Temperature, productivity, and habitat characteristics collectively drive lake food web structure.

While many efforts have been devoted to understand variations in food web structure among terrestrial and aquatic ecosystems, the environmental factors influencing food web structure at large spatial scales remain hardly explored. Here, we compiled biodiversity inventories to infer food web structure of 67 French lakes using an allometric niche-based model and tested how environmental variables (temperature, productivity, and habitat) influence them. By applying a multivariate analysis on 20 metrics of food web topology, we found that food web structural variations are represented by two distinct complementary and independent structural descriptors. The first is related to the overall trophic diversity, whereas the second is related to the vertical structure. Interestingly, the trophic diversity descriptor was mostly explained by habitat size (26.7% of total deviance explained) and habitat complexity (20.1%) followed by productivity (dissolved organic carbon: 16.4%; nitrate: 9.1%) and thermal variations (10.7%). Regarding the vertical structure descriptor, it was mostly explained by water thermal seasonality (39.0% of total deviance explained) and habitat depth (31.9%) followed by habitat complexity (8.5%) and size (5.5%) as well as annual mean temperature (5.6%). Overall, we found that temperature, productivity, and habitat characteristics collectively shape lake food web structure. We also found that intermediate levels of productivity, high levels of temperature (mean and seasonality), as well as large habitats are associated with the largest and most complex food webs. Our findings, therefore, highlight the importance of focusing on these three components especially in the context of global change, as significant structural changes in aquatic food webs could be expected under increased temperature, pollution, and habitat alterations.

Bayesian inference of physicochemical quality elements of tropical lagoon Nokoué (Benin).

In view of the very strong degradation of aquatic ecosystems, it is urgent to set up monitoring systems that are best able to report on the effects of the stresses they undergo. This is particularly true in developing countries, where specific and relevant quality standards and funding for monitoring programs are lacking. The objective of this study was to make a relevant and objective choice of physicochemical parameters informative of the main stressors occurring on African lakes and to identify their alteration thresholds. Based on statistical analyses of the relationship between several driving forces and the physicochemical parameters of the Nokoué lagoon, relevant physicochemical parameters were selected for its monitoring. An innovative method based on Bayesian statistical modeling was used. Eleven physicochemical parameters were selected for their response to at least one stressor and their threshold quality standards also established: Total Phosphorus (<4.5mg/L), Orthophosphates (<0.2mg/L), Nitrates (<0.5 mg/L), TKN (<1.85 mg/L), Dry Organic Matter (<5 mg/L), Dissolved Oxygen (>4 mg/L), BOD (<11.6 mg/L), Salinity (7.6 ‰), Water Temperature (<28.7 °C), pH (>6.2), and Transparency (>0.9 m). According to the System for the Evaluation of Coastal Water Quality, these thresholds correspond to "good to medium" suitability classes, except for total phosphorus. One of the original features of this study is the use of the bounds of the credibility interval of the fixed-effect coefficients as local weathering standards for the characterization of the physicochemical status of this anthropized African ecosystem.

Otolith shape analysis and daily increment validation during ontogeny of larval and juvenile European chub Squalius cephalus.

This assesses features of otoliths from laboratory-reared embryos, larvae and juvenile European chub Squalius cephalus from hatching to 180 days post-hatching (dph). We observed the development of the three pairs of otoliths (lapilli, sagittae and asterisci) and more precisely shape changes, as well as timing and deposition rate of increments of the lapilli. The lapilli and the sagittae were present at hatching, whereas the asterisci formed between 20 and 30 dph. The lapillus and sagitta shapes were round until 20 dph. From 60 dph the anterior and the posterior rostra of the sagittae were well developed, but very thin, making this otolith too fragile to manipulate for further studies of shape and validation of otolith increment deposition rate. The lapilli provided reliable age estimates for free embryos, larvae and juveniles up to 120 dph. However, caution should be taken when ageing fish older than 150 dph as an underestimation was noticeable. The regression of the number of otolith increments on age showed a slope and an intercept not significantly different from 1 and 0, respectively, which indicated that otolith growth increments were deposited on a daily basis, with the first microincrement occurring at hatching. Increment counts were consistent between three interpreters, indicating a consistent and reliable age estimate. This study validates that the otolith increment deposition rate can be used to assess hatching dates and daily growth of wild S. cephalus under 150 dph and in environments similar to the conditions used in this study.

Next-generation monitoring of aquatic biodiversity using environmental DNA metabarcoding.

Global biodiversity in freshwater and the oceans is declining at high rates. Reliable tools for assessing and monitoring aquatic biodiversity, especially for rare and secretive species, are important for efficient and timely management. Recent advances in DNA sequencing have provided a new tool for species detection from DNA present in the environment. In this study, we tested whether an environmental DNA (eDNA) metabarcoding approach, using water samples, can be used for addressing significant questions in ecology and conservation. Two key aquatic vertebrate groups were targeted: amphibians and bony fish. The reliability of this method was cautiously validated in silico, in vitro and in situ. When compared with traditional surveys or historical data, eDNA metabarcoding showed a much better detection probability overall. For amphibians, the detection probability with eDNA metabarcoding was 0.97 (CI = 0.90-0.99) vs. 0.58 (CI = 0.50-0.63) for traditional surveys. For fish, in 89% of the studied sites, the number of taxa detected using the eDNA metabarcoding approach was higher or identical to the number detected using traditional methods. We argue that the proposed DNA-based approach has the potential to become the next-generation tool for ecological studies and standardized biodiversity monitoring in a wide range of aquatic ecosystems.

Lake hydromorphology assessment in Europe: Where are we 20 years after the adoption of the Water Framework Directive?

The characterization of lake hydromorphology is crucial to understand the dynamics of biodiversity. In Europe, it is also a regulatory requirement of the Water Framework Directive. However, according to the literature, few methods include this characterization. The aim of this study is to review the state of the art of the methods currently used or under development in European countries to assess lake hydromorphological status for the implementation of the WFD. Our analysis is based on responses to a questionnaire distributed to national experts on hydromorphology of the 28 countries implementing the WFD. Our results highlighted significant progress in the assessment of hydromorphological features and processes. Water level regime, through the range of water flow or existing water management, and structure of the shore zone through macrophytes and substrate characteristics or measurement of lateral connectivity, are the most frequently assessed features. Stratification, surface/groundwater connection and planform pattern are the lake features most frequently omitted from the methods. However, in most of the countries, the development of methods was still in progress to meet the WFD requirement. Definition of reference condition is a central component of all WFD compliant assessment tools but this is a challenge particularly in the assessment of hydromorphological alteration of reservoirs. Similarly, demonstrating strong links between hydromorphological indicators and biological quality elements remains a challenge with many knowledge gaps still evident. These results highlight the need for rapid collection of new environmental data and the need for conceptual and applied research to make methodological progress in assessing lake hydromorphology and ensuring habitat quality.

Can size distributions of European lake fish communities be predicted by trophic positions of their fish species?

An organism's body size plays an important role in ecological interactions such as predator-prey relationships. As predators are typically larger than their prey, this often leads to a strong positive relationship between body size and trophic position in aquatic ecosystems. The distribution of body sizes in a community can thus be an indicator of the strengths of predator-prey interactions. The aim of this study was to gain more insight into the relationship between fish body size distribution and trophic position in a wide range of European lakes. We used quantile regression to examine the relationship between fish species' trophic position and their log-transformed maximum body mass for 48 fish species found in 235 European lakes. Subsequently, we examined whether the slopes of the continuous community size distributions, estimated by maximum likelihood, were predicted by trophic position, predator-prey mass ratio (PPMR), or abundance (number per unit effort) of fish communities in these lakes. We found a positive linear relationship between species' maximum body mass and average trophic position in fishes only for the 75% quantile, contrasting our expectation that species' trophic position systematically increases with maximum body mass for fish species in European lakes. Consequently, the size spectrum slope was not related to the average community trophic position, but there were negative effects of community PPMR and total fish abundance on the size spectrum slope. We conclude that predator-prey interactions likely do not contribute strongly to shaping community size distributions in these lakes.

Defining chlorophyll-a reference conditions in European lakes.

The concept of "reference conditions" describes the benchmark against which current conditions are compared when assessing the status of water bodies. In this paper we focus on the establishment of reference conditions for European lakes according to a phytoplankton biomass indicator--the concentration of chlorophyll-a. A mostly spatial approach (selection of existing lakes with no or minor human impact) was used to set the reference conditions for chlorophyll-a values, supplemented by historical data, paleolimnological investigations and modelling. The work resulted in definition of reference conditions and the boundary between "high" and "good" status for 15 main lake types and five ecoregions of Europe: Alpine, Atlantic, Central/Baltic, Mediterranean, and Northern. Additionally, empirical models were developed for estimating site-specific reference chlorophyll-a concentrations from a set of potential predictor variables. The results were recently formulated into the EU legislation, marking the first attempt in international water policy to move from chemical quality standards to ecological quality targets.

Impacts of multiple stressors on freshwater biota across spatial scales and ecosystems.

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 variable. These results vindicate the traditional focus of lake restoration and management on nutrient stress, while highlighting that river management requires more bespoke management solutions.

Evidence that niche specialization explains species-energy relationships in lake fish communities.

1. Interspecific niche differences have long been identified as a major explanation for the occurrence of species-rich communities. However, much fieldwork studying variation in local species richness has focused upon physical habitat attributes or regional factors, such as the size of the regional species pool. 2. We applied indices of functional diversity and niche overlap to data on the species niche to examine the importance of interspecific niche differentiation for species richness in French lake fish communities. We combined this information with environmental data to test generalizations of the physiological tolerance and niche specialization hypotheses for species-energy relationships. 3. We found evidence for a largely non-saturating relationship (relative to random expectation) between species richness and functional evenness (evenness of spacing between species in niche space), while functional richness (volume of niche space occupied) peaked at moderate levels of species richness and niche overlap showed an initial decrease followed by saturation. This suggests that increased niche specialization may have allowed species to coexist in the most species-rich communities. 4. We tested for evidence that increased temperature, local habitat area, local habitat diversity and immigration affected species richness via increased niche specialization. Temperature explained by far the largest amount of variation in species richness, functional diversity and niche overlap. These results, combined with the largely non-saturating species richness-functional evenness relationship, suggest that increased temperature may have permitted increased species richness by allowing increased niche specialization. 5. These results emphasize the importance of niche differences for species coexistence in species-rich communities, and indicate that the conservation of functional diversity may be vital for the maintenance of species diversity in biological communities. Our approach may be applied readily to many types of community, and at any scale, thus providing a flexible means of testing niche-based hypotheses for species richness gradients.

Does niche overlap control relative abundance in French lacustrine fish communities? A new method incorporating functional traits.

1. The mechanisms that structure biological communities hold the key to understanding ecosystem functioning and the maintenance of biodiversity. Patterns of species abundances have been proposed as a means of differentiation between niche-based and neutral processes, but abundance information alone cannot provide unequivocal discrimination. 2. We combined species niche information and species' relative abundances to test the effects of two opposing structuring mechanisms (environmental filtering and niche complementarity) on species' relative abundances in French lacustrine fish communities. The test involved a novel method comparing the abundance-weighted niche overlap within communities against that expected when relative abundances were randomized among species within the community. 3. Observed overlap was consistently significantly lower than expected at random for two (swimming ability and trophic status) of four primary niche axes across lakes of differing physical environments. Thus, for these niche axes, pairs of abundant species tended to have relatively low niche overlap, while rare species tended to have relatively high niche overlap with abundant species. 4. This suggests that niche complementarity may have acted to enhance ecosystem function and that it is important for species coexistence in these fish communities. The method used may be easily applied to any sort of biological community and thus may have considerable potential for determining the generality of niche complementarity effects on community structure.

Response of fish communities to multiple pressures: Development of a total anthropogenic pressure intensity index.

Lakes in Europe are subject to multiple anthropogenic pressures, such as eutrophication, habitat degradation and introduction of alien species, which are frequently inter-related. Therefore, effective assessment methods addressing multiple pressures are needed. In addition, these systems have to be harmonised (i.e. intercalibrated) to achieve common management objectives across Europe. Assessments of fish communities inform environmental policies on ecological conditions integrating the impacts of multiple pressures. However, the challenge is to ensure consistency in ecological assessments through time, across ecosystem types and across jurisdictional boundaries. To overcome the serious comparability issues between national assessment systems in Europe, a total anthropogenic pressure intensity (TAPI) index was developed as a weighted combination of the most common pressures in European lakes that is validated against 10 national fish-based water quality assessment systems using data from 556 lakes. Multi-pressure indices showed significantly higher correlations with fish indices than single-pressure indices. The best-performing index combines eutrophication, hydromorphological alterations and human use intensity of lakes. For specific lake types also biological pressures may constitute an important additional pressure. The best-performing index showed a strong correlation with eight national fish-based assessment systems. This index can be used in lake management for assessing total anthropogenic pressure on lake ecosystems and creates a benchmark for comparison of fish assessments independent of fish community composition, size structure and fishing-gear. We argue that fish-based multiple-pressure assessment tools should be seen as complementary to single-pressure tools offering the major advantage of integrating direct and indirect effects of multiple pressures over large scales of space and time.

Functional redundancy and sensitivity of fish assemblages in European rivers, lakes and estuarine ecosystems.

The impact of species loss on ecosystems functioning depends on the amount of trait similarity between species, i.e. functional redundancy, but it is also influenced by the order in which species are lost. Here we investigated redundancy and sensitivity patterns across fish assemblages in lakes, rivers and estuaries. Several scenarios of species extinction were simulated to determine whether the loss of vulnerable species (with high propensity of extinction when facing threats) causes a greater functional alteration than random extinction. Our results indicate that the functional redundancy tended to increase with species richness in lakes and rivers, but not in estuaries. We demonstrated that i) in the three systems, some combinations of functional traits are supported by non-redundant species, ii) rare species in rivers and estuaries support singular functions not shared by dominant species, iii) the loss of vulnerable species can induce greater functional alteration in rivers than in lakes and estuaries. Overall, the functional structure of fish assemblages in rivers is weakly buffered against species extinction because vulnerable species support singular functions. More specifically, a hotspot of functional sensitivity was highlighted in the Iberian Peninsula, which emphasizes the usefulness of quantitative criteria to determine conservation priorities.

Quantified biotic and abiotic responses to multiple stress in freshwater, marine and ground waters.

We reviewed 219 papers and built an inventory of 532 items of ecological evidence on multiple stressor impacts in rivers, lakes, transitional and coastal waters, as well as groundwaters. Our review revealed that, despite the existence of a huge conceptual knowledge base in aquatic ecology, few studies actually provide quantitative evidence on multi-stress effects. Nutrient stress was involved in 71% to 98% of multi-stress situations in the three types of surface water environments, and in 42% of those in groundwaters. However, their impact manifested differently along the groundwater-river-lake-transitional-coastal continuum, mainly determined by the different hydro-morphological features of these ecosystems. The reviewed papers addressed two-stressor combinations most frequently (42%), corresponding with the actual status-quo of pressures acting on European surface waters as reported by the Member States in the WISE WFD Database (EEA, 2015). Across all biological groups analysed, higher explanatory power of the stress-effect models was discernible for lakes under multi-stressor compared to single stressor conditions, but generally lower for coastal and transitional waters. Across all aquatic environments, the explanatory power of stress-effect models for fish increased when multi-stressor conditions were taken into account in the analysis, qualifying this organism group as a useful indicator of multi-stress effects. In contrast, the explanatory power of models using benthic flora decreased under conditions of multiple stress.

Managing aquatic ecosystems and water resources under multiple stress--an introduction to the MARS project.

Water resources globally are affected by a complex mixture of stressors resulting from a range of drivers, including urban and agricultural land use, hydropower generation and climate change. Understanding how stressors interfere and impact upon ecological status and ecosystem services is essential for developing effective River Basin Management Plans and shaping future environmental policy. This paper details the nature of these problems for Europe's water resources and the need to find solutions at a range of spatial scales. In terms of the latter, we describe the aims and approaches of the EU-funded project MARS (Managing Aquatic ecosystems and water Resources under multiple Stress) and the conceptual and analytical framework that it is adopting to provide this knowledge, understanding and tools needed to address multiple stressors. MARS is operating at three scales: At the water body scale, the mechanistic understanding of stressor interactions and their impact upon water resources, ecological status and ecosystem services will be examined through multi-factorial experiments and the analysis of long time-series. At the river basin scale, modelling and empirical approaches will be adopted to characterise relationships between multiple stressors and ecological responses, functions, services and water resources. The effects of future land use and mitigation scenarios in 16 European river basins will be assessed. At the European scale, large-scale spatial analysis will be carried out to identify the relationships amongst stress intensity, ecological status and service provision, with a special focus on large transboundary rivers, lakes and fish. The project will support managers and policy makers in the practical implementation of the Water Framework Directive (WFD), of related legislation and of the Blueprint to Safeguard Europe's Water Resources by advising the 3rd River Basin Management Planning cycle, the revision of the WFD and by developing new tools for diagnosing and predicting multiple stressors.

Assessing the ecological status in the context of the European Water Framework Directive: where do we go now?

The Water Framework Directive (WFD) is now well established as the key management imperative in river basins across Europe. However, there remain significant concerns with the way WFD is implemented and there is now a need for water managers and scientists to communicate better in order to find solutions to these concerns. To address this, a Science-Policy Interface (SPI) activity was launched in 2010 led by Directorate-General for Research and Innovation and Onema (the French national agency for water and aquatic ecosystems), which provided an interactive forum to connect scientists and WFD end-users. One major aim of the SPI activity was to establish a list of the most crucial research and development needs for enhancing WFD implementation. This paper synthesises the recommendations from this event highlighting 10 priority issues relating to ecological status. For lakes, temporary streams and transitional and coastal waters, WFD implementation still suffers from a lack of WFD-compliant bioassessment methods. For rivers, special attention is required to assess the ecological impacts of hydromorphological alterations on biological communities, notably those affecting river continuity and riparian covering. Spatial extrapolation tools are needed in order to evaluate ecological status for water bodies for which no data are available. The need for more functional bioassessment tools as complements to usual WFD-compliant tools, and to connect clearly good ecological state, biodiversity and ecosystem services when implementing WFD were also identified as crucial issues.

A new freshwater biodiversity indicator based on fish community assemblages.

Biodiversity has reached a critical state. In this context, stakeholders need indicators that both provide a synthetic view of the state of biodiversity and can be used as communication tools. Using river fishes as model, we developed community indicators that aim at integrating various components of biodiversity including interactions between species and ultimately the processes influencing ecosystem functions. We developed indices at the species level based on (i) the concept of specialization directly linked to the niche theory and (ii) the concept of originality measuring the overall degree of differences between a species and all other species in the same clade. Five major types of originality indices, based on phylogeny, habitat-linked and diet-linked morphology, life history traits, and ecological niche were analyzed. In a second step, we tested the relationship between all biodiversity indices and land use as a proxy of human pressures. Fish communities showed no significant temporal trend for most of these indices, but both originality indices based on diet- and habitat- linked morphology showed a significant increase through time. From a spatial point of view, all indices clearly singled out Corsica Island as having higher average originality and specialization. Finally, we observed that the originality index based on niche traits might be used as an informative biodiversity indicator because we showed it is sensitive to different land use classes along a landscape artificialization gradient. Moreover, its response remained unchanged over two other land use classifications at the global scale and also at the regional scale.

Functional characters combined with null models reveal inconsistency in mechanisms of species turnover in lacustrine fish communities.

Functional characters have the potential to act as indicators of species turnover between local communities. Null models provide a powerful statistical approach to test for patterns using functional character information. A combined null model/functional character approach provides the ability to distinguish between the effect of competition and environmental filtering on species turnover. We measured 13 functional characters relating directly to resource use for the fish species found in French lakes. We combined this functional character data with a null model approach to test whether co-occurring species overlapped more or less than expected at random for four primary niche axes. We used an environmentally constrained null model approach to determine if the same mechanisms were responsible for species turnover at different sections of the altitudinal gradient. Functional diversity indices were used to examine the variation in functional character diversity with altitude, as a test of the hypothesis that competitive intensity decreases with increasing environmental adversity. The unconstrained null model showed that environmental filtering was the dominant influence on species turnover between lakes. In the constrained null model, there was much less evidence for environmental filtering, emphasising the strong effect of altitude on turnover in functional character values between local communities. Different results were obtained for low-altitude and high-altitude lake subsets, with more evidence for the effect of environmental filtering being found in the high-altitude lakes. This demonstrates that different processes may influence species turnover throughout an environmental gradient. Functional diversity values showed a slight decrease with altitude, indicating that there was only weak evidence that competitive intensity decreased with increasing altitude. Variation resource availability and environmental stress probably cause the observed turnover in functional characters along the altitudinal gradient, though the effects of dispersal limitation and species introductions in high-altitude lakes cannot be ruled out.