Consistent stoichiometric long-term relationships between nutrients and chlorophyll-a across shallow lakes.

Aquatic ecosystems are threatened by eutrophication from nutrient pollution. In lakes, eutrophication causes a plethora of deleterious effects, such as harmful algal blooms, fish kills and increased methane emissions. However, lake-specific responses to nutrient changes are highly variable, complicating eutrophication management. These lake-specific responses could result from short-term stochastic drivers overshadowing lake-independent, long-term relationships between phytoplankton and nutrients. Here, we show that strong stoichiometric long-term relationships exist between nutrients and chlorophyll a (Chla) for 5-year simple moving averages (SMA, median R² = 0.87) along a gradient of total nitrogen to total phosphorus (TN:TP) ratios. These stoichiometric relationships are consistent across 159 shallow lakes (defined as average depth < 6 m) from a cross-continental, open-access database. We calculate 5-year SMA residuals to assess short-term variability and find substantial short-term Chla variation which is weakly related to nutrient concentrations (median R² = 0.12). With shallow lakes representing 89% of the world's lakes, the identified stoichiometric long-term relationships can globally improve quantitative nutrient management in both lakes and their catchments through a nutrient-ratio-based strategy.

Cascading climate effects in deep reservoirs: Full assessment of physical and biogeochemical dynamics under ensemble climate projections and ways towards adaptation.

We coupled twenty-first century climate projections with a well-established water quality model to depict future ecological changes of Rappbode Reservoir, Germany. Our results document a chain of climate-driven effects propagating through the aquatic ecosystem and interfering with drinking water supply: intense climate warming (RCP8.5 scenario) will firstly trigger a strong increase in water temperatures, in turn leading to metalimnetic hypoxia, accelerating sediment nutrient release and finally boosting blooms of the cyanobacterium Planktothrix rubescens. Such adverse water quality developments will be suppressed under RCP2.6 and 6.0 indicating that mitigation of climate change is improving water security. Our results also suggested surface withdrawal can be an effective adaptation strategy to make the reservoir ecosystem more resilient to climate warming. The identified consequences from climate warming and adaptation strategies are relevant to many deep waters in the temperate zone, and the conclusion should provide important guidances for stakeholders to confront potential climate changes.

Insect communities under skyglow: diffuse night-time illuminance induces spatio-temporal shifts in movement and predation.

Artificial light at night (ALAN) is predicted to have far-reaching consequences for natural ecosystems given its influence on organismal physiology and behaviour, species interactions and community composition. Movement and predation are fundamental ecological processes that are of critical importance to ecosystem functioning. The natural movements and foraging behaviours of nocturnal invertebrates may be particularly sensitive to the presence of ALAN. However, we still lack evidence of how these processes respond to ALAN within a community context. We assembled insect communities to quantify their movement activity and predation rates during simulated Moon cycles across a gradient of diffuse night-time illuminance including the full range of observed skyglow intensities. Using radio frequency identification, we tracked the movements of insects within a fragmented grassland Ecotron experiment. We additionally quantified predation rates using prey dummies. Our results reveal that even low-intensity skyglow causes a temporal shift in movement activity from day to night, and a spatial shift towards open habitats at night. Changes in movement activity are associated with indirect shifts in predation rates. Spatio-temporal shifts in movement and predation have important implications for ecological networks and ecosystem functioning, highlighting the disruptive potential of ALAN for global biodiversity and the provision of ecosystem services. This article is part of the theme issue 'Light pollution in complex ecological systems'.

Artificial light at night decreases plant diversity and performance in experimental grassland communities.

Artificial light at night (ALAN) affects many areas of the world and is increasing globally. To date, there has been limited and inconsistent evidence regarding the consequences of ALAN for plant communities, as well as for the fitness of their constituent species. ALAN could be beneficial for plants as they need light as energy source, but they also need darkness for regeneration and growth. We created model communities composed of 16 plant species sown, exposed to a gradient of ALAN ranging from 'moonlight only' to conditions like situations typically found directly underneath a streetlamp. We measured plant community composition and its production (biomass), as well as functional traits of three plant species from different functional groups (grasses, herbs, legumes) in two separate harvests. We found that biomass was reduced by 33% in the highest ALAN treatment compared to the control, Shannon diversity decreased by 43% and evenness by 34% in the first harvest. Some species failed to establish in the second harvest. Specific leaf area, leaf dry matter content and leaf hairiness responded to ALAN. These responses suggest that plant communities will be sensitive to increasing ALAN, and they flag a need for plant conservation activities that consider impending ALAN scenarios. This article is part of the theme issue 'Light pollution in complex ecological systems'.

Skyglow relieves a crepuscular bird from visual constraints on being active.

Artificial light at night significantly alters the predictability of the natural light cycles that most animals use as an essential Zeitgeber for daily activity. Direct light has well-documented local impacts on activity patterns of diurnal and nocturnal organisms. However, artificial light at night also contributes to an indirect illumination of the night sky, called skyglow, which is rapidly increasing. The consequences of this wide-spread form of artificial night light on the behaviour of animals remain poorly understood, with only a few studies performed under controlled (laboratory) conditions. Using animal-borne activity loggers, we investigated daily and seasonal flight activity of a free-living crepuscular bird species in response to nocturnal light conditions at sites differing dramatically in exposure to skyglow. We find that flight activity of European Nightjars (Caprimulgus europaeus) during moonless periods of the night is four times higher in Belgium (high skyglow exposure) than in sub-tropical Africa and two times higher than in Mongolia (near-pristine skies). Moreover, clouds darken the sky under natural conditions, but skyglow can strongly increase local sky brightness on overcast nights. As a result, we find that nightjars' response to cloud cover is reversed between Belgium and sub-tropical Africa and between Belgium and Mongolia. This supports the hypothesis that cloudy nights reduce individual flight activity in a pristine environment, but increase it when the sky is artificially lit. Our study shows that in the absence of direct light pollution, anthropogenic changes in sky brightness relieve nightjars from visual constraints on being active. Individuals adapt daily activities to artificial night-sky brightness, allowing them more time to fly than conspecifics living under natural light cycles. This modification of the nocturnal timescape likely affects behavioural processes of most crepuscular and nocturnal species, but its implications for population dynamics and interspecific interactions remain to be investigated.

Reservoir water quality deterioration due to deforestation emphasizes the indirect effects of global change.

Deforestation is currently a widespread phenomenon and a growing environmental concern in the era of rapid climate change. In temperate regions, it is challenging to quantify the impacts of deforestation on the catchment dynamics and downstream aquatic ecosystems such as reservoirs and disentangle these from direct climate change impacts, let alone project future changes to inform management. Here, we tackled this issue by investigating a unique catchment-reservoir system with two reservoirs in distinct trophic states (meso­ and eutrophic), both of which drain into the largest drinking water reservoir in Germany. Due to the prolonged droughts in 2015-2018, the catchment of the mesotrophic reservoir lost an unprecedented area of forest (exponential increase since 2015 and ca. 17.1% loss in 2020 alone). We coupled catchment nutrient exports (HYPE) and reservoir ecosystem dynamics (GOTM-WET) models using a process-based modeling approach. The coupled model was validated with datasets spanning periods of rapid deforestation, which makes our future projections highly robust. Results show that in a short-term time scale (by 2035), increasing nutrient flux from the catchment due to vast deforestation (80% loss) can turn the mesotrophic reservoir into a eutrophic state as its counterpart. Our results emphasize the more prominent impacts of deforestation than the direct impact of climate warming in impairment of water quality and ecological services to downstream aquatic ecosystems. Therefore, we propose to evaluate the impact of climate change on temperate reservoirs by incorporating a time scale-dependent context, highlighting the indirect impact of deforestation in the short-term scale. In the long-term scale (e.g. to 2100), a guiding hypothesis for future research may be that indirect effects (e.g., as mediated by catchment dynamics) are as important as the direct effects of climate warming on aquatic ecosystems.

Design and implementation of an illumination system to mimic skyglow at ecosystem level in a large-scale lake enclosure facility.

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 0.01 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.

Phenological shifts in lake stratification under climate change.

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.

Lake heatwaves under climate change.

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.

Unravelling winter diatom blooms in temperate lakes using high frequency data and ecological modeling.

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.

Ensemble warming projections in Germany's largest drinking water reservoir and potential adaptation strategies.

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.

The formation of a metalimnetic oxygen minimum exemplifies how ecosystem dynamics shape biogeochemical processes: A modelling study.

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.

Planktonic events may cause polymictic-dimictic regime shifts in temperate lakes.

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.

Temperature and photoperiod interactions with phosphorus-limited growth and competition of two diatoms.

In lakes, trophic change and climate change shift the relationship between nutrients and physical factors, like temperature and photoperiod, and interactions between these factors should affect the growth of phytoplankton species differently. We therefore determined the relationship between P-limited specific growth rates and P-quota (biovolume basis) of Stephanodiscus minutulus and Nitzschia acicularis (diatoms) at or near light saturation in axenic, semi-continuous culture at 10, 15 and 20 °C and at 6, 9 and 12 h d(-1) photoperiod. Photoperiod treatments were performed at constant daily light exposure to allow comparison. Under these conditions, we also performed competition experiments and estimated relative P-uptake rates of the species. Temperature strongly affected P-limited growth rates and relative P uptake rates, whereas photoperiod only affected maximum growth rates. S. minutulus used internal P more efficiently than N. acicularis. N. acicularis was the superior competitor for P due to a higher relative uptake rate and its superiority increased with increasing temperature and photoperiod. S. minutulus conformed to the Droop relationship but N. acicularis did not. A model with a temperature-dependent normalised half-saturation coefficient adequately described the factor interactions of both species. The temperature dependence of the quota model reflected each species' specific adaptation to its ecological niche. The results demonstrate that increases in temperature or photoperiod can partially compensate for a decrease in P-quota under moderately limiting conditions, like during spring in temperate lakes. Thus warming may counteract de-eutrophication to some degree and a relative shift in growth factors can influence the phytoplankton species composition.