

The physical response of lakes to climate warming is regionally variable and highly dependent on individual lake characteristics, making generalizations about their development difficult. To qualify the role of individual lake characteristics in their response to regionally homogeneous warming, we simulated temperature, ice cover, and mixing in four intensively studied German lakes of varying morphology and mixing regime with a one-dimensional lake model. We forced the model with an ensemble of 12 climate projections (RCP4.5) up to 2100. The lakes were projected to warm at 0.10–0.11 ∘C decade−1, which is 75 %–90 % of the projected air temperature trend. In simulations, surface temperatures increased strongly in winter and spring, but little or not at all in summer and autumn. Mean bottom temperatures were projected to increase in all lakes, with steeper trends in winter and in shallower lakes. Modelled ice thaw and summer stratification advanced by 1.5–2.2 and 1.4–1.8 days decade−1 respectively, whereas autumn turnover and winter freeze timing was less sensitive. The projected summer mixed-layer depth was unaffected by warming but sensitive to changes in water transparency. By mid-century, the frequency of ice and stratification-free winters was projected to increase by about 20 %, making ice cover rare and shifting the two deeper dimictic lakes to a predominantly monomictic regime. The polymictic lake was unlikely to become dimictic by the end of the century. A sensitivity analysis predicted that decreasing transparency would dampen the effect of warming on mean temperature but amplify its effect on stratification. However, this interaction was only predicted to occur in clear lakes, and not in the study lakes at their historical transparency. Not only lake morphology, but also mixing regime determines how heat is stored and ultimately how lakes respond to climate warming. Seasonal differences in climate warming rates are thus important and require more attention., Hydrology and Earth System Sciences, 23 (3), ISSN:1027-5606, ISSN:1607-7938

In this paper the concept of resilience is discussed on the base of 13 case studies from the German branch of the International Long-Term Ecological Research Program. In the introduction the resilience approach is presented as one possibility to describe ecosystem dynamics. The relations with the concepts of adaptability and ecological integrity are discussed and the research questions are formulated. The focal research objectives are related to the conditions of resilient behaviour of ecosystems, the role of spatio-temporal scales, the differences between short- or long-term dynamics, the basic methodological requirements to exactly define resilience, the role of the reference state and indicators and the suitability of resilience as a management concept. The main part of the paper consists of 13 small case study descriptions, which demonstrate phase transitions and resilient dynamics of several terrestrial and aquatic ecosystems at different time scales. In the discussion, some problems arising from the interpretation of the time series are highlighted and discussed. The topics of discussion are the conceptual challenges of the resilience approach, methodological problems, the role of indicator selection, the complex interactions between different disturbances, the significance of time scales and a comparison of the case studies. The article ends with a conclusion which focuses on the demand to link resilience with adaptability, in order to support the long-term dynamics of ecosystem development. published

The effectiveness of controlling nitrogen (N) to manage lake eutrophication is debated. Long‐term, whole‐lake case studies are required to determine whether diazotrophic cyanobacteria can fix sufficient N to offset a reduction of N‐inputs. We document the recovery of shallow, productive Lake Müggelsee (Germany) over 37 yr (sampling interval 1–2 weeks) during a decrease of N and phosphorus (P) loading of 79% and 69%, respectively. Nitrogen concentrations in the lake responded immediately to loading reduction whereas P concentrations remained elevated for about 20 yr. Total nitrogen (TN) in the lake was always lower than TN in the inflow. Accordingly, estimated denitrification and N‐burial rates substantially exceeded N2 fixation rates in the long term. Phosphorus was growth limiting in spring whereas N was clearly limiting in summer due to high sediment P‐release. TN : TP ratios, normalized to phytoplankton biovolume by regression, were 25.5 (weight) in spring and 3.3 in summer. During the study period, dissolved inorganic N (DIN) concentrations in summer decreased and the duration of low DIN concentrations increased by ca. 100 d. The biovolume of cyanobacteria and total phytoplankton decreased by 89% and 76%, respectively. The proportion of N2‐fixing cyanobacteria during summer decreased from 36% to 14% of the total phytoplankton biovolume. The total concentration of heterocysts and estimated total N2 fixation did not change over time. In the long term, decreasing N‐inputs effectively controlled summer cyanobacteria including N2‐fixing taxa, which did not compensate for the N‐deficit. A P‐only control strategy would not have been as successful.

Rewetting of long-term drained fens often results in the formation of eutrophic shallow lakes with an average water depth of less than 1 m. This is accompanied by a fast vegetation shift from cultivated grasses via submerged hydrophytes to helophytes. As a result of rapid plant dying and decomposition, these systems are highly dynamic wetlands characterised by a high mobilisation of nutrients and elevated emissions of CO2 and CH4. However, the impact of specific plant species on these phenomena is not clear. Therefore we investigated the CO2 and CH4 production due to the subaqueous decomposition of shoot biomass of five selected plant species which represent different rewetting stages (Phalaris arundinacea, Ceratophyllum demersum, Typha latifolia, Phragmites australis and Carex riparia) during a 154 day mesocosm study. Beside continuous gas flux measurements, we performed bulk chemical analysis of plant tissue, including carbon, nitrogen, phosphorus and plant polymer dynamics. Plant-specific mass losses after 154 days ranged from 25% (P. australis) to 64% (C. demersum). Substantial differences were found for the CH4 production with highest values from decomposing C. demersum (0.4 g CH4 kg−1 dry mass day) that were about 70 times higher than CH4 production from C. riparia. Thus, we found a strong divergence between mass loss of the litter and methane production during decomposition. If C. demersum as a hydrophyte is included in the statistical analysis solely nutrient contents (nitrogen and phosphorus) explain varying greenhouse gas production of the different plant species while lignin and polyphenols demonstrate no significant impact at all. Taking data of annual biomass production as important carbon source for methanogens into account, high CH4 emissions can be expected to last several decades as long as inundated and nutrient-rich conditions prevail. Different restoration measures like water level control, biomass extraction and top soil removal are discussed in the context of mitigation of CH4 emissions from rewetted fens.

When severely degraded fens are rewetted, they often become shallow lakes with an average water depth of less than 1 m. The additional high nutrient availability in highly decomposed peat soils of these newly formed ecosystems favours the fast establishment of a small number of helophytes while the return of lost target species like low sedges and brown mosses could be delayed for decades. We hypothesise that the phosphorus (P) uptake of the newly developed vegetation substantially influences the P cycle in rewetted fens. Therefore, we investigated how much of the P released in upper degraded peat soils is pumped across the redox-interface between the soil and surface water (=‘P barrier’) during the growing season (∼150 days) by six helophytes ( Phragmites australis , Typha latifolia , Glyceria maxima , Carex acutiformis , Carex riparia , and Phalaris arundinacea ) in five rewetted fens. We then assessed how this would affect the different plant-available P fractions in the rooted degraded peat layers. The highest P uptake during the growing season (duration 150 days from May to September) was recorded for T. latifolia and G. maxima (3.0 and 2.8 g m −2 , respectively). Overall, the P uptake was in the range of the P mobilisation rates we measured in highly decomposed peat soils (range: 0.8–15.6 g P m −2 , n = 30), but four to 10-fold higher than diffusive net P fluxes at the interface between soil and surface water. Accordingly, helophytes are able to compensate for the high P mobilisation in degraded peat soils during the growing season, by incorporating this P into biomass. On the other hand a large part of the plant-P stock is released after die back through leaching and mineralisation, which increases the P load of these newly formed shallow lakes and possibly also of adjacent water courses. We estimated that it would still take 20–50 years to exhaust the large pool of plant-available P in highly decomposed peat soils if aboveground biomass was removed. Without any further management apart from fen rewetting it is unlikely that the fens will return to low nutrient levels within a human life time.

Summary We investigated the combined effects of thermal pollution from a nuclear power plant (NPP) and regional climate warming on the thermal regime of a lake. For this purpose, we used the lake model FLake and analyzed 50 years of temperature data from Lake Stechlin, Germany, which served as the cooling water reservoir for the Rheinsberg NPP from 1966 until 1990. Both modeling and statistical data analysis revealed a strong influence of the NPP cooling water discharge on the lake water temperatures and the vertical stability of the water column. A remarkable effect of thermal pollution consisted of strong vertical mixing in winter produced by the discharge of warm water into the lake when ambient water temperatures were below 4 °C. This effect caused a significant increase in the deep hypolimnion temperatures and a corresponding decrease of the vertical stability in the summer. In turn, climate warming had the opposite effect on the summer stability by increasing lake surface temperatures. Both the thermal pollution and climate change increased the duration of the summer stratification period. Our results suggest that industrial thermal pollution in temperate lakes during winter is stored in the deep water column until the next winter, whereas heat added in the summer dissipates relatively rapidly into the atmosphere. Accordingly, the winter thermal pollution could have a long-lasting effect on the lake ecology by affecting benthic biogeochemical processes.

The hydrology of the lake-rich Tibetan Plateau is important for the global climate yet little is known about the thermal regime of Tibetan lakes due to scant data. We (i) investigated the characteristic seasonal temperature patterns and recent trends in thermal and stratification regime of lakes on the Tibetan Plateau and (ii) tested theperformance of the one-dimensional lake parameterization scheme FLake for the Tibetan lake system. For this purpose we combined three years of in situ lake temperature measurements, several decadesof satellite observations and the global reanalysis data. We chose the two largest freshwater Tibetan lakes – Ngoring and Gyaring – as study sites. The lake model FLake faithfully reproduced the specific features of the high-altitude lakes and was subsequently applied to reconstruct the vertically resolved heat transport in both lakes during the last four decades. The modelsuggested Ngoring and Gyaring were ice-covered forabout 6 months and stratified in summer for about 4 months per year with a short spring overturn and longer autumn overturn. In summer the surface mixed boundary layer extended to 6–8 m depth and was about 20 % shallower in the more turbid Gyaring. The thermal regime of transparent Ngoring responded more strongly to atmospheric forcing than Gyaring, where the higher turbidity dampened the response. According to reanalysis data, air temperatures and humidity increased, whereas solar radiation decreased since the 1970s. Surprisingly, mean lake temperatures did not change, nor did the phenology of ice cover or stratification. Lake surface temperatures in summerincreased only marginally. The reason is that the increase in air temperature was offset by the decrease in radiation, probably due to increasing humidity. This study demonstrates that air temperature trends are not directly coupled to lake temperatures and underscores the importance of short-wave radiation for the thermal regime of high-altitude lakes.

We measured specific growth rates of Stephanodiscus minutulus, Nitzschia acicularis (diatoms), and Limnothrix redekei (cyanobacterium) under fluctuating and constant light in semi-continuous culture at 10uC, 15uC, and 20uC and under photoperiods of 6 h d21 and 12 h d21. Fluctuating light regimes simulated regular vertical mixing in lakes with a ratio of euphotic to mixed depth (zeu:zmix) of 1 and 0.5 on a cloudless day. Light fluctuations at zeu:zmix 5 1 decreased the growth rates of S. minutulus, N. acicularis, and L. redekei by 18% ,3 3%, and 29%, respectively, compared to constant light at the same daily light supply. Temperature had no effect on this decrease. Halving zeu:zmix (simulating deep mixing) had the same effect on growth as halving the photoperiod, demonstrating that these factors are cumulative. We introduce a simple empirical factor to adjust growth rates measured under constant light to account for fluctuating light. This factor is independent of temperature and photoperiod, applies over a range of zeu:zmix, and accurately describes present and published growth rates of several species. We show how to account for temporal variability of the light supply at different temperatures and photoperiods when predicting growth rates of phytoplankton. Phytoplankton experience a steadily changing light supply, due to, for exampl e, the course of sunlight throughout the day, varying cloud cover, wave reflections, and vertical transport within the mixed layer. Thus, light is a complex resource for phytoplankton, with both temporal components, like daylength and intensity fluctuations, and a quantitative component, the amount of light energy. In terms of phytoplankton growth, ‘‘light limitation’’ generally refers to limitation by the amount of light energy; however, this may not be so simple. During spring, light and temperature generally limit algal growth in lakes before nutrient limitation sets in (Sommer et al. 1986). However, there is evidence that it is not the daily amount of light

Global climate change alters the relationship between temperature and light in aquatic ecosystems, which is expected to affect the success of different phytoplankton species. To examine this, the interactions between temperature, photoperiod and light exposure (LE) (integral daily light supply) on specific growth rates were analysed for Limnothrix redekei, Planktothrix agardhii (cyanobacteria), Nitzschia acicularis and Stephanodiscus minutulus (diatoms). A model of factor interactions was developed based on new (P. agardhii and St. minutulus) and previously published laboratory studies. It describes the measured data with high precision. Temperature and photoperiod affect the parameters of the light-growth response curve differently, but these effects are the same for all species. The link between functions for temperature and photoperiod is more species-specific. Using meteorological data, the model developed here was used to study the interplay of these factors during a spring bloom in Lake Muggelsee (Berlin). It was found that while all three factors influenced phytoplankton growth, temperature and photoperiod were more important than LE. Both the intensities of the factors and the interactions between them influenced each species to a different degree. The results may help improve our understanding and ability to predict shifts in phytoplankton communities caused by weather patterns and climate change.

Water transparency affects the thermal structure of lakes and within certain lake depth ranges, it can determine whether a lake mixes regularly (polymictic regime) or stratifies continuously (dimictic regime) from spring through summer. Phytoplankton biomass can influence transparency but the effect of its seasonal pattern on stratification is unknown. Therefore we analysed long term field data from two lakes of similar depth, transparency and climate but one polymictic and one dimictic and simulated a conceptual lake with a hydrodynamic model. Transparency in the study lakes was typically low during spring and summer blooms and high in between during the clear water phase (CWP), caused when zooplankton graze the spring bloom. The effect of variability of transparency on thermal structure was stronger at intermediate transparency and stronger during a critical window in spring when the rate of lake warming is highest. Whereas the spring bloom strengthened stratification in spring, the CWP weakened it in summer. The presence or absence of the CWP influenced stratification duration and under some conditions determined the mixing regime. Therefore seasonal plankton dynamics, including biotic interactions that suppress the CWP, can influence lake temperatures, stratification duration and potentially also the mixing regime.

Hutchinson and Loffler's (1956) classification of lakes based on the seasonal thermal mixing regime has become a cornerstone of any analysis of lakes as elements of the earth surface. Until now however the lake classification has lacked a physically sound quantitative criterion distinguishing between two fundamental lake types: thermally stratified during a large portion of the year (mono- and dimictic) and predominantly mixed to the bottom (polymictic). Using the mechanistic balance between potential and kinetic energy we review the different formulations of the Richardson number to derive a generalized scaling for seasonal stratification in a closed lake basin. The scaling parameter is the critical mean basin depth, Hcrit, that delineates lakes that mix regularly from those that stratify seasonally based on lake water transparency, lake length, and an annual mean estimate for the Monin-Obukhov length. We validate the scaling on available data of lakes worldwide using logistic regression. The scaling criterion consistently described the mixing regime significantly better than either the conventional unbounded basin scaling or a simple depth threshold. Thus, the generalized scaling is universal for freshwater lakes and allows the seasonal mixing regime to be estimated without numerically solving the heat transport equations.

Model ensembles have several benefits compared to single-model applications but are not frequently used within the lake modelling community. Setting up and running multiple lake models can be challenging and time consuming, despite the many similarities between the existing models (forcing data, hypsograph, etc.). Here we present an R package, LakeEnsemblR, that facilitates running ensembles of five different vertical one-dimensional hydrodynamic lake models (FLake, GLM, GOTM, Simstrat, MyLake). The package requires input in a standardised format and a single configuration file. LakeEnsemblR formats these files to the input required by each model, and provides functions to run and calibrate the models. The outputs of the different models are compiled into a single file, and several post-processing operations are supported. LakeEnsemblR's workflow standardisation can simplify model benchmarking and uncertainty quantification, and improve collaborations between scientists. We showcase the successful application of LakeEnsemblR for two different lakes.