Timing matters: Sampling frequency for early-warning indicators across food web components in a virtual lake

Shallow lakes are known for sudden shifts between a desired clear and an undesired turbid state despite only incremental changes in the underlying drivers. Such sudden shifts are a major challenge for lake managers who can be confronted with abrupt losses of desired ecosystem services without easily observable warning signals. Predictive tools for the loss of ecosystem resilience are vital to respond with timely mitigation measures and avert a shift to the undesired state. Early-warning indicators (EWIs) have faithfully preceded critical transitions in minimal models but have proven more elusive in real-world data, suggesting a mismatch between measurement strategy and the detectability of EWIs. Here, we capitalize on data simulated using the aquatic ecosystem model PCLake+ which represents real systems more closely than reductionistic models and which allows the generation of critical transitions in response to gradual changes in phosphorus load. We tested the effect of different sampling intervals (daily to yearly) on the detection of three often-used EWIs across a range of food web and nutrient-related variables. Moreover, we included one integrated sampling interval (yearly average of daily measurements) to represent time-integrated measurements. EWIs generally performed better at shorter intervals (daily, weekly) but integrated measurements over the year also proved suitable to detect oncoming state shifts. We propose that lake managers should aim for high-frequency measurements of variables that can be easily and cheaply measured (e.g. oxygen, Secchi) or, alternatively, focus on integrated approaches using passive samplers or sedimented material

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.</p

Exploring how cyanobacterial traits affect nutrient loading thresholds in shallow lakes: A modelling approach

Globally, many shallow lakes have shifted from a clear macrophyte-dominated state to a turbid phytoplankton-dominated state due to eutrophication. Such shifts are often accompanied by toxic cyanobacterial blooms, with specialized traits including buoyancy regulation and nitrogen fixation. Previous work has focused on how these traits contribute to cyanobacterial competitiveness. Yet, little is known on how these traits affect the value of nutrient loading thresholds of shallow lakes. These thresholds are defined as the nutrient loading at which lakes shift water quality state. Here, we used a modelling approach to estimate the effects of traits on nutrient loading thresholds. We incorporated cyanobacterial traits in the process-based ecosystem model PCLake+, known for its ability to determine nutrient loading thresholds. Four scenarios were simulated, including cyanobacteria without traits, with buoyancy regulation, with nitrogen fixation, and with both traits. Nutrient loading thresholds were obtained under N-limited, P-limited, and colimited conditions. Results show that cyanobacterial traits can impede lake restoration actions aimed at removing cyanobacterial blooms via nutrient loading reduction. However, these traits hardly affect the nutrient loading thresholds for clear lakes experiencing eutrophication. Our results provide references for nutrient loading thresholds and draw attention to cyanobacterial traits during the remediation of eutrophic water bodies

Warming and CO2 effects under oligotrophication on temperate phytoplankton 2 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

Alternative stable states in large shallow lakes?

Many lakes worldwide are experiencing great change due to eutrophication. Consequently, species composition changes, toxic algal blooms proliferate, and drinking water supplies dwindle. The transition to the deteriorated state can be catastrophic with an abrupt change from macrophyte to phytoplankton domination. This has been shown repeatedly in small lakes. Whether such alternative stable states also exist in large shallow lakes is less clear, however. Here we discuss the characteristics that give rise to alternative stable states in large shallow lakes either in the lake as whole or restricted to specific regions of the lake. We include the effect of lake size, spatial heterogeneity and internal connectivity on a lake's response along the eutrophication axis. As a case study, we outline the eutrophication history of Lake Taihu (China) and illustrate how lake size, spatial heterogeneity and internal connectivity can explain the observed spatial presence of different states. We discuss whether these states can be alternatively stable by comparing the data with model output (PCLake). These findings are generalised for other large, shallow lakes. We conclude that locations with prevailing size effects generally lack macrophytes; and, therefore, alternative stable states are unlikely to occur there. However, most large shallow lakes have macrophytes whose presence remains unexplained when only size effect is taken into account. By including spatial heterogeneity in the analysis, the presence of macrophytes and alternative stable states in large shallow lakes is better understood. Finally, internal connectivity is important because a high internal connectivity reduces the stability of alternative states

Warming and CO2 effects under oligotrophication on temperate phytoplankton 2 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

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

Exploring, exploiting and evolving diversity of aquatic ecosystem models: A community perspective

Here, we present a community perspective on how to explore, exploit and evolve the diversity in aquatic ecosystem models. These models play an important role in understanding the functioning of aquatic ecosystems, filling in observation gaps and developing effective strategies for water quality management. In this spirit, numerous models have been developed since the 1970s. We set off to explore model diversity by making an inventory among 42 aquatic ecosystem modellers, by categorizing the resulting set of models and by analysing them for diversity. We then focus on how to exploit model diversity by comparing and combining different aspects of existing models. Finally, we discuss how model diversity came about in the past and could evolve in the future. Throughout our study, we use analogies from biodiversity research to analyse and interpret model diversity. We recommend to make models publicly available through open-source policies, to standardize documentation and technical implementation of models, and to compare models through ensemble modelling and interdisciplinary approaches. We end with our perspective on how the field of aquatic ecosystem modelling might develop in the next 5–10 years. To strive for clarity and to improve readability for non-modellers, we include a glossary