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© Copyright 2022. Sociedad Meteorológica Estadounidense (AMS). Para obtener permiso para reutilizar cualquier parte de este Trabajo, comuníquese con permisos@ametsoc. org . Cualquier uso del material en este Trabajo que se determine como "uso justo" según la Sección 107 de la Ley de derechos de autor de EE. UU. (17 Código de EE. UU. § 107) o que cumpla las condiciones especificadas en la Sección 108 de la Ley de derechos de autor de EE. ) no requiere el permiso de la AMS. La republicación, la reproducción sistemática, la publicación en forma electrónica, como en un sitio web o en una base de datos de búsqueda, u otros usos de este material, excepto los exentos de la declaración anterior, requieren un permiso por escrito o una licencia de la AMS. Todas las publicaciones periódicas y monográficas de AMS están registradas en el Centro de Autorización de Derechos de Autor (https://www.copyright.com ). Se proporcionan detalles adicionales en la declaración de política de derechos de autor de AMS, disponible en el sitio web de AMS ( https://www.ametsoc.org/PUBSCopyrightPolicy ) ., In 2021, both social and economic activities began to return towards the levels preceding the COVID-19 pandemic for some parts of the globe, with others still experiencing restrictions. Meanwhile, the climate has continued to respond to the ongoing increase in greenhouse gases and resulting warming. La Niña, a phenomenon which tends to depress global temperatures while changing rainfall patterns in many regions, prevailed for all but two months of the year. Despite this, 2021 was one of the six-warmest years on record as measured by global mean surface temperature with an anomaly of between +0.21° and +0.28°C above the 1991–2020 climatology., Lake surface water temperatures from satellite data have been generated within the GloboLakes project funded by the UK National Environment Research Council (NE/J023345/2), with extensions funded by the EU Copernicus Climate Change Service (C3S) programme...

Quantifying the potential for abrupt non-linear changes in ecological communities is a key managerial goal, leading to a significant body of research aimed at identifying indicators of approaching regime shifts. Most of this work has built on the theory of bifurcations, with the assumption that critical transitions are a common feature of complex ecological systems. This has led to the development of a suite of often inaccurate early warning signals (EWSs), with more recent techniques seeking to overcome their limitations by analysing multivariate time series or applying machine learning. However, it remains unclear whether regime shifts and/or critical transitions are common occurrences in natural systems, and – if they are present – whether classic and second-generation EWS methods predict rapid community change. Here, using multitrophic data on nine lakes from around the world, we both identify the type of transition a lake is exhibiting, and the reliability of classic and second generation EWSs methods to predict whole ecosystem change. We find few instances of critical transitions in our lake dataset, with different trophic levels often expressing different forms of abrupt change. The ability to predict this change is highly technique dependant, with multivariate EWSs generally classifying correctly, classical rolling window univariate EWSs performing not better than chance, and recently developed machine learning techniques performing poorly. Our results suggest that predictive ecology should start to move away from the concept of critical transitions and develop methods suitable for predicting change in the absence of the strict bounds of bifurcation theory.

Floating solar photovoltaic (FPV) deployments are increasing globally as the switch to renewable energy intensifies, representing a considerable water surface transformation. FPV installations can potentially impact aquatic ecosystem function, either positively or negatively. However, these impacts are poorly resolved given the challenges of collecting empirical data for field or modelling experiments. In particular, there is limited evidence on the response of phytoplankton to changes in water body thermal dynamics and light climate with FPV. Given the importance of understanding phytoplankton biomass and species composition for managing ecosystem services, we use an uncertainty estimation approach to simulate the effect of FPV coverage and array siting location on a UK reservoir. FPV coverage was modified in 10 % increments from a baseline with 0 % coverage to 100 % coverage for three different FPV array siting locations based on reservoir circulation patterns. Results showed that FPV coverage significantly impacted thermal properties, resulting in highly variable impacts on phytoplankton biomass and species composition. The impacts on phytoplankton were often dependent on array siting location as well as surface coverage. Changes to phytoplankton species composition were offset by the decrease in phytoplankton biomass associated with increasing FPV coverage. We identified that similar phytoplankton biomass reductions could be achieved with less FPV coverage by deploying the FPV array on the water body's faster-flowing area than the central or slower flowing areas. The difference in response dependent on siting location could be used to tailor phytoplankton management in water bodies. Simulation of water body-FPV interactions efficiently using an uncertainty approach is an essential tool to rapidly develop understanding and ultimately inform FPV developers and water body managers looking to minimise negative impacts and maximise co-benefits.

Managing ecosystems to effectively preserve function and services requires reliable tools that can infer changes in the stability and dynamics of a system. Conceptually, functional diversity (FD) appears as a sensitive and viable monitoring metric stemming from suggestions that FD is a universally important measure of biodiversity and has a mechanistic influence on ecological processes. It is however unclear whether changes in FD consistently occur prior to state responses or vice versa, with no current work on the temporal relationship between FD and state to support a transition towards trait-based indicators. There is consequently a knowledge gap regarding when functioning changes relative to biodiversity change and where FD change falls in that sequence. We therefore examine the lagged relationship between planktonic FD and abundance-based metrics of system state (e.g. biomass) across five highly monitored lake communities using both correlation and cutting edge non-linear empirical dynamic modelling approaches. Overall, phytoplankton and zooplankton FD display synchrony with lake state but each lake is idiosyncratic in the strength of relationship. It is therefore unlikely that changes in plankton FD are identifiable before changes in more easily collected abundance metrics. These results highlight the power of empirical dynamic modelling in disentangling time lagged relationships in complex multivariate ecosystems, but suggest that FD cannot be generically viable as an early indicator. Individual lakes therefore require consideration of their specific context and any interpretation of FD across systems requires caution. However, FD still retains value as an alternative state measure or a trait representation of biodiversity when considered at the system level.

Managing ecosystems to effectively preserve function and services requires reliable tools that can infer changes in the stability and dynamics of a system. Conceptually, functional diversity (FD) appears a viable monitoring metric due to its mechanistic influence on ecological processes, but it is unclear whether changes in FD occur prior to state responses or vice versa. We examine the lagged relationship between planktonic FD and abundance-based metrics of system state (e.g. biomass) across five highly monitored lake communities using both correlation and non-linear causality approaches. Overall, phytoplankton and zooplankton FD display synchrony with lake state but each lake is idiosyncratic in the strength of relationship. It is therefore unlikely that changes in plankton FD are identifiable before changes in more easily collected abundance metrics. This suggests that FD is unlikely to be a viable early indicator, but has value as an alternative state measure if considered at the lake level.Graphical AbstractData accessibility statement:Lake Kinneret and Lake Kasumigaura data are available on request, with all other data publicly available and referenced throughout. All code for analysis is available in the Zenodo record (to be released) and the associated GitHub repository (https://github.com/duncanobrien/plankton-FD).

Floating solar photovoltaics (FPV) are deployed on aquatic systems worldwide as an alternative to ground- and roof-mounted installations. FPV installations represent a considerable water surface transformation, and the consequent threats and opportunities for hosting aquatic systems are poorly understood. Moreover, we must consider any impacts within the context of a changing climate, given FPV operational lifetimes.Impacts on aquatic systems may be significant given that FPV can perturb two key drivers of water body function - wind shear stress and solar radiation intensity. The potential impacts of changes on water body function are wide-ranging. For example, FPV may beneficially reduce the occurrence of nuisance algal blooms or could detrimentally lead to anoxic conditions, leading to the release of heavy metals from bed sediments. However, impacts are likely to be highly water body-specific, dependent on deployment configuration and be contingent on future climate conditions.To better understand FPV effects on aquatic ecosystem processes, which underpin ecosystem services, we extended an existing lake model to simulate FPV installations under future climate scenarios on a UK reservoir. We examined plausible changes to a range of meteorological variables, water temperatures, reservoir inflow and depth. We found that FPV alters key water quality properties, including water temperature and phytoplankton community composition. Depending on the conditions, the implications are positive or negative. Our analysis shows that FPV can partially mitigate the impacts of climate change by reducing water temperature. The extended lake model will help inform policymakers and practitioners on best practices for deploying FPV, minimising detrimental impacts and maximising co-benefits.

In many regions across the globe, extreme weather events such as storms have increased in frequency, intensity, and duration due to climate change. Ecological theory predicts that such extreme events should have large impacts on ecosystem structure and function. High winds and precipitation associated with storms can affect lakes via short‐term runoff events from watersheds and physical mixing of the water column. In addition, lakes connected to rivers and streams will also experience flushing due to high flow rates. Although we have a well‐developed understanding of how wind and precipitation events can alter lake physical processes and some aspects of biogeochemical cycling, our mechanistic understanding of the emergent responses of phytoplankton communities is poor. Here we provide a comprehensive synthesis that identifies how storms interact with lake and watershed attributes and their antecedent conditions to generate changes in lake physical and chemical environments. Such changes can restructure phytoplankton communities and their dynamics, as well as result in altered ecological function (e.g., carbon, nutrient and energy cycling) in the short‐ and long‐term. We summarize the current understanding of storm‐induced phytoplankton dynamics, identify knowledge gaps with a systematic review of the literature, and suggest future research directions across a gradient of lake types and environmental conditions., Our understanding of ecosystem‐scale responses to storm events is fragmented, and terminological variability in defining a “storm” hinders resolution of generalizable impacts. We provide a comprehensive synthesis of the interactions between physical properties and biological responses in lakes that demonstrates the context‐dependency of storm impacts; lake/watershed attributes and their antecedent conditions mediate the extent to which storms impact lake ecosystems. We develop a framework which conceptualizes how abrupt, storm‐induced changes in the lake environment influence phytoplankton community dynamics through functional traits, identifies current limitations, and highlights several avenues of research to narrow knowledge gaps in synthetic and interdisciplinary ways.

This article contributes to work in creative geographies through the lens of river spaces and a multimodal practice among an ensemble of artists/poets/scientists. The collaboration created two collage series with both planned and unplanned visual, textual, and audio-visual media. Orchestrated in a time of global pandemic, learnings from the ensemble’s online creative practice are shared and discussed, including: the methodological possibilities of online collaboration and the Internet’s ability to facilitate distanced and sustained co-creation, as well as the potential of collage and poetry to reimagine relationships with rivers.

Land use and climate change are anticipated to affect phytoplankton of lakes worldwide. The effects will depend on the magnitude of projected land use and climate changes and lake sensitivity to these factors. We used random forests fit with long-term (1971-2016) phytoplankton and cyanobacteria abundance time series, climate observations (1971-2016), and upstream catchment land use (global Clumondo models for the year 2000) data from 14 European and 15 North American lakes basins. We projected future phytoplankton and cyanobacteria abundance in the 29 focal lake basins and 1567 lakes across focal regions based on three land use (sustainability, middle of the road, and regional rivalry) and two climate (RCP 2.6 and 8.5) scenarios to mid-21st century. On average, lakes are expected to have higher phytoplankton and cyanobacteria due to increases in both urban land use and temperature, and decreases in forest habitat. However, the relative importance of land use and climate effects varied substantially among regions and lakes. Accounting for land use and climate changes in a combined way based on extensive data allowed us to identify urbanization as the major driver of phytoplankton development in lakes located in urban areas, and climate as major driver in lakes located in remote areas where past and future land use changes were minimal. For approximately one-third of the studied lakes, both drivers were relatively important. The results of this large scale study suggest the best approaches for mitigating the effects of human activity on lake phytoplankton and cyanobacteria will depend strongly on lake sensitivity to long-term change and the magnitude of projected land use and climate changes at a given location. Our quantitative analyses suggest local management measures should focus on retaining nutrients in urban landscapes to prevent nutrient pollution from exacerbating ongoing changes to lake ecosystems from climate change. ISSN:1354-1013 ISSN:1365-2486

Emergent properties of ecosystems are community attributes, such as structure and function, that arise from connections and interactions (e.g., predator–prey, competition) among populations, species, or assemblages that, when viewed together, provide a holistic representation that is more than the sum of its individual parts. Climate change is altering emergent properties of aquatic ecosystems through component responses, a combination of shifts in species range, phenology, distribution, and productivity, which lead to novel ecosystems that have no historical analog. The reshuffling, restructuring, and rewiring of aquatic ecosystems due to climate impacts are of high concern for natural resource management and conservation as these changes can lead to species extinctions and reductions in ecosystem services. Overall, we found that substantial progress has been made to advance our understanding of how climate change is affecting emergent properties of aquatic ecosystems. However, responses are incredibly complex, and high uncertainty remains for how systems will reorganize and function over the coming decades. This cross-system perspective summarizes the state of knowledge of climate-driven emergent properties in aquatic habitats with case studies that highlight mechanisms of change, observed or anticipated outcomes, as well as insights into confounding non-climate effects, research tools, and management approaches to advance the field.

Anthropogenic eutrophication caused by excess loading of nutrients, especially phosphorus (P), from catchments is a major cause of lake water quality degradation. The release of P from bed sediments to the water column, termed internal loading, can exceed catchment P load in eutrophic lakes, especially those that stratify during warm summer periods. Managing internal P loading is challenging, and although a range of approaches have been implemented, long-term success is often limited, requiring lake-specific solutions. Here, we assess the manipulation of lake residence time to inhibit internal loading in Elterwater, a shallow stratifying lake in the English Lake District, UK. Since 2016, additional inflowing water has been diverted into the inner basin of Elterwater to reduce its water residence time, with the intention of limiting the length of the stratified period and reducing internal loading. Combining eight years of field data in a Before-After-Control-Impact study with process-based hydrodynamic modelling enabled the quantification of the residence time intervention effects on stratification length, water column stability, and concentrations of chlorophyll a and P. Annual water residence time was reduced during the study period by around 40% (4.9 days). Despite this change, the lake continued to stratify and developed hypolimnetic anoxia. As a result, there was little significant change in phosphorus (as total or soluble reactive phosphorus) or chlorophyll a concentrations. Summer stratification length was 2 days shorter and 7% less stable with the intervention. Our results suggest that the change to water residence time in Elterwater was insufficient to induce large enough physical changes to improve water quality. However, the minor physical changes suggest the management measure had some impact and that larger changes in water residence time may have the potential to induce reductions in internal loading. Future assessments of management requirements should combine multi-year observations and physical lake modelling to provide improved understanding of the intervention effect size required to alter the physical structure of the lake, leading to increased hypolimnetic oxygen and reduced potential for internal loading.

Climate warming has caused the seasonal timing of many components of ecological food chains to advance (Thackeray et al. 2010, 2016). Differential shifts lead to phenological asynchrony, often referred to as trophic mismatch when it is detrimental for consumers (Cushing 1990). In the context of trophic interactions, it has been suggested that consumers will shift their phenology to adapt to shifts in the availability of their food source (Visser and Both 2005), but they rarely do so in practice (Thackeray et al. 2016; Kharouba et al. 2018). Whether such unequal shifts are detrimental or not is unresolved (Johansson and Jonzén 2012; Reed et al. 2013a; Samplonius et al. 2016; Radchuk et al. 2019; Visser and Gienapp 2019). At present there has been no consistent analysis of the links between temperature change, phenological asynchrony, and individual-to-population level impacts across taxa, trophic levels and biomes at a global scale. Instead, many of our insights into mismatch and its impacts stem from a handful of independent single-system studies, varying greatly in their conceptual basis and methodological approach. Here, we propose five criteria that all need to be met to demonstrate that temperature-mediated trophic mismatch poses a growing risk to consumers. These criteria are: 1) an ephemeral resource contributes a large proportion of the consumer’s diet; 2) asynchrony between phenology of consumer and resource is increasing over time; 3) interannual variation in asynchrony is driven by interannual variation in temperature; 4) asynchrony reduces consumer fitness, 5) mismatch impacts negatively on consumer population size or growth. We conduct a literature review of 109 papers studying 132 taxa, and find that for most taxa only two of the five criteria are met. Moreover, all five criteria are only assessed for two taxa. The most commonly-tested criteria are 1 and 2, and few studies further examined evidence for criteria 4 or 5. Furthermore, effects of mismatch are heavily skewed towards juvenile stages rather than adults. Crucially, nearly every study was conducted in Europe or North America, and most studies were on terrestrial secondary consumers. We thus lack a robust evidence base from which to draw general conclusions about the risk that climate-mediated trophic mismatch may pose to populations worldwide.

Embargo until February 26, 2023 Despite advances in conceptual understanding, single-stressor abatement approaches remain common in the management of fresh waters, even though they can produce unexpected ecological responses when multiple stressors interact. Here we identify limitations restricting the development of multiple-stressor management strategies and address these, bridging theory and practice, within a novel empirical framework. Those critical limitations include that (i) monitoring schemes fall short of accounting for theory on relationships between multiple-stressor interactions and ecological responses, (ii) current empirical modelling approaches neglect the prevalence and intensity of multiple-stressor interactions, and (iii) mechanisms of stressor interactions are often poorly understood. We offer practical recommendations for the use of empirical models and experiments to predict the effects of freshwater degradation in response to changes in multiple stressors, demonstrating this approach in a case study. Drawing on our framework, we offer practical recommendations to support the development of effective management strategies in three general multiple-stressor scenarios.

In an era of looming land scarcity and environmental degradation, the development of low carbon energy systems without adverse impacts on land and land-based resources is a global challenge. ‘Floatovoltaic’ energy systems—comprising floating photovoltaic (PV) panels over water—are an appealing source of low carbon energy as they spare land for other uses and attain greater electricity outputs compared to land-based systems. However, to date little is understood of the impacts of floatovoltaics on the hosting water body. Anticipating changes to water body processes, properties and services owing to floatovoltaic deployment represents a critical knowledge gap that may result in poor societal choices and water body governance. Here, we developed a theoretically-derived hierarchical effects framework for the assessment of floatovoltaic impacts on freshwater water bodies, emphasising ecological interactions. We describe how the presence of floatovoltaic systems may dramatically alter the air-water interface, with subsequent implications for surface meteorology, air-water fluxes and physical, chemical and biological properties of the recipient water body. We apply knowledge from this framework to delineate three response typologies—‘magnitude’, those for which the direction and magnitude of effect can be predicted; ‘direction’, those for which only the direction of effect can be predicted; and ‘uncertain’, those for which the response cannot be predicted—characterised by the relative importance of levels in the effects hierarchy. Illustrative decision trees are developed for an example water body response within each typology, specifically, evaporative water loss, cyanobacterial biomass, and phosphorus release from bed sediments, and implications for ecosystem services, including climate regulation, are discussed. Finally, the potential to use the new understanding of likely ecosystem perturbations to direct floatovoltaic design innovations and identify future research priorities is outlined, showcasing how inter-sectoral collaboration and environmental science can inform and optimise this low carbon, land-sparing renewable energy for ecosystem gains.

Plans are currently being drafted for the next decade of action on biodiversity—both the post‐2020 Global Biodiversity Framework of the Convention on Biological Diversity (CBD) and Biodiversity Strategy of the European Union (EU). Freshwater biodiversity is disproportionately threatened and underprioritized relative to the marine and terrestrial biota, despite supporting a richness of species and ecosystems with their own intrinsic value and providing multiple essential ecosystem services. Future policies and strategies must have a greater focus on the unique ecology of freshwater life and its multiple threats, and now is a critical time to reflect on how this may be achieved. We identify priority topics including environmental flows, water quality, invasive species, integrated water resources management, strategic conservation planning, and emerging technologies for freshwater ecosystem monitoring. We synthesize these topics with decades of first‐hand experience and recent literature into 14 special recommendations for global freshwater biodiversity conservation based on the successes and setbacks of European policy, management, and research. Applying and following these recommendations will inform and enhance the ability of global and European post‐2020 biodiversity agreements to halt and reverse the rapid global decline of freshwater biodiversity. We thank the organizers of the ALTER‐Net/EKLIPSE Post‐2020 Biodiversity Workshop for discussions that led to this collaboration. CBvR was supported by a Fulbright Early Career Scholar Award from the Fulbright Spain Commission, SJT by the NERC Highlight Topic “Hydroscape” (NE/N006437/1), SCJ and GK by the “GLANCE” project (01LN1320A) from the German Federal Ministry of Education and Research (BMBF), HPG by the BMBF “BIBS” project (01LC1501G1), KAW by the Rural & Environment Science & Analytical Services Division of the Scottish Government (2016–2021 Strategic Research programme), SD by the Leibniz Competition (J45/2018), AIL by FCT (CESAM; UID/AMB/50017/2019), IJ by the J. E. Purkyně Fellowship of the Czech Academy of Science, and VH by a Ramon y Cajal Contract (RYC‐2013‐13979).

Aim: Hotspots of human activity are focal points for ecosystem disturbance and non‐native introduction, from which invading populations disperse and spread. As such, connectivity to locations used by humans may influence the likelihood of invasion. Moreover, connectivity in freshwater ecosystems may follow the hydrological network. Here we tested whether multiple forms of connectivity to human recreational activities promotes biological invasion of freshwater ecosystems. Location: England, UK. Time period: 1990–2018. Major taxa studied: One hundred and twenty‐six non‐native freshwater birds, crustaceans, fish, molluscs and plants. Methods: Machine learning was used to predict spatial gradients in human recreation and two high risk activities for invasion (fishing and water sports). Connectivity indices were developed for each activity, in which human influence decayed from activity hotspots according to Euclidean distance (spatial connectivity) or hydrological network distance (downstream, upstream and along‐channel connectivity). Generalized linear mixed models identified the connectivity type most associated to invasive species richness of each group, while controlling for other anthropogenic and environmental drivers. Results: Connectivity to humans generally had stronger positive effects on invasion than all other drivers except recording effort. Recreation had stronger influence than urban land cover, and for most groups high risk activities had stronger effects than general recreation. Downstream human connectivity was most important for invasion by most of the groups, potentially reflecting predominantly hydrological dispersal. An exception was birds, for which spatial connectivity was most important, possibly because of overland dispersal capacity. Main conclusions: These findings support the hypothesis that freshwater invasion is partly determined by an interaction between human activity and species dispersal in the hydrological network. By comparing alternative connectivity types for different human activities, our approach could enable robust inference of specific pathways and spread mechanisms associated with particular taxa. This would provide evidence to support better prioritization of surveillance and management for invasive non‐native species.

Water temperature is critical for the ecology of lakes. However, the ability to predict its spatial and seasonal variation is constrained by the lack of a thermal classification system. Here we define lake thermal regions using objective analysis of seasonal surface temperature dynamics from satellite observations. Nine lake thermal regions are identified that mapped robustly and largely contiguously globally, even for small lakes. The regions differed from other global patterns, and so provide unique information. Using a lake model forced by 21st century climate projections, we found that 12%, 27% and 66% of lakes will change to a lower latitude thermal region by 2080–2099 for low, medium and high greenhouse gas concentration trajectories (Representative Concentration Pathways 2.6, 6.0 and 8.5) respectively. Under the worst-case scenario, a 79% reduction in the number of lakes in the northernmost thermal region is projected. This thermal region framework can facilitate the global scaling of lake-research., Water temperature is a critical variable for lakes, but its spatial and temporal patterns are not well characterised globally. Here, the authors use surface temperature dynamics to define lake thermal regions that group lakes with similar patterns, and show how these regions shift under climate change.

The concentration of dissolved organic matter (DOM) in freshwaters is increasing in large areas of the world. In addition to carbon, DOM contains nitrogen and phosphorus and there is growing concern that these organic nutrients may be bioavailable and contribute to eutrophication. However, relatively few studies have assessed the potential for dissolved organic nitrogen (DON) or dissolved organic phosphorus (DOP) compounds to be bioavailable to natural river phytoplankton communities at different locations or times. Temporal and spatial variations in uptake, relative to environmental characteristics were examined at six riverine sites in two contrasting catchments in the UK. This study also examined how the uptake by riverine phytoplankton of four DON and four DOP compounds commonly found in rivers, varied with concentration. Total nitrogen (TN) and phosphorus (TP) concentrations, the proportion of inorganic nutrient species, and nutrient limitation varied temporally and spatially, as did the potential for DON and DOP uptake. All eight of the DOM compounds tested were bioavailable, but to different extents. Organic nutrient use depended on the concentration of the organic compound supplied, with simple compounds (urea and glucose-6-phosphate) supporting algal growth even at very low concentrations. DON use was negatively correlated with the TN and ammonia concentration and DOP use was negatively correlated with soluble reactive phosphorus (SRP) and dissolved organic carbon (DOC) concentration. The evidence indicates that DOM in rivers has been overlooked as a potential source of nutrients to phytoplankton and therefore as an agent of eutrophication.

Freshwater ecosystems are heavily impacted by multiple stressors, and a freshwater biodiversity crisis is underway. This realization has prompted calls to integrate global freshwater ecosystem data, including traditional taxonomic and newer types of data (e.g., eDNA, remote sensing), to more comprehensively assess change among systems, regions, and organism groups. We argue that data integration should be done, not only with the important purpose of filling gaps in spatial, temporal, and organismal representation, but also with a more ambitious goal: to study fundamental cross-scale biological phenomena. Such knowledge is critical for discerning and projecting ecosystem functional dynamics, a realm of study where generalizations may be more tractable than those relying on taxonomic specificity. Integration could take us beyond cataloging biodiversity losses, and toward predicting ecosystem change more broadly. Fundamental biology questions should be central to integrative, interdisciplinary research on causal ecological mechanisms, combining traditional measures and more novel methods at the leading edge of the biological sciences. We propose a conceptual framework supporting this vision, identifying key questions and uncertainties associated with realizing this research potential. Our framework includes five interdisciplinary "complementarities." First, research approaches may provide comparative complementarity when they offer separate realizations of the same focal phenomenon. Second, for translational complementarity, data from one research approach is used to translate that from another, facilitating new inferences. Thirdly, causal complementarity arises when combining approaches allows us to "fill in" cause-effect relationships. Fourth, contextual complementarity is realized when together research methodologies establish the wider ecological and spatiotemporal context within which focal biological responses occur. Finally, integration may allow us to cross inferential scales through scaling complementarity. Explicitly identifying the modes and purposes of integrating research approaches, and reaching across disciplines to establish appropriate collaboration will allow researchers to address major biological questions that are more than the sum of the parts.

The idea that living organisms may contribute to turbulence and mixing in lakes and oceans (biomixing) dates to the 1960s, but has attracted increasing attention in recent years. Recent modeling and experimental studies suggest that marine organisms can enhance turbulence as much as winds and tides in oceans, with an impact on mixing. However, other studies show opposite and contradictory results, precluding definitive conclusions regarding the potential importance of biomixing. For lakes, only models and lab studies are available. These generally indicate that small zooplankton or passive bodies generate turbulence but different levels of mixing depending on their abundance. Nevertheless, biogenic mixing is a complex problem, which needs to be explored in the field, to overcome limitations arising from numerical models and lab studies, and without altering the behavior of the animals under study.

Climate and land-use change drive a suite of stressors that shape ecosystems and interact to yield complex ecological responses (that is, additive, antagonistic and synergistic effects). We know little about the spatial scales relevant for the outcomes of such interactions and little about effect sizes. These knowledge gaps need to be filled to underpin future land management decisions or climate mitigation interventions for protecting and restoring freshwater ecosystems. This study combines data across scales from 33 mesocosm experiments with those from 14 river basins and 22 cross-basin studies in Europe, producing 174 combinations of paired-stressor effects on a biological response variable. Generalized linear models showed that only one of the two stressors had a significant effect in 39% of the analysed cases, 28% of the paired-stressor combinations resulted in additive effects and 33% resulted in interactive (antagonistic, synergistic, opposing or reversal) effects. For lakes, the frequencies of additive and interactive effects were similar for all spatial scales addressed, while for rivers these frequencies increased with scale. Nutrient enrichment was the overriding stressor for lakes, with effects generally exceeding those of secondary stressors. For rivers, the effects of nutrient enrichment were dependent on the specific stressor combination and biological response vari- able. These results vindicate the traditional focus of lake restoration and management on nutrient stress, while highlighting that river management requires more bespoke management solutions. This work was supported by the MARS project (Managing Aquatic Ecosystems and Water Resources under Multiple Stress) funded under the 7th EU Framework Programme, Theme 6 (Environment including Climate Change), contract no. 603378 (http://www.mars-project.eu). Further support was received through the ILES (SAW- 2015-IGB-1) and BIBS (BMBF 01LC1501G) projects. Partner organizations provided 25% cofunding through their institutional budgets. We thank J. Strackbein, J. Lorenz and L. Mack for their support. This work was supported by the MARS project (Managing Aquatic Ecosystems and Water Resources under Multiple Stress) funded under the 7th EU Framework Programme, Theme 6 (Environment including Climate Change), contract no. 603378 (http://www.mars-project.eu). Further support was received through the ILES (SAW- 2015-IGB-1) and BIBS (BMBF 01LC1501G) projects. Partner organizations provided 25% cofunding through their institutional budgets. We thank J. Strackbein, J. Lorenz and L. Mack for their support.

Zooplankton diel vertical migration (DVM) is an ecologically important process, affecting nutrient transport and trophic interactions. Available measurements of zooplankton displacement velocity during the DVM in the field are rare; therefore, it is not known which factors are key in driving this velocity. We measured the velocity of the migrating layer at sunset (upward bulk velocity) and sunrise (downwards velocity) in summer 2015 and 2016 in a lake using the backscatter strength (VBS) from an acoustic Doppler current profiler. We collected time series of temperature, relative change in light intensity chlorophyll-a concentration and zooplankton concentration. Our data show that upward velocities increased during the summer and were not enhanced by food, light intensity or by VBS, which is a proxy for zooplankton concentration and size. Upward velocities were strongly correlated with the water temperature in the migrating layer, suggesting that temperature could be a key factor controlling swimming activity. Downward velocities were constant, likely because Daphnia passively sink at sunrise, as suggested by our model of Daphnia sinking rate. Zooplankton migrations mediate trophic interactions and web food structure in pelagic ecosystems. An understanding of the potential environmental determinants of this behaviour is therefore essential to our knowledge of ecosystem functioning.

Global warming has advanced the timing of biological events, potentially leading to disruption across trophic levels. The potential importance of phenological change as a driver of population trends has been suggested. To fully understand the possible impacts, there is a need to quantify the scale of these changes spatially and according to habitat type. We studied the relationship between phenological trends, space and habitat type between 1965 and 2012 using an extensive UK dataset comprising 269 aphid, bird, butterfly and moth species. We modelled phenologies using generalized additive mixed models that included covariates for geographical (latitude, longitude, altitude), temporal (year, season) and habitat terms (woodland, scrub, grassland). Model selection showed that a baseline model with geographical and temporal components explained the variation in phenologies better than either a model in which space and time interacted or a habitat model without spatial terms. This baseline model showed strongly that phenologies shifted progressively earlier over time, that increasing altitude produced later phenologies and that a strong spatial component determined phenological timings, particularly latitude. The seasonal timing of a phenological event, in terms of whether it fell in the first or second half of the year, did not result in substantially different trends for butterflies. For moths, early season phenologies advanced more rapidly than those recorded later. Whilst temporal trends across all habitats resulted in earlier phenologies over time, agricultural habitats produced significantly later phenologies than most other habitats studied, probably because of nonclimatic drivers. A model with a significant habitat‐time interaction was the best‐fitting model for birds, moths and butterflies, emphasizing that the rates of phenological advance also differ among habitats for these groups. Our results suggest the presence of strong spatial gradients in mean seasonal timing and nonlinear trends towards earlier seasonal timing that varies in form and rate among habitat types.

Vertical mixing in lakes is a key driver of transport of ecologically important dissolved constituents, such as oxygen and nutrients. In this study we focus our attention on biomixing, which refers to the contribution of living organisms towards the turbulence and mixing of oceans and lakes. While several studies of biomixing in the ocean have been conducted, no in situ studies exist that assess the turbulence induced by freshwater zooplanktonic organisms under real environmental conditions. Here, turbulence is sampled during three different sampling days during the sunset diel vertical migration of Daphnia spp. in a small man-made lake. This common genus may create hydrodynamic disturbances in the lake interior where the thermal stratification usually suppresses the vertical diffusion. Concurrent biological sampling assessed the zooplankton vertical concentration profile. An acoustic-Doppler current profiler was also used to track zooplankton concentration and migration via the backscatter strength. Our datasets do not show biologically-enhanced dissipation rates of temperature variance and turbulent kinetic energy in the lake interior, despite Daphnia concentrations as high as 60 org. L−1. No large and significant turbulent patches were created within the migrating layer to generate irreversible mixing. This suggests that Daphnia do not affect the mixing in the lake at the organism concentrations observed here.

Through cascading effects within lake food webs, commercial and recreational fisheries may indirectly affect the abundances of organisms at lower trophic levels, such as phytoplankton, even if they are not directly consumed. So far, interactive effects of fisheries, changing trophic state and climate upon lake ecosystems have been largely overlooked. Here we analyse case studies from five European lake basins of differing trophic states (Lake Võrtsjärv, two basins of Windermere, Lake Geneva and Lake Maggiore) with long-term limnological and fisheries data. Decreasing phosphorus concentrations (re-oligotrophication) and increasing water temperatures have been reported in all five lake basins, while phytoplankton concentration has decreased only slightly or even increased in some cases. To examine possible ecosystem-scale effects of fisheries, we analysed correlations between fish and fisheries data, and other food web components and environmental factors. Re-oligotrophication over different ranges of the trophic scale induced different fish responsesIn the deeper lakes Geneva and Maggiore, we found a stronger link between phytoplankton and planktivorous fish and thus a more important cascading top-down effect than in other lakes. This connection makes careful ecosystem-based fisheries management extremely important for maintaining high water quality in such systems. We also demonstrated that increasing water temperature might favour piscivores at low phosphorus loading, but suppresses them at high phosphorus loading and might thus either enhance or diminish the cascading top-down control over phytoplankton with strong implications for water quality.

1. Anthropogenic pressures, including climate change, are causing nonlinear changes in ecosystems globally. The development of reliable early warning indicators (EWIs) to predict these changes is vital for the adaptive management of ecosystems and the protection of biodiversity, natural capital and ecosystem services. Increased variance and autocorrelation are potential early warning indicators and can be readily estimated from ecological time series. Here, we undertook a comprehensive test of the consistency between early warning indicators and nonlinear abundance change across species, trophic levels and ecosystem types. 2. We tested whether long-term abundance time series of 55 taxa (126 data sets) across multiple trophic levels in marine and freshwater ecosystems showed (i) significant nonlinear change in abundance ‘turning points’ and (ii) significant increases in variance and autocorrelation (‘early warning indicators’). For each data set, we then quantified the prevalence of three cases: true positives (early warning indicators and associated turning point), false negatives (turning point but no associated early warning indicators) and false positives (early warning indicators but no turning point). 3. True positives were rare, representing only 9% (16 of 170) of cases using variance, and 13% (19 of 152) of cases using autocorrelation. False positives were more prevalent than false negatives (53% vs. 38% for variance; 47% vs. 40% for autocorrelation). False results were found in every decade and across all trophic levels and ecosystems. 4. Time series that contained true positives were uncommon (8% for variance; 6% for autocorrelation), with all but one time series also containing false classifications. Coherence between the types of early warning indicators was generally low with 43% of time series categorized differently based on variance compared to autocorrelation. 5. Synthesis and applications. Conservation management requires effective early warnings of ecosystem change using readily available data, and variance and autocorrelation in abundance data have been suggested as candidates. However, our study shows that they consistently fail to predict nonlinear change. For early warning indicators to be effective tools for preventative management of ecosystem change, we recommend that multivariate approaches of a suite of potential indicators are adopted, incorporating analyses of anthropogenic drivers and process-based understanding.

Ecological resilience is developing into a credible paradigm for policy development and environmental management for preserving natural capital in a rapidly changing world. However, resilience emerges from complex interactions, limiting the translation of theory into practice. Main limitations include the following: (i) difficulty in quantification and detection of changes in ecological resilience, (ii) a lack of empirical evidence to support preventative orproactive management and (iii) difficulties in managing processes operating across socio-ecological systems that vary in space and time. We highlight recent research with the potential to address these limitations including new and/or improved indicators of resilience and tools to assess scale as a driver of resilience.Synthesis and applications. Effective resilience-based management must be adaptive in nature. To support this, we propose an operational model using resilience-based iterative management actions operating across scales.Effective resilience-based management must be adaptive in nature. To support this, we propose an operational model using resilience-based iterative management actions operating across scales.

Lakes are highly connected systems that are affected by a hierarchy of stressors operating at different scales, making them particularly sensitive to anthropogenic perturbation. Traditionally, lakes are studied as a whole system ‘from physics to fish’ and long-term monitoring programmes were initiated on this basis, some starting over a century ago. This chapter describes the long-term monitoring programme on the Cumbrian lakes, UK, how it is operated and how its scientific value is increased by combining it with additional activities. Case-studies are presented on the advances long-term research has made to testing ecological theory and understanding teleconnexions and phenology. Automatic high-frequency measurements are an important complementary approach that has been made possible by technological revolutions in computing, and telecommunications. They provide a window into the true dynamic nature of lakes that cannot be achieved by manual sampling. The large volume of data produced can now be quality controlled and analysed by bespoke software that has been developed in recent years by a global network of lake and data scientists. Finally, lake models constructed using the insights from monitoring, as well as experiments, are powerful ways to identify knowledge gaps and allow forecasts to be made of future responses to environmental change or management intervention. As other approaches become incorporated into lake research, such as Earth Observation and citizen science, the scale of knowledge about the system will increase, improving our ability to provide robust scientific advice for the sustainable management of these fragile, but important ecosystems.

There is a pressing need to apply stability and resilience theory to environmental management to restore degraded ecosystems effectively and to mitigate the effects of impending environmental change. Lakes represent excellent model case studies in this respect and have been used widely to demonstrate theories of ecological stability and resilience that are needed to underpin preventative management approaches. However, we argue that this approach is not yet fully developed because the pursuit of empirical evidence to underpin such theoretically grounded management continues in the absence of an objective probability framework. This has blurred the lines between intuitive logic (based on the elementary principles of probability) and extensional logic (based on assumption and belief) in this field.

There are many useful metrics currently available to explain ecological variation within phytobenthic communities. However, most metrics are challenging to use (requiring specialist taxonomic skills and knowledge), limiting their widespread applicability. Furthermore, because most metrics have been developed to represent ecological responses to single pressures, no single metric in isolation can effectively describe complex changes in the state of communities responding to multiple environmental pressures. Understanding of such changes therefore requires the use of multiple metrics to account for the impacts of many environmental pressures. This study explores the potential of functional and morphological classifications to explain phytobenthic community responses to differences in nutrient concentration, current velocity, simulated high flow disturbances and invertebrate grazers. Three previously used phytobenthic classifications and a new metric developed from a phytoplankton classification were tested using two datasets from streams in the north west of England in 2010. A combination of the newly applied morphological classification (using maximum linear dimension, surface area and volume) and a functional classification (using life-forms) showed great potential for aiding the understanding of phytobenthic community responses to environmental pressures. Furthermore, it is suggested that, with further testing, this new classification, which requires less specialist knowledge, could be widely implemented and would potentially give great insight into the ecology of the entire phytobenthic community.

Phenological changes have been observed globally for marine, freshwater and terrestrial species, and are an important element of the global biological ‘fingerprint’ of climate change. Differences in rates of change could desynchronize seasonal species interactions within a food web, threatening ecosystem functioning. Quantification of this risk is hampered by the rarity of long-term data for multiple interacting species from the same ecosystem and by the diversity of possible phenological metrics, which vary in their ecological relevance to food web interactions. We compare phenological change for phytoplankton (chlorophyll a), zooplankton (Daphnia) and fish (perch, Perca fluviatilis) in two basins of Windermere over 40 years and determine whether change has differed among trophic levels, while explicitly accounting for among-metric differences in rates of change. Though rates of change differed markedly among the nine metrics used, seasonal events shifted earlier for all metrics and trophic levels: zooplankton advanced most, and fish least, rapidly. Evidence of altered synchrony was found in both lake basins, when combining information from all phenological metrics. However, comparisons based on single metrics did not consistently detect this signal. A multimetric approach showed that across trophic levels, earlier phenological events have been associated with increasing water temperature. However, for phytoplankton and zooplankton, phenological change was also associated with changes in resource availability. Lower silicate, and higher phosphorus, concentrations were associated with earlier phytoplankton growth, and earlier phytoplankton growth was associated with earlier zooplankton growth. The developing trophic mismatch detected between the dominant fish species in Windermere and important zooplankton food resources may ultimately affect fish survival and portend significant impacts upon ecosystem functioning.We advocate that future studies on phenological synchrony combine data from multiple phenological metrics, to increase confidence in assessments of change and likely ecological consequences.

Physico-chemical gradients and local biotic interactions combine at different scales to cause spatial heterogeneity among lake zooplankton. However, current understanding is based overwhelmingly on natural systems and there have been few assessments of zooplankton spatial pattern in artificial lakes subjected to intensive management. Here, we investigate spatial structure among crustacean zooplankton at scales ranging from 1 m to 2700 m in the newly-formed Cardiff Bay (UK). We hypothesised that zooplankton spatial structuring in Cardiff Bay would be weakened by artificial aeration that was expected to create intense, continuous disturbance that homogenised physico-chemical conditions. Water quality across Cardiff Bay varied only moderately, and variations in zooplankton community composition were weakly related to temperature, oxygen, chlorophyll-a concentration and salinity at scales of around 2 km. However, spatial pattern explained over 60 % of variance in zooplankton composition at the smallest scales (0-5 m), implying that inter-specific interactions were involved. Additionally, the numerically dominant zooplankton (Daphnia hyalina (Leydig 1860), Eurytemora affinis (Poppe 1880) and Eucyclops agilis (Koch 1838)) had highly contagious distributions in warmer locations with increased chlorophyll-a reflecting aggregated response to abundant food. These data suggest that local biotic interactions among zooplankton and broader-scale links between dominant species and their algal food are sufficiently intense to persist despite the homogenising effects of artificial aeration and lake mixing. Such spatial pattern and scale-dependence has ramifications for zooplankton sampling while illustrating the self-organising capacity of predator-prey interactions even in novel, disturbed environments.

Plant phenology research has gained increasing attention because of the sensitivity of phenology to climate change and its consequences for ecosystem function. Recent technological development has made it possible to gather invaluable data at a variety of spatial and ecological scales. Despite our ability to observe phenological change at multiple scales, the mechanistic basis of phenology is still not well understood. Integration of multiple disciplines, including ecology, evolutionary biology, climate science, and remote sensing, with long-term monitoring data across multiple spatial scales is needed to advance understanding of phenology. We review the mechanisms and major drivers of plant phenology, including temperature, photoperiod, and winter chilling, as well as other factors such as competition, resource limitation, and genetics. Shifts in plant phenology have significant consequences on ecosystem productivity, carbon cycling, competition, food webs, and other ecosystem functions and services. We summarize recent advances in observation techniques across multiple spatial scales, including digital repeat photography, other complementary optical measurements, and solar-induced fluorescence, to assess our capability to address the importance of these scale-dependent drivers. Then, we review phenology models as an important component of earth system modeling. We find that the lack of species-level knowledge and observation data leads to difficulties in the development of vegetation phenology models at ecosystem or community scales. Finally, we recommend further research to advance understanding of the mechanisms governing phenology and the standardization of phenology observation methods across networks. With the opportunity for “big data” collection for plant phenology, we envision a breakthrough in process-based phenology modeling.

Changes in the seasonal timing of recurring biological events are considered to be a major component of the global “fingerprint” of climate change. One effect of these changes is that ecologically important seasonal species interactions could become desynchronised as a result of these shifts (i.e. phenological mismatching), leading to reductions in fitness for some or all of the organisms concerned. One important, but unresolved, issue is the extent to which variations in voltinism (the number of generations a population of a species produces per year) may serve to exacerbate, or confer resilience to, the effects of seasonal shifts. Univoltine organisms (those with one generation per year) will always suffer the deleterious consequences of phenological mismatch, whereas multivoltine species are likely to experience at least some relief from these negative effects in generations that occur later in the season. Conversely, univoltine species will experience continual selection to adapt to changing seasonality, whereas multivoltine species will experience reduced or no selection during those generations that occur later in the season. Here, we present a new theoretical model to explore the population consequences of scenarios of changing seasonality and varying voltinism in clonal species. We find that organisms that undergo multiple generations per year show greater resilience to phenological mismatching in the spring and adapt better to changing seasonality, because of the recovery of population size and genetic diversity after each spring mismatching event. These results have clear implications for management and conservation of populations that are threatened by the effects of mismatch.

Accumulating scientific evidence has demonstrated widespread shifts in the biological seasons. These shifts may modify seasonal interspecific interactions, with consequent impacts upon reproductive success and survival. However, current understanding of these impacts is based upon a limited number of studies that adopt a simplified ‘bottom-up’ food-chain paradigm, at a local scale. I argue that there is much insight to be gained by widening the scope of phenological studies to incorporate food-web interactions and landscape-scale processes across a diversity of ecosystem types, with the ultimate goal of developing a generic understanding of the systems most vulnerable to synchrony effects in the future. I propose that co-location of predator and prey phenological monitoring at sentinel sites, acting as research platforms for detailed food-web studies, experimentation and match-up with earth observation data, would be an important first step in this endeavour.

Peer-reviewed. Copyright: © 2016 Woolway et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.PLoS ONE 11(3): e0152466. doi:10.1371/journal.pone.0152466, Ecological and biogeochemical processes in lakes are strongly dependent upon water temperature. Long-term surface warming of many lakes is unequivocal, but little is known about the comparative magnitude of temperature variation at diel timescales, due to a lack of appropriately resolved data. Here we quantify the pattern and magnitude of diel temperature variability of surface waters using high-frequency data from 100 lakes. We show that the near-surface diel temperature range can be substantial in summer relative to long-term change and, for lakes smaller than 3 km2, increases sharply and predictably with decreasing lake area. Most small lakes included in this study experience average summer diel ranges in their near-surface temperatures of between 4 and 7°C. Large diel temperature fluctuations in the majority of lakes undoubtedly influence their structure, function and role in biogeochemical cycles, but the full implications remain largely unexplored., This project was funded by OTT Hydrometry Ltd and a University College London impact award awarded to RIW and is part of the Natural Environment Research Council (NERC) Sensor Network project United Kingdom Lake Ecological Observatory Network (UKLEON;NE/I007407/1). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

This work was funded by Natural Environment Research Council (NERC) grant NE/J02080X/1. The Rothamsted Insect Survey (RIS) is a National Capability strategically funded by BBSRC. Differences in phenological responses to climate change among species can desynchronise ecological interactions and thereby threaten ecosystem function. To assess these threats, we must quantify the relative impact of climate change on species at different trophic levels. Here, we apply a Climate Sensitivity Profile approach to 10,003 terrestrial and aquatic phenological data sets, spatially matched to temperature and precipitation data, to quantify variation in climate sensitivity. The direction, magnitude and timing of climate sensitivity varied markedly among organisms within taxonomic and trophic groups. Despite this variability, we detected systematic variation in the direction and magnitude of phenological climate sensitivity. Secondary consumers showed consistently lower climate sensitivity than other groups. We used mid-century climate change projections to estimate that the timing of phenological events could change more for primary consumers than for species in other trophic levels (6.2 versus 2.5–2.9 days earlier on average), with substantial taxonomic variation (1.1–14.8 days earlier on average). Postprint

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