Process-based modeling for ecosystem service provisioning: Non-linear responses to restoration efforts in a quarry lake under climate change.

Healthy freshwater ecosystems can provide vital ecosystem services (ESs), and this capacity may be hampered due to water quality deterioration and climate change. In the currently available ES modeling tools, ecosystem processes are either absent or oversimplified, hindering the evaluation of impacts of restoration measures on ES provisioning. In this study, we propose an ES modeling tool that integrates lake physics, ecology and service provisioning into a holistic modeling framework. We applied this model to a Dutch quarry lake, to evaluate how nine ESs respond to technological-based (phosphorus (P) reduction) and nature-based measures (wetland restoration). As climate change might be affecting the future effectiveness of restoration efforts, we also studied the climate change impacts on the outcome of restoration measures and provisioning of ESs, using climate scenarios for the Netherlands in 2050. Our results indicate that both phosphorus reduction and wetland restoration mitigated eutrophication symptoms, resulting in increased oxygen concentrations and water transparency, and decreased phytoplankton biomass. Delivery of most ESs was improved, including swimming, P retention, and macrophyte habitat, whereas the ES provisioning that required a more productive system was impaired (sport fishing and bird watching). However, our modeling results suggested hampered effectiveness of restoration measures upon exposure to future climate conditions, which may require intensification of restoration efforts in the future to meet restoration targets. Importantly, ESs provisioning showed non-linear responses to increasing intensity of restoration measures, indicating that effectiveness of restoration measures does not necessarily increase proportionally. In conclusion, the ecosystem service modeling framework proposed in this study, provides a holistic evaluation of lake restoration measures on ecosystem services provisioning, and can contribute to development of climate-robust management strategies.

Long-term exposure to experimental light affects the ground-dwelling invertebrate community, independent of light spectra.

Our planet endures a progressive increase in artificial light at night (ALAN), which affects virtually all species, and thereby biodiversity. Mitigation strategies include reducing its intensity and duration, and the adjustment of light spectrum using modern light emitting diode (LED) light sources. Here, we studied ground-dwelling invertebrate (predominantly insects, arachnids, molluscs, millipedes, woodlice and worms) diversity and community composition after 3 or 4 years of continued nightly exposure (every night from sunset to sunrise) to experimental ALAN with three different spectra (white-, and green- and red-dominated light), as well as for a dark control, in natural forest-edge habitat. Diversity of pitfall-trapped ground-dwelling invertebrates, and the local contribution to beta diversity, did not differ between the dark control and illuminated sites, or between the different spectra. The invertebrate community composition, however, was significantly affected by the presence of light. Keeping lights off during single nights did show an immediate effect on the composition of trapped invertebrates compared to illuminated nights. These effects of light on species composition may impact ecosystems by cascading effects across the food web. This article is part of the theme issue 'Light pollution in complex ecological systems'.

Regime shifts in shallow lakes explained by critical turbidity.

Worldwide, water quality managers target a clear, macrophyte-dominated state over a turbid, phytoplankton-dominated state in shallow lakes. The competition mechanisms underlying these ecological states were explored in the 1990s, but the concept of critical turbidity seems neglected in contemporary water quality models. In particular, a simple mechanistic model of alternative stable states in shallow lakes accounting for resource competition mechanisms and critical turbidity is lacking. To this end, we combined Scheffer's theory on critical turbidity with insights from nutrient and light competition theory founded by Tilman, Huisman and Weissing. This resulted in a novel graphical and mathematical model, GPLake-M, that is relatively simple and mechanistically understandable and yet captures the essential mechanisms leading to alternative stable states in shallow lakes. The process-based PCLake model was used to parameterize the model parameters and to test GPLake-M using a pattern-oriented strategy. GPLake-M's application range and position in the model spectrum are discussed. We believe that our results support the fundamental understanding of regime shifts in shallow lakes and provide a starting point for further mechanistic and management-focused explorations and model development. Furthermore, the concept of critical turbidity and the relation between light-limited submerged macrophytes and nutrient-limited phytoplankton might provide a new focus for empirical aquatic ecological research and water quality monitoring programs.

Making waves: Lessons learned from the COVID-19 anthropause in the Netherlands on urban aquatic ecosystem services provisioning and management.

The anomalous past two years of the COVID-19 pandemic have been a test of human response to global crisis management as typical human activities were significantly altered. The COVID-instigated anthropause has illustrated the influence that humans and the biosphere have on each other, especially given the variety of national mobility interventions that have been implemented globally. These local COVID-19-era restrictions influenced human-ecosystem interactions through changes in accessibility of water systems and changes in ecosystem service demand. Four urban aquatic case studies in the Netherlands demonstrated shifts in human demand during the anthropause. For instance, reduced boat traffic in Amsterdam canals led to improved water clarity. In comparison, ongoing service exploitation from increased recreational fishing, use of bathing waters and national parks visitation are heightening concerns about potential ecosystem degradation. We distilled management lessons from both the case studies as well as from recent literature pertaining to ecological intactness and social relevance. Equally important to the lessons themselves, however, is the pace at which informed management practices are established after the pandemic ends, particularly as many communities currently recognize the importance of aquatic ecosystems and are amenable to their protection.

Towards climate-robust water quality management: Testing the efficacy of different eutrophication control measures during a heatwave in an urban canal.

Harmful algal blooms are symptomatic of eutrophication and lead to deterioration of water quality and ecosystem services. Extreme climatic events could enhance eutrophication resulting in more severe nuisance algal blooms, while they also may hamper current restoration efforts aimed to reduce nutrient loads. Evaluation of restoration measures on their efficacy under climate change is essential for effective water management. We conducted a two-month mesocosm experiment in a hypertrophic urban canal focussing on the reduction of sediment phosphorus (P)-release. We tested the efficacy of four interventions, measuring phytoplankton biomass, nutrients in water and sediment. The measures included sediment dredging, water column aeration and application of P-sorbents (lanthanum-modified bentonite - Phoslock® and iron-lime sludge, a by-product from drinking water production). An extreme heatwave (with the highest daily maximum air temperature up to 40.7 °C) was recorded in the middle of our experiment. This extreme heatwave was used for the evaluation of heatwave-induced impacts. Dredging and lanthanum modified bentonite exhibited the largest efficacy in reducing phytoplankton and cyanobacteria biomass and improving water clarity, followed by iron-lime sludge, whereas aeration did not show an effect. The heatwave negatively impacted all four measures, with increased nutrient releases and consequently increased phytoplankton biomass and decreased water clarity compared to the pre-heatwave phase. We propose a conceptual model suggesting that the heatwave locks nutrients within the biological P loop, which is the exchange between labile P and organic P, while the P fraction in the chemical P loop will be decreased. As a consequence, the efficacy of chemical agents targeting P-reduction by chemical binding will be hampered by heatwaves. Our study indicates that current restoration measures might be challenged in a future with more frequent and intense heatwaves.

Monitoring biological water quality by volunteers complements professional assessments.

Progressively more community initiatives have been undertaken over last decades to monitor water quality. Biological data collected by volunteers has been used for biodiversity and water quality studies. Despite the many citizen science projects collecting and using macroinvertebrates, the number of scientific peer-reviewed publications that use this data, remains limited. In 2018, a citizen science project on biological water quality assessment was launched in the Netherlands. In this project, volunteers collect macroinvertebrates from a nearby waterbody, identify and count the number of specimens, and register the catch through a web portal to instantaneously receive a water quality score based on their data. Water quality monitoring in the Netherlands is traditionally the field of professionals working at water authorities. Here, we compare the data from the citizen science project with the data gathered by professionals. We evaluate information regarding type and distribution of sampled waterbodies and sampling period, and compare general patterns in both datasets with respect to collected animals and calculated water quality scores. The results show that volunteers and professionals seldomly sample the same waterbody, that there is some overlap in sampling period, and that volunteers more frequently sampled urban waters and smaller waterbodies. The citizen science project is thus yielding data about understudied waters and this spatial and temporal complementarity is useful. The character and thoroughness of the assessments by volunteers and professionals are likely to differentiate. Volunteers collected significantly lower numbers of animals per sample and fewer animals from soft sediments like worms and more mobile individuals from the open water column such as boatsmen and beetles. Due to the lack of simultaneous observations at various locations by volunteers and professionals, a direct comparison of water quality scores is impossible. However, the obtained patterns from both datasets show that the water quality scores between volunteers and professionals are dissimilar for the different water types. To bridge these differences, new tools and processes need to be further developed to increase the value of monitoring biological water quality by volunteers for professionals.

What is the pollution limit? Comparing nutrient loads with thresholds to improve water quality in Lake Baiyangdian.

Ecological thresholds are useful indicators for water quality managers to define limits to nutrient pollution. A common approach to estimating ecological thresholds is using critical nutrient loads. Critical nutrient loads are typically defined as the loads at which the phytoplankton chlorophyll-a exceeds a certain concentration. However, national policies, such as in China, use chemical indicators (nitrogen and phosphorus concentrations) rather than ecological indicators (phytoplankton chlorophyll-a) to assess water quality. In this study, we uniquely define the critical nutrient loads based on maximum allowable nutrient concentrations for lake Baiyangdian. We assess whether current and future nutrient loads in this lake comply with the Chinese Water Quality standards. To this end, we link two models (MARINA-Lakes and PCLake+). The PCLake+ model was applied to estimate the critical nutrient loads related to ecological thresholds for total nitrogen, total phosphorus and chlorophyll-a. The current (i.e., 2012) and future (i.e., 2050) nutrient loads were derived from the water quality MARINA-Lakes model. Nitrogen loads exceeded the nitrogen threshold in 2012. Phosphorus loads were below all ecological thresholds in 2012. Ecological thresholds are exceeded in 2050 with limited environmental policies, and urbanization may increase nutrient loads above the ecological thresholds in 2050. Recycling and reallocating animal manure is needed to avoid future water pollution in Lake Baiyangdian. Our study highlights the need for effective policies for clean water based on policy-relevant indicators.

Characterizing 19 thousand Chinese lakes, ponds and reservoirs by morphometric, climate and sediment characteristics.

Chinese lakes, including ponds and reservoirs, are increasingly threatened by algal blooms. Yet, each lake is unique, leading to large inter-lake variation in lake vulnerability to algal blooms. Here, we aim to assess the effects of unique lake characteristics on lake vulnerability to algal blooms. To this end, we built a novel and comprehensive database of lake morphometric, climate and sediment characteristics of 19,536 Chinese lakes, including ponds and reservoirs (>0.1 km2). We assessed lake characteristics for nine stratification classes and show that lakes, including ponds and reservoirs, in eastern China typically have a warm stratification class (Tavg>4 °C) and are slightly deeper than those in western China. Model results for representative lakes suggest that the most vulnerable lakes to algal blooms are in eastern China where pollution levels are also highest. Our characterization provides an important baseline to inform policymakers in what regions lakes are potentially most vulnerable to algal blooms.

The value of novel ecosystems: Disclosing the ecological quality of quarry lakes.

Intense sand and gravel mining has created numerous man-made lakes around the world in the past century. These small quarry lakes (1-50 ha) are usually hydrologically isolated, often deep (6-40 m) and stratify during summer and in cold winters. Due to their small size, these deep man-made lakes are usually not included in the regular monitoring campaigns, e.g. as required for the European Water Framework Directive (WFD). Therefore, not much is known about the ecological functioning of these novel ecosystems. During two summers, we determined the macrophyte diversity and measured a range of physico-chemical and biological parameters in 51 quarry lakes in the catchment area of the rivers Meuse and Rhine. We compared the results of this campaign to the chemical and macrophyte sampling as performed for the WFD in the immediate surrounding shallow standing waters. Alpha (local) and beta diversity (regional), and local contribution to beta diversity were calculated for the whole region of which beta diversity was further partitioned into a true species replacement and richness difference component. Quarry lakes contain higher water quality reflected by lower nutrient and chlorophyll-a concentration compared with shallow water bodies. Additionally, quarry lakes contribute significantly to the regional macrophyte diversity pool by harboring distinctly different macrophyte communities (beta diversity - replacement). Specifically quarry lakes with a total phosphorus concentration in the water column below 35 µg P/l contribute most to beta diversity among quarry lakes. Novel ecosystems such as deep quarry lakes are often perceived as less valuable ecosystems, with strong implications regarding their management. Our results show that quarry lakes are in general of better chemical and biological quality compared with shallow standing waters. We therefore call for a more integrated assessment of the quality of quarry lakes and corresponding management strategy of these waters by water managers.

Warming and CO2 effects under oligotrophication on temperate phytoplankton communities.

Eutrophication, global warming, and rising carbon dioxide (CO2) levels are the three most prevalent pressures impacting the biosphere. Despite their individual effects are well-known, it remains untested how oligotrophication (i.e. nutrients reduction) can alter the planktonic community responses to warming and elevated CO2 levels. Here, we performed an indoor mesocosm experiment to investigate the warming × CO2 interaction under a nutrient reduction scenario (40%) mediated by an in-lake management strategy (i.e. addition of a commercial solid-phase phosphorus sorbent -Phoslock®) on a natural freshwater plankton community. Biomass production increased under warming × CO2 relative to present-day conditions; however, a Phoslock®-mediated oligotrophication reduced such values by 30-70%. Conversely, the warming × CO2 × oligotrophication interaction stimulated the photosynthesis by 20% compared to ambient nutrient conditions, and matched with higher resource use efficiency (RUE) and nutrient demand. Surprisingly, at a group level, we found that the multi-stressors scenario increased the photosynthesis in eukaryotes by 25%, but greatly impaired in cyanobacteria (ca. -25%). This higher cyanobacterial sensitivity was coupled with a reduced light harvesting efficiency and compensation point. Since Phoslock®-induced oligotrophication unmasked a strong negative warming × CO2 effect on cyanobacteria, it becomes crucial to understand how the interplay between climate change and nutrient abatement actions may alter the, ecosystems functioning. With an integrative understanding of these processes, policy makers will design more appropriate management strategies to improve the ecological status of aquatic ecosystems without compromising their ecological attributes and functioning.

A Generically Parameterized model of Lake eutrophication (GPLake) that links field-, lab- and model-based knowledge.

Worldwide, eutrophication is threatening lake ecosystems. To support lake management numerous eutrophication models have been developed. Diverse research questions in a wide range of lake ecosystems are addressed by these models. The established models are based on three key approaches: the empirical approach that employs field surveys, the theoretical approach in which models based on first principles are tested against lab experiments, and the process-based approach that uses parameters and functions representing detailed biogeochemical processes. These approaches have led to an accumulation of field-, lab- and model-based knowledge, respectively. Linking these sources of knowledge would benefit lake management by exploiting complementary information; however, the development of a simple tool that links these approaches was hampered by their large differences in scale and complexity. Here we propose a Generically Parameterized Lake eutrophication model (GPLake) that links field-, lab- and model-based knowledge and can be used to make a first diagnosis of lake water quality. We derived GPLake from consumer-resource theory by the principle that lacustrine phytoplankton is typically limited by two resources: nutrients and light. These limitations are captured in two generic parameters that shape the nutrient to chlorophyll-a relations. Next, we parameterized GPLake, using knowledge from empirical, theoretical, and process-based approaches. GPLake generic parameters were found to scale in a comparable manner across data sources. Finally, we show that GPLake can be applied as a simple tool that provides lake managers with a first diagnosis of the limiting factor and lake water quality, using only the parameters for lake depth, residence time and current nutrient loading. With this first-order assessment, lake managers can easily assess measures such as reducing nutrient load, decreasing residence time or changing depth before spending money on field-, lab- or model- experiments to support lake management.

Modelling induced bank filtration effects on freshwater ecosystems to ensure sustainable drinking water production.

Induced bank filtration (IBF) is a water abstraction technology using different natural infiltration systems for groundwater recharge, such as river banks and lake shores. It is a cost-effective pre-treatment method for drinking water production used in many regions worldwide, predominantly in urban areas. Until now, research concerning IBF has almost exclusively focussed on the purification efficiency and infiltration capacity. Consequently, knowledge about the effects on source water bodies is lacking. Yet, IBF interrupts groundwater seepage and affects processes in the sediment potentially resulting in adverse effects on lake or river water quality. Securing sufficient source water quality, however, is important for a sustainable drinking water production by IBF. In this study, we analysed the effects of five predicted mechanisms of IBF on shallow lake ecosystems using the dynamic model PCLake: declining CO2 and nutrient availability, as well as increasing summer water temperatures, sedimentation rates and oxygen penetration into sediments. Shallow lake ecosystems are abundant worldwide and characterised by the occurrence of alternative stable states with either clear water and macrophyte dominance or turbid, phytoplankton-dominated conditions. Our results show that IBF in most scenarios increased phytoplankton abundance and thus had adverse effects on shallow lake water quality. Threshold levels for critical nutrient loading inducing regime shifts from clear to turbid conditions were up to 80% lower with IBF indicating a decreased resilience to eutrophication. The effects were strongest when IBF interrupted the seepage of CO2 rich groundwater resulting in lower macrophyte growth. IBF could also enhance water quality, but only when interrupting the seepage of groundwater with high nutrient concentrations. Higher summer water temperatures increased the share of cyanobacteria in the phytoplankton community and thus the risk of toxin production. In relative terms, the effects of changing sedimentation rates and oxygen penetration were small. Lake depth and size influenced the effect of IBF on critical nutrient loads, which was strongest in shallower and smaller lakes. Our model results stress the need of a more comprehensive ecosystem perspective including an assessment of IBF effects on threshold levels for regime shifts to prevent high phytoplankton abundance in the source water body and secure a sustainable drinking water supply.

Effect of river restoration on life-history strategies in fish communities.

Assessments of river restoration outcomes are mostly based on taxonomic identities of species, which may not be optimal because a direct relationship to river functions remains obscure and results are hardly comparable across biogeographic borders. The use of ecological species trait information instead of taxonomic units may help to overcome these challenges. Abundance data for fish communities were gathered from 134 river restoration projects conducted in Switzerland, Germany and Finland, monitored for up to 15 years. These data were related to a dataset of 22 categories of ecological traits describing fish life-history strategies to assess the outcome of the restoration projects. Restoration increased trait functional diversity and evenness in projects that were situated in the potamal zone of rivers. Restoration effect increased with the length of the restored river reaches. In areas with low levels of anthropogenic land use, the peak of the restoration effect was reached already within one to five years after the restoration and effect receded thereafter, while communities responded later in areas with higher levels of anthropogenic land use. In the lower potamal zone, a shift towards opportunistic life-history strategists was observed. In the upper rhithral zone, in contrast, species with an opportunistic life-history strategy increased only in the first five years of restoration, followed by a shift towards equilibrium strategists at restorations older than 5 years. This pattern was more pronounced in rivers with higher level of anthropogenic land use and longer restored river reaches. Restoration reduced the variability in community trait composition between river reaches suggesting that community trait composition within these zones converges when rivers are restored. This study showed how ecological traits are suitable to analyse restoration outcomes and how such an approach can be used for the evaluation and comparison of environmental management actions across geographical regions.

An affordable and reliable assessment of aquatic decomposition: Tailoring the Tea Bag Index to surface waters.

Litter decomposition is a vital part of the global carbon cycle as it determines not only the amount of carbon to be sequestered, but also how fast carbon re-enters the cycle. Freshwater systems play an active role in the carbon cycle as it receives, and decomposes, terrestrial litter material alongside decomposing aquatic plant litter. Decomposition of organic matter in the aquatic environment is directly controlled by water temperature and nutrient availability, which are continuously affected by global change. We adapted the Tea Bag Index (TBI), a highly standardized methodology for determining soil decomposition, for lakes by incorporating a leaching factor. By placing Lipton pyramid tea bags in the aquatic environment for 3 h, we quantified the period of intense leaching which usually takes place prior to litter (tea) decomposition. Standard TBI methodology was followed after this step to determine how fast decomposition takes place (decomposition rate, k1) and how much of the material cannot be broken down and is thus sequestered (stabilization factor, S). A Citizen Science project was organized to test the aquatic TBI in 40 European lakes located in four climate zones, ranging from oligotrophic to hypereutrophic systems. We expected that warmer and/or eutrophic lakes would have a higher decomposition rate and a more efficient microbial community resulting in less tea material to be sequestered. The overall high decomposition rates (k1) found confirm the active role lakes play in the global carbon cycle. Across climate regions the lakes in the warmer temperate zone displayed a higher decomposition rate (k1) compared to the colder lakes in the continental and polar zones. Across trophic states, decomposition rates were higher in eutrophic lakes compared to oligotrophic lakes. Additionally, the eutrophic lakes showed a higher stabilization (S), thus a less efficient microbial community, compared to the oligotrophic lakes, although the variation within this group was high. Our results clearly show that the TBI can be used to adequately assess the decomposition process in aquatic systems. Using "alien standard litter" such as tea provides a powerful way to compare decomposition across climates, trophic states and ecosystems. By providing standardized protocols, a website, as well as face to face meetings, we also showed that collecting scientifically relevant data can go hand in hand with increasing scientific and environmental literacy in participants. Gathering process-based information about lake ecosystems gives managers the best tools to anticipate and react to future global change. Furthermore, combining this process-based information with citizen science, thus outreach, is in complete agreement with the Water Framework Directive goals as set in 2010.

Local Functioning, Landscape Structuring: Drivers of Soil Microbial Community Structure and Function in Peatlands.

Agricultural peatlands are essential for a myriad of ecosystem functions and play an important role in the global carbon (C) cycle through C sequestration. Management of these agricultural peatlands takes place at different spatial scales, ranging from local to landscape management, and drivers of soil microbial community structure and function may be scale-dependent. Effective management for an optimal biogeochemical functioning thus requires knowledge of the drivers on soil microbial community structure and functioning, as well as the spatial scales upon which they are influenced. During two field campaigns, we examined the importance of different drivers (i.e., soil characteristics, nutrient management, vegetation composition) at two spatial scales (local vs. landscape) for, respectively, the soil microbial community structure (determined by PLFA) and soil microbial community functional capacity (as assessed by CLPP) in agricultural peatlands. First, we show by an analysis of PLFA profiles that the total microbial biomass changes with soil moisture and relative C:P nutrient availability. Secondly, we showed that soil communities are controlled by a distinct set of drivers at the local, as opposed to landscape, scale. Community structure was found to be markedly different between areas, in contrast to community function which showed high variability within areas. We further found that microbial structure appears to be controlled more at a landscape scale by nutrient-related variables, whereas microbial functional capacity is driven locally through plant community feedbacks. Optimal management strategies within such peatlands should therefore consider the scale-dependent action of soil microbial community drivers, for example by first optimizing microbial structure at the landscape scale by targeted areal management, and then optimizing soil microbial function by local vegetation management.

Managing Successional Stage Heterogeneity to Maximize Landscape-Wide Biodiversity of Aquatic Vegetation in Ditch Networks.

The presence of a high diversity of different successional stages in a landscape may help to conserve and promote landscape-wide biodiversity. A strategy to achieve this is using Cyclic Rejuvenation through Management (CRM), an approach employed in a variety of different ecosystems. CRM periodically resets the successional stages in a landscape. For aquatic systems this constitutes vegetation removal and dredging. For this approach to be useful (a) successional stages are required to be different in community composition and (b) these differences need to be caused by true replacement of species between stages. While potentially valid, these assumptions are not generally tested prior to application of CMR. In this study we test these assumptions to explore the usefulness of managing on successional stage heterogeneity for maximizing landscape-wide aquatic plant diversity. We carried out vegetation surveys in the ditch networks of 21 polder landscapes in Netherlands, each containing 24 ditch reaches. Using a clustering approach combined with insight from literature on vegetation succession in these systems we assigned our sampled communities to defined successional stages. After partitioning landscape diversity into its alpha and beta components, we quantified the relative importance of replacement among successional stages. Next, through scenario analyses based on simulations we studied the effects of reducing successional stage heterogeneity on landscape-wide biodiversity. Results showed that differences in community composition among successional stages were a potentially important factor contributing to landscape diversity. Early successional stages were characterized by higher replacement of species compared to late successional stages. In a scenario of gradual decrease of heterogeneity through the systematic loss of the earliest successional stages we found 20% of the species richness in a polder was lost, pointing toward the importance of maintaining early successional stages in a polder. This makes a compelling case for application of CRM within agricultural drainage ditch landscapes to maximize regional aquatic plant diversity. While applied to drainage ditch systems, our data-driven approach is broadly applicable to other systems and may help in providing first indications of the potential of the CRM approach. We argue that CRM may maintain and promote regional biodiversity without compromising the hydrological function of the systems.

Temperature Effects Explain Continental Scale Distribution of Cyanobacterial Toxins.

Insight into how environmental change determines the production and distribution of cyanobacterial toxins is necessary for risk assessment. Management guidelines currently focus on hepatotoxins (microcystins). Increasing attention is given to other classes, such as neurotoxins (e.g., anatoxin-a) and cytotoxins (e.g., cylindrospermopsin) due to their potency. Most studies examine the relationship between individual toxin variants and environmental factors, such as nutrients, temperature and light. In summer 2015, we collected samples across Europe to investigate the effect of nutrient and temperature gradients on the variability of toxin production at a continental scale. Direct and indirect effects of temperature were the main drivers of the spatial distribution in the toxins produced by the cyanobacterial community, the toxin concentrations and toxin quota. Generalized linear models showed that a Toxin Diversity Index (TDI) increased with latitude, while it decreased with water stability. Increases in TDI were explained through a significant increase in toxin variants such as MC-YR, anatoxin and cylindrospermopsin, accompanied by a decreasing presence of MC-LR. While global warming continues, the direct and indirect effects of increased lake temperatures will drive changes in the distribution of cyanobacterial toxins in Europe, potentially promoting selection of a few highly toxic species or strains.

Response of Submerged Macrophyte Communities to External and Internal Restoration Measures in North Temperate Shallow Lakes.

Submerged macrophytes play a key role in north temperate shallow lakes by stabilizing clear-water conditions. Eutrophication has resulted in macrophyte loss and shifts to turbid conditions in many lakes. Considerable efforts have been devoted to shallow lake restoration in many countries, but long-term success depends on a stable recovery of submerged macrophytes. However, recovery patterns vary widely and remain to be fully understood. We hypothesize that reduced external nutrient loading leads to an intermediate recovery state with clear spring and turbid summer conditions similar to the pattern described for eutrophication. In contrast, lake internal restoration measures can result in transient clear-water conditions both in spring and summer and reversals to turbid conditions. Furthermore, we hypothesize that these contrasting restoration measures result in different macrophyte species composition, with added implications for seasonal dynamics due to differences in plant traits. To test these hypotheses, we analyzed data on water quality and submerged macrophytes from 49 north temperate shallow lakes that were in a turbid state and subjected to restoration measures. To study the dynamics of macrophytes during nutrient load reduction, we adapted the ecosystem model PCLake. Our survey and model simulations revealed the existence of an intermediate recovery state upon reduced external nutrient loading, characterized by spring clear-water phases and turbid summers, whereas internal lake restoration measures often resulted in clear-water conditions in spring and summer with returns to turbid conditions after some years. External and internal lake restoration measures resulted in different macrophyte communities. The intermediate recovery state following reduced nutrient loading is characterized by a few macrophyte species (mainly pondweeds) that can resist wave action allowing survival in shallow areas, germinate early in spring, have energy-rich vegetative propagules facilitating rapid initial growth and that can complete their life cycle by early summer. Later in the growing season these plants are, according to our simulations, outcompeted by periphyton, leading to late-summer phytoplankton blooms. Internal lake restoration measures often coincide with a rapid but transient colonization by hornworts, waterweeds or charophytes. Stable clear-water conditions and a diverse macrophyte flora only occurred decades after external nutrient load reduction or when measures were combined.